@prefix vivo: . @prefix edm: . @prefix ns0: . @prefix dcterms: . @prefix skos: . vivo:departmentOrSchool "Science, Faculty of"@en, "Zoology, Department of"@en ; edm:dataProvider "DSpace"@en ; ns0:degreeCampus "UBCV"@en ; dcterms:creator "Zhang, Junxia"@en ; dcterms:issued "2012-05-23T22:44:57Z"@en, "2012"@en ; vivo:relatedDegree "Doctor of Philosophy - PhD"@en ; ns0:degreeGrantor "University of British Columbia"@en ; dcterms:description "The Euophryinae is one of the largest subfamilies of jumping spiders (Salticidae) with worldwide distribution. As the only currently recognized salticid subfamily that has diversified almost evenly in both the Old and New World, its historical biogeography is particularly interesting. To clarify the phylogeny of Euophryinae, I amplified and sequenced four genes (nuclear: 28S rDNA, Actin 5C; mitochondrial: 16S-ND1, COI) from 261 jumping spider species, most euophryines, covering all major distribution areas of this subfamily. The molecular phylogeny strongly supports the monophyly of euophryines. Diolenius and its relatives are shown to be euophryines. The phylogeny also indicates euophryines from different continents tend to form their own clades with few cases of mixture. Temporal divergence of Euophryinae was investigated to understand its historical biogeography. The results suggest rapid radiations early during their evolutionary history, with most divergences after the Eocene. Given the age, several intercontinental dispersal events are required to explain the distribution of euophryines. The suggested tolerance to cold may have facilitated early dispersals between the Old and New World through the Antarctic land bridge. I also extensively studied morphological characteristics of a broad range of euophryine genera and species in order to extend our phylogenetic understanding beyond those taxa sampled for molecular data. Systematics of Euophryinae is discussed and a full list of euophryine genera is provided with 122 genera included (34 genera before this study). Euophryine generic groups and redefined delimitations for some genera are reviewed in detail, with 22 new synonyms of genera and 191 new combinations of species proposed. Photographs and illustrations of 173 euophryine species are provided. In addition, 14 new genera and 96 new species of euophryines are described. Correlated evolution between female copulatory duct and male embolus of euophryines was studied in a phylogenetic context. Intra-specific variation of these traits was also examined. The study reveals a positive correlation between the lengths of female copulatory duct and male embolus among euophryine species. However, the inter- and intra-specific variation patterns are not sufficient to tell whether this correlation results from sexual selection or species recognition mechanisms."@en ; edm:aggregatedCHO "https://circle.library.ubc.ca/rest/handle/2429/42354?expand=metadata"@en ; skos:note "!PHYLOGENY AND SYSTEMATICS OF THE JUMPING SPIDER SUBFAMILY EUOPHRYINAE (ARANEAE: SALTICIDAE), WITH CONSIDERATION OF BIOGEOGRAPHY AND GENITALIC EVOLUTION! by Junxia Zhang A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in The Faculty of Graduate Studies (Zoology) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) May 2012 © Junxia Zhang, 2012 ii! Abstract The Euophryinae is one of the largest subfamilies of jumping spiders (Salticidae) with worldwide distribution. As the only currently recognized salticid subfamily that has diversified almost evenly in both the Old and New World, its historical biogeography is particularly interesting. To clarify the phylogeny of Euophryinae, I amplified and sequenced four genes (nuclear: 28S rDNA, Actin 5C; mitochondrial: 16S-ND1, COI) from 261 jumping spider species, most euophryines, covering all major distribution areas of this subfamily. The molecular phylogeny strongly supports the monophyly of euophryines. Diolenius and its relatives are shown to be euophryines. The phylogeny also indicates euophryines from different continents tend to form their own clades with few cases of mixture. Temporal divergence of Euophryinae was investigated to understand its historical biogeography. The results suggest rapid radiations early during their evolutionary history, with most divergences after the Eocene. Given the age, several intercontinental dispersal events are required to explain the distribution of euophryines. The suggested tolerance to cold may have facilitated early dispersals between the Old and New World through the Antarctic land bridge. I also extensively studied morphological characteristics of a broad range of euophryine genera and species in order to extend our phylogenetic understanding beyond those taxa sampled for molecular data. Systematics of Euophryinae is discussed and a full list of euophryine genera is provided with 122 genera included (34 genera before this study). Euophryine generic groups and redefined delimitations for some genera are reviewed in detail, with 22 new synonyms of genera and 191 new combinations of species proposed. Photographs and illustrations of 173 euophryine species are provided. In addition, 14 new genera and 96 new species of euophryines are described. Correlated evolution between female copulatory duct and male embolus of euophryines was studied in a phylogenetic context. Intra-specific variation of these traits was also examined. The study reveals a positive correlation between the lengths of female copulatory duct and male embolus among euophryine species. However, the inter- and intra-specific variation patterns are not sufficient to tell whether this correlation results from sexual selection or species recognition mechanisms. iii! Preface The sequence of research chapters in this thesis is determined by their relevance to the core of this project (phylogeny and systematics of Euophryinae). However, the sequence of publishing these research chapters will be different, with the chapters describing new taxa (Chapters 5-8) being published first, because they provide names for terminals in the molecular phylogeny. The next one that will be published is Chapter 2, because the molecular phylogeny presented in this chapter forms a framework for Chapter 3 and Chapter 4. Chapter 3 will be published before Chapter 4 because taxonomic placements of the two species involved in Chapter 4 are corrected in this chapter. A version of Chapter 2 will be submitted for publication: Zhang, J. and Maddison, W. P., Molecular phylogeny and temporal scale divergence of the subfamily Euophryinae (Araneae: Salticidae), with implications on its historical biogeography. I participated in the field work during which some material for this study was collected, carried out laboratory work and data analyses, and wrote the manuscript. Wayne P. Maddison collected some specimens for this study, provided insights on taxon sampling and analyses, and contributed to writing the manuscript. A version of Chapter 3 will be submitted for publication: Zhang, J. and Maddison, W. P., Generic review of euophryine jumping spiders (Araneae: Salticidae), with a phylogeny from combined molecular and morphological data. I participated in the field work during which some material for this study was collected, conducted morphological studies including illustrating and photographing, carried out data analyses, and wrote the manuscript. Wayne P. Maddison collected some specimens for this study, provided insights on choosing morphological characters for phylogenetic analyses, and contributed to writing the manuscript. A version of Chapter 4 will be submitted for publication: Zhang, J. and Maddison, W. P., Intersexual correlated evolution of genitalia in euophryine jumping spiders (Araneae: Salticidae): sexual selection or “lock-and-key”? I participated in the field work during which some material for this study was collected, conducted laboratory work in collecting data, carried out data analyses, and wrote the manuscript. Wayne P. Maddison collected some specimens for this iv! study, provided insights on data collecting and analyses, and contributed to writing the manuscript. A version of Chapter 5 has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from the Dominican Republic and Puerto Rico (Araneae: Salticidae: Euophryinae). I participated in the field work during which material for this study was collected, prepared species descriptions, diagnostic character illustrations and photographs of preserved specimens and some living specimens, and wrote the manuscript. Wayne P. Maddison collected some specimens for this study, provided most of living spider photos and contributed to writing the manuscript. A version of Chapter 6 has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from Papua New Guinea (Araneae: Salticidae: Euophryinae). I prepared species descriptions, diagnostic character illustrations and photographs of preserved specimens, and wrote the manuscript. Wayne P. Maddison collected most of the specimens for this study, provided most of living spider photos and contributed to writing the manuscript. A version of Chapter 7 has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from Central America and South America (Araneae: Salticidae: Euophryinae). I prepared species descriptions, diagnostic character illustrations and photographs of preserved specimens, and wrote the manuscript. Wayne P. Maddison collected most of the specimens for this study, provided living spider photos and contributed to writing the manuscript. A version of Chapter 8 has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from Southeast Asia and Africa (Araneae: Salticidae: Euophryinae). I collected some specimens for this study, prepared species descriptions, diagnostic character illustrations and photographs of preserved specimens, and wrote the manuscript. Wayne P. Maddison collected some specimens for this study, provided living spider photos and contributed to writing the manuscript. v! Table of Contents Abstract ....................................................................................................................................ii Preface .....................................................................................................................................iii Table of Contents .....................................................................................................................v List of Tables ..........................................................................................................................xv List of Figures .......................................................................................................................xvi Acknowledgements .............................................................................................................xxiv 1 Introduction ........................................................................................................................1 1.1 Taxonomy and phylogeny of the family Salticidae.......................................................1 1.2 Taxonomy and phylogeny of the subfamily Euophryinae ............................................2 1.3 Biogeography of the family Salticidae and subfamily Euophryinae ............................4 1.4 Correlated evolution of female and male genitalia........................................................5 1.5 Objectives of the thesis..................................................................................................7 2 Molecular phylogeny and temporal scale divergence of the subfamily Euophryinae (Araneae: Salticidae), with implications on its historical biogeography ...........................12 2.1 Synopsis.......................................................................................................................12 2.2 Introduction .................................................................................................................12 2.3 Material and methods ..................................................................................................14 2.3.1 Taxon sampling ...................................................................................................14 2.3.2 DNA extraction, amplification and sequencing...................................................15 2.3.3 Sequence alignment .............................................................................................17 2.3.4 Phylogenetic analyses..........................................................................................18 2.3.4.1 Model selection............................................................................................18 2.3.4.2 Maximum likelihood (ML) analysis............................................................18 2.3.4.3 Bayesian (BI) analysis .................................................................................19 2.3.4.4 Maximum parsimony (MP) analysis ...........................................................20 2.3.5 Divergence time analyses ....................................................................................20 2.3.5.1 Calibrations..................................................................................................20 2.3.5.2 BEAST (Bayesian Evolutionary Analysis Sampling Trees) .......................22 2.3.5.3 Penalized likelihood implemented in r8s.....................................................23 vi! 2.4 Results .........................................................................................................................23 2.4.1 Phylogenetic analyses..........................................................................................23 2.4.1.1 All genes combined .....................................................................................23 2.4.1.2 28S ...............................................................................................................24 2.4.1.3 Actin 5C.......................................................................................................25 2.4.1.4 16S-ND1......................................................................................................25 2.4.1.5 COI ..............................................................................................................26 2.4.2 Divergence time analyses ....................................................................................26 2.5 Discussion....................................................................................................................27 2.5.1 Phylogeny of Euophryinae ..................................................................................27 2.5.1.1 Monophyly and content of Euophryinae .....................................................28 2.5.1.2 Diolenius and its relatives are euophryines .................................................29 2.5.1.3 “Bathippus” pahang is not a euophryine ....................................................29 2.5.1.4 Position of Euophryinae in jumping spider phylogeny ...............................30 2.5.2 Implications on taxonomy of Euophryinae .........................................................30 2.5.3 Temporal divergence and implication on biogeography of Euophryinae ............31 2.5.3.1 Diversification in the southern hemisphere .................................................31 2.5.3.2 Dispersal is important in the diversification of euophryines .......................33 2.5.3.2.1 Dispersals across Wallace-line ........................................................33 2.5.3.2.2 Dispersals in north hemisphere........................................................33 2.5.3.2.3 Dispersals between Neotropic and Nearctic ....................................34 2.5.3.2.4 Dispersals into Africa ......................................................................34 2.5.3.3 Two “hot-spots” of euophryine diversity ....................................................34 2.5.3.3.1 Divergences in New Guinea ............................................................34 2.5.3.3.2 Divergences in Caribbean................................................................35 2.5.4 Other implications from the molecular phylogeny..............................................35 2.6 Conclusions .................................................................................................................36 3 Generic review of euophryine jumping spiders (Araneae: Salticidae), with a phylogeny from combined molecular and morphological data ...........................................................50 3.1 Synopsis.......................................................................................................................50 3.2 Introduction .................................................................................................................50 3.3 Material and methods ..................................................................................................52 3.3.1 Techniques for morphological study ...................................................................52 vii! 3.3.2 Phylogenetic study...............................................................................................53 3.3.2.1 Taxon sampling ..........................................................................................53 3.3.2.2 Morphological characters ...........................................................................54 3.3.2.3 DNA sequences ..........................................................................................54 3.3.2.4 Phylogenetic analyses.................................................................................54 3.3.2.4.1 Morphological matrix ......................................................................54 3.3.2.4.2 DNA matrix .....................................................................................55 3.3.2.4.3 Combined DNA and morphology matrix ........................................55 3.3.2.5 Character optimization ...............................................................................56 3.4 Results .........................................................................................................................56 3.4.1 Morphological study on a broad range of euophryine taxa .................................56 3.4.2 Phylogenetic study...............................................................................................56 3.4.2.1 Morphological dataset ................................................................................56 3.4.2.2 Molecular dataset........................................................................................57 3.4.2.3 Combined DNA and morphology dataset ..................................................58 3.4.2.4 Character optimization ...............................................................................58 3.5 Discussion....................................................................................................................58 3.5.1 Phylogenetic analyses..........................................................................................58 3.5.1.1 DNA vs. morphology in phylogenetic reconstruction................................58 3.5.1.2 Role of morphology in phylogeny of Euophryinae ....................................60 3.5.2 Taxonomic review of Euophryinae .....................................................................61 3.5.2.1 Content of Euophryinae..............................................................................61 3.5.2.2 Major euophryine clades from the New World ..........................................62 3.5.2.2.1 Anasaitis-Corythalia Clade ..............................................................62 3.5.2.2.2 Antillattus Clade: Antillattus, Petemathis, Truncattus, and possibly Allodecta, Caribattus.........................................................................................64 3.5.2.2.3 Agobardus Clade: Agobardus, Bythocrotus, Compsodecta, Parasaitis ...........................................................................................................................66 3.5.2.2.4 Naphrys-Corticattus Clade ...............................................................68 3.5.2.2.5 Sidusa Clade .....................................................................................69 3.5.2.2.6 Mopiopia-Saphrys Clade ..................................................................71 3.5.2.2.7 Chapoda-Maeota Clade: Chapoda, Maeota, Commoris ..................73 3.5.2.2.8 Amphidraus-Marma Clade ...............................................................75 viii! 3.5.2.2.9 Coryphasia Clade: Coryphasia and possibly Semnolius ..................76 3.5.2.2.10 Pensacola-Mexigonus Clade ..........................................................78 3.5.2.2.11 Neonella Clade: Neonella and Darwinneon ...................................78 3.5.2.2.12 Belliena Clade: Belliena, Saitidops and possibly Stoidis................78 3.5.2.2.13 Soesilarishius Clade: Soesilarishius and possibly Rhyphelia .........79 3.5.2.2.14 Other genera from the New World: Ecuadattus, Ilargus, Popcornella, Tylogonus ..........................................................................................................79 3.5.2.3 Major euophryine clades from the Old World .............................................80 3.5.2.3.1 Bathippus-Canama Clade: Bathippus, Canama and possibly Spilargis ...........................................................................................................................80 3.5.2.3.2 Omoedus Clade .................................................................................81 3.5.2.3.3 Bulolia-Coccorchestes Clade: Bulolia, Coccorchestes, Leptathamas, Variratina and possibly Athamas......................................................................82 3.5.2.3.4 Diolenius Clade: Chalcolecta, Chalcolemia, Diolenius, Efate, Furculattus, Ohilimia, Paraharmochirus, Sobasina, Tarodes, Udvardya........83 3.5.2.3.5 Pristobaeus Clade: Pristobaeus, Opisthoncana, Ergane and Ascyltus ...........................................................................................................................83 3.5.2.3.6 Phasmolia Clade: Bindax, Lakarobius, Phasmolia, Araneotanna....85 3.5.2.3.7 Parabathippus-Parvattus Clade........................................................85 3.5.2.3.8 Euophrys Clade: Chalcoscirtus, Euophrys, Pseudeuophrys, Talavera and Featheroides...............................................................................................85 3.5.2.3.9 Saitis Clade: Saitis and possibly Margaromma ................................86 3.5.2.3.10 Laufeia Clade: Laufeia and possibly Magyarus .............................89 3.5.2.3.11 Colyttus Clade .................................................................................90 3.5.2.3.12 Cytaea-Euryattus Clade: Cytaea, Euryattus, possibly Aruattus and Charippus..........................................................................................................91 3.5.2.3.13 Thiania Clade..................................................................................92 3.5.2.3.14 Emathis-Lepidemathis Clade ..........................................................92 3.5.2.3.15 Thyenula Clade: Thyenula, possibly Tanzania, Lophostica and Pseudemathis ....................................................................................................93 3.5.2.3.16 Other genera from the Old World: Chalcotropis, Chinophrys, Foliabitus, Lagnus, Servaea, Sigytes, Thorelliola, Viribestus, Xenocytaea, Zabkattus...........................................................................................................94 ix! 3.6 Conclusions .................................................................................................................95 4 Intersexual correlated evolution of genitalia in euophryine jumping spiders (Araneae: Salticidae): sexual selection or “lock-and-key”? ...............................................................160 4.1 Synopsis.....................................................................................................................160 4.2 Introduction ...............................................................................................................161 4.3 Material and methods ................................................................................................163 4.3.1 Intersexual correlated evolution ........................................................................163 4.3.2 Intra-specific variation.......................................................................................164 4.4 Results .......................................................................................................................166 4.4.1 Intersexual correlation of embolus length and copulatory duct length..............166 4.4.2 Intra-specific variation.......................................................................................167 4.5 Discussion..................................................................................................................168 4.5.1 Intersexual correlated evolution of embolus and copulatory duct.....................168 4.5.2 “Lock-and-key” or sexual selection...................................................................169 4.5.3 Sexual selection mechanisms ............................................................................171 4.6 Conclusions and future directions .............................................................................173 5 New euophryine jumping spiders from the Dominican Republic and Puerto Rico (Araneae: Salticidae: Euophryinae) ...................................................................................187 5.1 Synopsis.....................................................................................................................187 5.2 Introduction ...............................................................................................................187 5.3 Material and methods ................................................................................................188 5.4 Taxonomy..................................................................................................................189 5.4.1 Genus Agobardus Keyserling, 1885..................................................................189 5.4.1.1 Agobardus bahoruco sp. nov. ..................................................................190 5.4.1.2 Agobardus cordiformis sp. nov. ..............................................................191 5.4.1.3 Agobardus gramineus sp. nov. .................................................................192 5.4.1.4 Agobardus oviedo sp. nov. ......................................................................193 5.4.1.5 Agobardus phylladiphilus sp. nov. ..........................................................194 5.4.2 Genus Anasaitis Bryant, 1950 ...........................................................................195 5.4.2.1 Anasaitis adorabilis sp. nov. ...................................................................196 5.4.2.2 Anasaitis brunnea sp. nov. ......................................................................197 5.4.2.3 Anasaitis hebetata sp. nov. ......................................................................198 5.4.2.4 Anasaitis laxa sp. nov. .............................................................................199 x! 5.4.3 Genus Antillattus Bryant, 1943 .........................................................................200 5.4.3.1 Antillattus applanatus sp. nov. ................................................................201 5.4.4 Genus Bythocrotus Simon, 1903 .......................................................................202 5.4.4.1 Bythocrotus crypticus sp. nov. .................................................................202 5.4.5 Genus Corticattus new genus ............................................................................203 5.4.5.1 Corticattus guajataca sp. nov. .................................................................204 5.4.5.2 Corticattus latus sp. nov. .........................................................................205 5.4.6 Genus Corythalia C. L. Koch, 1850..................................................................206 5.4.6.1 Corythalia broccai sp. nov. .....................................................................206 5.4.6.2 Corythalia bromelicola sp. nov. ..............................................................207 5.4.6.3 Corythalia coronai sp. nov. .....................................................................208 5.4.6.4 Corythalia peblique sp. nov. ....................................................................209 5.4.7 Genus Popcornella new genus ..........................................................................211 5.4.7.1 Popcornella furcata sp. nov. ...................................................................211 5.4.7.2 Popcornella nigromaculata sp. nov. .......................................................212 5.4.7.3 Popcornella spiniformis sp. nov. .............................................................213 5.4.7.4 Popcornella yunque sp. nov. ...................................................................214 5.4.8 Genus Truncattus new genus ............................................................................215 5.4.8.1 Truncattus cachotensis sp. nov. ...............................................................216 5.4.8.2 Truncattus dominicanus sp. nov. .............................................................217 5.4.8.3 Truncattus flavus sp. nov. ........................................................................218 6 New euophryine jumping spiders from Papua New Guinea (Araneae: Salticidae: Euophryinae).........................................................................................................................267 6.1 Synopsis.....................................................................................................................267 6.2 Introduction ...............................................................................................................267 6.3 Material and methods ................................................................................................268 6.4 Taxonomy..................................................................................................................269 6.4.1 Genus Bathippus Thorell, 1892 .........................................................................269 6.4.1.1 Bathippus directus sp. nov. ......................................................................269 6.4.1.2 Bathippus gahavisuka sp. nov. ................................................................271 6.4.1.3 Bathippus korei sp. nov. ..........................................................................272 6.4.1.4 Bathippus madang sp. nov. ......................................................................273 6.4.2 Genus Canama Simon, 1903 .............................................................................273 xi! 6.4.2.1 Canama extranea sp. nov. .......................................................................274 6.4.2.2 Canama fimoi sp. nov. .............................................................................275 6.4.2.3 Canama triramosa sp. nov. .....................................................................277 6.4.3 Genus Chalcolemia new genus..........................................................................278 6.4.3.1 Chalcolemia nakanai sp. nov. .................................................................278 6.4.4 Genus Omoedus Thorell, 1881 ..........................................................................279 6.4.4.1 Omoedus brevis sp. nov. ..........................................................................281 6.4.4.2 Omoedus darleyorum sp. nov. .................................................................282 6.4.4.3 Omoedus meyeri sp. nov. .........................................................................283 6.4.4.4 Omoedus omundseni sp. nov. ..................................................................284 6.4.4.5 Omoedus papuanus sp. nov. ....................................................................285 6.4.4.6 Omoedus swiftorum sp. nov. ....................................................................287 6.4.4.7 Omoedus tortuosus sp. nov. .....................................................................288 6.4.5 Genus Paraharmochirus Szombathy, 1915.......................................................289 6.4.5.1 Paraharmochirus tualapaensis sp. nov. ..................................................289 6.4.6 Genus Phasmolia new genus .............................................................................291 6.4.6.1 Phasmolia elegans sp. nov. .....................................................................291 6.4.7 Genus Sobasina Simon, 1898............................................................................292 6.4.7.1 Sobasina wanlessi sp. nov. ......................................................................293 6.4.8 Genus Thorelliola Strand, 1942.........................................................................294 6.4.8.1 Thorelliola aliena sp. nov. .......................................................................295 6.4.8.2 Thorelliola crebra sp. nov. ......................................................................296 6.4.8.3 Thorelliola joannae sp. nov. ....................................................................297 6.4.8.4 Thorelliola squamosa sp. nov. .................................................................298 6.4.8.5 Thorelliola tamasi sp. nov. ......................................................................299 6.4.8.6 Thorelliola tualapa sp. nov. ....................................................................300 6.4.8.7 Thorelliola zabkai sp. nov. ......................................................................301 6.4.9 Genus Variratina new genus .............................................................................303 6.4.9.1 Variratina minuta sp. nov. .......................................................................303 6.4.10 Genus Viribestus new genus............................................................................304 6.4.10.1 Viribestus suyanensis sp. nov. ...............................................................304 6.4.11 Genus Xenocytaea Berry, Beatty & Prószy!ski, 1998 ....................................305 6.4.11.1 Xenocytaea agnarssoni sp. nov. ............................................................306 xii! 6.4.11.2 Xenocytaea albomaculata sp. nov. ........................................................307 6.4.11.3 Xenocytaea proszynskii sp. nov. ............................................................308 6.4.12 Genus Zabkattus new genus ............................................................................309 6.4.12.1 Zabkattus brevis sp. nov. .......................................................................309 6.4.12.2 Zabkattus furcatus sp. nov. ....................................................................310 6.4.12.3 Zabkattus richardsi sp. nov. ..................................................................311 6.4.12.4 Zabkattus trapeziformis sp. nov. ............................................................312 7 New euophryine jumping spiders from Central America and South America (Araneae: Salticidae: Euophryinae)......................................................................................................370 7.1 Synopsis.....................................................................................................................370 7.2 Introduction ...............................................................................................................370 7.3 Material and methods ................................................................................................371 7.4 Taxonomy..................................................................................................................371 7.4.1 Genus Amphidraus Simon, 1900 .......................................................................371 7.4.1.1 Amphidraus complexus sp. nov. ..............................................................372 7.4.2 Genus Belliena Simon, 1902 .............................................................................373 7.4.2.1 Belliena ecuadorica sp. nov. ...................................................................373 7.4.3 Genus Chapoda Peckham & Peckham, 1896....................................................374 7.4.3.1 Chapoda angusta sp. nov. .......................................................................375 7.4.3.2 Chapoda fortuna sp. nov. ........................................................................376 7.4.3.3 Chapoda gitae sp. nov. ............................................................................377 7.4.4 Genus Ecuadattus new genus ............................................................................378 7.4.4.1 Ecuadattus elongatus sp. nov. ..................................................................379 7.4.4.2 Ecuadattus napoensis sp. nov. .................................................................379 7.4.4.3 Ecuadattus pichincha sp. nov. .................................................................380 7.4.4.4 Ecuadattus typicus sp. nov. .....................................................................381 7.4.5 Genus Ilargus Simon, 1901 ...............................................................................382 7.4.5.1 Ilargus foliosus sp. nov. ...........................................................................383 7.4.5.2 Ilargus galianoae sp. nov. ........................................................................384 7.4.5.3 Ilargus macrocornis sp. nov. ....................................................................385 7.4.5.4 Ilargus moronatigus sp. nov. ....................................................................386 7.4.5.5 Ilargus pilleolus sp. nov. ..........................................................................387 7.4.5.6 Ilargus serratus sp. nov. ...........................................................................388 xiii! 7.4.6 Genus Maeota Simon, 1901 ...............................................................................389 7.4.6.1 Maeota dorsalis sp. nov. ...........................................................................389 7.4.6.2 Maeota flava sp. nov. ...............................................................................390 7.4.6.3 Maeota simoni sp. nov. .............................................................................391 7.4.7 Genus Soesilarishius Makhan, 2007 ..................................................................392 7.4.7.1 Soesilarishius micaceus sp. nov. ..............................................................392 7.4.7.2 Soesilarishius ruizi sp. nov. ......................................................................393 7.4.8 Genus Tylogonus Simon, 1902...........................................................................394 7.4.8.1 Tylogonus parvus sp. nov. ........................................................................395 7.4.8.2 Tylogonus yanayacu sp. nov. ....................................................................395 8 New euophryine jumping spiders from Southeast Asia and Africa (Araneae: Salticidae: Euophryinae).........................................................................................................................423 8.1 Synopsis.....................................................................................................................423 8.2 Introduction ...............................................................................................................423 8.3 Material and methods ................................................................................................424 8.4 Taxonomy..................................................................................................................424 8.4.1 Genus Chinophrys new genus ............................................................................424 8.4.1.1 Chinophrys pengi sp. nov. ........................................................................425 8.4.2 Genus Colyttus Thorell, 1891.............................................................................426 8.4.2.1 Colyttus robustus sp. nov. .........................................................................427 8.4.3 Genus Emathis Simon, 1899 ..............................................................................427 8.4.3.1 Emathis gombak sp. nov. ..........................................................................428 8.4.4 Genus Foliabitus new genus...............................................................................429 8.4.4.1 Foliabitus longzhou sp. nov. .....................................................................429 8.4.5 Genus Lagnus L. Koch, 1879 .............................................................................430 8.4.5.1 Lagnus edwardsi sp. nov. .........................................................................431 8.4.6 Genus Laufeia Simon, 1889 ...............................................................................432 8.4.6.1 Laufeia concava sp. nov. ..........................................................................432 8.4.6.2 Laufeia eximia sp. nov. .............................................................................434 8.4.7 Genus Parabathippus new genus .......................................................................435 8.4.7.1 Parabathippus cuspidatus sp. nov. ...........................................................436 8.4.7.2 Parabathippus kiabau sp. nov. .................................................................437 8.4.7.3 Parabathippus magnus sp. nov. ...............................................................438 xiv! 8.4.8 Genus Parvattus new genus ...............................................................................439 8.4.8.1 Parvattus zhui sp. nov. .............................................................................439 8.4.9 Genus Thiania C. L. Koch, 1846........................................................................440 8.4.9.1 Thiania latibola sp. nov. ...........................................................................441 8.4.9.2 Thiania tenuis sp. nov. ..............................................................................441 8.4.10 Genus Thyenula Simon, 1902...........................................................................442 8.4.10.1 Thyenula laxa sp. nov. ............................................................................442 8.4.10.2 Thyenula nelshoogte sp. nov. .................................................................443 8.4.10.3 Thyenula wesolowskae sp. nov. ..............................................................444 9 Conclusions and future directions.................................................................................464 9.1 Major conclusions......................................................................................................464 9.2 Future directions ........................................................................................................467 References..............................................................................................................................470 Appendices ............................................................................................................................487 Appendix 1: List of species used in phylogenetic analyses................................................487 Appendix 2: List of morphological characters scored for phylogenetic analyses ..............501 Appendix 3: Scored morphological matrix ........................................................................511 xv! List of Tables Table 2.1. List of primers and primer sequences used in gene amplification and sequencing.... ..................................................................................................................................................38 Table 2.2. Summary of annealing temperatures for gene amplification using different pairs of primers ......................................................................................................................................39 Table 2.3. Substitution models selected by ModelTest for each individual gene region and partition.....................................................................................................................................40 Table 2.4. Summary of calibration points used in divergence time analyses..........................41 Table 2.5. Estimates of divergence times (in millions of years) for nodes in Fig. 2.7 .............42 Table 3.1. Substitution models selected by ModelTest for each molecular partition..............96 Table 3.2. Optimization of morphological chatacters for major clades of euophryines on different phylogenies ................................................................................................................97 Table 3.3. Full list of euophryine genera..................................................................................99 Table 4.1. Intra-specific variation of somatic and genitalic traits in Chapoda recondita and Antillattus cambridgei ............................................................................................................175 xvi! List of Figures Figure 1.1. Spider eyes ..............................................................................................................9 Figure 1.2. Summary of phylogeny of jumping spiders (Salticidae).......................................10 Figure 1.3. Typical genitalic structures of euophryine, as in Parabathippus shelfordi ..........11 Figure 2.1. Summary of phylogenetic analyses on combined matrix of all genes (28S, Actin 5C, 16S-ND1 and COI) ...................................................................................................................43 Figure 2.2. The best tree from ML analysis on combined matrix of all genes (28S, Actin 5C, 16S-ND1 and COI) ...................................................................................................................44 Figure 2.3. Summary of ML and MP analyses on 28S.............................................................45 Figure 2.4. Summary of ML and MP analyses on Actin 5C ....................................................46 Figure 2.5. Summary of ML and MP analyses on 16S-ND1 ...................................................47 Figure 2.6. Summary of ML and MP analyses on COI............................................................48 Figure 2.7. Chronogram of euophryine divergence.................................................................49 Figure 3.1. Phylogeny of Euophryinae ..................................................................................102 Figure 3.2. Strict consensus of four equally parsimonious trees (score=1141) from the morphological dataset that found in TNT, without constraining euophryine taxa as a monophyletic group................................................................................................................103 Figure 3.3. Strict consensus of seven equally parsimonious trees (score=1145) from the morphological dataset that found in TNT, with euophryine taxa being enforced as a monophyletic group................................................................................................................104 Figure 3.4. The best tree from ML analysis on the DNA dataset (28S, Actin 5C, 16S-ND1 and COI) ........................................................................................................................................105 Figure 3.5. The best tree from ML analysis on the combined morphology and DNA dataset (morphology, 28S, Actin 5C, 16S-ND1 and COI) .................................................................106 Figure 3.6. Anasaitis-Corythalia Clade ..................................................................................107 Figure 3.7. Anasaitis-Corythalia Clade ..................................................................................108 Figure 3.8. Antillattus Clade...................................................................................................109 Figure 3.9. Antillattus Clade...................................................................................................110 Figure 3.10. Antillattus Clade.................................................................................................111 Figure 3.11. Antillattus Clade.................................................................................................112 Figure 3.12. Agobardus Clade ................................................................................................113 xvii! Figure 3.13. Agobardus Clade ................................................................................................114 Figure 3.14. Agobardus Clade ................................................................................................115 Figure 3.15. Naphrys-Corticattus Clade.................................................................................116 Figure 3.16. Sidusa Clade.......................................................................................................117 Figure 3.17. Mopiopia-Saphrys Clade....................................................................................118 Figure 3.18. Chapoda-Maeota Clade .....................................................................................119 Figure 3.19. Chapoda-Maeota Clade .....................................................................................120 Figure 3.20. Amphidraus-Marma Clade.................................................................................121 Figure 3.21. Coryphasia Clade...............................................................................................122 Figure 3.22. Pensacola-Mexigonus Clade ..............................................................................123 Figure 3.23. Neonella Clade ...................................................................................................124 Figure 3.24. Belliena Clade ....................................................................................................125 Figure 3.25. Soesilarishius Clade ...........................................................................................126 Figure 3.26. Ecuadattus..........................................................................................................127 Figure 3.27. Ilargus ................................................................................................................128 Figure 3.28. Popcornella ........................................................................................................129 Figure 3.29. Tylogonus ...........................................................................................................130 Figure 3.30. Bathippus-Canama Clade ..................................................................................131 Figure 3.31. Omoedus Clade ..................................................................................................132 Figure 3.32. Bulolia-Coccorchestes Clade .............................................................................133 Figure 3.33. Bulolia-Coccorchestes Clade .............................................................................134 Figure 3.34. Diolenius Clade ..................................................................................................135 Figure 3.35. Diolenius Clade ..................................................................................................136 Figure 3.36. Diolenius Clade ..................................................................................................137 Figure 3.37. Pristobaeus Clade ..............................................................................................138 Figure 3.38. Phasmollia Clade ...............................................................................................139 Figure 3.39. Parabathippus-Parvattus Clade.........................................................................140 Figure 3.40. Euophrys Clade ..................................................................................................141 Figure 3.41. Euophrys Clade ..................................................................................................142 Figure 3.42. Saitis Clade.........................................................................................................143 Figure 3.43. Saitis Clade.........................................................................................................144 Figure 3.44. Laufeia Clade .....................................................................................................145 Figure 3.45. Colyttus Clade ....................................................................................................146 xviii! Figure 3.46. Cytaea-Euryattus Clade .....................................................................................147 Figure 3.47. Thiania Clade .....................................................................................................148 Figure 3.48. Emathis-Lepidemathis Clade .............................................................................149 Figure 3.49. Thyenula Clade...................................................................................................150 Figure 3.50. Chalcotropis .......................................................................................................151 Figure 3.51. Chinophrys .........................................................................................................152 Figure 3.52. Foliabitus ...........................................................................................................153 Figure 3.53. Lagnus ................................................................................................................154 Figure 3.54. Thorelliola..........................................................................................................155 Figure 3.55. Viribestus............................................................................................................156 Figure 3.56. Xenocytaea .........................................................................................................157 Figure 3.57. Zabkattus ............................................................................................................158 Figure 3.58. Expanded male left palp.....................................................................................159 Figure 4.1. Variation in the lengths of embolus and copulatory duct among euophryines ....178 Figure 4.2. Comparison of sexual dimorphism in Chapoda recondita and Antillattus cambridgei ................................................................................................................................................179 Figure 4.3. Non-genitalic traits measured for Antillattus cambridgei (same for Chapoda recondita)................................................................................................................................180 Figure 4.4. Genitalic traits measured......................................................................................181 Figure 4.5. Correlated evolution of the lengths of male embolus (left) and female copulatory duct (right) in euophryine jumping spiders, both standardized by carapace length ...............182 Figure 4.6. Independent contrasts of male embolus length (x) vs. female copulatory duct length (y) are positively correlated in euophryine species ................................................................183 Figure 4.7. Linear regression of absolute lengths of male embolus (x) vs. female copulatory duct (y) in euophryine species, not corrected for phylogeny .........................................................184 Figure 4.8. Allometric relationships of genitalic and non-genitalic traits in Chapoda recondita, using carapace length as the indicator of body size................................................................185 Figure 4.9. Allometric relationships of genitalic and non-genitalic traits in Antillattus cambridgei, using carapace length as the indicator of body size ...........................................186 Figure 5.1. Agobardus bahoruco sp. nov. .............................................................................220 Figure 5.2. Agobardus bahoruco sp. nov. .............................................................................221 Figure 5.3. Agobardus cordiformis sp. nov. ..........................................................................222 Figure 5.4. Agobardus cordiformis sp. nov. ..........................................................................223 xix! Figure 5.5. Agobardus gramineus sp. nov. ............................................................................224 Figure 5.6. Agobardus gramineus sp. nov. ............................................................................225 Figure 5.7. Agobardus oviedo sp. nov. ..................................................................................226 Figure 5.8. Agobardus oviedo sp. nov. ..................................................................................227 Figure 5.9. Agobardus phylladiphilus sp. nov. ......................................................................228 Figure 5.10. Agobardus phylladiphilus sp. nov. ....................................................................229 Figure 5.11. Anasaitis adorabilis sp. nov. .............................................................................230 Figure 5.12. Anasaitis adorabilis sp. nov. .............................................................................231 Figure 5.13. Anasaitis brunnea sp. nov. ................................................................................232 Figure 5.14. Anasaitis brunnea sp. nov. ................................................................................233 Figure 5.15. Anasaitis hebetata sp. nov. ................................................................................234 Figure 5.16. Anasaitis laxa sp. nov. ......................................................................................235 Figure 5.17. Anasaitis laxa sp. nov. ......................................................................................236 Figure 5.18. Antillatus applanatus sp. nov. ...........................................................................237 Figure 5.19. Antillatus applanatus sp. nov. ...........................................................................238 Figure 5.20. Bythocrotus crypticus sp. nov. ..........................................................................239 Figure 5.21. Bythocrotus crypticus sp. nov. ..........................................................................240 Figure 5.22. Corticattus guajataca sp. nov. ..........................................................................241 Figure 5.23. Corticattus guajataca sp. nov. ..........................................................................242 Figure 5.24. Corticattus latus sp. nov. ..................................................................................243 Figure 5.25. Corticattus latus sp. nov. ..................................................................................244 Figure 5.26. Corythalia broccai sp. nov. ...............................................................................245 Figure 5.27. Corythalia broccai sp. nov. ...............................................................................246 Figure 5.28. Corythalia bromelicola sp. nov. ........................................................................247 Figure 5.29. Corythalia bromelicola sp. nov. ........................................................................248 Figure 5.30. Corythalia coronai sp. nov. ..............................................................................249 Figure 5.31. Corythalia coronai sp. nov. ..............................................................................250 Figure 5.32. Corythalia peblique sp. nov. .............................................................................251 Figure 5.33. Corythalia peblique sp. nov. .............................................................................252 Figure 5.34. Popcornella furcata sp. nov. .............................................................................253 Figure 5.35. Popcornella furcata sp. nov. .............................................................................254 Figure 5.36. Popcornella nigromaculata sp. nov. .................................................................255 Figure 5.37. Popcornella nigromaculata sp. nov. .................................................................256 xx! Figure 5.38. Popcornella spiniformis sp. nov. ......................................................................257 Figure 5.39. Popcornella spiniformis sp. nov. ......................................................................258 Figure 5.40. Popcornella yunque sp. nov. .............................................................................259 Figure 5.41. Popcornella yunque sp. nov. .............................................................................260 Figure 5.42. Truncattus cachotensis sp. nov. ........................................................................261 Figure 5.43. Truncattus cachotensis sp. nov. ........................................................................262 Figure 5.44. Truncattus dominicanus sp. nov. ......................................................................263 Figure 5.45. Truncattus dominicanus sp. nov. ......................................................................264 Figure 5.46. Truncattus flavus sp. nov. .................................................................................265 Figure 5.47. Truncattus flavus sp. nov. .................................................................................266 Figure 6.1. Bathippus directus sp. nov. .................................................................................314 Figure 6.2. Bathippus directus sp. nov. .................................................................................315 Figure 6.3. Bathippus gahavisuka sp. nov. ............................................................................316 Figure 6.4. Bathippus gahavisuka sp. nov. ............................................................................317 Figure 6.5. Bathippus korei sp. nov. ......................................................................................318 Figure 6.6. Bathippus korei sp. nov. ......................................................................................319 Figure 6.7. Bathippus madang sp. nov. .................................................................................320 Figure 6.8. Canama extranea sp. nov. ...................................................................................321 Figure 6.9. Canama extranea sp. nov. ...................................................................................322 Figure 6.10. Canama fimoi sp. nov. ......................................................................................323 Figure 6.11. Canama fimoi sp. nov. ......................................................................................324 Figure 6.12. Canama triramosa sp. nov. ...............................................................................325 Figure 6.13. Canama triramosa sp. nov. ...............................................................................326 Figure 6.14. Chalcolemia nakanai sp. nov. ...........................................................................327 Figure 6.15. Omoedus brevis sp. nov. ...................................................................................328 Figure 6.16. Omoedus darleyorum sp. nov. ..........................................................................329 Figure 6.17. Omoedus darleyorum sp. nov. ..........................................................................330 Figure 6.18. Omoedus meyeri sp. nov. ..................................................................................331 Figure 6.19. Omoedus meyeri sp. nov. ..................................................................................332 Figure 6.20. Omoedus omundseni sp. nov. ............................................................................333 Figure 6.21. Omoedus omundseni sp. nov. ............................................................................334 Figure 6.22. Omoedus papuanus sp. nov. ..............................................................................335 Figure 6.23. Omoedus papuanus sp. nov. ..............................................................................336 xxi! Figure 6.24. Omoedus swiftorum sp. nov. .............................................................................337 Figure 6.25. Omoedus swiftorum sp. nov. .............................................................................338 Figure 6.26. Omoedus tortuosus sp. nov. ..............................................................................339 Figure 6.27. Omoedus tortuosus sp. nov. ..............................................................................340 Figure 6.28. Paraharmochirus tualapaensis sp. nov. ............................................................341 Figure 6.29. Paraharmochirus tualapaensis sp. nov. ............................................................342 Figure 6.30. Phasmolia elegans sp. nov. ...............................................................................343 Figure 6.31. Phasmolia elegans sp. nov. ...............................................................................344 Figure 6.32. Sobasina wanlessi sp. nov. ................................................................................345 Figure 6.33. Thorelliola aliena sp. nov. ................................................................................346 Figure 6.34. Thorelliola aliena sp. nov. ................................................................................347 Figure 6.35. Thorelliola crebra sp. nov. ................................................................................348 Figure 6.36. Thorelliola crebra sp. nov. ................................................................................349 Figure 6.37. Thorelliola joannae sp. nov. .............................................................................350 Figure 6.38. Thorelliola squamosa sp. nov. ..........................................................................351 Figure 6.39. Thorelliola tamasi sp. nov. ................................................................................352 Figure 6.40. Thorelliola tamasi sp. nov. ................................................................................353 Figure 6.41. Thorelliola tualapa sp. nov. ..............................................................................354 Figure 6.42. Thorelliola tualapa sp. nov. ..............................................................................355 Figure 6.43. Thorelliola zabkai sp. nov. ................................................................................356 Figure 6.44. Thorelliola zabkai sp. nov. ................................................................................357 Figure 6.45. Variratina minuta sp. nov. ................................................................................358 Figure 6.46. Variratina minuta sp. nov. ................................................................................359 Figure 6.47. Viribestus suyanensis sp. nov. ...........................................................................360 Figure 6.48. Viribestus suyanensis sp. nov. ...........................................................................361 Figure 6.49. Xenocytaea agnarssoni sp. nov. ........................................................................362 Figure 6.50. Xenocytaea albomaculata sp. nov. ....................................................................363 Figure 6.51. Xenocytaea proszynskii sp. nov. ........................................................................364 Figure 6.52. Zabkattus brevis sp. nov. ...................................................................................365 Figure 6.53. Zabkattus brevis sp. nov. ...................................................................................366 Figure 6.54. Zabkattus furcatus sp. nov. ...............................................................................367 Figure 6.55. Zabkattus richardsi sp. nov. ..............................................................................368 Figure 6.56. Zabkattus trapeziformis sp. nov. .......................................................................369 xxii! Figure 7.1. Amphidraus complexus sp. nov. ..........................................................................397 Figure 7.2. Belliena ecuadorica sp. nov. ...............................................................................398 Figure 7.3. Chapoda angusta sp. nov. ...................................................................................399 Figure 7.4. Chapoda fortuna sp. nov. ....................................................................................400 Figure 7.5. Chapoda gitae sp. nov. ........................................................................................401 Figure 7.6. Ecuadattus elongatus sp. nov. .............................................................................402 Figure 7.7. Ecuadattus napoensis sp. nov. ............................................................................403 Figure 7.8. Ecuadattus pichincha sp. nov. ............................................................................404 Figure 7.9. Ecuadattus typicus sp. nov. .................................................................................405 Figure 7.10. Ilargus foliosus sp. nov. ....................................................................................406 Figure 7.11. Ilargus galianoae sp. nov. .................................................................................407 Figure 7.12. Ilargus macrocornis sp. nov. ............................................................................408 Figure 7.13. Ilargus moronatigus sp. nov. ............................................................................409 Figure 7.14. Ilargus pilleolus sp. nov. ...................................................................................410 Figure 7.15. Ilargus serratus sp. nov. ....................................................................................411 Figure 7.16. Maeota dorsalis sp. nov. ...................................................................................412 Figure 7.17. Maeota flava sp. nov. ........................................................................................413 Figure 7.18. Maeota simoni sp. nov. .....................................................................................414 Figure 7.19. Soesilarishius micaceus sp. nov. .......................................................................415 Figure 7.20. Soesilarishius ruizi sp. nov. ..............................................................................416 Figure 7.21. Tylogonus parvus sp. nov. .................................................................................417 Figure 7.22. Tylogonus yanayacu sp. nov. ............................................................................418 Figure 7.23. Living spider photos of Chapoda gitae sp. nov. ...............................................419 Figure 7.24. Living spider photos of Ecuadattus pichincha sp. nov. ....................................420 Figure 7.25. Living spider photos of Ecuadattus typicus sp. nov. ........................................421 Figure 7.26. Living spider photos ..........................................................................................422 Figure 8.1. Chinophrys pengi sp. nov. ...................................................................................446 Figure 8.2. Colyttus robustus sp. nov. ...................................................................................447 Figure 8.3. Emathis gombak sp. nov. ....................................................................................448 Figure 8.4. Foliabitus longzhou sp. nov. ...............................................................................449 Figure 8.5. Lagnus edwardsi sp. nov. ....................................................................................450 Figure 8.6. Laufeia concava sp. nov. .....................................................................................451 Figure 8.7. Laufeia eximia sp. nov. .......................................................................................452 xxiii! Figure 8.8. Parabathippus cuspidatus sp. nov. .....................................................................453 Figure 8.9. Parabathippus kiabau sp. nov. ............................................................................454 Figure 8.10. Parabathippus magnus sp. nov. ........................................................................455 Figure 8.11. Parvattus zhui sp. nov. ......................................................................................456 Figure 8.12. Thiania latibola sp. nov. ...................................................................................457 Figure 8.13. Thiania tenuis sp. nov. ......................................................................................458 Figure 8.14. Thyenula laxa sp. nov. ......................................................................................459 Figure 8.15. Thyenula nelshoogte sp. nov. ............................................................................460 Figure 8.16. Thyenula wesolowskae sp. nov. ........................................................................461 Figure 8.17. Living spider photos ..........................................................................................462 Figure 8.18. Living spider photos ..........................................................................................463 xxiv! Acknowledgements First and foremost, I want to thank my PhD supervisor, Dr. Wayne P. Maddison. Without his patient guidance, generous input of insight, time and funding throughout my PhD period, I would not have been able to finish this project smoothly and successfully. I also would like to thank the other members of my supervisory committee: Dr. Mary Berbee, Dr. Leticia Avilés, and Dr. Brian Leander for their time, critiques and advice on my work. Sincere gratitude is to Karen Needham, former or current members of spider lab (Dr. Ingi Agnarsson, Dr. Peter Midford, Dr. Damian O. Elias, Dr. Gustavo R. S. Ruiz, Melissa R. Bodner, Gwylim Blackburn, Edyta Piascik, Mauricio Vega, Dr. Maxence Salomon and Jennifer Guevara) for constructive discussions on my work, their friendship, encouragement and support during my PhD period. I am very grateful to people who generously loaned me specimens in museums or private collections for molecular and/or morphological studies, without which this study would have been impossible: Dr. Charles Griswold (California Academy of Sciences, USA), Dr. G. B. Edwards (Florida State Collection of Arthropods, USA), Dr. Martín J. Ramírez and Mr. Cristian J. Grismado (Museo Argentino de Ciencias Naturales, Argentina), Ms. Laura Leibensperger (Museum of Comparative Zoology, USA), Mr. Louis N. Sorkin and Dr. Norman I. Platnick (American Museum of Natural History, USA), Dr. Christine Rollard and Ms. Elise-Anne Leguin (Muséum National d’Histoire Naturelle, France), Dr. Rudy Jocqué (Koninklijk Museum, Belgium), Dr. Domir De Bakker (Royal Museum for Central Africa, Tervuren, Belgium), Dr. Gustzvo R. S. Ruiz (Instituto Butantan, São Paulo, Brazil), Dr. Alexander Riedel (Staatliches Museum für Naturkunde, Germany), Dr. Marek Zakba, Dr. Rosemary G. Gillespie, Dr. Damian O. Elias, Mr. Ken Schneider. Acknowledgements are due to Dr. Stephen Richards, Mr. J. Brocca, Dr. Ingi Agnarsson, Dr. D. Li, Prof. Mingsheng Zhu, Dr. Michael L. Draney and Dr. Petra Sierwald for their effort in organizing field trips during which some specimens used in this study were collected. Additional assistance in the field was also provided by N. Corona, J. Brocca, G. B. Edwards, Gustavo R. S. Ruiz, Bruce Beehler, Modi Pontio, Victoria Niesi, William H. Thomas, Max Kuduk, Muse Opiang, Banak Gamui, Jim Robins, Ingi Agnarsson, Jeremy Woon, W. G. Lian and H. Q. Ma. xxv! I also would like to express my gratitude to Dr. Gitanjali S. S. Bodner for her kindness in sharing her knowledge and expertise in euophryine jumping spiders; to Dr. Tamas Szüts for extensive discussions on morphological characters and classification of jumping spiders, and help in obtaining some early literature; to Dr. G. B. Edwards and Dr. Charles Griswold for constructive feedback on this work; and to Dr. William G. Eberhard and Bernhard A. Huber for their help in the study of genitalic evolution in euophryine jumping spiders. Thanks are also due to my Chinese colleagues (Dr. Feng Zhang, Dr. Baoshi Zhang and Dr. Zizhong Yang) for their help in obtaining literature on Chinese salticid fauna. Finally, I would like to thank my parents and sisters for their love, support and encouragement in my educational endeavors. Special gratitude is to my sisters who have always taken good care of my parents when I am away from home.! 1! 1 Introduction Spiders (Arthropoda: Chelicerata: Arachnida: Araneae) currently include more than 40,000 species (Platnick 2011). They are distributed all over the world and have conquered all ecological environments with the only exceptions of the air and open sea (Foelix 1996). Being extraordinary because of their use of silk, their carnivory without exception, and the male palp modified into an indirect copulatory organ, these organisms have long drawn the attention of biologists. 1.1 Taxonomy and phylogeny of the family Salticidae The Salticidae (jumping spiders) is the most diverse spider family, containing 5368 described species in 574 genera (Platnick 2011), almost 1/8 of all spiders. Jumping spiders are most diverse in tropical regions, but can be found in almost all habitats from rainforest to desert. They are easily distinguishable by their two large anterior median eyes (Fig. 1.1), which give them acute vision, enabling them to evolve complex mating behaviors and predatory strategies (Foelix 1996). Although they are easy to tell apart from other spider groups, the classification within the jumping spider family has long been considered problematical, mainly due to its enormous species diversity, varied body forms and the generally simple male palp, which usually provides insufficient information for a detailed classification. The first comprehensive classification of this family was proposed by Eugène Simon (1901a; 1903), in which he placed salticid genera into three groups (Pluridentati, Unidentati, Fissidentati) based on the teeth of the cheliceral retromargin, and then further divided them into 69 subgroups. This classification certainly is inadequate and somewhat artificial, but it shows great insights into the family nonetheless. Jerzy Prószy!ski (1976) made another major advance in salticid systematics and redelimited 11 subfamilies (Aelurillinae, Dendryphantinae, Euophryinae, Heliophaninae, Hyllinae, Marpissinae, Pelleninae, Plexippinae, Salticinae, Sitticinae, Synemosyninae) in a partial classification. Unlike Eugène Simon, he concentrated mainly on the male palp and internal structures of the epigynum, which have been shown to be informative for resolving phylogenetic relationships of various spider groups (e.g. Wang 2002; Agnarsson 2004; Kuntner 2006). 2! During the last ten years, we have made great progress in understanding the phylogeny of Salticidae. Maddison and Hedin (2003a) proposed the first molecular phylogeny of salticids using 5 genes (nuclear 28S, EF1-a; mitochondrial 16S, COI, ND1) from 81 salticid genera and 5 outgroups. Even though the taxon sampling in that study was relatively sparse compared to jumping spiders’ enormous biodiversity, it shed light on the basic phylogenetic structure of this family and provided insights on the traditional classifications based on morphological traits. For instance, their results refined the concepts of the subfamilies Pelleninae and Plexippinae, and discovered several major clades such as the Marpissoida, Plexippoida and Amycoida. After gathering more molecular data from additional salticid taxa, the phylogeny of this family was subsequently improved and more major clades were recognized (Maddison et al. 2008). Other studies have also contributed to the phylogenetic relationships of various salticid groups using either molecular data or morphological characters (Hedin & Maddison 2001a, b; Maddison & Hedin 2003b; Benjamin 2004; Maddison & Needham 2006; Maddison et al. 2007; Bodner 2009; Edwards & Benjamin 2009). In addition to helping us form an adequate classification of this family, those phylogenetic studies have also gradually built up a foundation for conducting studies of evolutionary processes, for instance character evolution and biogeography. A diagram summarizing the current understanding about the phylogeny of salticids from previous studies is shown in Fig. 1.2. In sum, several groups of Salticidae are usually called “basal salticids”, including the subfamilies Lyssomaninae, Spartaeinae, Lapsiinae, Hisponinae and Eupoa Zabka. The remaining of salticids (almost 90% of biodiversity) fall into a clade called the Salticoida (“typical salticids”). Within the Salticoida, amycoids (including Amycinae, Thiodinae, Sitticinae etc.) are the sister group to the rest; the remaining salticoids include several major clades (Astioida, Marpissoida, Aelurilloida, Plexippoida, Euophryinae, Heliophaninae etc.), among and within which most relationships are not yet well resolved. Thus, in spite of the progress that has been made so far, much more work remains on the taxonomy and phylogeny of this large family. 1.2 Taxonomy and phylogeny of the subfamily Euophryinae Initially erected by Eugène Simon in 1901, the Euophryinae was considered as a group in Unidentati in Simon’s classification (Simon 1901a; 1903), and was comprised three genera: Akela, Euophrys and Rhyphelia. However, among them only Euophrys is now widely accepted 3! as euophryine, and most of the genera currently considered as euophryines were scattered among more than 20 groups of Pluridentati, Unidentati and Fissidentati (Simon 1901a; 1903). Jerzy Prószy!ski (1976) clarified the delimitation of the Euophryinae for the first time: the presence of a coiled embolus at the distal end of the palpal tegulum. Thirteen genera were included as euophryines in his partial classification of salticids, although two of them are now placed elsewhere: Admestina is now considered a marpissoid (Maddison & Hedin 2003a) and Marchena a heliophanine (Maddison 1987). The content of Euophryinae was considerably extended by Maddison and Hedin (2003a), who listed 34 genera in this subfamily. In addition, the delimitation was further revised to specify the particular form of the genitalia in euophryines (Fig. 1.3): the plane of the spiral of the embolus is more or less parallel to the longitudinal axis of the palp and a loop in the sperm duct projects towards the centre of the tegulum, and the female epigynum commonly shows two spiral grooves that frame circular areas of relatively transparent and flat integument (Maddison & Hedin 2003a). Based on the work presented here, the Euophryinae has about 1000 described species, and is one of the largest subfamilies in jumping spiders. The majority of euophryine species are found in the tropics of both the Old World and the New World. Spiders of this group show a great variety of body forms: some are large and slender (e.g. Bathippus spp.), some are relatively compact (e.g. Omoedus piceus Simon), some are flattened and pseudoscorpion-lke (e.g. Thiania spectrum Simon), some are weevil-like (Coccorchestes spp.), and some are ant-like (Sobasina spp. and Paraharmochirus spp.). In contrast, the genitalic organs within this subfamily are relatively conservative and show little variation. Similar body forms in different euophryine lineages may have arisen by convergence, and thus contain little phylogenetic signal to resolve relationships among euophryine groups. Consequently, the classification within Euophryinae has been very difficult. Most work on this subfamily has concentrated on describing species or genera (e.g. Bryant 1943; Balogh 1980b; Berry et al. 1996, 1997, 1998; Edwards 2002; Logunov & Azarkina 2008, etc.), and there is little systematic study on the group as a whole. Although a few genera have been reviewed in the past (Prószy!ski 1971; Galiano 1985; Zabka 1987; Jendrzejewska 1995; Gardzinska & Patoleta 1997; Logunov 1998; Bodner 2002; Zabka & Pollard 2002; Logunov & Kronestedt 2003; Richardson & Zabka 2007), the delimitations of most genera are still not clear, e.g. Corythalia and Bathippus. Therefore, the taxonomy within 4! this subfamily is still confused, and it is difficult to identify even some commonly collected species to genus. To date, twelve euophryine genera have been sampled in molecular phylogenetic studies of jumping spiders (Maddison & Hedin 2003a; Maddison et al. 2008) and their monophyly was supported. However, those genera represent only a very small fraction of euophryine biodiversity, and a much more dense sampling of euophryine genera in phylogenetic study is essential to determine if all or most morphologically-recognized euophryines are a monophyletic group. G. S. S. Bodner (2002) presented the only phylogenetic work on the Euophryinae to date. She reconstructed the group's phylogeny using 66 morphological characters and 93 euophryine species of 43 genera. Although the phylogeny was not well resolved, this study has provided some characters that are potentially informative for phylogenetic reconstruction and genera delimitation of this group. . In summary, the Euophryinae is a poorly studied group, and much work remains in its systematics. 1.3 Biogeography of the family Salticidae and subfamily Euophryinae Biogeography is the study of the distribution of organisms in space and time (Wiley 1981). With growing interest in understanding the historical biogeography of various groups, more and more effort has been focused on inferring the temporal and spatial evolution of organisms in a phylogenetic context (Ree et al. 2005; Hines 2008; Ree & Smith 2008; Sanmartín et al. 2008; Clayton et al. 2009; Lamm & Redelings 2009, etc.). Results from molecular phylogenetic studies on jumping spiders indicated that most major salticid groups are primarily restricted to one continental region (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009). For instance, amycoids are numerous in genera and species in the New World but represented in the Old World only by Sitticus (Maddison & Hedin 2003a), whereas astioids are very diverse in Australasia of the Old World and have only a few species in the New World (Maddison et al. 2008). This finding suggests that much of salticid 5! diversification occurred after the separation of continents of the Old and New World (Maddison & Hedin 2003a; Maddison et al. 2008). However, euophryines are a notable exception. They are well represented in both the Old and New World: among the 96 valid genera (see Chapter 3), 60 are from the Old World and 36 are from the New World. This scenario either indicates that the Euophryinae is a rather old lineage and has diversified before the separation of the Old and New World continents, or suggests that members of this subfamily are relatively good “dispersers” and they have achieved one or more intercontinental dispersals in history. The regional isolation of major salticid clades was supported by a recent study of temporal divergence of jumping spiders, which showed that most major salticid clades diversified in the late Eocene or Oligocene when the continents were largely separated (Bodner 2009). This result implied euophryines diversified after the continents separated, possibly requiring intercontinental dispersal to explain their distribution pattern. However, based on the fact that there are two centers of diversity in this subfamily, one in Neotropical area and the other in Australasian region, Hill (2009b) indicated that the Antarctic land bridge might have served as a passageway to enable euophryines to disperse between South America and Australia. This suggests euophryines may have diversified before the last connection of the Old World and New World continents through the Antarctic land bridge broke up. In sum, the picture of historical biogeography of salticids, and euophryines in particular, is still far from complete. A detailed phylogeny and well-estimated divergence time would be important to achieve a more complete understanding of their biogeography. 1.4 Correlated evolution of female and male genitalia Animal genitalia usually tend to undergo a relatively rapid divergence and therefore are often species-specific, showing more differences among closely related species than other somatic traits such as legs and eyes. Thus, genital traits have long been used as diagnostic criteria in taxonomic studies of various animal groups (Eberhard 1985; 2010), especially in spiders (Foelix 1996). 6! Several hypotheses have been postulated in the past to explain the rapid divergence of genitalia. The classic explanation is the “lock and key” theory, which suggests that the divergence of genitalic evolution results from the hybridization avoidance; that is, only males of the same species have the right key to fit the female lock (Hosken & Stockley 2004). The other non- sexual selection mechanism is the pleiotropy hypothesis, which suggests that female and male genitalia diverge via pleiotropic effects of selection on other traits (Hosken & Stockley 2004). The two currently most popular sexual selection hypotheses are sexually antagonistic coevolution and sexual selection by cryptic female choice (Eberhard 2010). The sexually antagonistic coevolution theory focuses on the intersexual conflict in the events associated with reproduction, and indicates that the divergence of female and male genitalia occurs through the evolutionary arms race for adaptations to control over processes such as copulation, insemination and fertilization (Arnqvist & Rowe 2002a; Eberhard 2010). The female cryptic choice hypothesis proposes that the divergence of genitalia is driven by female choice for males with most “favorable” genitalia; that is, females favor males with genitalia that best stimulate them during copulation and give them more opportunity in post-copulatory processes such as sperm transport, oviposition and remating, etc. (Eberhard 1996; 2010). More and more evidence has accumulated to support the role of sexual selection in genital evolution. However, which one of the sexual selection hypotheses is more important to explain the evolution of genitalic organs is still debatable (Hosken & Stockley 2004; Eberhard 2010). Recent studies have revealed the strongly correlated evolution of female and male genitalia in insects (Arnqvist & Rowe 2002b; McPeek et al. 2008; Joly & Schiffer 2010) and web-building spiders (Ramos et al. 2005; Kuntner et al. 2009). But no work has been conducted in jumping spiders. The female copulatory duct and male embolus are important parts of the genitalia of jumping spiders. During copulation, the male inserts the embolus into the female genitalic opening, which enters the copulatory duct and deposits sperms into the spermathecae (Foelix 1996). We would expect if females of a species have a long copulatory duct the corresponding males would have a similarly long embolus. However, the correlated evolution of the lengths of female copulatory duct and male embolus has never been tested explicitly in a phylogenetic context. Genitalia in the Euophryinae are relatively simple in shape, and the lengths of female copulatory 7! duct and male embolus account for most of the inter-specific variation (see Chapter 4), making these characters in euophryines an excellent system to test the above theory. 1.5 Objectives of the thesis The overall goal of the thesis is to understand the phylogeny and to clarify the systematics of the subfamily Euophryinae. On the basis of phylogeny, the historical biogeography and the intersexual correlated evolution of genitalia in euophryine jumping spiders are also explored. Chapter 2 is focused on the phylogeny of euophryine jumping spiders using molecular data. DNA sequences of four gene regions (nuclear: 28S, Actin 5C; mitochondrial: 16S-ND1, COI) are collected from an extensive sample of euophryine taxa representing all continents well. I aim to test the monophyly of euophryines and to resolve phylogenetic relationships within the Euophryinae. The temporal divergence of Euophryinae is also studied, calibrated with known fossil records of jumping spiders, in order to understand the historical biogeography of this subfamily. Chapter 3 aims to clarify the systematics of Euophryinae. In addition to molecular data, I also extensively study morphological traits of a broad range of euophryine genera and species in this thesis. I intend to place all the species not sampled for molecules on the euophryine phylogeny through the assessment of a set of shared morphological characteristics. Also a better understanding of intergeneric and intrageneric morphological variation provides more clear generic delimitations and clarifies problems in the traditional taxonomy of Euophryinae. The performance of morphological traits compared to molecules in resolving the phylogeny of euophryines is also investigated. Chapter 4 focuses on the evolution of genitalia in euophryine jumping spiders. I investigate the correlated evolutionary pattern between the lengths of male embolus and female copulatory duct under the phylogenetic context. In an attempt to narrow down the possible mechanisms causing a positively correlated intersexual evolution of genitalia, I also study the intra-specific variation of traits in two euophryine species that show considerable difference in sexual dimorphism: Chapoda recondita and Antillattus cambridgei. 8! Chapters 5-8 present descriptions of new genera and species of euophryine jumping spiders discovered during this study. ! 9! Figure 1.1. Spider eyes. A. salticid; B. araneid. AME, anterior median eye; ALE, anterior lateral eye; PME, posterior median eye; PLE, posterior lateral eye. 10! Figure 1.2. Summary of phylogeny of jumping spiders (Salticidae). Modified from Maddison et al. 2008 and Bodner 2009. Lyssomaninae Spartaeinae Lapsiinae Eupoa Hisponinae Other amycoids Amycinae Thiodinae Sitticinae Astioida Marpissoida Heliophaninae Hasarieae Aelurilloida Euophryinae Plexippoida Salticus Other salticoids Philaeus group * Salticidae Salticoida Amycoida 11! ! Figure 1.3. Typical genitalic structures of euophryine, as in Parabathippus shelfordi. A. male left palp, ventral view; B. female epigynum, ventral view; C. cleared epigynum, dorsal view. ! ! 12! 2 Molecular phylogeny and temporal scale divergence of the subfamily Euophryinae (Araneae: Salticidae), with implications on its historical biogeography 2.1 Synopsis I investigate phylogenetic relationships of the subfamily Euophryinae, the only jumping spider subfamily that has diversified almost evenly in both the Old and the New World. DNA sequence data of four gene regions (nuclear: 28S, Actin 5C; mitochondrial: 16S-ND1, COI) were collected from 261 jumping spider species. The molecular phylogeny strongly supports the monophyly of Euophryinae, whose content is dramatically extended to 85 genera. Diolenius and its relatives are shown to be euophryines. Euophryines from different continental regions generally form their own clades on the phylogeny with few cases of mixture. Temporal divergence of Euophryinae was also studied, calibrated by known fossils of jumping spiders. The results suggest euophryines radiated rapidly early during their evolutionary history, with most divergences after the Eocene. Given the divergence times, several intercontinental dispersal events are required to explain the distribution of euophryines. Early transitions of continental distribution between the Old and New World may have been facilitated by the Antarctic land bridge, which euophryines may have been uniquely able to exploit compared to other salticid groups of similar age because of their apparent cold tolerance. Two hot-spots of diversity of euophryines are discovered: New Guinea and the Caribbean Islands. Evolution of relatively unusual genitalic forms and myrmecophagy within Euophryinae, as well as implications of molecular phylogeny on taxonomy of Euophryinae are also briefly discussed. 2.2 Introduction The subfamily Euophryinae as here delimited is a major clade of Salticidae (jumping spiders) with about 1000 described species. It belongs to the Salticoida, a lineage comprising the vast majority of jumping spider species (Maddison & Hedin 2003a). Although the classification of Salticidae has been challenging and problematic due to its enormous species diversity (5368 species in 574 genera, Platnick 2011), varied body forms and a generally simple male palp, work on molecular phylogeny of jumping spiders has achieved great progress during the last ten years (Hedin & Maddison 2001a, b; Maddison & Hedin 2003a, b; Maddison & Needham 2006; Maddison et al. 2007; Su et al. 2007; Maddison et al. 2008; Bodner 2009). Multiple clades 13! within Salticoida, such as Amycoida, Astioida, Marpissoida, Plexippoida, Aelurilloida, and Thiratoscirtines, have been discovered or clarified (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009). So far, twelve euophryine genera have been sampled in molecular phylogenetic studies, and monophyly of Euophryinae has been supported (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009). The subfamily Euophryinae was initially erected by Eugène Simon (1901a) using the name “Evophrydeae”. Its delimitation was first clarified by Jerzy Prószy!ski (1976), characterizing it by the presence of a coiled embolus at the distal end of the palpal tegulum. This delimitation was further revised by Maddison and Hedin (2003a): in euophryines, the plane of the spiral of the embolus is more or less parallel to the longitudinal axis of the palp, and a loop of the sperm duct projects towards the centre of the tegulum; the female epigynum commonly shows two spiral grooves that frame two circular areas of relatively transparent and flat integument. Maddison and Hedin (2003a) also considerably extended the content of Euophryinae and listed 34 genera as members of the subfamily, which makes Euophryinae one of the largest groups in jumping spiders. The only phylogenetic work focused on Euophryinae to date is that of G. S. S. Bodner (2002), who reconstructed the phylogeny using 66 morphological characters from 93 euophryine species of 43 genera. Although most parts of the phylogeny were not resolved, this study has provided valuable information on the basic phylogenetic structure, sampling priorities, and potential phylogenetically informative characters of this group. The Euophryinae is particularly interesting for understanding biogeography of jumping spiders. The molecular phylogeny of jumping spiders indicates most major salticid groups are primarily restricted to one continental region, with few or no representative in the other (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009). However, among the shallower groups, Euophryinae is the only notable exception. They are well represented in both the Old World and the New World: among the 85 currently recognized genera (see discussion), 54 are mainly from the Old World and 31 are from the New World. A recent study of divergence times of jumping spiders showed that most divergences of major salticid clades were in the late Eocene or Oligocene when the continents were largely separated 14! (Bodner 2009). This result was concordant with the regional isolation of major salticid clades, and implied euophryines diversified after the continents separated, possibly requiring intercontinental dispersal to explain their distribution pattern. However, David E. Hill (2009b) indicated that the Antarctic land bridge might have served as a passageway that enabled euophryines to disperse between South America and Australia based on the fact that there are two centers of diversity in this subfamily, one in the Neotropics and the other in the Australasia. This suggests euophryines may have diversified before the last connection of the Old World and New World continents through the Antarctic land bridge broke up. Until the phylogeny is resolved and divergence times estimated, we cannot determine whether or not the euophryines diversified before the continents separated. If they diversified after the continents separated, their even distribution could be explained by free and frequent dispersals between the Old and New World continents, or by a few dispersals followed by radiation. If the latter were true, the Euophryinae would not be so unique in distribution in jumping spiders. In this study, I collected molecular data from an extensive sampling of euophryine taxa to test if all or most of euophryines are a monophyletic group, to reveal the continental distribution pattern of euophryines, and to resolve the phylogenetic relationships within the Euophryinae. I also took advantage of the molecular phylogeny with worldwide sampling to explore the temporal evolution of this subfamily calibrated by fossil records of salticids. Implications of the divergence times on biogeography are discussed. Insights from the molecular phylogeny on euophryine classification and on the evolution of unusual genitalic forms and myrmecophagy are briefly introduced. 2.3 Material and methods 2.3.1 Taxon sampling In total, I collected molecular data from 261 jumping spider species, most of them euophryine or potential euophryine species, with the sampling covering all areas of euophryine biodiversity: Eurasia, Africa, Australasia, North America, Central and South America, and the Caribbean Islands. Diolenius and its relatives, and Bristowia were also included because their position on the jumping spider phylogeny had been a mystery (Maddison & Hedin 2003a; Maddison et al. 2008). 15! In addition, sequences of 28 salticid species were obtained from previous studies (Maddison & Hedin 2003a, b; Maddison & Needham 2006; Maddison et al. 2008; Bodner 2009) to represent all other major groups of jumping spiders: Lyssomaninae (1 species), Spartaeinae (2 species), Lapsiinae (2 species), Hisponinae (1 species), Amycoida (4 species), Astioida (2 species), Ballinae (2 species), Marpissoida (2 species), Hasarieae (1 species), Heliophaninae (1 species), Leptorchesteae (1 species), Aelurilloida (2 species), Plexippoida (3 species), Philaeus group (1 species), Cheliceroides (1 species), Nannenus (1 species) and Salticus (1 species). One thomisid species was also included to represent an outgroup of Salticidae. A full list of species included in this study with sequence and locality information is given in Appendix 1. All voucher specimens are preserved in 95% ethanol and stored at -20 ºC, and deposited in the Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM). 2.3.2 DNA extraction, amplification and sequencing For total genomic DNA extraction, usually legs were used. For small specimens, cephalothorax (female) or abdomen (male) or the whole body except genitalia was used to obtain enough amount of total genomic DNA for gene amplification. The rest of each specimen was kept as a voucher. The Puregene DNA Purification Kit (Gentra Systems) was used for total genomic DNA extraction. Agarose gels were run to assess the concentration of the total genomic DNA extracted. Some samples were diluted before amplification. Four gene regions were amplified and sequenced for phylogenetic analyses: the nuclear 28S and Actin 5C, and the mitochondrial 16S-ND1 and COI. Most sequences were amplified and sequenced with primers that have been widely used in jumping spider molecular phylogenetic studies (Maddison & Hedin 2003a, b; Maddison & Needham 2006; Maddison et al. 2007; Maddison et al. 2008; Bodner 2009). However, about 30% of 16S-ND1 sequences could not be amplified by the commonly used primers. For these, two pairs of internal primers (16SND1- WPM-F1/R3 and 16SND1-WPM-F2/R2) were designed and proved to be successful. Three pairs of internal primers (28S-WPM-F1, F2, F3/28S-WPM-R1, R2, R3) were also designed to amplify or sequence a small number of difficult 28S fragments. A summary of all primers used in this study is shown in Table 2.1. 16! The polymerase chain reaction (PCR) was run using either Taq DNA Polymerase (Invitrogen) or Paq5000 DNA Polymerase (Agilent Technologies), with their respective buffers, the dNTPs supplied by Invitrogen and the primers supplied by Oligo. In general, when Paq5000 DNA Polymerase was used the annealing temperature usually had to be increased in order to obtain clean PCR product. The PCR conditions to amplify 28S, Actin 5C and COI were as follows: initial denaturation at 95°C for 2 min; 35 cycles of 45 sec at 95°C, 45 sec of annealing at variable temperature (see Table 2.2), 1 min at 72°C; followed by a 10 min extension at 72°C. The PCR conditions to amplify 16S-ND1 were: initial denaturation at 94°C for 2 min; 35 cycles of 35 sec at 94°C, 35 sec of annealing at variable temperature (see Table 2.2), 70 sec at 65°C; followed by a 10 min extension at 65°C (Maddison & Hedin 2003a, b; Maddison & Needham 2006; Maddison et al. 2008; Bodner 2009). Sequencing reactions were usually conducted with the pair of primers used for PCR. About half of the PCR products were sent to Macrogen Inc. (Korea) and sequenced with the 3730xl DNA analyzer. All other PCR products were purified with the QIAquick PCR purification Kit (QIAGEN Inc.) and visualized on agarose gels stained with ethidium bromide. Purified PCR products were cycle-sequenced with Applied Biosystems BigDye v3.1 terminator chemistry. The sequencing reaction conditions: initial denaturation at 96°C for 1 min; 25 cycles of 10 sec at 96°C, 5 sec at 52°C, and 4 min at 60°C. Cycle-sequenced products were further purified by Sephadex TM G-50 fine (GE Healthcare Bio-Sciences Corp.) using microspin columns (GE Healthcare Bio-Sciences Corp.). Finally they were sent to the University of British Columbia NAPS facility and sequenced on the Applied Biosystems 3730 DNA Analyzer. The chromaseq package (Maddison & Maddison 2010c) for Mesquite (Maddison & Maddison 2010a) was used to obtain sequences from the “.ab1” chromatogram files by Phred (Ewing & Green 1998; Ewing et al. 1998; Green & Ewing 2002) and Phrap (Green 1999), and to further proofread the sequences by comparing against the chromatograms (Maddison & Needham 2006). DNA sequences obtained are listed in Appendix 1. In total, 942 sequences (261 of 28SrDNA, 258 of Actin 5C, 258 of 16S-ND1 and 165 of COI) from 261 jumping spider species were amplified and sequenced, and combined with 102 sequences obtained from previous work (Maddison & Hedin 2003a, b; Maddison & Needham 2006; Maddison et al. 2008; Bodner 2009). 17! 2.3.3 Sequence alignment Multiple sequence alignments were carried out using the Opalescent package (Wheeler & Kececioglu 2007) in Mesquite 2.73 (Maddison & Maddison 2010a). To find the boundary of the intron region, Actin 5C sequences were aligned with the cDNA sequence of Paraphidippus aurantius (Lucas, 1883) (GeneBank Accession No. EU293228; see Vink et al. 2008). The intron data were highly variable and very difficult to align, and thus excluded from analyses (Vink et al. 2008). To find the boundary of ND1 within the 16S-ND1 region, I used amino acid translation in comparison with ND1 sequences of Habronattus oregonensis (Peckham & Peckham) from the complete mitochondrial genome (GeneBank Accession No. AY571145; see Masta & Boore 2004). The protein-coding data (Actin 5C exon region, ND1 and COI) were manually aligned in Mesquite (Maddison & Maddison 2010a) with reference to the amino acid translation using the “Color Nucleotide by Amino Acid” option. Although Wheeler and Kececioglu determined the defaults in the Opalescent package on the basis of careful study of performance of different settings (T. J. Wheeler, pers. comm.), I further explored the performance of ten combinations of gap open/gap extension costs (200/100, 260/69 [default], 300/100, 300/200, 400/100, 400/200, 400/300, 500/300, 600/200, 800/200) for alignment of the non-coding regions, including 28S and 16S (plus the adjacent tRNA). The preliminary ML searches on different alignments and the concatenation of all alignments were conducted using GARLI0.96b8 (Zwickl 2006) with 5 search replicates and the GTR invariant- gamma model (GTR+I+G). Apparent misalignments were recognized by the “Highlight Apparently Slightly Misaligned Regions” tool and manually edited in Mesquite. The ten different gap open/gap extension costs resulted in distinct 28S+16S alignments with the aligned lengths ranging from 2018 to 2287 bp and the parsimony-informative sites ranging from 1055 to 1119 bp. The phylogenetic tree from the “260/69” (default in Opalescent) alignment was most topologically congruent with that from the concatenated alignments. Hence the elision matrix 18! approach (Hedin & Maddison 2001a) is in accord with Opalescent's defaults, and thus I utilized the “260/69” alignment matrices for more comprehensive analyses. The matrix with all genes combined contained 292 sequences and had 4311 sites. The 28S data had 293 sequences and the alignment resulted in 1443 sites. The Actin 5C alignment had 279 sequences and 717 sites. The 16S-ND1 alignment contained 282 sequences and 1161 sites. The COI alignment was of 190 sequences and 990 sites. 2.3.4 Phylogenetic analyses Phylogenetic analyses were performed on the individual gene matrices (28S, Actin 5C, 16S- ND1 and COI) and a combined matrix with all genes concatenated. In the combined matrix, the data were further divided into eight partitions: 28S; Actin 5C first, second and third codon positions; 16S; ND1+COI first, second and third codon positions. 2.3.4.1 Model selection Modeltest 3.7 (Posada & Crandall 1998; Posada & Buckley 2004) in combination with PAUP* 4.0b10 (Swofford 2002) was used to choose the appropriate substitution model for each dataset and each partition via the Akaike Information Criterion (AIC). Most of the datasets had the GTR+I+G as the best-fit model. However, for the individual gene matrices “Actin 5C” and “16S-ND1”, the TVM+I+G model was chosen, and for the partitions in the combined matrix, HKY+G was chosen for the “Actin 5C 2 nd codon position” partition, K81uf+I+G was chosen for the “16S” partition, and TVM+G was chosen for the “ND1+COI 3rd codon position” partition (see Table 2.3). 2.3.4.2 Maximum likelihood (ML) analysis GARLI0.96b8 (Zwickl 2006) was used to perform maximum likelihood analyses on the individual gene matrices, each with 100 search replicates. All the settings were defaults except the model was specified as that chosen in Modeltest (Table 2.3). The model parameters were optimized during the tree-searching process. Bootstrap analyses were also carried out to calculate the replicability of clades in a separate GARLI run with 500 bootstrap replicates. The “genthreshfortopoterm” was set to 10000 (20000 in default) and the “treerejectionthreshold” was set to 20 (50 in default) as suggested in the online manual (https://www.nescent.org/wg_garli/Manual). For each bootstrap replicate, only one search was 19! executed to decrease the computational time. Trees from bootstrap analyses were read into Mesquite for consensus calculation. For the combined matrix, a test version of GARLI (GARLI-partition-r601, https://www.nescent.org/wg_garli/Partition_testing_version) was used for the maximum likelihood tree search because it allows each partition to have its own model or model parameters (see Table 2.3 for models selected for each partition by Modeltest). GARLI search parameters were explored by running preliminary analyses on a combined matrix with fewer outgroup taxa (17 instead of 30) in order to find the best configuration for the partitioned dataset. Totally five different combinations of the parameters “topoweight/modweight/brlenweight/scorethreshforterm” were tried: 0.01/0.005/0.002/0.001; 0.05/0.009/0.002/0.001; 0.1/0.01/0.01/0.001; 0.05/0.009/0.002/0.01; 0.1/0.02/0.02/0.01. The results indicated that the setting “0.05/0.009/0.002/0.001” performed best in finding the best score tree, and thus it was chosen to run the analysis on the final combined matrix with more outgroup taxa. Ten separate GARLI runs each with 50 search replicates (500 search replicates in total) were carried out to find the best ML tree. Because the GARLI run on the combined matrix with eight partitions took about 30 hours to complete each search replicate, it would have been too time expensive to do an extensive bootstrap analysis using GARLI. Instead, 1000 replicates of bootstrap analysis on the combined matrix were completed in RAxML 7.0.4 (Stamatakis 2006; Stamatakis et al. 2008) with a GTRCAT model for each partition and model parameters permitted to differ among data partitions (command line as “./raxmlHPC -f d -# 1000 -b 3 -s CombinedMatrixW30OutPartition.phy -n 1000boots.out -q partition -m GTRCAT”). 2.3.4.3 Bayesian (BI) analysis MrBayes v. 3.1.2 (Huelsenbeck & Ronquist 2001; Ronquist & Huelsenbeck 2003; Altekar et al. 2004) was used to perform a Bayesian analysis on the combined matrix with the data divided into 8 partitions (28S; Actin 5C first, second and third codon positions; 16S; ND1+COI first, second and third codon positions). Each partition was allowed to use the model selected by Modeltest. In case that the model selected by Modeltest was not implemented in MrBayes, the next more complex model available in the program was used instead (see Table 2.3). The analysis was run using the following parameters: mcmcp ngen= 200,000,000 printfreq=1000 samplefreq=1000 nchains=8 savebrlens=yes. However, it was terminated at 120,000,000 generations as the stdDev of clade frequencies already reached 0.007. The outputs from both 20! runs in the analysis were imported in the program Tracer v1.5 (Rambaut & Drummond 2007) to determine when the likelihood has stabilized. In run one, the likelihood became stabilized after 50,000,000 generations, and thus trees sampled before that generation were discarded as “burn- in”; in run two, the likelihood stabilized after 10,000,000 generations, and thus trees sampled before that were removed as “burn-in”. The remaining trees from these two runs were collected and input for the majority rule consensus using PAUP* to count the frequency of various clades. 2.3.4.4 Maximum parsimony (MP) analysis MP analyses were conducted on the individual gene matrices and the combined matrix. TNT 1.1 (Goloboff et al. 2008) was run to find most parsimonious trees using “New Technology Search” method with default settings except (1) sectorial search with XSS, CSS and RSS all selected; (2) rachet and drift selected with default settings; (3) tree fusing with “Do global fuse every 2 hits” and “Dump fused trees to prevent clogging” selected; (4) the option to find the minimum length 20 times for all the matrices except for Actin 5C (set to find the minimum length 50 times instead). To find more equally parsimonious trees, the trees found by TNT were imported into PAUP* 4.0b10 (Swofford 2002) and swapped on using tree bisection reconnection (TBR) branch swapping without constraint except by MAXTREES of 1,000,000. The strict consensus tree of all the equally most parsimonious trees was built in PAUP. 2.3.5 Divergence time analyses Divergence time analyses were conducted on the combined all genes matrix in which 262 euophryine sequences and 30 outgroup sequences were included. To avoid the negative effect on global estimation of divergence time caused by the abnormally long branch in the Athamas clade, the three Athamas species (JXZ142, JXZ181, JXZ345) were deleted. 2.3.5.1 Calibrations Four calibration points were employed in the analyses based on the fossil data of spiders (see detailed summary in Bodner 2009): Salticidae, Salticoida, Lyssomaninae/Spartaeinae, Euophryinae/Sister-group. The minimum divergence age of Salticidae was set at 44 Mya because the oldest known fossil salticids are from Baltic amber which is estimated to be 44-49 Mya old (Weitschat & Wichard 2002 as cited in Penney 2008), and are comprised of members of the extant subfamily 21! Hisponinae (referred to as Gorgopsininae in Petrunkevitch 1950, 1958; Wunderlich 2004) and other basal salticids that cannot be placed in any extant group (Wunderlich 2004). Since there is currently no salticid fossil record from the Cretaceous period (Penney 2008), I hypothesized that the divergence of Salticidae was at most 100 Mya. Salticoids are abundant in the Dominican Republic’s amber deposits (estimated to be 16 Mya; Penney 2008), so the minimum divergence age of Salticoida was set to 16 Mya. No salticoid fossil is currently known from Baltic amber or earlier amber deposits (Penney 2008). Based on this information, I constrained that the divergence of Salticoida as no earlier than 49 Mya. A Lyssomanes species was identified and described by García-Villafuerte and Penney (2003) from the Mexican Chiapas amber deposit dating to 22-26 Mya (Berggren & van Couvering 1974). However, as discussed in Bodner (2009), the taxonomic placement is unclear because of the absence of a synapomorphy linking this species to any extant lyssomanine genus. Thus, I placed the calibration point at the base of the Lyssomaninae/Spaetaeinae node on the tree, and constrained it to be no younger than 22 Mya old (the minimum age of Chiapas amber) and no older than 100 Mya (the maximum divergence time for Salticidae). Six euophryine fossil species have been reported from the Dominican Republic amber: Corythalia ocululiter; C. pilosa; C. scissa; Pensacolatus coxalis; P. spinipes; P. tibialis (see Penney 2008). Among them, the holotypes of Pensacolatus coxalis (SMF Be 938) and P. spinipes (SMF Be 930) were borrowed from the Research Institute and Natural History Museum Senckenberg (Germany). These two species are confirmed to be euophryines based on the typical palp structure. However, it is difficult at this point to place them in any clade within euophryines because of the absence of obvious synapomorphies. As such, the calibration point was put at the base of split between Euophryinae and its sister group instead of at the base of Euophryinae. I set the minimum age of Euophryinae/Sister-group at 16 Mya because the Dominican Republic amber was estimated to be 16 Mya (Penney 2008), and the maximum age at 49 Mya, which is the estimated oldest age of Baltic amber. The maximum constraints to the two calibration points, Salticoida and Euophryinae/Sister- group, are based on the absence of fossils of these groups in Baltic amber. It could be argued that salticoids and euophryines may be much older, but simply absent from the Baltic fauna. 22! With that in consideration, I also ran an independent analysis with the maximum constraints of the Salticoida and the Euophryinae/Sister-group loosened (set to 100 Mya instead of 49 Mya). A summary of the two sets of constraints in the divergence time analyses is given in Table 2.4. Divergence times were mainly estimated by Bayesian MCMC framework using BEAST v1.5.3 (Drummond & Rambaut 2007). The penalized likelihood criterion (Sanderson 2002) implemented in r8s v1.70 (Sanderson 2003; 2004) was also used as an independent corroboration. 2.3.5.2 BEAST (Bayesian Evolutionary Analysis Sampling Trees) The estimation of divergence times by the Bayesian criterion was conducted in BEAST v1.5.3. BEAUti v1.5.3 (Drummond & Rambaut 2007) was used to generate a BEAST XML file. The XML file was manually edited before importing into BEAST for analysis. The dataset was also divided into eight partitions (28S; Actin 5C first, second and third codon positions; 16S; ND1+COI first, second and third codon positions). The model assignment for each partition was the same as in the MrBayes analysis (see Table 2.3). Four MRCA (most recent common ancestor) groups were established: Salticidae, Salticoida, Lyssomaninae/Spartaeinae, Euophryinae/Sister-group, with none of them restricted to be monophyletic. The upper and lower constraints of the calibration points (see Table 2.4 for details of the two sets of constraints) were specified as tMRCA (time to most recent common ancestor) prior to estimating the age of divergence using the relaxed (uncorrelated lognormal) molecular clock model. The best tree found by a preliminary ML search using GARLI0.96b8 (5 search replicates and GTR+I+G model) was modified into an ultrametric tree in Mesquite (Maddison & Maddison 2010a) and used as the starting tree for the BEAST analysis. Analysis under each constraint was run for 200,000,000 generations (trees sampled at every 1000 generations). Tracer v1.5 (Rambaut & Drummond 2007) was used to check when the MCMC chains had reached to the stationary distribution by visual inspection of plotted posterior estimates. Using the LogCombiner v1.5.3 program (Drummond & Rambaut 2007), trees sampled during the first 50,000,000 generations (25%) were removed as burn-in and the remaining trees were resampled at lower frequency (every 2000 generations) to generate a smaller tree file for annotation. For easy comparison with the results from r8s analyses, these trees from the BEAST analyses were summarized on the best ML tree from the GARLI all-genes partitioned analysis in 23! TreeAnnotator v1.5.3 (Drummond & Rambaut 2007) using the “User target tree” option, and then displayed with age in millions of years using Mesquite 2.74 (Maddison & Maddison 2010a). In addition, the estimates from the BEAST analyses were also summarized into a Maximum Credibility Tree in TreeAnnotator v1.5.3 (Drummond & Rambaut 2007) using “keep target height” option, and then displayed in FigTree v1.3.1 (Rambaut 2009) or Mesquite 2.74 (Maddison & Maddison 2010a). 2.3.5.3 Penalized likelihood implemented in r8s The best ML tree from the GARLI all-genes partitioned analysis was used for r8s analyses. Calibration constraints were the same as in the BEAST analyses. Cross-validation was used to choose between the Langley-Fitch method (LF, strict molecular clock, Langley & Fitch 1974) and the penalized likelihood method (PL, relaxed molecular clock, Sanderson 2002), each with the TN algorithm and additive penalty. The best smoothing parameter for the PL method was also found during the cross-validation procedure with the command “divtime method=PL algorithm=tn crossv=yes cvstart=0 cvinc=0.5 cvnum=8;”. This procedure found that the best smoothing parameter for the PL method was 3.2 in Analysis One and 10 in Analysis Two. The correctness of solutions was checked by the “CheckGradient” command (set checkGradient=yes; set activeEpsilon=0.001;). Ten initial starting conditions were explored using the command “set number_time_guesses=10;”. The chronogram obtained from r8s was imported and visualized in Mesquite 2.74 (Maddison & Maddison 2010a). 2.4 Results 2.4.1 Phylogenetic analyses 2.4.1.1 All genes combined The results from the ML, MP and BI analyses are summarized in Fig. 2.1. The best tree found by ML analysis (Fig. 2.2) has lnL -181734.801. The tree shows the monophyly of Euophryinae, which is strongly supported by the ML bootstrap analysis (bootstrap value=0.989). On the phylogeny, euophryine taxa from different continental regions tend to form their own clades with few cases of mixture (Fig. 2.1). The phylogeny suggests the Euophryinae is the sister to a clade with many salticoid groups (Nannenus, Cheliceroides, Heliophanus, etc.). However, this relationship is not supported by the ML bootstrap analysis. 24! Diolenius and its close relatives fall into a strongly supported clade (ML bootstrap value=1.0) within the subfamily Euphryinae. However, Bristowia and “Bathippus” pahang Zhang, Song & Li fall outside of the Euophryinae. The tree recovers most of euophryine genera when multiple species of a genus were included in the analysis, and they are well supported by the ML bootstrap analysis, e.g. bootstrap support value is 1.0 for Tylogonus and 0.997 for Euryattus. In addition, the phylogeny shows close relationships of some genera. For instance, the genera Variratina, Bulolia, Leptathamas and Coccorchestes from Papua New Guinea falls into one clade with ML bootstrap value as 0.983. The consensus tree from the 180,000 trees retained from the two runs of the MrBayes analysis is similar to the ML tree, but with some clades collapsed. Many resolved clades have high posterior probability support values, e.g. 1.0 for Euophryinae, 1.0 for the clade with Diolenius and its relatives, 1.0 for Agobardus spp., 1.0 for Bathippus plus Canama. The result of MrBayes analysis also strongly supports (posterior probability value=1.0) the clade with Euophryinae and other salticid groups as shown in the ML tree (Fig. 2.2). However, the placement of Euophryinae as the sister to the remaining members is only moderately supported (posterior probability value=0.9). The TNT analysis was stopped after the best score was hit 10 times, and 12 equally parsimonious trees (score=44516) were saved. Swapping on the 12 trees in PAUP did not improve the score, but found an additional eight trees of the same score. The strict consensus tree of the 20 equally parsimonious trees is similar to the ML tree except that some of the relatively deeper branches are unresolved. 2.4.1.2 28S Fig. 2.3 summarizes results from the ML and MP analyses. The ML analysis best tree has lnL -51870.560. The phylogeny also strongly supports the monophyly of the Euophryinae (ML bootstrap value=0.824). The tree recovers most of the genera or closely related genera, with the exceptions of the genus Corticattus (JXZ305, 337), and the African euophryines with Thyenula spp. (JXZ103, 104, 107, 108, 149, 192) and two misplaced Saitis species (JXZ105, 106). However, only some of the recovered clades are well supported by the ML bootstrap analysis, e.g. 0.902 for the clade with Diolenius and its relatives. 25! Also, the ML tree indicates an abnormally long branch for Athamas nitidus (JXZ142). Since no 28S rDNA sequences were obtained from the other two Athamas species included in this study (JXZ182, 345), the three Athamas species were excluded in the combined all genes analyses in order to avoid the effect of the abnormally long branch and large portion of missing data on resolving the phylogeny. The TNT analysis was stopped when the best score was hit 20 times. Sixty-five equally parsimonious trees (score=11293) were saved. Swapping on those trees in PAUP found more equally parsimonious trees (1,000,000) but without improvement on the score. The strict consensus tree of the 1,000,000 equally parsimonious trees recovers most genera and closely related genera, but many of the relatively deeper relationships are unresolved. 2.4.1.3 Actin 5C Fig. 2.4 summarizes results from the ML and MP analyses. The best ML tree has lnL= -13886.791, with many genera and closely related genera recovered, such as Agobardus spp. and the Bathippus-Canama clade. However, the ML bootstrap analysis shows low levels of clade support except those corresponding to closely related species. Seven equally parsimonious trees (score=2687) were saved from the TNT analysis after the best score was hit four times. Swapping on those trees in PAUP found 1,000,000 equally parsimonious trees of the same score. The strict consensus tree of the 1,000,000 equally parsimonious trees shows poor resolution at the deeper branches. 2.4.1.4 16S-ND1 The results from the ML and MP analyses are summarized in Fig. 2.5. The ML analysis best tree has lnL -69623.304. Similar to 28S and Actin 5C, most of the clades corresponding to a genus or closely related genera are recognized, but many of them are poorly supported by the ML bootstrap analysis. The TNT analysis was stopped after the best score was hit two times and five equally parsimonious trees (score=18014) were saved. Swapping on those trees in PAUP found an 26! additional 41 equally parsimonious trees of the same score. The strict consensus tree of the 46 equally parsimonious trees shows little resolution except on some of the nodes corresponding to closely related species or genera. 2.4.1.5 COI The results from the ML and MP analyses are summarized in Fig. 2.6. The best ML tree found has lnL -43516.919. Compared to the results from other genes, fewer clades corresponding to genus and related genera are recovered with low level of clade support from the ML bootstrap analysis. The TNT analysis found six equally parsimonious trees (score=11664) after the best score was hit four times. Swapping on those trees in PAUP found additional 29 trees of the same score. The strict consensus tree of the 35 equally parsimonious trees shows little resolution. 2.4.2 Divergence time analyses A summary of the age estimates for major nodes of euophryines from the BEAST and r8s analyses under different set of constraints is shown in Table 2.5. The divergence time chronogram of euophryines and outgroups using the ML topology is presented in Fig. 2.7, with the branch length as the median age estimated from BEAST Analysis Two (with the maximum constraints to Salticoida and Euophryinae/Sister-group inactive). BEAST analysis with maximum constraints on Salticoida and Euophryinae/Sister-group active (Analysis One) indicates the first divergence within Euophryinae (node 1) happened in the Oligocene (median = 30.19 mya; 95% highest posterior density (HPD) = 37.84-28.93 mya); and most of the subsequent divergences into major extant genera happened before the late Miocene (10 mya). Loosening the maximum constraints to the nodes Salticoida and Euophryinae/Sister- group (Analysis Two) results in similar but slightly earlier median age estimate for all nodes, but the 95% HPD interval is rather wide. For instance, the BEAST Analysis Two estimates the median age for the first divergence within Euophryinae (node 1) is 33.84 mya and the 95% HPD is 55.52-23.10 mya. In addition, the BEAST analyses estimate the family Salticidae is about 50- 60 mya (median age estimate = 53.28 mya from BEAST Analysis One; median age estimate = 60.03 mya from BEAST Analysis Two). 27! In BEAST Analysis One, the maximum credibility tree is very similar to the ML tree from the partitioned GARLI analysis on the combined all genes matrix, except minor differences in groupings within a neotropical clade and the relative position of Phasmolia elegans (JXZ225) within the Papua New Guinea Clade. However, the maximum credibility tree from the BEAST Analysis Two is quite different from the ML all genes tree, and some groupings contradict the MrBayes result. For instance, the MrBayes analysis strongly supports the clade with Parabathippus and the Holarctic euophryines including Euophrys, Talavera, Chalcocirtus, Pseudeuophrys (posterior probability value = 1.0), whereas in the maximum credibility tree from the BEAST Analysis Two, the clade with Parabathippus is sister to the Neotropical Cobanus/Sidusa Clade. Compared to the BEAST analyses, the r8s analyses result in older age estimates for all nodes especially when the maximum constraints for the two calibration points, Salticoida and Euophryinae/Sister-group, were inactive. For example, the r8s Analysis One estimates the divergence of Euophryinae (node 1) started at 39.34 mya (upper boundary of 95% HPD is 37.84 from the BEAST Analysis One); whereas the estimation for the same node from the r8s Analysis Two is 66.53 mya (upper boundary of 95% HPD is 55.52 from the BEAST Analysis Two). In addition, the posterior probability values for some clades were lower in the BEAST analyses than in the unconstrained MrBayes analysis. For instance, the clade (node 2) with all euophryines excluding the Anasaitis-Corythalia Clade (node 3) has posterior probability value 0.99 from the unconstrained MrBayes analysis, but only 0.75 from the BEAST Analysis One. 2.5 Discussion 2.5.1 Phylogeny of Euophryinae I synthesize the results from various phylogenetic analyses in Fig. 2.1. Although some relationships, especially those at relatively deeper levels are still unresolved, this study provides a basic phylogenetic framework of euophryines, and resolves some clusters of euophryine genera. However, detailed discussion of euophryine generic groups is beyond the scope of this 28! Chapter, and will be reviewed in detail in Chapter 3. The following discussion is based mainly on the results from the ML and BI analyses. 2.5.1.1 Monophyly and content of Euophryinae Previous work (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009) supported the monophyly of Euophryinae with a small sample of euophryine genera. The present study dramatically extends the sampling, including more than 200 species of suspected euophryines. All fall into a monophyletic group with the exception of “Bathippus” pahang Zhang, Song & Li (JXZ028, see discussion of implications on taxonomy of Euophryinae), and thus the monophyly of Euophryinae is further strongly supported. In addition, the unusual Diolenius and relatives, which have not previously been associated with euophryines, are here shown to be derived euophryines. Maddison and Hedin (2003a) clearly indicated 34 genera as euophryines and suggested the euophryines are recognizable by a typical male palp structure: the embolus is free and coiled at the distal end of the tegulum, with the plane of spiral more or less parallel to the longitudinal axis of the tegulum; the sperm duct forms a retrolateral loop projecting to the centre of the tegulum; and the epigynum usually has a “window”-like structure. Almost all of them are sampled in this study except Ascyltus and Spilargis, both of which are no doubt euophryines based on the genitalia. This study adds to Maddison and Hedin's list of euophryines another 53 genera. The full list of genera belonging to Euophryinae which have been supported by the molecular phylogeny is as follows: Agobardus, Amphidraus, Anasaitis, Antillattus, Asaphobelis, Athamas, Bathippus, Belliena, Bulolia, Bythocrotus, Canama, Chalcolecta, Chalcolemia, Chalcoscirtus, Chalcotropis, Chapoda, Chinophrys, Cobanus, Coccorchestes, Colyttus, Compsodecta, Corticattus, Coryphasia, Corythalia, Cytaea, Dinattus, Diolenius, Donoessus, Ecuadattus, Efate, Emathis, Euophrys, Euryattus, Foliabitus, Hypoblemum, Ilargus, Jotus, Junxattus, Lagnus, Laufeia, Lepidemathis, Leptathamas, Lycidas, Maeota, Maileus, Maratus, Marma, Mexigonus, Naphrys, Nebridia, Neonella, Ohilimia, Omoedus, Orcevia, Palpelius, Parabathippus, Paraharmochirus, Parvattus, Pensacola, Petemathis, Phasmolia, Popcornella, Pristobaeus, Prostheclina, Pseudeuophrys, Pystira (=Omoedus, see Chapter 6), Saitis, Servaea, Sidusa, Siloca, Sobasina, Soesilarishius, Talavera, Tariona, Thiania, Thianitara, Thorelliola, Thyenula, Truncattus, Tylogonus, Variratina, Viribestus, Xenocytaea, Zabkattus, Zenodorus 29! (=Omoedus, see Chapter 6). More genera can be included in the subfamily based on their similar morphology with the above genera, which will be addressed in Chapter 3. 2.5.1.2 Diolenius and its relatives are euophryines The results show Diolenius, Chalcolecta and Ohilimia fall into a clade within the subfamily Euophryinae with Chalcolemia, Efate, Paraharmochirus and Sobasina. This clade is strongly supported by the combined all genes analyses as well as by 28S. Spiders of this clade are elongate and usually have very long first legs armed with many ventral macrosetae on tibiae and metatarsi. Some have abnormally long trochanters of first legs (e.g. Diolenius and Ohilimia); some are more or less ant-like (e.g. species of the genera Efate, Paraharmochirus and Sobasina). Genitalia of spiders in this clade are usually quite different from the typical euophryine type (see discussion on other implications from the molecular phylogeny). Diolenius and its relatives were once placed in the subfamily Dioleninae (Simon 1901a, 1903; Gardzinska & Zabka 2005). The present study suggests they belong within the subfamily Euophryinae. The genera Lystrocteisa Simon and Ligonipes Karsch were once considered as diolenines, but have been placed in the newly erected group Astioida based on molecular data (Maddison et al. 2008). Bristowia is a genus described from the Oriental and Afrotropical regions. Szüts (2004) indicated the general body form of Bristowia resembles Diolenius, whereas the copulatory organs and the spination of the first legs are similar to Phintella Strand. An unidentified Bristowia species from Gabon was included in this study, and the results show that it is not closely related to Diolenius and its relatives but falls outside of the euophryine clade. However, its position on the jumping spider phylogeny is still uncertain. 2.5.1.3 “Bathippus” pahang is not a euophryine “Bathippus” pahang was described from Malaysia (Zhang et al. 2003). Although its body form closely resembles euophryine, the molecular phylogeny indicates that it is unrelated to the other Southeast Asian “Bathippus” spp. (transferred to Parabathippus, see Chapter 8), and indeed falls outside of the euophryine clade and clusters with Nannenus lyriger (d105). The male palp of B. pahang has free and coiled embolus. However, the plane of the spiral is perpendicular to the longitudinal axis of the bulb and the tegulum lacks retrolateral sperm duct loop, which are not consistent with the typical euophryine palp (Maddison & Hedin 2003a). Its palpal structure 30! is also different from Nannenus spp. (Prószy!ski 1984; 1987), which has fixed embolus. Thus, B. pahang may represent a novel origin of a coiled embolus. 2.5.1.4 Position of Euophryinae in jumping spider phylogeny The first jumping spider molecular phylogeny (Maddison & Hedin 2003a) showed that Euophryinae was a clade within Salticoida and suggested it might be sister to the Plexippoida, while the recently revised phylogeny by Maddison et al. (2008) suggested it was the sister to the Plexippoida plus Philaeus group and Salticus. In the analysis of Bodner (2009), the Euophryinae fell in a large clade with some salticoid groups such as Plexippoida, Aelurilloida, Heliophaninae, Philaeus group, etc. This study focuses mainly on phylogenetic relationships within the Euophryinae, and thus other salticid clades were not densely sampled. However, the ML tree from the combined all genes data proposed a similar position for the Euophryinae as that of Bodner (2009), as the sister to a clade with many salticoid groups including the unclassified genera Nannenus, “Bathippus” pahang, Bristowia, Cheliceroides, Salticus, and the already classified genera Chinattus (Hasarieae), Heliophanus (Heliophaninae), Yllenus (Leptorchesteae), Aelurillus and Freya (Aelurilloida), Philaeus (Philaeus group), Plexippus, Habronattus and Havaika (Plexippoida). However, this relationship is only moderately supported by the MrBayes analysis of the combined all genes dataset, and thus more data are needed to further clarify the exact position of the Euophryinae on the jumping spider phylogeny. 2.5.2 Implications on taxonomy of Euophryinae The molecular phylogeny reveals many problems in the taxonomy of Euophryinae, which will be reviewed in detail in Chapter 3 and thus only are briefly addressed here. The monophyly of the genera Chalcoscirtus, Cobanus, Colyttus, Corythalia, Cytaea, Euophrys, Hypoblemum, Jotus, Lycidas, Pensacola, Saitis, Sidusa, Siloca, and Thiania is challenged (Fig. 2.1). The results suggest some species in Cobanus, Corythalia, Euophrys, Margaromma, Pensacola, Saitis, Sidusa, Siloca and Wallaba are wrongly placed and need to be transferred into either existing or newly erected genera (Fig. 2.1). The phylogeny supports separating the Southeast Asian species once placed in Bathippus into the newly erected genus Parabathippus 31! (see Chapter 8) and synonymizing Pystira and Zenodorus with Omoedus (see Chapter 6). One species once placed in Pystira (Omoedus ephippigera, JXZ022), one species of Omoedus (Omoedus cf. piceus, JXZ206) and five species once placed in Zenodorus (Omoedus cf. danae, JXZ274; Omoedus cf. durvillei, JXZ294; Omoedus cf. metallescens, JXZ154; Omoedus cf. ponapensis, JXZ138; Omoedus orbiculatus, JXZ088, 136) were included in the analyses, and the results suggest Pystira and Omoedus are embedded within Zenodorus, which makes Zenodorus paraphyletic. 2.5.3 Temporal divergence and implication on biogeography of Euophryinae Even though the present study is focused on the subfamily Euophryinae with other major salticid clades only sparsely sampled, it obtained similar age estimates for the family Salticidae and its main groups as the previous study by Bodner (2009). We both date an early Paleogene origin for Salticidae. This is consistent with the prediction for the age of Salticidae by Penney et al. (2003, also see Penney 2010) based on paleontological evidence. Implications of divergence time estimates on the historical biogeography of Euophryinae are discussed below, based mainly on the BEAST analyses. 2.5.3.1 Diversification in the southern hemisphere Euophryines are most diverse in the Neotropical and Australasian regions. This might suggest they have a Gondwanaland origin (Hill 2009b). After the initial break-up of Gondwana in the Jurassic (165-150 mya), different continental blocks such as India, Africa and Tasmantis (including New Zealand and New Caledonia), broke away successively and moved towards their current positions. By the late Cretaceous (ca. 75 mya), only the southern Gondwana composed of Australia, South America and Antarctica remained together. Australia and South America remained in contact through an Antarctic land bridge until the end of the Eocene (35 mya), when the Australian plate finally separated from Antarctica and drifted rapidly towards Asia. The contact between southern South America and Antarctica was broken by the Oligocene (30-28 mya) (Sanmartín & Ronquist 2004). The BEAST analyses result in a wide range of 95% HPD for the very early divergence of euophryines with the upper boundary earlier than 35 mya. This indicates euophryines may be old enough for the southern Gondwanaland origin. However, similar to findings in other salticid lineages (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009), the reconstructed phylogeny shows that the Old World and New World 32! euophryines are usually grouped in their own clades with very few cases of mixing together — most divergences within Euophryinae appear to be intracontinental. This pattern suggests even if ancestors of extant euophryines originated in southern Gondwana, most diversification within the Euophryinae happened after Australia, South America and Antarctica broke up; otherwise, we would expect taxa with Old World and New World distributions should be more mixed on the phylogeny. After the break-up of the connection between Australia and South America through the Antarctic land bridge (Sanmartín & Ronquist 2004), Antarctica may still have served as a stepping-stone to enable the faunal interaction between Australia and South America for a period since the Antarctic ice cap did not form until the middle Miocene (14-11 mya) (Kennett 1977). A recent study also indicates that the last vestige of a tundra community, comprising diatoms, palynomorphs, mosses, ostracodes and insects, persisted in Antarctica till 13.8 mya before this continent started to have a full polar climate (Lewis et al. 2008). Thus, the Antarctic land bridge may have played an important role in facilitating the faunal exchange between southern South America and Australia in early evolutionary history of euophryines. However, if euophryines used the Antarctic land bridge early in their history to cross between South America and Australasia, why did not other major salticoid clades in those continents and of about the same age successfully cross Antarctica and diversify in both hemispheres? Two major salticoid clades, Amycoida (mostly in the Neotropics) and Astioida (mainly in Australasia), are highly diversified in one of these two regions but have very few representatives in the other. According to Bodner (2009), the Amycoida (33.4 mya) and Astioida (30.5 mya) are similar in age, slightly older than the Euophryinae (26.9 mya). Divergence time analyses of the present study, with greater taxon sampling, estimates a slightly older age for Euophryinae (see results), about as old as Amycoida and Astioida. If Amycoida and Astioida are as old as Euophryinae, why did not they cross Antarctica? At the dated time of the early diversification of euophryines, a significant worldwide temperature decline led to a switch from a “greenhouse” to an “icehouse” world, and probably also caused a dramatic faunal turnover on the earth with warm tropical taxa being replaced by cooler temperate taxa (Zanazzi et al. 2007; Hines 2008). This implies that euophryines would have needed cold tolerance if they were to have crossed through the polar Antarctica. There are hints that extant euophryines may be cold tolerant. First, even though euophryines and amycoids are found throughout most of South America, amycoids tend to dominate diversity at lower 33! elevations, euophryines at higher elevations (Maddison, personal observation). Second, by far the most common jumping spiders in cold and wet areas of southern South America are a group of euophryines described as “Euophrys” (e.g. “Euophrys” a-notata, “Euophrys” patagonica, “Euophrys” tehuelche, etc.). Tolerance to low temperature would be especially crucial for spiders to cross Antarctica after the late Eocene when the temperature dramatically dropped in this continent. Therefore, one likely scenario would be that in the Oligocene when the globe switched from a “greenhouse” to an “icehouse” world, most amycoids and astioids retreated towards the equator, but more euophryines persisted further south, with some crossing Antarctica to flourish in the other hemisphere. 2.5.3.2 Dispersal is important in the diversification of euophryines Previous studies have suggested vicariance as the major mechanism in the historical biogeography of animals, especially in southern hemisphere (Sanmartín & Ronquist 2004; Bossuyt et al. 2006; Gamble et al. 2008; Toon et al. 2010, etc.). However, recent studies indicate dispersal may be more important in shaping the biogeographic pattern of some animals than we have expected (e.g. Braby et al. 2007; Hines 2008; Kelly et al. 2009). Dispersal have played an important role in the historical biogeography of euophryines because several clades with mixed euophryine lineages from different continents are younger than the continental splits. In addition to the possible trans-Antarctic dispersals between southern South America and Australia during the early evolutionary history of euophryines, other dispersal routes may have existed. 2.5.3.2.1 Dispersals across Wallace-line The faunal interactions across the Wallace-line between Australasia and Eurasia may have been facilitated by the collision of the Australian and Asian plates at the end of the Oligocene (about 25 mya, Hall 2002). Many genera, such as Thiania, Saitis, Thorelliola, Cytaea, Euryattus, Palpelius, Zenodorus (=Omoedus), etc., have successfully crossed the Wallace-line and leaked into the other region (Hill 2010). 2.5.3.2.2 Dispersals in north hemisphere Dispersals between North America and Eurasia are most likely through the Bering land bridge, which has been suggested to be important for intercontinental faunal exchange (Sanmartín et al. 2001). Dispersals through this route can be detected in the clade with Euophrys, Chalcoscirtus, Talavera and Pseudeuophrys, which are northerly, with a Holarctic distribution pattern (node 7). 34! 2.5.3.2.3 Dispersals between Neotropic and Nearctic A few elements in the Nearctic euophryine fauna apparently are associated with Neotropical euophryine clades, such as Anasaitis canosa (Walckenaer), Naphrys spp., Mexigonus spp. and Neonella vinnula Gertsch. Faunal exchange between these two regions was possibly through the rows of islands in the Caribbean Sea and the Panama Island Arc, and later via the Panama Isthmus (Sanmartín & Ronquist 2004; Iturralde-Vinent 2006). 2.5.3.2.4 Dispersals into Africa The divergence of African euophryines is dated no early than the late Eocene, and the African clade tends to be not at the basal position of the euophryine phylogeny. This suggests they are likely radiations (dispersals) from other continents. However, whether they are radiations from Eurasia after Africa collided with the Eurasian plate in Paleocene (60 mya) (Sanmartín & Ronquist 2004), or from other disjunctive continents via across-oceanic dispersal is still uncertain. 2.5.3.3 Two “hot-spots” of euophryine diversity Euophryines have independently developed two “hot spots” of diversity: New Guinea (Old World) and the Caribbean Islands (New World). Euophryine spiders are more dominant in both species and specimen diversity in the jumping spider community of these two regions compared to any other areas in the Neotropics and Asian tropics (also see Maddison & Zhang 2011). For example, during a jumping spider expedition to the Dominican Republic and Puerto Rico in 2009, we counted roughly that at least 80% of salticid species and specimens collected were euophryines. 2.5.3.3.1 Divergences in New Guinea Most euophryines from New Guinea fall into one clade and represent a single radiation (Fig. 2.7, node 18), with the only exceptions being the Cytaea-Euryattus Clade and the Thorelliola Clade. This large New Guinea clade contains the most diverse body forms of euophryines. Some of them, such as the ant-like form (Sobasina and Paraharmochirus), the beetle-like form (Coccorchestes), and the weird forms of Diolenius and Leptathamas, are not found anywhere else in the world but New Guinea. The age for this radiation is estimated as 27-32 mya, which implies euophryines were probably among the earliest salticoid immigrants in New Guinea and 35! explains why they are so abundant in the local salticid community. The other two clades that are also well-known from New Guinea, Thorelliola and Cytaea/Euryattus, are much younger (16-19 mya), and they seem to represent independent radiations later into New Guinea. 2.5.3.3.2 Divergences in Caribbean In contrast, the Caribbean euophryines are embedded within two big clades, and seem to represent as many as five independent radiations, most likely from South America (Fig. 2.7). These radiations are dated from the late Eocene to the middle Miocene when various rows of islands existed in the Caribbean Sea (Iturralde-Vinent 2006). Euophryines in the Caribbean Islands have all the body forms that appear in the other areas of Neotropics. However, the body form diversity from this region is not comparable to that of New Guinea. 2.5.4 Other implications from the molecular phylogeny Although most of the genera suggested as Euophryinae have typical palp and epigynum as described above (and Maddison & Hedin 2003a), some of them have other forms of genitalia. For instance, the embolus of Diolenius’s palp comes from the proximal end of the bulb instead of the distal end; the sperm duct of Chalcolecta does not form the retrolateral loop; the plane of the embolic spiral of some Coccorchestes species is perpendicular to the longitudinal axis of the tegulum but not parallel; Tylogonus has fixed embolus; Anasaitis has no obvious window in the epigynum. However, these genera are closely related to others with typical euophryine-like genitalia, or clearly derived from within a clade with typical genitalia. This indicates that these abnormal genitalic forms are derived from the typical euophryine-like genitalic structures. Myrmecophagy (ant-eating) is relatively rare in jumping spiders probably due to the ants’ defenses of powerful mandibles, poison-injecting sting and formic acid (Jackson & Li 2001). However, some salticid species routinely feed on ants using ant-specific prey-capture tactics. Among more than 20 salticids that have been thoroughly studied and appear to be ant-feeding specialists, about half of them are euophryines: Chalcotropis (six species), Xenocytaea (two species), Zenodorus (=Omoedus, three species), Anasaitis (one species), Naphrys (one species) (Jackson et al. 1998; Clark et al. 2000; Jackson & Li 2001). The scattered placement of these lineages on the phylogeny seems to suggest the myrmecophagic behavior may have evolved not only independently in the Old World and the New World, but also separately in different lineages in each continent. However, more ecological data on whether or not other euophryines 36! are ant-feeding specialists are needed to understand the evolution of myrmecophagy in Euophryinae. Within the big New Guinea radiation (node 18), the Diolenius-Clade (node 19) contains species that resemble ants (e.g. Sobasina and Paraharmochirus), and the Omoedus-Clade (node 20) has members specialized in ant-feeding (e.g. Omoedus durvillei (Walckenaer), see Jackson & Li 2001 as Zenodorus durvillei). An interesting finding from this study is that these two lineages are both about 19-22 mya. Their similarity in age implies that their ancestors followed two completely different evolutionary paths after encountering ants: one went for ant-mimicry, and the other for ant-eating. 2.6 Conclusions The molecular phylogeny from an extensive sampling of euophryine taxa strongly supports the monophyly of the Euophryinae. An additional 53 genera are shown to be euophryines, including Diolenius and its relatives, which have unique body forms and genitalic forms different than typical euophryines. Within the Euophryinae, relationships at relatively shallow levels on the phylogeny are usually well resolved, and some genera, such as Anasaitis and Corythalia, Canama and Bathippus, etc., are suggested to be closely related. The molecular phylogeny also reveals that euophryines from different continental regions generally form their own clades with few cases of mixture. The estimation of temporal divergence of Euophryinae calibrated by known fossils suggests rapid radiations early during their evolutionary history, with most divergences after the Eocene. Given the age, several intercontinental dispersal events are required to explain the distribution of euophryines. Ancestors of extant euophryines may have been able to survive better under low temperature than other salticids, which may have been crucial to their early dispersals especially through the Antarctic land bridge and their diversification in both the Old World and New World. Euophryines have independently developed two “hot-spots” of diversity: New Guinea (Old World) and the Caribbean Islands (New World). Most euophryines from New Guinea fall into one clade and represent a single radiation, with the only exceptions being the Cytaea-Euryattus clade and the Thorelliola clade. In contrast, the Caribbean euophryines are embedded within two big clades, and seem to represent as many as five independent radiations. Some genera that fall in the clade Euophryinae show unusual genitalic forms (e.g. Diolenius) compared to typical euophryines. The molecular 37! phylogeny indicates that these abnormal genitalic forms are derived from the typical euophryine genitalic structures. Some lineages of euophryines appear to be ant-feeding specialists. The scattered pattern of these lineages on the phylogeny seems to suggest independent origins of the myrmecophagic behavior not only in the Old World and the New World, but also in each continent. However, the evolution of myrmecophagy in Euophryinae is still uncertain and more ecological data on whether or not other euophryines are ant-feeding specialists are needed. 38! Table 2.1. List of primers and primer sequences used in gene amplification and sequencing. “*” indicates the primer is for sequencing reactions only. Gene region Primer name Primer sequence Reference 28S rDNA ZX1 (forward) 5' ACCCGCTGAATTTAAGCATAT 3' van der Auwera et al. 1994 (with the 11th bp changed from a Y to T) 28SO (forward) 5' GAAACTGCTCAAAGGTAAACGG 3' Hedin & Maddison 2001a 28SC (reverse) 5' GGTTCGATTAGTCTTTCGCC 3' Hedin & Maddison 2001a 28S-WPM-F1 (forward) 5' GGTCGCGGGAAATGTGGCG 3' this study *28S-WPM-F2 (forward) 5' GTAGCGGGTGCGAGGCCCATAG 3' this study 28S-WPM-F3 (forward) 5' AGTCGGGTTGCTTGGGAGTGC 3' this study 28S-WPM-R1 (reverse) 5' GCACATGTTAGACTCCTTGGTC 3' this study 28S-WPM-R2 (reverse) 5' CCAGAGTTTCCTCTGGCCTCG 3' this study *28S-WPM-R3 (reverse) 5' GATCCGTCCCGGCGATTCG 3' this study Actin 5C ActMBF2 (forward) 5' GCTCCYTTRAATCCHAAAG 3' Bodner 2009 ActinR1B (reverse) 5' TTNGADATCCACATTTGTTGGAA 3' Vink et al. 2008 16S-ND1 N1-J-12261 (forward) 5' TCRTAAGAAATTATTTGAGC 3' Hedin 1997 LR-N-13398 (reverse) 5' CGCCTGTTTAACAAAAACAT 3' Simon et al. 1994 16SND1-WPM-F1 (forward) 5' GCRTCTCTRAAAGGTTG 3' this study 16SND1-WPM-F2 (forward) 5' GCRTCTCTRAAGGGTTG 3' this study 16SND1-WPM-R2 (reverse) 5' GTGCTAAGGTAGCATAATA 3' this study 16SND1-WPM-R3 (reverse) 5' CGCCTGTTTAACAAAAAC 3' this study COI C1-J-1718 (forward) 5' GGAGGATTTGGAAATTGATTAGTTCC 3' Simon et al. 1994 C1-N-2776 (reverse) 5' GGATAATCAGAATATCGTCGAGG 3' Hedin & Maddison 2001a 39! Table 2.2. Summary of annealing temperatures for gene amplification using different pairs of primers. Gene region Primer pair DNA Polymerase Annealing temperature 28S rDNA ZX1/28SC Taq Paq5000 48-50°C 54-58°C 28SO/28SC Taq Paq5000 48°C 52-55°C 28S-WPM-F1/28S-WPM-R1 Paq5000 62°C 28S-WPM-F3 /28S-WPM-R2 Paq5000 64°C Actin5C ActMBF2/ ActinR1B Paq5000 55-57°C 16S-ND1 N1-J-12261/ LR-N-13398 Taq Paq5000 44°C 44-46°C 16SND1-WPM-F1/16SND1-WPM-R3 Paq5000 46-55°C 16SND1-WPM-F2/16SND1-WPM-R2 Paq5000 52-55°C COI C1-J-1718/ C1-N-2776 Taq Paq5000 42-44°C 45-50°C 40! Table 2.3. Substitution models selected by ModelTest for each individual gene region and partition. Gene region or partition Sites included (bp) Number of sequences included Model selected by ModelTest Model in Bayesian and BEAST analyses 28S 1443 293 GTR+I+G - Actin 5C 717 279 TVM+I+G - 16S-ND1 1161 282 TVM+I+G - COI 990 190 GTR+I+G - Combined dataset 4311 292 GTR+I+G - Partitions in the combined dataset 28S 1443 292 GTR+I+G GTR+I+G Actin 5C 1 st codon position 239 292 GTR+I+G GTR+I+G Actin 5C 2 nd codon position 239 292 HKY+G HKY+G Actin 5C 3 rd codon position 239 292 GTR+I+G GTR+I+G 16S 783 292 K81uf+I+G GTR+I+G ND1+COI 1 st codon position 456 292 GTR+I+G GTR+I+G ND1+COI 2 nd codon position 456 292 GTR+I+G GTR+I+G ND1+COI 3 rd codon position 456 292 TVM+G GTR+G 41! Table 2.4. Summary of calibration points used in divergence time analyses. Calibration point Analysis One Analysis Two Salticidae max. 100 Mya; min. 44 Mya max. 100 Mya; min. 44 Mya Salticoida max. 49 Mya; min. 16 Mya max. 100 Mya; min. 16 Mya Lyssomaninae/Spartaeinae max. 100 Mya; min. 22 Mya max. 100 Mya; min. 22 Mya Euophryinae/Sister-group max. 49 Mya; min. 16 Mya max. 100 Mya; min. 16 Mya 42! Table 2.5. Estimates of divergence times (in millions of years) for nodes in Fig. 2.7. Divergence times are calculated using a Bayesian relaxed molecular clock (implemented in BEAST) and independently using Penalized Likelihood and rate smoothing (PLRS, implemented in r8s), with two sets of constraints (Analysis One and Analysis Two; see Table 2.3) respectively. Analysis One Analysis Two BEAST BEAST # Node Name Median 95% HPD r8s Median 95% HPD r8s 1 Euophryinae 30.19 [37.84, 28.93] 39.34 33.84 [55.52, 23.10] 66.53 2 29.26 [36.61, 23.19] 38.25 32.75 [53.75, 22.31] 64.65 3 Anasaitis- Corythalia Clade 22.27 [28.35, 16.82] 30.81 24.99 [41.39, 17.00] 51.90 4 Caribbean radiation one 19.06 [24.83, 14.20] 27.31 21.45 [35.59, 14.19] 45.85 5 Caribbean radiation two 10.35 [13.87, 7.37] 13.85 11.80 [19.63, 7.21] 23.33 6 Parabathippus- Parvattus Clade 20.54 [26.96, 15.16] 28.74 23.13 [38.42, 14.87] 48.57 7 Euophrys Clade 15.69 [20.73, 11.42] 21.46 17.39 [28.68, 11.24] 36.23 8 Caribbean radiation three 19.74 [25.49, 14.79] 27.24 22.34 [36.83, 14.77] 45.97 9 Caribbean radiation four 14.25 [19.85, 9.17] 20.53 16.44 [27.90, 9.15] 34.64 10 Caribbean radiation five 23.52 [29.70, 18.10] 32.84 26.46 [43.24, 17.75] 55.46 11 25.96 [32.94, 20.44] 35.53 29.53 [47.93, 21.22] 60.02 12 26.98 [33.96, 21.10] 36.53 30.36 [49.51, 21.68] 61.72 13 26.26 [33.14, 20.73] 35.95 29.14 [47.94, 20.02] 60.72 14 25.06 [31.60, 19.69] 34.36 27.93 [46.08, 19.15] 58.04 15 Papua New Guinea radiation one (Thorelliola) 16.96 [21.79, 12.68] 23.24 18.99 [31.33, 12.62] 39.34 16 African radiation 22.75 [29.50, 17.03] 31.31 25.25 [41.68, 16.51] 52.82 17 Papua New Guinea radiation two (Cytaea-Euryattus Clade) 16.27 [21.30, 11.79] 22.11 18.31 [30.32, 11.48] 37.25 18 Papua New Guinea radiation three 27.74 [35.09, 21.52] 35.83 31.68 [52.18, 21.65] 60.57 19 Diolenius Clade 19.67 [25.47, 14.73] 25.28 22.30 [36.78, 14.59] 42.75 20 Omoedus Clade 19.57 [25.11, 14.98] 24.96 22.04 [36.08, 14.58] 42.21 43! Figure 2.1. Summary of phylogenetic analyses on combined matrix of all genes (28S, Actin 5C, 16S-ND1 and COI). Tree shown is the best tree from the ML analysis with all non-euophryine taxa trimmed off. Approximate position of euophryine genera is indicated on the right side of the tree; continental distribution of taxa is indicated in colored blocks in front of the taxon names. Numbers for major clades is consistent with those of Table 2.5. Anasaitis adorabilis Anasaitis brunnea Anasaitis hebetata 'Corythalia' cf. canalis Anasaitis canosa [USA] Anasaitis canosa [Panama] 'Corythalia' banksi 'Stoidis' placida 'Corythalia' gloriae Anasaitis laxa 'Corythalia' elegantissima 'Corythalia' locuples Anasaitis sp. [Peblique] Corythalia porphyra Corythalia cf. albicincta Corythalia sulfurea Corythalia broccai Dinattus minor Corythalia peblique Corythalia bromelicola Corythalia coronai 'Wallaba' decora Corythalia bicincta Corythalia sp. [PuertoLopez] Corythalia cf. latipes Corythalia cf. valida Corythalia electa Corythalia sp. [Baeza] Corythalia sp. [JatunSacha] Parvattus zhui Parabathippus kiabau Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Euophrys frontalis Euophrys cf. proszynskii Euophrys monodnock Chalcoscirtus alpicola Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Cobanus sp. [Ecuador] Cobanus cf. electus Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Cobanus cf. cambridgei Cobanus mandibularis Cobanus sp. [CostaRica] Cobanus unicolor Cobanus extensus Cobanus sp. [Panama] Pensacola signata Mexigonus morosus Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Petemathis minuta Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis 'Cobanus' cambridgei 'Pensacola' darlingtoni 'Pensacola' maxillosa 'Siloca' electa Naphrys pulex Naphrys xerophila Corticattus guajataca Corticattus latus Popcornella furcata Popcornella spiniformis Popcornella nigromaculata Popcornella yunque Compsodecta haytiensis Compsodecta peckhami Bythocrotus cf. crypticus Bythocrotus crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo cf. Chapoda sp. [Arizona] 'Sidusa' recondita Chapoda gitae Chapoda angusta Chapoda sp. [Fortuna] Chapoda cf. fortuna [CostaRica] Chapoda fortuna [Panama] Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [Cutucú] Maeota dichrura Maeota sp. [Napo] Maeota flava Maeota simoni Maeota sp. [MoronaSantiago] 'Pensacola' tuberculotibiata Marma nigritarsis Amphidraus complexus Nebridia cf. semicana Tylogonus cf. viridimicans Tylogonus yanayacu Tylogonus cf. auricapillus Tylogonus parvus Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Tariona cf. bruneti 'Euophrys' a-notata 'Euophrys' cf. patagonica cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Asaphobelis physonychus Siloca cf. campestrata Siloca cf. sanguiniceps Neonella vinnula Ecuadattus napoensis Ecuadattus pichincha Belliena sp. [Yanayacu] Belliena sp. [JatunSacha] Belliena ecuadorica (F) Belliena ecuadorica (M) Ilargus galianoae Ilargus pilleolus Ilargus coccineus Ilargus serratus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Servaea vestita Jotus auripes Prostheclina sp. cf. Jotus sp. [Queensland] Lycidas cf. vittatus Maileus cf. fuscus cf. Saitis sp. [WesternAustralia] Saitis barbipes Maratus cf. amabilis Maratus sp. [SouthAustralia] Lycidas cf. griseus Hypoblemum sp. [NewSouthWales] Hypoblemum cf. albovittatum Lycidas cf. karschi Emathis gombak Lepidemathis haemorrhoidalis Chalcotropis cf. caeruleus Chalcotropis luceroi Colyttus robustus Colyttus bilineatus Donoessus striatus Euophryine sp. [GentingHighlands] Lagnus edwardsi Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola ensifera Thorelliola tamasi Thorelliola tualapa Thorelliola cf. mahunkai Thorelliola mahunkai Foliabitus longzhou Foliabitus sp. [Malaysia] Laufeia concava Junxattus daiqini Laufeia eximia Orcevia keyserlingi Thiania cf. viscaensis Thianitara spectrum Thiania latibola Thiania cf. suboppressa Thiania bhamoensis Thiania tenuis Chinophrys pengi Thyenula laxa Thyenula nelshoogte 'Saitis' leighi Thyenula sp. [SouthAfrica] Thyenula wesolowskae 'Saitis' cf. mundus Thyenula cf. aurantiaca Cytaea oreophila Cytaea mitellata Cytaea cf. rai Cytaea sp. [Wewak] Cytaea cf. sinuata Cytaea nimbata Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Euryattus cf. venustus Euryattus bleekeri Euryattus cf. porcellus Euryattus sp. [Queensland] Phasmolia elegans Zabkattus furcatus Zabkattus richardsi Zabkattus brevis Zabkattus trapeziformis Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Bathippus macrognathus Bathippus korei Bathippus madang Bathippus gahavisuka Bathippus cf. directus [Suyang] Bathippus directus [Tualapa] Pristobaeus cf. jocosus Palpelius beccarii Palpelius cf. discedens Xenocytaea agnarssoni Xenocytaea albomaculata Xenocytaea proszynskii Chalcolemia nakanai Sobasina wanlessi Efate albobicinctus Paraharmochirus tualapaensis Chalcolecta prensitans Ohilimia scutellata Diolenius cf. decorus Diolenius varicus Omoedus cf. piceus Omoedus ephippigera Omoedus cf. metallescens Omoedus orbiculatus Omoedus brevis 'Margaromma' cf. semirasum Omoedus swiftorum Omoedus cf. durvillei Omoedus cf. ponapensis 'Margaromma' cf. torquatum Omoedus cf. danae Omoedus sp. [Adalbert] Omoedus darleyorum Omoedus papuanus Omoedus tortuosus Omoedus sp. [Gahavisuka] Omoedus meyeri Omoedus omundseni Euophryinae Anasaitis Corythalia Dinattus Parvattus Parabathippus Euophrys Chalcoscirtus Talavera Pseudeuophrys Cobanus Sidusa Pensacola Mexigonus Petemathis Truncattus Antillattus Naphrys Corticattus Popcornella Compsodecta Bythocrotus Agobardus Chapoda Maeota Marma Amphidraus Nebridia Tylogonus Soesilarishius Tariona South American ‘Euophrys’ Coryphasia Asaphobelis Siloca Neonella Ecuadattus Belliena Ilargus Servaea Jotus Maileus Prostheclina Lycidas Saitis Maratus Hypoblemum Emathis Lepidemathis Chalcotropis Colyttus Donoessus Lagnus Thorelliola Foliabitus Laufeia Junxattus Orcevia Thiania Thianitara Chinophrys Thyenula Cytaea Euryattus Phasmolia Zabkattus Viribestus Variratina Bulolia Leptathamas Coccorchestes Canama Bathippus Pristobaeus Palpelius Xenocytaea Omoedus Chalcolemia Sobasina Efate Paraharmochirus Chalcolecta Ohilimia Diolenius Neotropics Nearctic Australasia Eurasia Africa Continental distribution Support by analysis All Genes Individual Gene: if the clade present in ML tree of individual gene 28 S 16 S- ND 1 Ac tin 5C CO I ML bo ot str ap po ste rio r p ro ba bil ity if t he cl ad e p res en t in st ric t co ns en su s o f M P t ree s va lue va lue 1. 0 0. 9- 0. 99 0. 8- 0. 89 0. 7- 0. 79 < 0. 7 pr es en t pr es en t pr es en t no t pr es en t no t 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 19 18 20 44! Figure 2.2. The best tree from ML analysis on combined matrix of all genes (28S, Actin 5C, 16S-ND1 and COI) with lnL = -181734.801. Xysticus sp. Lyssomanes viridis Galianora bryicola Thrandina parocula Holcolaetis sp. Portia labiata Tomocyrba sp. cf. Acragus Hurius vulpinus Cotinusa sp. Jollas sp. Arasia mollicoma Neon nelli Attidops youngi Ghelna canadensis Mantisatta longicauda Pachyballus sp. [S.Afr.] 'Bathippus' pahang Nannenus lyriger Bristowia sp. [Gabon] Cheliceroides sp. [China] Chinattus parvulus Heliophanus cupreus Yllenus arenarius Aelurillus cf. ater Freya decorata Philaeus chrysops Salticus scenicus Plexippus paykulli Habronattus decorus Havaika sp. Anasaitis adorabilis Anasaitis brunnea Anasaitis hebetata 'Corythalia' cf. canalis Anasaitis canosa [USA] Anasaitis canosa [Panama] 'Corythalia' banksi 'Stoidis' placida 'Corythalia' gloriae Anasaitis laxa 'Corythalia' elegantissima 'Corythalia' locuples Anasaitis sp. [Peblique] Corythalia porphyra Corythalia cf. albicincta Corythalia sulfurea Corythalia broccai Dinattus minor Corythalia peblique Corythalia bromelicola Corythalia coronai 'Wallaba' decora Corythalia bicincta Corythalia sp. [PuertoLopez] Corythalia cf. latipes Corythalia cf. valida Corythalia electa Corythalia sp. [Baeza] Corythalia sp. [JatunSacha] Parvattus zhui Parabathippus kiabau Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Euophrys frontalis Euophrys cf. proszynskii Euophrys monodnock Chalcoscirtus alpicola Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Cobanus sp. [Ecuador] Cobanus cf. electus Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Cobanus cf. cambridgei Cobanus mandibularis Cobanus sp. [CostaRica] Cobanus unicolor Cobanus extensus Cobanus sp. [Panama] Pensacola signata Mexigonus morosus Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Petemathis minuta Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis 'Cobanus' cambridgei 'Pensacola' darlingtoni 'Pensacola' maxillosa 'Siloca' electa Naphrys pulex Naphrys xerophila Corticattus guajataca Corticattus latus Popcornella furcata Popcornella spiniformis Popcornella nigromaculata Popcornella yunque Compsodecta haytiensis Compsodecta peckhami Bythocrotus cf. crypticus Bythocrotus crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo cf. Chapoda sp. [Arizona] 'Sidusa' recondita Chapoda gitae Chapoda angusta Chapoda sp. [Fortuna] Chapoda cf. fortuna [CostaRica] Chapoda fortuna [Panama] Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [Cutucú] Maeota dichrura Maeota sp. [Napo] Maeota flava Maeota simoni Maeota sp. [MoronaSantiago] 'Pensacola' tuberculotibiata Marma nigritarsis Amphidraus complexus Nebridia cf. semicana Tylogonus cf. viridimicans Tylogonus yanayacu Tylogonus cf. auricapillus Tylogonus parvus Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Tariona cf. bruneti 'Euophrys' a-notata 'Euophrys' cf. patagonica cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Asaphobelis physonychus Siloca cf. campestrata Siloca cf. sanguiniceps Neonella vinnula Ecuadattus napoensis Ecuadattus pichincha Belliena sp. [Yanayacu] Belliena sp. [JatunSacha] Belliena ecuadorica (F) Belliena ecuadorica (M) Ilargus galianoae Ilargus pilleolus Ilargus coccineus Ilargus serratus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Servaea vestita Jotus auripes Prostheclina sp. cf. Jotus sp. [Queensland] Lycidas cf. vittatus Maileus cf. fuscus cf. Saitis sp. [WesternAustralia] Saitis barbipes Maratus cf. amabilis Maratus sp. [SouthAustralia] Lycidas cf. griseus Hypoblemum sp. [NewSouthWales] Hypoblemum cf. albovittatum Lycidas cf. karschi Emathis gombak Lepidemathis haemorrhoidalis Chalcotropis cf. caeruleus Chalcotropis luceroi Colyttus robustus Colyttus bilineatus Donoessus striatus Euophryine sp. [GentingHighlands] Lagnus edwardsi Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola ensifera Thorelliola tamasi Thorelliola tualapa Thorelliola cf. mahunkai Thorelliola mahunkai Foliabitus longzhou Foliabitus sp. [Malaysia] Laufeia concava Junxattus daiqini Laufeia eximia Orcevia keyserlingi Thiania cf. viscaensis Thianitara spectrum Thiania latibola Thiania cf. suboppressa Thiania bhamoensis Thiania tenuis Chinophrys pengi Thyenula laxa Thyenula nelshoogte 'Saitis' leighi Thyenula sp. [SouthAfrica] Thyenula wesolowskae 'Saitis' cf. mundus Thyenula cf. aurantiaca Cytaea oreophila Cytaea mitellata Cytaea cf. rai Cytaea sp. [Wewak] Cytaea cf. sinuata Cytaea nimbata Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Euryattus cf. venustus Euryattus bleekeri Euryattus cf. porcellus Euryattus sp. [Queensland] Phasmolia elegans Zabkattus furcatus Zabkattus richardsi Zabkattus brevis Zabkattus trapeziformis Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Bathippus macrognathus Bathippus korei Bathippus madang Bathippus gahavisuka Bathippus cf. directus [Suyang] Bathippus directus [Tualapa] Pristobaeus cf. jocosus Palpelius beccarii Palpelius cf. discedens Xenocytaea agnarssoni Xenocytaea albomaculata Xenocytaea proszynskii Chalcolemia nakanai Sobasina wanlessi Efate albobicinctus Paraharmochirus tualapaensis Chalcolecta prensitans Ohilimia scutellata Diolenius cf. decorus Diolenius varicus Omoedus cf. piceus Omoedus ephippigera Omoedus cf. metallescens Omoedus orbiculatus Omoedus brevis 'Margaromma' cf. semirasum Omoedus swiftorum Omoedus cf. durvillei Omoedus cf. ponapensis 'Margaromma' cf. torquatum Omoedus cf. danae Omoedus sp. [Adalbert] Omoedus darleyorum Omoedus papuanus Omoedus tortuosus Omoedus sp. [Gahavisuka] Omoedus meyeri Omoedus omundseni 0.1 Euophryinae 45! Figure 2.3. Summary of ML and MP analyses on 28S. Tree shown is the best tree from the ML analysis with lnL = -51870.560. Xysticus sp. Lyssomanes viridis Galianora bryicola Portia labiata Holcolaetis sp. Thrandina parocula Tomocyrba sp. Cotinusa sp. Jollas sp. cf. Acragus Hurius vulpinus Arasia mollicoma Neon nelli Attidops youngi Ghelna canadensis Mantisatta longicauda Pachyballus sp. [S.Afr.] Heliophanus cupreus 'Bathippus' pahang Nannenus lyriger Bristowia sp. [Gabon] Cheliceroides sp. [China] Chinattus parvulus Yllenus arenarius Aelurillus cf. ater Freya decorata Salticus scenicus Plexippus paykulli Habronattus decorus Havaika sp. Philaeus chrysops Servaea vestita Jotus auripes Prostheclina sp. cf. Jotus sp. [Queensland] Lycidas cf. vittatus Maileus cf. fuscus Lycidas cf. griseus Maratus cf. amabilis Maratus sp. [SouthAustralia] cf. Saitis sp. [WesternAustralia] Saitis barbipes Hypoblemum sp. [NewSouthWales] Hypoblemum cf. albovittatum Lycidas cf. karschi Emathis gombak Chalcotropis cf. caeruleus Chalcotropis luceroi Euophryine sp. [GentingHighlands] Lagnus edwardsi Athamas nitidus Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola ensifera Thorelliola tamasi Thorelliola tualapa Thorelliola cf. mahunkai Thorelliola mahunkai Foliabitus longzhou Foliabitus sp. [Malaysia] Colyttus robustus Colyttus bilineatus Donoessus striatus Lepidemathis haemorrhoidalis Laufeia concava Orcevia keyserlingi Laufeia eximia Junxattus daiqini Thiania cf. viscaensis Thianitara spectrum Thiania latibola Thiania cf. suboppressa Thiania bhamoensis Thiania tenuis Marma nigritarsis Amphidraus complexus Nebridia cf. semicana cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Asaphobelis physonychus Siloca cf. campestrata Siloca cf. sanguiniceps Neonella vinnula Tariona cf. bruneti 'Euophrys' a-notata 'Euophrys' cf. patagonica Belliena sp. [Yanayacu] Belliena sp. [JatunSacha] Belliena ecuadorica (F) Belliena ecuadorica (M) Ilargus pilleolus Ilargus galianoae Ilargus coccineus Ilargus serratus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Ecuadattus napoensis Ecuadattus pichincha Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Thyenula laxa Thyenula nelshoogte Cytaea oreophila Cytaea cf. sinuata Cytaea nimbata Cytaea cf. rai Cytaea mitellata Cytaea sp. [Wewak] Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Euryattus bleekeri Euryattus cf. venustus Euryattus cf. porcellus Euryattus sp. [Queensland] Cobanus cf. cambridgei Cobanus mandibularis Cobanus extensus Cobanus sp. [Panama] Cobanus unicolor Cobanus sp. [CostaRica] Cobanus sp. [Ecuador] Cobanus cf. electus Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Phasmolia elegans Zabkattus brevis Zabkattus trapeziformis Zabkattus furcatus Zabkattus richardsi Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Bathippus macrognathus Bathippus cf. directus [Suyang] Bathippus directus [Tualapa] Bathippus gahavisuka Bathippus korei Bathippus madang Chalcolemia nakanai Sobasina wanlessi Chalcolecta prensitans Efate albobicinctus Paraharmochirus tualapaensis Ohilimia scutellata Diolenius cf. decorus Diolenius varicus Pristobaeus cf. jocosus Palpelius beccarii Palpelius cf. discedens Xenocytaea albomaculata Xenocytaea agnarssoni Xenocytaea proszynskii Omoedus papuanus Omoedus sp. [Gahavisuka] Omoedus meyeri Omoedus omundseni Omoedus tortuosus Omoedus sp. [Adalbert] Omoedus darleyorum Omoedus cf. durvillei Omoedus brevis Omoedus cf. danae 'Margaromma' cf. torquatum Omoedus cf. ponapensis 'Margaromma' cf. semirasum Omoedus swiftorum Omoedus ephippigera Omoedus cf. piceus Omoedus cf. metallescens Omoedus orbiculatus Corticattus latus Naphrys pulex Naphrys xerophila cf. Maeota sp. [Panama] cf. Chapoda sp. [Arizona] 'Sidusa' recondita Chapoda gitae Chapoda angusta Chapoda sp. [Fortuna] Chapoda cf. fortuna [CostaRica] Chapoda fortuna [Panama] Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [Cutucú] Maeota dichrura Maeota sp. [Napo] Maeota flava Maeota simoni Maeota sp. [MoronaSantiago] 'Pensacola' tuberculotibiata Antillattus gracilis 'Cobanus' cambridgei 'Siloca' electa 'Pensacola' maxillosa 'Pensacola' darlingtoni Truncattus dominicanus Truncattus cachotensis Truncattus flavus Antillattus cf. applanatus Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Petemathis tetuani Petemathis minuta Corticattus guajataca Popcornella furcata Popcornella spiniformis Popcornella nigromaculata Popcornella yunque Compsodecta haytiensis Compsodecta peckhami Bythocrotus cf. crypticus Bythocrotus crypticus Agobardus bahoruco Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus oviedo Agobardus gramineus Agobardus phylladiphilus Pensacola signata Mexigonus morosus Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Chinophrys pengi Tylogonus cf. viridimicans Tylogonus yanayacu Tylogonus cf. auricapillus Tylogonus parvus Thyenula wesolowskae 'Saitis' cf. mundus Thyenula cf. aurantiaca 'Saitis' leighi Thyenula sp. [SouthAfrica] Parvattus zhui Parabathippus kiabau Parabathippus cf. macilentus Parabathippus magnus Parabathippus cuspidatus Parabathippus shelfordi Chalcoscirtus infimus Chalcoscirtus diminutus Chalcoscirtus alpicola Pseudeuophrys erratica Talavera minuta Euophrys frontalis Euophrys cf. proszynskii Euophrys monodnock Anasaitis adorabilis 'Corythalia' cf. canalis Anasaitis brunnea Anasaitis hebetata Anasaitis canosa [USA] Anasaitis canosa [Panama] 'Corythalia' gloriae 'Stoidis' placida 'Corythalia' banksi 'Corythalia' locuples Anasaitis sp. [Peblique] Anasaitis laxa 'Corythalia' elegantissima Corythalia porphyra Corythalia cf. albicincta Corythalia sulfurea Corythalia broccai Dinattus minor Corythalia peblique Corythalia bromelicola Corythalia coronai 'Wallaba' decora Corythalia cf. latipes Corythalia bicincta Corythalia sp. [PuertoLopez] Corythalia cf. valida Corythalia electa Corythalia sp. [Baeza] Corythalia sp. [JatunSacha] Euophryinae * * * * * * * * * * * * * * * * * * * * * * * * * ** * * * * * * * * * * * * * * * * * * * * * ** * ** * * ** * * * * * ** * *** * * * * * * * * * * * * * ** * * ** * * * * * * * * * * ** * * * ** ** * * * * ** * * * * * * * * * * * * * * * * * * * * ** * * * * * * * * * * * * * * * * * * * ** * * * * * * * * * * ** * * * * 0.1 * ML bootstrap value for clade >=0.7 clade also present in strict consensus of MP trees 46! Figure 2.4. Summary of ML and MP analyses on Actin 5C. Tree shown is the best tree from the ML analysis with lnL = -13886.791. Xysticus sp. Holcolaetis sp. Lyssomanes viridis Galianora bryicola Thrandina parocula Ghelna canadensis Philaeus chrysops Salticus scenicus Heliophanus cupreus Yllenus arenarius Arasia mollicoma Bythocrotus cf. crypticus Cotinusa sp. Pachyballus sp. [S.Afr.] 'Bathippus' pahang Aelurillus cf. ater Nannenus lyriger Freya decorata Bristowia sp. [Gabon] Cheliceroides sp. [China] Habronattus decorus Plexippus paykulli Thyenula sp. [SouthAfrica] Thyenula laxa Thyenula wesolowskae Thyenula nelshoogte 'Saitis' leighi 'Saitis' cf. mundus Thyenula cf. aurantiaca Phasmolia elegans Thiania bhamoensis Thiania cf. suboppressa Thiania latibola Chinophrys pengi Xenocytaea agnarssoni Xenocytaea albomaculata Xenocytaea proszynskii Ilargus serratus Ilargus coccineus Neonella vinnula Chalcoscirtus alpicola Chalcoscirtus infimus Chalcoscirtus diminutus Talavera minuta Ilargus pilleolus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Ilargus galianoae Tomocyrba sp. Cobanus cf. electus Cobanus cf. cambridgei Cobanus sp. [CostaRica] Cobanus sp. [Panama] Cobanus extensus Cobanus unicolor Sidusa sp.2 [FrenchGuiana] Cobanus mandibularis Sidusa sp.1 [FrenchGuiana] Amphidraus complexus Tylogonus cf. auricapillus Tylogonus parvus Tylogonus cf. viridimicans Tylogonus yanayacu Ecuadattus napoensis Ecuadattus pichincha Popcornella spiniformis Naphrys pulex Naphrys xerophila Corticattus guajataca Lepidemathis haemorrhoidalis Corticattus latus Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Petemathis minuta Truncattus dominicanus Truncattus cachotensis Truncattus flavus Antillattus cf. applanatus Antillattus gracilis 'Cobanus' cambridgei 'Pensacola' darlingtoni 'Pensacola' maxillosa Chalcotropis luceroi Lagnus edwardsi Marma nigritarsis Dinattus minor Corythalia broccai Corythalia peblique Corythalia coronai Corythalia bromelicola 'Wallaba' decora Anasaitis adorabilis Anasaitis brunnea Anasaitis hebetata 'Corythalia' cf. canalis Anasaitis canosa [USA] Anasaitis canosa [Panama] 'Corythalia' banksi 'Stoidis' placida 'Corythalia' gloriae Anasaitis laxa 'Corythalia' elegantissima 'Corythalia' locuples Anasaitis sp. [Peblique] Corythalia sulfurea Corythalia cf. albicincta Corythalia porphyra Corythalia sp. [PuertoLopez] Corythalia cf. latipes Corythalia electa Corythalia sp. [JatunSacha] Corythalia cf. valida Corythalia sp. [Baeza] Belliena sp. [Yanayacu] Servaea vestita cf. Coryphasia sp. [Brazil] Asaphobelis physonychus Coryphasia fasciiventris Siloca cf. campestrata Siloca cf. sanguiniceps Tariona cf. bruneti 'Euophrys' a-notata 'Euophrys' cf. patagonica Parvattus zhui Parabathippus cf. macilentus Parabathippus cuspidatus Parabathippus magnus Parabathippus kiabau Parabathippus shelfordi Compsodecta haytiensis Compsodecta peckhami Pensacola signata Mexigonus morosus Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Agobardus oviedo Agobardus gramineus Agobardus cordiformis Agobardus phylladiphilus Agobardus bahoruco Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Popcornella nigromaculata Popcornella yunque cf. Maeota sp. [Panama] Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami Chapoda sp. [Fortuna] Chapoda fortuna [Panama] Euophrys monodnock Euryattus sp. [Queensland] Chapoda cf. fortuna [CostaRica] Thiania tenuis Euophrys cf. proszynskii Euophrys frontalis cf. Chapoda sp. [Arizona] 'Sidusa' recondita Chapoda gitae Chapoda angusta Maeota dorsalis Maeota flava Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [Cutucú] Maeota dichrura Maeota sp. [Napo] Maeota sp. [MoronaSantiago] Maeota simoni 'Pensacola' tuberculotibiata Jotus auripes Soesilarishius cf. amrishi Chinattus parvulus Soesilarishius micaceus Soesilarishius ruizi cf. Jotus sp. [Queensland] Belliena sp. [JatunSacha] Belliena ecuadorica (F) Belliena ecuadorica (M) cf. Saitis sp. [WesternAustralia] Prostheclina sp. Maratus sp. [SouthAustralia] Maratus cf. amabilis Saitis barbipes Hypoblemum cf. albovittatum Lycidas cf. karschi Hypoblemum sp. [NewSouthWales] Lycidas cf. griseus Chalcotropis cf. caeruleus Maileus cf. fuscus Junxattus daiqini Laufeia concava Laufeia eximia Orcevia keyserlingi Emathis gombak Viribestus suyanensis Euophryine sp. [GentingHighlands] Colyttus bilineatus Colyttus robustus Donoessus striatus Thiania cf. viscaensis Thianitara spectrum Foliabitus longzhou Foliabitus sp. [Malaysia] Cytaea oreophila Cytaea mitellata Cytaea cf. rai Cytaea sp. [Wewak] Cytaea cf. sinuata Cytaea nimbata Euryattus sp.2 [Gahavisuka] Euryattus sp.1 [Gahavisuka] Euryattus cf. venustus Euryattus bleekeri Euryattus cf. porcellus Thorelliola crebra Thorelliola aliena Thorelliola Joannae Thorelliola ensifera Thorelliola tamasi Thorelliola tualapa Thorelliola cf. mahunkai Thorelliola mahunkai Canama cf. forceps Canama hinnulea Canama extranea Canama fimoi Canama triramosa Bathippus macrognathus Bathippus korei Bathippus madang Bathippus gahavisuka Bathippus cf. directus [Suyang] Bathippus directus [Tualapa] Athamas cf. whitmeei Athamas nitidus Athamas sp. Zabkattus brevis Zabkattus trapeziformis Zabkattus furcatus Zabkattus richardsi Nebridia cf. semicana Variratina minuta Leptathamas paradoxus Bulolia excentrica Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Pristobaeus cf. jocosus Palpelius beccarii Palpelius cf. discedens Sobasina wanlessi Chalcolemia nakanai Efate albobicinctus Paraharmochirus tualapaensis Ohilimia scutellata Chalcolecta prensitans Diolenius cf. decorus Diolenius varicus Omoedus sp. [Adalbert] Omoedus darleyorum Omoedus cf. piceus Omoedus ephippigera Omoedus cf. metallescens Omoedus orbiculatus Omoedus meyeri Omoedus papuanus Omoedus sp. [Gahavisuka] Omoedus omundseni Omoedus brevis 'Margaromma' cf. semirasum Omoedus swiftorum Omoedus cf. durvillei Omoedus cf. danae 'Margaromma' cf. torquatum Omoedus cf. ponapensis * * * * * * * * * ** * * * * * * * **** * * * * * * * * * * * ** * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * 0.02 * ML bootstrap value for clade >=0.7 clade also present in strict consensus of MP trees 47! Figure 2.5. Summary of ML and MP analyses on 16S-ND1. Tree shown is the best tree from the ML analysis with lnL = -69623.304. Xysticus sp. Galianora bryicola Lyssomanes viridis Thrandina parocula Portia labiata 'Bathippus' pahang Nannenus lyriger Freya decorata Philaeus chrysops Aelurillus cf. ater Neon nelli Mantisatta longicauda Jollas sp. Plexippus paykulli Salticus scenicus Havaika sp. Cheliceroides sp. [China] Habronattus decorus Naphrys pulex Naphrys xerophila Corticattus guajataca Corticattus latus Phasmolia elegans Chalcoscirtus alpicola Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Euophrys cf. proszynskii Euophrys frontalis Euophrys monodnock Attidops youngi Amphidraus complexus Nebridia cf. semicana Cytaea oreophila Cytaea mitellata Cytaea cf. rai Cytaea sp. [Wewak] Cytaea cf. sinuata Cytaea nimbata Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Euryattus cf. venustus Euryattus bleekeri Euryattus cf. porcellus Euryattus sp. [Queensland] Pensacola signata Emathis gombak Lepidemathis haemorrhoidalis Mexigonus morosus Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Compsodecta haytiensis Compsodecta peckhami Chalcolemia nakanai Sobasina wanlessi Efate albobicinctus Paraharmochirus tualapaensis Ohilimia scutellata Chalcolecta prensitans Diolenius cf. decorus Diolenius varicus Zabkattus furcatus Zabkattus trapeziformis Zabkattus brevis Zabkattus richardsi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Parabathippus kiabau Parabathippus shelfordi Omoedus cf. durvillei Omoedus cf. piceus Omoedus ephippigera Omoedus cf. metallescens Omoedus orbiculatus Omoedus cf. ponapensis Omoedus darleyorum Omoedus cf. danae 'Margaromma' cf. torquatum Omoedus sp. [Adalbert] Omoedus sp. [Gahavisuka] Omoedus meyeri Omoedus omundseni Omoedus papuanus Omoedus tortuosus Omoedus brevis 'Margaromma' cf. semirasum Omoedus swiftorum Coryphasia fasciiventris Asaphobelis physonychus Siloca cf. campestrata Siloca cf. sanguiniceps Euophryine sp. [GentingHighlands] Anasaitis canosa [USA] Anasaitis canosa [Panama] Anasaitis adorabilis 'Corythalia' cf. canalis Anasaitis brunnea Anasaitis hebetata 'Corythalia' banksi 'Stoidis' placida 'Corythalia' gloriae Corythalia broccai Corythalia peblique Corythalia bromelicola Dinattus minor Corythalia coronai 'Wallaba' decora Anasaitis laxa 'Corythalia' locuples Anasaitis sp. [Peblique] Corythalia porphyra Corythalia cf. albicincta Corythalia sulfurea Corythalia bicincta Corythalia sp. [PuertoLopez] Corythalia cf. latipes Corythalia sp. [Baeza] Corythalia sp. [JatunSacha] Corythalia cf. valida Corythalia electa Ecuadattus napoensis Ecuadattus pichincha 'Euophrys' a-notata 'Euophrys' cf. patagonica Tariona cf. bruneti cf. Coryphasia sp. [Brazil] Belliena sp. [JatunSacha] Belliena ecuadorica (F) Belliena ecuadorica (M) Ilargus galianoae Ilargus serratus Ilargus pilleolus Ilargus moronatigus Ilargus coccineus Ilargus foliosus Ilargus macrocornis Marma nigritarsis cf. Acragus Hurius vulpinus Belliena sp. [Yanayacu] Arasia mollicoma Thyenula sp. [SouthAfrica] 'Saitis' cf. mundus Thyenula cf. aurantiaca Thyenula wesolowskae 'Saitis' leighi Thyenula laxa Thyenula nelshoogte Chinophrys pengi Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. ildikoae Coccorchestes cf. aiyura Coccorchestes cf. inermis Cobanus cf. electus Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Cobanus sp. [CostaRica] Cobanus cf. cambridgei Cobanus mandibularis Cobanus unicolor Cobanus sp. [Panama] Cobanus extensus Cobanus sp. [Ecuador] Canama cf. forceps Canama hinnulea Bathippus macrognathus Canama fimoi Canama extranea Canama triramosa Bathippus korei Bathippus madang Bathippus gahavisuka Bathippus cf. directus [Suyang] Bathippus directus [Tualapa] Colyttus bilineatus Colyttus robustus Donoessus striatus Maileus cf. fuscus Saitis barbipes Jotus auripes Prostheclina sp. cf. Jotus sp. [Queensland] Lycidas cf. vittatus cf. Saitis sp. [WesternAustralia] Maratus cf. amabilis Maratus sp. [SouthAustralia] Hypoblemum cf. albovittatum Lycidas cf. griseus Hypoblemum sp. [NewSouthWales] Lycidas cf. karschi Laufeia concava Junxattus daiqini Laufeia eximia Orcevia keyserlingi Tylogonus cf. viridimicans Tylogonus yanayacu Tylogonus cf. auricapillus Tylogonus parvus Chalcotropis luceroi Palpelius beccarii Palpelius cf. discedens Pristobaeus cf. jocosus Xenocytaea agnarssoni Xenocytaea albomaculata Xenocytaea proszynskii Thiania cf. viscaensis Thiania latibola Thiania cf. suboppressa Thiania bhamoensis Thiania tenuis Lagnus edwardsi Thianitara spectrum Foliabitus longzhou Foliabitus sp. [Malaysia] Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola tamasi Thorelliola ensifera Thorelliola cf. mahunkai Thorelliola mahunkai Thorelliola tualapa Agobardus bahoruco Agobardus gramineus Agobardus oviedo Agobardus cordiformis Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Parvattus zhui Servaea vestita Bythocrotus cf. crypticus Bythocrotus crypticus Popcornella yunque Popcornella furcata Popcornella spiniformis Athamas cf. whitmeei Athamas nitidus Athamas sp. Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus dominicanus Truncattus cachotensis Truncattus flavus Antillattus gracilis 'Cobanus' cambridgei Antillattus cf. applanatus 'Pensacola' darlingtoni 'Pensacola' maxillosa 'Siloca' electa Petemathis minuta cf. Chapoda sp. [Arizona] 'Sidusa' recondita Chapoda gitae Chapoda angusta Chapoda sp. [Fortuna] Chapoda cf. fortuna [CostaRica] Chapoda fortuna [Panama] Chapoda peckhami Chapoda cf. inermis [Panama] Chapoda cf. inermis [CostaRica] cf. Maeota sp. [Panama] Maeota dorsalis Maeota simoni Maeota flava Maeota sp. [Napo] Maeota dichrura Maeota sp. [Cutucú] Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [MoronaSantiago] 'Pensacola' tuberculotibiata * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * ** * * * ** ** * * * * * ** * * * * ** * * * * * * * * * * * * * * * * * * * * ** * * * * ** * * ** * * * * ** ** * * * * * * ** * * * * * ** * * * ** * * ** ** * * * * * * * * * 0.1 * ML bootstrap value for clade >=0.7 clade also present in strict consensus of MP trees 48! Figure 2.6. Summary of ML and MP analyses on COI. Tree shown is the best tree from the ML analysis with lnL = -43516.919. Xysticus sp. Hurius vulpinus Attidops youngi Salticus scenicus Jollas sp. Prostheclina sp. cf. Saitis sp. [WesternAustralia] Euophryine sp. [GentingHighlands] Thyenula sp. [SouthAfrica] Maileus cf. fuscus Popcornella nigromaculata Saitis barbipes Yllenus arenarius Hypoblemum cf. albovittatum Hypoblemum sp. [NewSouthWales] Lycidas cf. griseus Maratus cf. amabilis Maratus sp. [SouthAustralia] Jotus auripes Lycidas cf. vittatus Thianitara spectrum Servaea vestita Thorelliola cf. mahunkai Thorelliola ensifera Colyttus bilineatus Colyttus robustus Tylogonus cf. viridimicans Tylogonus yanayacu Tylogonus cf. auricapillus Tylogonus parvus Heliophanus cupreus Parvattus zhui Aelurillus cf. ater Belliena sp. [JatunSacha] Corticattus guajataca Corticattus latus Belliena sp. [Yanayacu] Naphrys pulex Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Corythalia bromelicola Corythalia coronai 'Wallaba' decora 'Corythalia' cf. canalis Anasaitis adorabilis Anasaitis canosa [USA] 'Corythalia' locuples 'Corythalia' banksi Corythalia porphyra Corythalia bicincta Corythalia sp. [Baeza] Corythalia electa Corythalia sp. [JatunSacha] Chalcotropis luceroi Laufeia concava Palpelius cf. discedens Thorelliola Joannae Arasia mollicoma Ohilimia scutellata Bythocrotus cf. crypticus Bythocrotus crypticus Amphidraus complexus Bulolia excentrica Nebridia cf. semicana Philaeus chrysops Marma nigritarsis Soesilarishius micaceus Soesilarishius ruizi Popcornella furcata Parabathippus shelfordi Parabathippus kiabau Parabathippus magnus Chalcolecta prensitans Diolenius cf. decorus Diolenius varicus Efate albobicinctus Paraharmochirus tualapaensis Chalcotropis cf. caeruleus Mantisatta longicauda Foliabitus longzhou Cobanus sp. [Ecuador] Sidusa sp.2 [FrenchGuiana] Cobanus sp. [CostaRica] Cobanus extensus Cobanus cf. electus Sidusa sp.1 [FrenchGuiana] Lepidemathis haemorrhoidalis 'Euophrys' a-notata 'Euophrys' cf. patagonica Lagnus edwardsi Asaphobelis physonychus Siloca cf. campestrata Siloca cf. sanguiniceps Ecuadattus napoensis Havaika sp. 'Margaromma' cf. semirasum Omoedus brevis Omoedus cf. ponapensis Omoedus papuanus Omoedus meyeri Omoedus cf. piceus Omoedus ephippigera Omoedus cf. metallescens Omoedus orbiculatus 'Saitis' cf. mundus Thyenula cf. aurantiaca Junxattus daiqini Orcevia keyserlingi Coccorchestes cf. aiyura Coccorchestes cf. inermis Thiania cf. viscaensis Thiania cf. suboppressa Thiania latibola Neon nelli Ilargus pilleolus Ilargus moronatigus Ilargus macrocornis Ilargus galianoae Ilargus serratus Athamas cf. whitmeei Athamas nitidus Thyenula nelshoogte Thyenula laxa Thyenula wesolowskae cf. Acragus Viribestus suyanensis Chinattus parvulus Compsodecta haytiensis 'Bathippus' pahang Tomocyrba sp. Galianora bryicola Thrandina parocula Lyssomanes viridis Holcolaetis sp. Portia labiata Chalcoscirtus alpicola Pseudeuophrys erratica Talavera minuta Leptathamas paradoxus Cytaea oreophila Donoessus striatus Cytaea sp. [Wewak] Cytaea cf. sinuata Cytaea nimbata Pristobaeus cf. jocosus Euryattus bleekeri Euryattus cf. porcellus Euryattus sp. [Queensland] Chapoda angusta 'Sidusa' recondita Chapoda cf. fortuna [CostaRica] Chapoda fortuna [Panama] Chapoda cf. inermis [Panama] Chapoda cf. inermis [CostaRica] Chapoda peckhami cf. Maeota sp. [Panama] cf. Chapoda sp. [Arizona] Maeota sp. [JatunSacha] Maeota simoni Maeota sp. [Manabi] Maeota sp. [Cutucú] Maeota dichrura Maeota sp. [Napo] Maeota flava Maeota sp. [MoronaSantiago] 'Pensacola' tuberculotibiata Plexippus paykulli Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus cachotensis Truncattus dominicanus Truncattus flavus 'Cobanus' cambridgei Antillattus cf. applanatus 'Pensacola' maxillosa 'Pensacola' darlingtoni Agobardus gramineus Agobardus cordiformis Agobardus oviedo Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Canama cf. forceps Canama extranea Bathippus gahavisuka Bathippus madang Bathippus korei Bathippus cf. directus [Suyang] Bathippus directus [Tualapa] * * * * * * ** * * * * * * * * * * * * * * ** * ** * * * ** * * * * * * ** * * * * * * * * * 0.1 * ML bootstrap value for clade >=0.7 clade also present in strict consensus of MP trees 49! Figure 2.7. Chronogram of euophryine divergence. Times shown are the median age estimates from the BEAST Analysis Two, with 95% HPD for major clades indicated as gray bars. Tree shown is the best tree from the ML analysis on all genes combined dataset with all non- euophryine taxa trimmed off. Continental distribution of taxa, and major Caribbean and Papua New Guinea Clades are indicated in front of the taxon names. Numbers for major clades are consistent with those of Table 2.5. Anasaitis adorabilis Anasaitis hebetata Anasaitis brunnea 'Corythalia' cf. canalis Anasaitis canosa [Panama] Anasaitis canosa [USA] 'Corythalia' banksi 'Stoidis' placida 'Corythalia' gloriae Anasaitis laxa 'Corythalia' elegantissima Anasaitis sp. [Peblique] 'Corythalia' locuples Corythalia porphyra Corythalia cf. albicincta Corythalia sulfurea Dinattus minor Corythalia broccai Corythalia peblique Corythalia bromelicola Corythalia coronai 'Wallaba' decora Corythalia sp. [PuertoLopez] Corythalia bicincta Corythalia cf. latipes Corythalia cf. valida Corythalia electa Corythalia sp. [JatunSacha] Corythalia sp. [Baeza] Parvattus zhui Parabathippus kiabau Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Euophrys frontalis Euophrys monodnock Euophrys cf. proszynskii Chalcoscirtus alpicola Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Cobanus sp. [Ecuador] Cobanus cf. electus Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Cobanus cf. cambridgei Cobanus mandibularis Cobanus sp. [CostaRica] Cobanus unicolor Cobanus extensus Cobanus sp. [Panama] Pensacola signata Mexigonus morosus Mexigonus arizonensis Mexigonus cf. minutus [Ecuador] Petemathis minuta Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis 'Cobanus' cambridgei 'Pensacola' darlingtoni 'Siloca' electa 'Pensacola' maxillosa Corticattus latus Corticattus guajataca Naphrys pulex Naphrys xerophila Popcornella spiniformis Popcornella furcata Popcornella yunque Popcornella nigromaculata Compsodecta haytiensis Compsodecta peckhami Bythocrotus crypticus Bythocrotus cf. crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo cf. Chapoda sp. [Arizona] 'Sidusa' recondita Chapoda gitae Chapoda angusta Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami Chapoda sp. [Fortuna] Chapoda fortuna [Panama] Chapoda cf. fortuna [CostaRica] cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [Cutucú] Maeota sp. [Napo] Maeota dichrura Maeota simoni Maeota flava Maeota sp. [MoronaSantiago] 'Pensacola' tuberculotibiata Marma nigritarsis Nebridia cf. semicana Amphidraus complexus Tylogonus cf. viridimicans Tylogonus yanayacu Tylogonus cf. auricapillus Tylogonus parvus Soesilarishius cf. amrishi Soesilarishius ruizi Soesilarishius micaceus Tariona cf. bruneti 'Euophrys' a-notata 'Euophrys' cf. patagonica cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Asaphobelis physonychus Siloca cf. campestrata Siloca cf. sanguiniceps Neonella vinnula Ecuadattus napoensis Ecuadattus pichincha Belliena sp. [Yanayacu] Belliena sp. [JatunSacha] Belliena ecuadorica (M) Belliena ecuadorica (F) Ilargus galianoae Ilargus pilleolus Ilargus serratus Ilargus coccineus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Servaea vestita Jotus auripes Prostheclina sp. Lycidas cf. vittatus cf. Jotus sp. [Queensland] Maileus cf. fuscus cf. Saitis sp. [WesternAustralia] Saitis barbipes Maratus cf. amabilis Maratus sp. [SouthAustralia] Lycidas cf. griseus Hypoblemum sp. [NewSouthWales] Hypoblemum cf. albovittatum Lycidas cf. karschi Emathis gombak Lepidemathis Chalcotropis Chalcotropis cf. caeruleus Colyttus robustus Colyttus bilineatus Donoessus striatus Lagnus edwardsi Euophryine sp. [GentingHighlands] Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola tamasi Thorelliola ensifera Thorelliola tualapa Thorelliola mahunkai Thorelliola cf. mahunkai Foliabitus sp. [Malaysia] Foliabitus longzhou Laufeia concava Junxattus daiqini Laufeia eximia Orcevia keyserlingi Thianitara spectrum Thiania cf. viscaensis Thiania latibola Thiania cf. suboppressa Thiania tenuis Thiania bhamoensis Chinophrys pengi Thyenula laxa Thyenula nelshoogte 'Saitis' leighi Thyenula sp. [SouthAfrica] Thyenula wesolowskae 'Saitis' cf. mundus Thyenula cf. aurantiaca Cytaea oreophila Cytaea mitellata Cytaea cf. sinuata Cytaea nimbata Cytaea sp. [Wewak] Cytaea cf. rai Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Euryattus cf. venustus Euryattus bleekeri Euryattus cf. porcellus Euryattus sp. [Queensland] Phasmolia elegans Zabkattus furcatus Zabkattus richardsi Zabkattus brevis Zabkattus trapeziformis Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Bathippus macrognathus Bathippus korei Bathippus madang Bathippus gahavisuka Bathippus directus [Tualapa] Bathippus cf. directus [Suyang] Pristobaeus cf. jocosus Palpelius cf. discedens Palpelius beccarii Xenocytaea agnarssoni Xenocytaea proszynskii Xenocytaea albomaculata Chalcolemia nakanai Sobasina wanlessi Paraharmochirus tualapaensis Efate albobicinctus Chalcolecta prensitans Ohilimia scutellata Diolenius varicus Diolenius cf. decorus Omoedus cf. piceus Omoedus ephippigera Omoedus orbiculatus Omoedus cf. metallescens Omoedus brevis Omoedus swiftorum 'Margaromma' cf. semirasum Omoedus cf. durvillei Omoedus cf. ponapensis 'Margaromma' cf. torquatum Omoedus cf. danae Omoedus darleyorum Omoedus sp. [Adalbert] Omoedus tortuosus Omoedus papuanus Omoedus sp. [Gahavisuka] Omoedus meyeri Omoedus omundseni 1 2 3 12 18 4 5 6 8 9 10 11 13 14 15 16 17 19 20 7 50 40 30 20 10 0 Millions of years Neotropics Nearctic Australasia Eurasia Africa Continental distribution Euophryinae 95% highest posterior distributions of node ages Caribbean radiation Papua New Guinea radiation 50! 3 Generic review of euophryine jumping spiders (Araneae: Salticidae), with a phylogeny from combined molecular and morphological data 3.1 Synopsis Morphological traits of a broad range of euophryine genera and species were extensively studied to clarify generic delimitations in the Euophryinae and to permit phylogenetic classification of genera lacking molecular data. A full list of euophryine genera is provided. Euophryine generic groups and the delimitation for some genera are reviewed in detail. In order to explore the effect of adding formal morphological data to previous molecular phylogenetic studies, and to find morphological synapomorphies, eighty-two morphological characters were scored for 203 euophryine species and seven outgroup species. The results show that the morphological dataset does not perform as well as the molecular dataset (genes 28S, Actin 5C; 16S-ND1, COI) in resolving the phylogeny of Euophryinae, probably because convergence and reversals of morphological traits are common in euophryines, diluting the phylogenetic signal of morphological characters. The formal morphological data were optimized on the phylogeny in order to seek synapomorphies, in hopes of extending the phylogeny to include taxa for which molecular data are not available. It was difficult to determine solid synapomorphies for euophryine clades due to repeated convergences and reversals. However, synapomorphies that work locally still help to delimit euophryine generic groups and genera. The following synonyms of euophryine genera are proposed: Maeotella with Anasaitis; Dinattus with Corythalia; Paradecta with Compsodecta; Cobanus, Chloridusa and Wallaba with Sidusa; Tariona with Mopiopia; Nebridia with Amphidraus; Asaphobelis and Siloca with Coryphasia; Ocnotelus with Semnolius; Palpelius with Pristobaeus; Lycidas, Maratus, Jotus, Hypoblemum and Maileus with Saitis; Junxattus with Laufeia; Donoessus with Colyttus; Nicylla, Pselcis and Thianitara with Thiania. The new genus Saphrys is erected to enclose misplaced species. 3.2 Introduction Euophryinae is one of the largest groups in jumping spiders (Salticidae), with about 1000 species (see Chapters 2, 5-8) reported from all over the world and many more undescribed. Most euophryine species are found in the tropics of both the Old and the New World. Spiders of this group have evolved a great variety of body forms: some are relatively large and slender with 51! long legs (e.g. Bathippus spp.), some are very robust with short legs and a high carapace (e.g. Omoedus piceus Simon), some are flattened (e.g. Thiania spectrum (Simon)); some are weevil- like (Coccorchestes spp.), some are ant-like (Sobasina spp. and Paraharmochirus spp.). The Euophryinae was initially erected by Eugène Simon (1901a) using the name “Evophrydeae”. In Simon’s classification (Simon 1901a; 1903), it was considered as a group in Unidentati and comprised three genera: Akela, Euophrys and Rhyphelia, of which Akela is apparently not closely related with the other two judging from its morphology (Galiano 1989: 49, figs. 1-4). However, most of the genera currently considered as euophryines were scattered among more than 20 groups of Pluridentati, Unidentati and Fissidentati in this classification (see Table 3.3). More than seventy years after the Euophryinae was erected, Jerzy Prószy!ski (1976) clarified the group for the first time by a delimiting feature: the presence of a coiled embolus at the distal end of palpal tegulum. Based on this delineation he included 13 genera (Admestina, Agobardus, Chalcoscirtus, Corythalia, Euophrys, Habrocestum, Laufeia, Marchena, Neonella, Saitis, Stoidis, Talavera, Thiania) as euophryines in his partial classification of salticids. Among these, Admestina is now considered a marpissoid (Maddison & Hedin 2003a) and Marchena a heliophanine (Maddison 1987). Maddison and Hedin (2003a) extended the content of the subfamily and explicitly indicated 34 genera as euophryines (see Table 3.3). They also revised the delimitation to further specify the particular form of the embolus and tegulum in euophryines: the plane of the spiral of the embolus is more or less parallel to the longitudinal axis of the palp and a loop in the sperm duct projects towards the centre of the tegulum. A recent molecular phylogenetic study on the Euophryinae further dramatically extended its content to 85 genera (see Chapter 2). Although the morphological delimitation for the subfamily Euophryinae is relatively clear, the taxonomy within the subfamily is a mess. Even though euophryine species show a variety of body forms, there are indications from the molecular phylogeny that similar body forms in different euophryine lineages have arisen by convergence, and therefore have resulted in polyphyletic genera (e.g., Bathippus and Parabathippus). In addition, genitalic organs within this subfamily are relatively simple and usually show little interspecific variation, hindering attempts to classify genera. Most work on this subfamily has concentrated on describing species and genera, or on a regional revision of some genera (e.g. Bryant 1940, 1943, 1950; Balogh 1980; Berry et al. 1996, 1997, 1998; Edwards 2002; Logunov & Azarkina 2008, etc.), and there 52! is little systematic work on the group as a whole. Although a few genera have been reviewed in the past (Prószy!ski 1971; Galiano 1985; Zabka 1987; Jendrzejewska 1995; Gardzinska & Patoleta 1997; Logunov 1998; Bodner 2002; Zabka & Pollard 2002; Logunov & Kronestedt 2003; Richardson & Zabka 2007), the delimitations of most genera are still not clear, e.g. Corythalia and Bathippus. Therefore, the group's taxonomy remains confused, and it is difficult to identify even some commonly collected species to genus. Unfortunately, the molecular phylogeny cannot completely resolve the taxonomic problems. The molecular phylogeny of Euophryinae is only skeletal, with a limited number of species sampled and sequenced. The majority of species of interest will have to be placed on this skeleton by their phenotypic (including morphological) characteristics (Maddison & Maddison 1992: 72). Thus, an extensive collection of morphological data of euophryine jumping spiders is vital not only for clarifying the taxonomy, but also for a more comprehensive phylogeny of Euophryinae (Maddison & Maddison 1992; Maddison 1996). The primary goal of this study is to resolve the phylogenetic placement of many euophryines, using morphology, and to achieve a broad taxonomic arrangement for the subfamily, based on the molecular phylogeny (see Chapter 2) and morphological studies. In addition, I formally scored morphological traits for 203 euophryine species and seven outgroup species for phylogenetic analyses. The performance of the morphological traits in resolving the phylogeny of euophryines was evaluated. In an attempt to find synapomorphies for euophryine clades, the evolution of morphological characters was also traced on the phylogeny. 3.3 Material and methods 3.3.1 Techniques for morphological study Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Preserved specimens were photographed under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0. Terminology is standard for Araneae. The following abbreviations are used: ALE, anterior lateral eye; AME, anterior median eye; PLE, posterior lateral eye; PME, posterior median eye (the \"small eyes\"); OAL, ocular area length; PS, primary spermatheca; SS, secondary 53! spermatheca; CD, copulatory duct; W, window of epigynum; MS, median septum; CO, copulatory opening; FD, fertilization duct; AG, accessory gland; RTA, retrolateral tibial apophysis; PTA, prolateral tibial apophysis; PA, patellar apophysis; FA, femoral apophysis; E, embolus; ED, embolic disc; DH, distal hematodocha; BH, basal hematodocha; T, tegulum; ST, subtegulum; SR, salticid radix; TL, tegular lobe; RSDL, retrolateral sperm duct loop; PSDL, prolateral sperm duct loop; VTB, ventral tibial bump; LE, lamella of embolus; LTS, lamella on tegular shoulder; PED, process on embolic disc. The specimens used in taxonomic and phylogenetic studies belong to the following institutions: Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM); Museum National d’Histoire Naturelle, Paris (MNHN); Museum of Comparative Zoology (MCZ); California Academy of Science (CAS); Florida State Collection of Arthropods (FSCA); Museo Argentino de Ciencias Naturales (MACN); Natural History Museum, London (NHM); American Museum of Natural History (AMNH); Royal Museum of Central Africa, Belgium (RMCA); Instituto Butantan, São Paulo, Brazil (IBSP). Many of the specimens are from recent expeditions (see Chapters 5-8). The habitat indications came from experience collecting during those expeditions. 3.3.2 Phylogenetic study 3.3.2.1 Taxon sampling In total, 210 salticid species of which DNA sequences were available (see Chapter 2) were included in phylogenetic analyses. Among them, 203 species are euophryines and seven species are outgroups. The seven outgroup species represent lineages closely related to Euophryinae (see Chapter 2; Maddison et al. 2008; Bodner 2009): Ghelna canadensis (Banks) (Marpissoida), Chinattus parvulus (Banks) (Hasarieae), Heliophanus cupreus (Walckenae) (Heliophaninae), Freya decorata (C.L. Koch) (Aelurilloida), Salticus scenicus (Clerck) (Salticinae), Plexippus paykulli (Audouin) (Plexippoida) and “Bathippus” pahang Zhang, Song & Li (Nanneninae). Morphological characters were scored for the same species as in the molecular phylogeny when specimens were available for morphological study. In a few cases when specimens for a species or a sex were not available for morphologicial study, a very closely related species of the same genus was used instead. 54! 3.3.2.2 Morphological characters Eighty-two morphological characters were formally scored for each species for phylogenetic analyses, of which 45 (Characters 1-45) are derived from somatic morphology, 26 (Characters 46-71) from male palpal structures, and 11 (Characters 72-82) from female copulatory organs. Characters were scored through direct observations on specimens. Descriptions of morphological characters are given in Appendix 2. 3.3.2.3 DNA sequences DNA sequences and alignments from Chapter 2 were used in phylogenetic analyses. Some taxa were trimmed to match the morphological dataset in order to compare the performance of DNA data and morphological data in resolving the phylogeny of Euophryinae. On this reduced set of taxa the non-coding regions, 28S and 16S (plus the adjacent tRNA), were realigned using the Opalescent package (Wheeler & Kececioglu 2007) in Mesquite 2.73 (Maddison & Maddison 2010) under the default gap open/gap extension costs: 260/69 (see Chapter 2). 3.3.2.4 Phylogenetic analyses Molecular sequences (28S, Actin 5C, 16SND1 and COI) were concatenated. Phylogenetic analyses were performed on the morphological matrix and the combined matrix of morphology and molecules. Independent analyses on the molecular dataset were also conducted because some taxa had been trimmed to match the morphological matrix. The phylogeny of Euophryinae from the combined matrix of all genes proposed in Chapter 2, with the taxa without morphological data for phylogenetic study pruned out, is shown in Fig. 3.1. This phylogeny is the basis of comparison with results from other datasets. 3.3.2.4.1 Morphological matrix Maximum parsimony (MP) analyses were conducted on the morphological matrix. TNT 1.1 (Goloboff et al. 2008) was run to find the most parsimonious trees using the “New Technology Search” method with the combination of sectorial search, rachet, drift and tree fusing techniques, and all under the default settings. The characters 14, 17, 33, 38-41, 46, 51, 58, 63, 70, 72, 79 and 82 were considered ordered and all other characters were treated unordered. The strict consensus tree was built in Mesquite 2.73 (Maddison & Maddison 2010a) from all the equally most 55! parsimonious trees. One analysis was run without constraining the monophyly of the Euophryinae; the other analysis was conducted with the Euophryinae being enforced to be monophyletic. The node support value was assessed by a TNT non-parametric bootstrap analysis (Felsenstein 1985) with 500 replicates. 3.3.2.4.2 DNA matrix Maximum likelihood (ML) analysis was conducted on the DNA matrix. The molecular data were divided into 8 partitions: 28S; Actin 5C first, second and third codon positions; 16S; ND1+COI first, second and third codon positions. Modeltest 3.7 (Posada & Crandall 1998; Posada & Buckley 2004) in combination with PAUP* 4.0b10 (Swofford 2002) was used to choose appropriate substitution model for each partition of the molecular data via the Akaike Information Criterion (AIC). The substitution model selected by ModelTest for each molecular partition is shown in Table 3.1. The maximum likelihood tree search was completed under the test version of GARLI (GARLI- Part-0.97, https://www.nescent.org/wg_garli/Partition_testing_version) because it allows each partition to have its own model or model parameters (see Table 3.1 for the model selected for each partition by ModelTest). GARLI parameters were the same as in the configuration template provided with the program, except for the following minor changes to reduce the computational time: attachmentspertaxon=50; startoptprec=5; minoptprec=1; numberofprecreductions=5; treerejectionthreshold=10.0; topoweight=0.05. Ten separate GARLI runs each with 30 search replicates (300 search replicates in total) were carried out to find the best ML tree. Five hundred replicates of bootstrap analysis was also conducted in GARLI to assess the node support value, each with one search replicate. 3.3.2.4.3 Combined DNA and morphology matrix Maximum likelihood (ML) analysis was conducted on the combined molecular and morphological data matrix with the data divided into nine partitions (morphology; 28S; Actin 5C first, second and third codon positions; 16S; ND1+COI first, second and third codon positions). The ML analysis was also conducted under the test version of GARLI (GARLI-Part-0.97, https://www.nescent.org/wg_garli/Partition_testing_version). It not only allows each molecular 56! partition to have its own model or model parameters, but also allows mixing of sequence and morphology data. The model selected by Modeltest (see Table 3.1) was assigned to each sequence partition, and the “Mkv” model (Lewis 2001) was used for the morphological characters. GARLI parameters were the same as in the configuration template provided with the program, except the same minor changes to reduce the computational time as in the GARLI analysis on the DNA matrix. Ten separate GARLI runs each with 50 search replicates (500 search replicates in total) were carried out to find the best ML tree. The node support value was assessed by a GARLI bootstrap analysis with 1000 replicates, each with one search replicate. 3.3.2.5 Character optimization MacClade 4.08 (Maddison & Maddison 2000) was used to study morphological character evolution on the phylogeny. The morphological traits were traced on the ML tree from the DNA dataset (see Chapter 2) with taxa of morphological data unavailable trimmed off (Fig. 3.1) and on the ML tree from the combined molecular and morphological dataset (Fig. 3.5). Unambiguous changes in the most parsimonious reconstructions on these two phylogenies were then compared. 3.4 Results 3.4.1 Morphological study on a broad range of euophryine taxa The results of morphological study on a broad range of euophryine genera and species are incorporated in the taxonomic review of Euophryinae presented in the Discussion. 3.4.2 Phylogenetic study 3.4.2.1 Morphological dataset The scored morphological matrix is shown in Appendix 3. The TNT analysis without constraining euophryine monophyly was stopped when the best score was hit twice and four equally parsimonious trees (score=1141) were saved. Strict consensus of these four trees is shown in Fig. 3.2. All of the most parsimonious trees failed to recover the clade Euophryinae. Most generic groups well supported by the molecular data were unrecognized, for instance, the Anasaitis-Corythalia Clade and the Bulolia-Coccorchestes Clade. 57! Some euophryine genera well defined by known morphological features were not recognized in the analysis, e.g. Anasaitis and Omoedus. The TNT analysis with euophryines enforced as a monophyletic group found seven equally parsimonious trees with tree length four steps longer than those from the unconstrained analysis (score=1145, stopped after the best score was hit three times). The recovered relationships within Euophryinae are more consistent with the molecular analysis and apparent morphological similarities when compared with the results from the unconstrained analysis. However, the strict consensus tree of the seven equally parsimonious trees (Fig. 3.3) recovered only a few clades (e.g. Diolenius and its relatives, Omoedus Clade etc.) but failed on most of the others (e.g. Anasaitis-Corythalia Clade, Agobardus Clade, Thiania Clade, etc.). The TNT bootstrap analysis suggests the morphological dataset is not informative in resolving the phylogenetic relationships within the Euophryinae (Fig. 3.2). Almost all clades were collapsed in the majority rule consensus tree of the 500 trees from the bootstrap replicates. Three genera (Bathippus, Coccorchestes and Parabathippus) were recovered in the bootstrap analysis, but only Coccorchestes and Parabathippus have bootstrap values higher than 70%. 3.4.2.2 Molecular dataset Realignment of the sequences resulted in 4311 sites. The best ML tree found in the GARLI analysis is shown in Fig. 3.4 (lnL = -135203.637, run 4, search replicate #13), with the topology very similar to the molecular phylogeny proposed in Chapter 2 (see Fig. 3.1). The Old World euophryine taxa also tend to cluster together rather than with the New World taxa, and vice versa. The major differences reside in the groupings at relatively deeper level within the Euophryinae, which may be due to the reduced sample of taxa in this study. For example, in the phylogeny proposed in Chapter 2, a New World clade (Anasaitis-Corythalia Clade) is sister to the rest, whereas in the tree shown in Fig. 3.4, the basal branches of the euophryine phylogeny are composed of Old World euophryines. All major euophryine generic groups and genera indicated in Fig. 3.1 are recovered, and most of them have bootstrap support values higher than 70%, with the exceptions of Thyenula, Popcornella and Petemathis. An unexpected outcome is that Chinophrys (from China) falls in a clade of 58! Neotropical euophryines (Amphidraus, Marma and Soesilarishius) rather than with the Old World euophryines. However, this grouping is not supported by the bootstrap analysis. 3.4.2.3 Combined DNA and morphology dataset The combined DNA and morphology dataset contained 4393 characters. The best ML tree has lnL -140981.450. The tree (Fig. 3.5) results in similar generic groups and genera of Euophryinae among the terminal clusters as the molecular phylogeny proposed in Chapter 2 (see Fig. 3.1). However, it shows a more mixed pattern of the Old World taxa with the New World taxa than the tree from DNA alone. For example, the New World Sidusa Clade clusters with the Parabathippus-Parvattus Clade from the Old World. In addition, the phylogeny does not put the two African Thyenula species, Thyenula laxa and T. nelshoogte in the same clade containing other Thyenula species. Compared to the DNA data alone, adding morphological data increased the ML bootstrap support value for some clades, but decreased it for others. For instance, the bootstrap value increased 15% for the Emathis-Lepidemathis Clade, 25% for Popcornella and 8.8% for Chapoda, but decreased 11.4% for the Agobardus Clade. 3.4.2.4 Character optimization The unambiguous changes among all most parsimonious reconstructions of character evolution on Fig. 3.1 and Fig. 3.5 are listed in Table 3.2. Character state changes are reconstructed differently on the two topologies for certain clades, e.g. Euophryinae Clade and Omoedus Clade. 3.5 Discussion I will first discuss the phylogenetic analyses, and then present the taxonomic review of Euophryinae. 3.5.1 Phylogenetic analyses 3.5.1.1 DNA vs. morphology in phylogenetic reconstruction The phylogeny from the molecular dataset with some taxa trimmed to match the morphological dataset recovers most of the clades indicated in Fig. 3.1. Most of the recovered clades have high 59! ML bootstrap support values. In contrast, very few clades indicated in Fig. 3.1 are recognized in the phylogeny from the morphological dataset, with even some easily recognized genera (e.g. Anasaitis) unrecovered. Most of the clades in the morphogical phylogeny are not well supported by the MP bootstrap analysis. In addition, the molecular phylogeny is more or less concordant with geography, such that the New World and Old World euophryines usually form their own clades with very few cases of mixure (see Chapter 2). However, this coherent geographic pattern falls apart in the morphological phylogeny, which has considerable mixing of euophryines from two hemispheres. Given the age of Euophryinae (see Chapter 2), the highly mixed geographic pattern shown in the morphological phylogeny would require many long-distance intercontinental dispersals. The greater resolution and geographic coherence suggest that the DNA dataset performs better than the morphological dataset in resolving the phylogeny of Euophryinae. The morphological characters were dramatically outnumbered by the DNA characters, and may have too few phylogenetically informative characters to resolve the relationships of the taxa. In addition, convergences and reversals in morphological characters may have diluted their phylogenetic signals, weakening their performance in the phylogenetic reconstruction (see below under “Role of morphology in phylogeny of Euophryinae”). The combined DNA and morphology tree differs from the DNA-only tree at a relatively deep level within the euophryines, showing more mixing of Old World and New World taxa. Because the molecular phylogeny from the full taxon sample (Fig. 3.1, also see Chapter 2) has the greatest geographic coherence and resolution of generic clades and genera, it is relied on as the basis for the taxonomic review of the Euophryinae. Besides the geographic mixing and lower resolution, there are hints that the addition of morphological data to the DNA degrades the phylogenetic signal through convergence. The Southeast Asian genus Parabathippus and the superficially-similar Neotropical genus Sidusa are grouped together in the combined analysis, but not with DNA alone. I suspect that this is an artifact due to convergence to similar body forms with elongate chelicerae in males. However, one of the BEAST analyses on the molecular data (see Chapter 2) also resulted in this grouping, suggesting that it could be correct. More data are needed to conclude definitively whether the similarity of Parabathippus and Sidusa is due to common ancestry, or to independent convergent evolution on different continents. 60! 3.5.1.2 Role of morphology in phylogeny of Euophryinae Using the molecular phylogeny (Fig. 3.1) as a basis, convergences and reversals in morphological traits are revealed to be common within the Euophryinae. For instance, the male palpal ventral tibial bump (ch. 48-1) has evolved at least seven times independently. Even though most euophryines have a retrolateral sperm duct loop on the male palp (ch. 56-0), this feature has been lost in several euophryine lineages, such as Neonella, Marma, Bulolia, and Sobasina. Such common convergence and reversal make it difficult to find solid synapomorphies for euophryine genera or generic groups. For some clades that are well supported by the phylogenetic analyses (also see Chapter 2), there are no unambiguous morphological synapomorphies seen in the character optimization, e.g. the Antillattus Clade. Nonetheless, the results of character optimization reveal morphological traits useful in delimiting euophryine clades. For instance, the lamella on the tegular shoulder of the male palp (ch. 49-1) is only seen in the Saitis Clade. Even characters that have evolved multiple times independently within Euophryinae can still be useful locally in distinguishing a euophryine clade from the others found in the same continental region. For example, even though the proximal or prolateral proximal sperm duct loop also appears in some New World euophryine lineages (e.g. Corticattus, Neonella, etc.), it is one of the defining features for the Bulolia- Coccorchestes Clade and distinguishes the clade from almost all other euophryines in New Guinea with the only exception of Xenocytaea albomaculata. The taxonomic review discusses other such useful morphological traits. In addition, extensive informal morphological studies on euophryine jumping spiders suggest some traits, even though not formally scored, are useful in uniting species, such as the foliate marking pattern and the body form in the Coryphasia Clade, and the extremely long and convoluted copulatory duct in the Omoedus Clade (see detailed discussion on taxonomic review). Therefore, even though morphology alone struggles to resolve phylogenetic relationships of Euophryinae, it helps in understanding the phylogenetic placement of many euophryine taxa and in extending the molecular phylogeny. For instance, the molecular phylogeny supports the monophyly of Euophryinae and provides a clear outline of the content of Euophryinae, with the vast majority of euophryines showing following characters: presence of retrolateral sperm duct loop, distal coiled embolus and epigynal window. Based on those morphological traits, I place 36 genera for which DNA data are unavailable within Euophryinae (Table 3.3). Among them, 61! 14 are grouped within generic groups and eight are considered as synonyms. An additional 13 genera are also tentatively placed on the phylogeny of Euophryinae, as discussed in the taxonomic review. 3.5.2 Taxonomic review of Euophryinae The generic content of Euophryinae is discussed below. Because of concerns that morphological convergences may distort phylogenetic relationships, I will rely on the molecular phylogeny from the full taxon sample (Fig. 3.1, also see Chapter 2) as a basis for the taxonomic discussion of clades and genera. When a synapomorphy provided for a clade resulted from character optimization on the molecular phylogeny (Fig. 3.1, Table 3.2), the character and state numbers are indicated in parentheses. When no number is indicated, it means the proposed diagnostic character for the clade is from informal morphological study but not from character optimization analyses. In a few cases, no clear morphological character has been discovered to define a clade and the grouping of genera or species is based purely on molecular phylogeny, e.g. the Parabathippus-Parvattus Clade. The phylogeny of Euophryinae (Fig. 3.1, also see Chapter 2) shows that most medium-sized clades are restricted to either Old or New World. Thus I divide my treatment by hemisphere in the review of euophryine clades (see below), which may be useful for those learning a regional euophryine fauna. 3.5.2.1 Content of Euophryinae Most euophryines have a curved or coiled embolus and a retrolateral sperm duct loop on the male palp, and a “window” structure on the female epigynum (Maddison & Hedin 2003a). The taxa grouped into the Euophryinae by the molecular data continue to support these delimitations, although some taxa (e.g. Diolenius and relatives) complicate them with highly derived genitalia (see Chapter 2). Based on these morphological delimitations, an additional 36 genera not sampled in the molecular phylogenetic analyses (see Chapter 2) are also classified as Euophryinae (see Table 3.3 for the full list of euophryine genera). The genera, Baviola, Gorgasella, Lauharulla, Lechia and Panysinus possibly also belong to the subfamily because their genitalic structures show some similarities with euophryines. Logunov and Azarkina (2008: 112, figs. 10-19) proposed the new genus Saaristattus and considered it as a euophryine. However, male palp of this genus lacks retrolateral sperm duct loop typical of euophryines, and the plane of embolic spiral is perpendicular to the longitudinal 62! axis of the palpal bulb, which is unusual for euophryines. Available information could support it either as closely related to Neon (Astioida, see Maddison et al. 2008), or as a euophryine. Thus, here I leave it out of Euophryinae pending more data. 3.5.2.2 Major euophryine clades from the New World 3.5.2.2.1 Anasaitis-Corythalia Clade (Figs 3.6-3.7) Anasaitis and Corythalia are included in this clade. Spiders of this clade are usually robust. Many species have iridescent scales. The chelicera usually has one biscuspid promarginal tooth (ch. 20-2, 22-3) and one unident retromarginal tooth. The male of some species has fringes on the first and second (ch. 35-1) legs. Male palp usually has a ventral bump on the tibia (ch. 48-1). The RTA is simple and finger-like. Anasaitis Bryant, 1950 Anasaitis Bryant, 1950: 168, type species: Saitis morgani (Peckham & Peckham, 1901). Maeotella Bryant, 1950: 186, type species: Saitis perplexus (Peckham & Peckham, 1901). New Synonmy Diagnosis. Species of Anasaitis are recognizable by the following genitalic characters: embolus usually short and not coiled; tegulum sometimes with a distinct proximal lobe; distal hematodocha highly reduced (ch. 70-1); epigynum without distinct window; copulatory duct usually short. Anasaitis are small to medium sized spiders. They are mainly ground dwellers and can be found on leaf litter, on rocks or in grass clumps. A few species can be found by beating low foliage. The following species are transferred to Anasaitis here. Most of them show the above genitalic features. However, Anasaitis banksi and A. elegantissima have a relatively long and slightly coiled embolus and relatively long copulatory ducts. Transferring them from Corythalia is based upon the molecular phylogeny in which these two species are embedded within Anasaitis (also see Chapter 2). Anasaitis perplexa (Peckham & Peckham, 1901) (from Maeotella, New Combination) Anasaitis banksi (Roewer, 1951) (from Corythalia, New Combination) Anasaitis canalis (Chamberlin, 1925) (from Corythalia, New Combination, Fig. 3.6H-N) 63! Anasaitis elegantissima (Simon, 1888) (from Corythalia, New Combination) Anasaitis gloriae (Petrunkevitch, 1930) (from Corythalia, New Combination) Anasaitis locuples (Simon, 1888) (from Corythalia, New Combination) Anasaitis arcuata (Franganillo, 1930) (from Corythalia, New Combination) Anasaitis cubana (Roewer, 1951) (from Corythalia, New Combination) Anasaitis emertoni (Bryant, 1940) (from Corythalia, New Combination) Anasaitis peckhami (Petrunkevitch, 1914) (from Corythalia, New Combination) Anasaitis squamata (Bryant, 1940) (from Corythalia, New Combination) Anasaitis placida (Bryant, 1947) (from Stoidis, New Combination, Fig. 3.6O-U) Notes. The results from phylogenetic analyses confirm Bryant’s (1950) concern that some species in Corythalia might have been misplaced and actually belonged to Anasaitis based on their palpal structure. I examined the type specimen of Stoidis placida Bryant (in MCZ, #22683) and determined this species has been misplaced and should belong to Anasaitis. In addition, the monotypic genus Maeotella Bryant from Jamaica is considered as a junior synonym of Anasaitis, based on it having the same genitalic pattern. Corythalia C. L. Koch, 1850 Corythalia C. L. Koch, 1850: 67, type species: Euophrys latipes C. L. Koch, 1846. Dinattus Bryant, 1943: 482, type species: Dinattus heros Bryant, 1943. New Synonym Diagnosis. Body form, color and marking pattern of Corythalia sometimes are similar to its sister, Anasaitis. Corythalia can be distinguished from Anasaitis by genitalia: embolus long and coiled with distinct embolic disc; distal hematodocha developed; epigynal window obvious with a median septum; opening to copulatory duct at anterior or posterior rim of the window; copulatory duct relatively long. Corythalia are small to medium sized, ground-dwelling or foliage-dwelling spiders. According to these features and the molecular phylogeny, the following species are transferred to Corythalia: Corythalia erebus (Bryant, 1943) (from Dinattus, New Combination) Corythalia heros (Bryant, 1943) (from Dinattus, New Combination) Corythalia minor (Bryant, 1943) (from Dinattus, New Combination, Fig. 3.7H-N) 64! Corythalia decora (Bryant, 1943) (from Wallaba, New Combination, Fig. 3.7O-U) Notes. Dinattus Bryant is a small genus with three species reported from Hispaniola (Bryant, 1943). It is well known by the prominent triangular lobe on the male carapace behind the ALEs (Bryant 1943). One species, Dinattus minor Bryant, 1943 was included in the molecular phylogenetic study, and the results indicate that it is embedded within Corythalia. Somatic and genitalic characters of Dinattus are like those of Corythalia, e.g. the presence of the distinct “window” of the epigynum and the coiled embolus (Fig. 3.7H-N). The molecular phylogeny indicates the lateral carapace extension in the male Dinattus is a derived feature within Corythalia, and carving out Dinattus based on that would leave Corythalia paraphyletic. Thus, Dinattus is considered as a junior synonym of Corythalia and all the three described species are transferred to Corythalia. Based on the results from morphological and phylogenetic studies, I also transfer Wallaba decora Bryant to Corythalia. The two other species of Wallaba, including the type species Wallaba metallica Mello-Leitão (Guyana) are not congeneric with W. decora according to their morphology (see the discussion under Sidusa Clade). 3.5.2.2.2 Antillattus Clade: Antillattus, Petemathis, Truncattus, and possibly Allodecta, Caribattus (Figs 3.8-3.11) This is a well-supported clade in the molecular phylogeny and is restricted to the Caribbean Islands. However, no unambiguous character change for this clade is resolved by character optimization. Most species in this clade have two promarginal teeth on the chelicera. The embolus of the male palp coils for no more than half a circle. The epigynum has a distinct window with a median septum. The opening to the copulatory duct is usually at median of the vulva. Copulatory duct is relatively short. This clade comprises Antillattus, Petemathis and Truncattus. Petemathis are small to medium sized spiders usually found on foliage, on branches or on tree trunks. They are similar in cheliceral teeth pattern to Emathis — promargin with two teeth and retromargin with a multi- cuspid fissident tooth — but can be easily distinguished by the presence of the median septum in the epigynum and the tegular lobe of the male palp. Petemathis differs from the other two genera in the same clade, Antillattus and Truncattus, by the following combination of characters: cheliceral retromargin with a multi-cuspid fissident tooth (ch. 21-3, ch. 23-4); male chelicera not elongate or with modifications, but usually with a depression on the posterior surface. 65! Truncattus (see Chapter 5 for detailed description) are small tree trunk dwelling spiders usually with one bicuspid retromarginal tooth on the chelicera (ch. 21-2, ch. 23-3) and a ventral bump on the male palpal tibia (ch. 48-1). The genus Antillattus is further discussed below. In addition, the two genera from Jamaica, Allodecta Bryant, 1950 (Bryant 1950: 166, figs. 1, 3, 8) and Caribattus Bryant, 1950 (Peckham & Peckham 1901: 10, figs. 4-4b; Bryant 1950: 177) may also belong to this clade based on similar morphology of body and genitalia. Antillattus Bryant, 1943 Antillattus Bryant, 1943: 461, type species: Antillattus gracilis Bryant, 1943. Diagnosis. The retromargin of the female chelicera has a unident tooth or a fissident tooth of more than two cusps. The male chelicera is elongate or has modifications such as projections or a flat front surface. The male endites are usually enlarged distally. Vulva usually has a pair of secondary spermathecae in addition to the primary spermathecae. Antillattus are medium sized spiders. Some species are relatively enlogate. Most species are found by beating foliage except A. applanatus, which lives on pine tree trunks (see Chapter 5). The following species are transferred to Antillattus here based on the features above and the molecular phylogeny: Antillattus darlingtoni (Bryant, 1943) (from Pensacola, New Combination, Fig. 3.9A-H) Antillattus electus (Bryant, 1943) (from Pensacola, New Combination) Antillattus maxillosus (Bryant, 1943) (from Pensacola, New Combination) Antillattus montanus (Bryant, 1943) (from Pensacola, New Combination) Antillattus peckhami (Bryant, 1943) (from Pensacola, New Combination) Antillattus cambridgei (Bryant, 1943) (from Cobanus, New Combination, Fig. 3.8H-N) Antillattus scutiformis New Name for Siloca electa Bryant, 1943, required by its being a junior secondary homonym because of its transfer to Antillatus, resulting in the New Combination Antillattus electus (Fig. 3.9I-P) Antillattus cubensis (Franganillo, 1935) (from Agobardus, New Combination) Antillattus keyserlingi (Bryant, 1940) (from Agobardus, New Combination) Antillattus mandibulatus (Bryant, 1940) (from Agobardus, New Combination) 66! Notes. The results show that the two Caribbean Pensacola species, P. maxillosa Bryant and P. darlingtoni Bryant do not fall into the same clade as the type species P. signata Peckham & Peckham from Guatemala. Instead they fall with two Antillattus species. Thus, here I transfer the Caribbean species described as Pensacola into Antillattus. Similarly, Siloca electa Bryant from Hispaniola also falls into the same clade with Antillattus rather than with other Siloca species from South America. Cobanus cambridgei (Bryant) was originally described as Amycus (Bryant 1943), and then Galiano (1968b) transferred it into Cobanus. The results from morphological and molecular phylogenetic studies indicate that it is neither an Amycus (Amycoida) nor a Cobanus, but should be placed in Antillattus. Transfering into Antillattus also removes it as a homonym with Cobanus cambridgei Chickering, 1946. In addition, three species described as Agobardus from Cuba (A. cubensis (Franganillo), A. keyserlingi Bryant, A. mandibulatus Bryant) are also transferred to Antillattus, because they have a multi-cuspid retromarginal tooth on the female chelicera and the male chelicerae are elongate, which do not fit in the delimitation of Agobardus, but are congruent with typical Antillattus. 3.5.2.2.3 Agobardus Clade: Agobardus, Bythocrotus, Compsodecta, Parasaitis (Figs 3.12- 3.14) Spiders of this clade are usually relatively robust. The embolus of the male palp coils for no more than one circle with a distinct embolic disc; the tegulum lacks a proximal lobe (ch. 47-0); The RTA is simple and finger-like. The epigynum has an obvious window structure with a median septum. The spermatheca is swollen. Four genera, Agobardus, Bythocrotus, Compsodecta and Parasaitis are included in this clade. Bythocrotus Simon can be easily distinguished from other euophryine genera by the broad cephalothorax, the swollen male palp, the small palpal bulb, the presence of a prolateral tibial apophysis (ch. 61-1) and numerous macrosetae on the male palpal tibia (ch. 62-1). Parasaitis Bryant, 1950 (197, figs. 30-32) from Jamaica was not sampled in the molecular phylogenetic analysis, but can also be included in this clade because it shows similarity in the epigynal form and the male palpal structure to Compsodecta. As indicated by Bryant (1950), Parasaitis has no tooth on both margins of the chelicera, whereas Compsodecta has two teeth on the promargin and one tooth on the retromargin. Agobardus and Compsodecta are further discussed below. 67! Agobardus Keyserling, 1885 Agobardus Keyserling, 1885: 519, type species: Agobardus anormalis Keyserling, 1885. Diagnosis. Spiders of Agobardus share the following characters: female chelicera with two promarginal teeth and one bicuspid retromarginal tooth (ch. 21-2); female and male first metatarsus with three pairs of ventral macrosetae (ch. 40-2, ch. 42-3); retrolateral sperm duct loop occupying at least half of the bulb width (ch. 58-2). The male chelicera of some species is relatively enlarged with modifications. The epigynal window is large or relatively small. The vulva lacks secondary spermathecae. Small to medium sized spiders with various habitats, such as on foliage, on ground (on leaf litter, on rocks or in grass clumps) and on tree trunks (see Chapter 5). According to the characters above, the following species are transferred to Agobardus here: Agobardus cubanus (Bryant, 1940) (from Siloca, New Combination) Agobardus minutus (Bryant, 1940) (from Siloca, New Combination) Agobardus modestus (Bryant, 1943) (from Commoris, New Combination) Notes. After comparing the descriptions and illustrations provided in the literature (Bryant 1940), two species originally described as Siloca (S. cubana Bryant and S. minuta Bryant) from Cuba are transferred to Agobardus. The type specimen of Commoris modesta Bryant from Hispaniola (in MCZ, #22130) was examined. I determine that it is not congeneric with the other two Commoris species including the type species, Commoris enoplognatha Simon (Galiano 1963: 329, pl. XV, figs. 4-6). Here I transfer it to Agobardus based on its similar genitalic and cheliceral forms with other Agobardus species. Compsodecta Simon, 1903 Compsodecta Simon, 1903: 678, type species: Cybele grisea Peckham & Peckham, 1901. Paradecta Bryant, 1950: 192, type species: Paradecta festiva Bryant, 1950. New Synonym Diagnosis. Compsodecta usually have enlarged male endites and modifications (parallel transverse ridges or flat surfaces or projections) on the front surface of the male chelicera. The male palp often has additional modifications (small projection, wide extension or flat surface) 68! on the tibia, patella and/or femur. Compsodecta are medium sized spiders usually found on foliage. According to the features above, the following species are transferred to Compsodecta here: Compsodecta darlingtoni (Bryant, 1950) (from Paradecta, New Combination) Compsodecta festiva (Bryant, 1950) (from Paradecta, New Combination, Fig. 3.14E-I) Compsodecta gratiosa (Bryant, 1950) (from Paradecta, New Combination) Compsodecta valida (Bryant, 1950) (from Paradecta, New Combination) Notes. After examining the type specimens of Paradecta festiva Bryant (in MCZ, #21302, 25818), the type species of Paradecta Bryant from Jamaica, and studying the morphology of other Paradecta species from the literature (Bryant 1950), I find their male palpal and cheliceral structures and female epigynal form show the same pattern as those of Compsodecta. Thus, Paradecta is considered as a junior synonym of Compsodecta here, and the Paradecta species are transferred to Compsodecta. 3.5.2.2.4 Naphrys-Corticattus Clade (Fig 3.15) The grouping of the Caribbean genus Corticattus Zhang & Maddison (see Chapter 5 for detailed description) with the North American genus Naphrys Edwards, 2002 (69, figs. 5-8) was unexpected but well supported by molecular phylogeny. Spiders in this clade are small sized and usually dull in color. They often have a small anterior-lateral cusp on the male endite (ch. 9-1); one bicuspid promarginal tooth on the chelicera (ch. 20-2, ch. 22-3); no more than two pairs of ventral macrosetae on the first tibia (ch. 38-1, ch. 39-1); and a ventral bump on the male palpal tibia (ch. 48-1). Their male palpal bulb is usually large with a proximal tegular lobe; the RTA is simple and finger-like. The epigynum has typical window structure with a median septum. Corticattus is found on tree trunks or by beating branches and has a body more or less flattened (see Chapter 5), while Naphrys is usually associated with leaf litter and has a high carapace. Also their genitalia are quite different: Corticattus has no retrolateral sperm duct loop but has a prolateral sperm duct loop on the male palpal bulb, while Naphrys has the retrolateral sperm duct loop but has no prolateral sperm duct loop. Corticattus has a triangular projection at the distal end of the tegulum, while this distal tegular projection is absent in Naphrys. Corticattus 69! usually has relatively long copulatory duct but very small spermatheca, while Naphrys often has very short copulatory duct and large spermatheca. 3.5.2.2.5 Sidusa Clade (Fig 3.16) Sidusa Peckham & Peckham, 1895 Sidusa Peckham & Peckham, 1895: 175, type species: Sidusa gratiosa Peckham & Peckham, 1895. Cobanus F. O. P.-Cambridge, 1900: 190, type species: Helorus extensus Peckham & Peckham, 1896. New Synonym Chloridusa Simon, 1902: 388, type species: Chloridusa viridiaurea Simon, 1902. New Synonym Wallaba Mello-Leitão, 1940: 191, type species: Wallaba metallica Mello-Leitão, 1940. New Synonym Diagnosis. Some females have paired dark markings on the dorsum of the abdomen, and most species, especially the males, are covered with iridescent green scales. They usually have two promarginal teeth and one bicuspid retromarginal tooth (ch. 21-2, ch. 23-3) on the chelicera. Some species have elongate chelicerae in males. First leg usually has four pairs of ventral macrosetae on the tibia (ch. 38-4, ch. 39-4) and three pairs on the metatarsus (ch. 40-2, ch. 41-3). Many species have a prespiracular bump on the venter of the male abdomen. The male palpal bulb is usually small without the proximal tegular lobe (ch. 47-0); the retrolateral sperm duct loop usually occupies less than half of the bulb width (ch. 58-0). Many species have a lamella beside the embolus on the male palp. The epigynal window is usually large with a median septum. Spiders of Sidusa are medium sized, mainly found on foliage with only a few species known living on tree trunks. The following species are transferred to Sidusa here based on the characters above and the molecular phylogeny: Sidusa beebei (Petrunkevitch, 1914) (from Cobanus, New Combination) Sidusa bifurcata (Chickering, 1946) (from Cobanus, New Combination) Sidusa cambridgei (Chickering, 1946) (from Cobanus, New Combination) Sidusa electa (Chickering, 1946) (from Cobanus, New Combination) Sidusa erythrocras (Chamberlin & Ivie, 1936) (from Cobanus, New Combination) 70! Sidusa extensa (Peckham & Peckham, 1896) (from Cobanus, New Combination, Fig. 3.16A-G) Sidusa flavens (Peckham & Peckham, 1896) (from Cobanus, New Combination) Sidusa guianensis (Caporiacco, 1947) (from Cobanus, New Combination) Sidusa incurva (Chickering, 1946) (from Cobanus, New Combination) Sidusa mandibularis (Peckham & Peckham, 1896) (from Cobanus, New Combination) Sidusa obscura (Chickering, 1946) (from Cobanus, New Combination) Sidusa perdita (Banks, 1898) (from Cobanus, New Combination) Sidusa scintillans (Crane, 1943) (from Cobanus, New Combination) Sidusa seclusa (Chickering, 1946) (from Cobanus, New Combination) Sidusa subfusca (F. O. P.-Cambridge, 1900) (from Cobanus, New Combination) Sidusa unicolor (F. O. P.-Cambridge, 1900) (from Cobanus, New Combination) Sidusa viridiaurea (Simon, 1902) (from Chloridusa, New Combination) Sidusa albopalpis (Peckham & Peckham, 1901) (from Wallaba, New Combination) Sidusa metallica (Mello-Leitão, 1940) (from Wallaba, New Combination) Notes. In the molecular phylogeny with full taxa, eight species of Cobanus and two species of Sidusa were included (see Chapter 2), and Cobanus is paraphyletic with Sidusa embedded within. One morphological character that has been used to distinguish these two genera is the male chelicera, elongate in Cobanus, not in Sidusa (Bodner 2002). Nevertheless, the length of male chelicera is quite variable both interspecifically and intraspecifically in Cobanus (Bodner 2002). Based on the phylogenetic results, Cobanus is considered a junior synonym of Sidusa. In addition, type specimens of Chloridusa viridiaurea Simon (in MNHN, #7720; type species of Chloridusa Simon) and Wallaba metallica Mello-Leitão (in NHM, #1930.12.17.117; type species of Wallaba Mello-Leitão) were examined. They both share the cheliceral teeth pattern, the ventral macrosetae pattern on the first tibia and metatarsus, and the genitalic form of Sidusa. Therefore, both Chloridusa and Wallaba are considered as junior synonyms of Sidusa. One species of Wallaba, W. decora has been moved to Corythalia (see discussion under the Anasaitis-Corythalia Clade). The other described species, W. albopalpis (Peckham & Peckham, 1901: 12, figs. 3-3b) is known from Jamaica and further study is still needed to clarify if it also belongs to Sidusa. 71! 3.5.2.2.6 Mopiopia-Saphrys Clade (Fig 3.17) A well-supported clade in the molecular phylogeny includes Mopiopia and the new genus Saphrys, although no unambiguous synapomophy is resolved in the character optimization. Spiders in this clade are small to medium sized. The endite usually has a small anterior-lateral cusp. The chelicera usually has two promarginal teeth and one retromarginal tooth. The femur of the male palp usually has two dorsal macrosetae. The male palp has a proximal tegular lobe and a relatively narrow embolic spiral. The RTA is simple and finger-like. Mopiopia Simon, 1902 Mopiopia Simon, 1902: 375, type species: Mopiopia comatula Simon, 1902. Tariona Simon, 1902: 383, type species: Tariona gounellei Simon, 1902. New Synonym Diagnosis. Spiders of Mopiopia are medium sized and usually relatively dark in color. The male palpal bulb is relatively narrow with a wide retrolateral sperm duct loop occupying more than half of the bulb width. The epigynum has typical window structure with a median septum. The spermatheca is relatively large and swollen. The following species are transferred to Mopiopia here: Mopiopia albibarbis (Mello-Leitão, 1947) (from Tariona, New Combination) Mopiopia bruneti (Simon, 1903) (from Tariona, New Combination, Fig. 3.17A-D) Mopiopia gounellei (Simon, 1902) (from Tariona, New Combination) Mopiopia maculata (Franganillo, 1930) (from Tariona, New Combination) Mopiopia mutica (Simon, 1903) (from Tariona, New Combination) Notes. Tariona Simon is mainly found from Brazil (Platnick 2011) with all the known Brazilian species, including the type species, T. gounellei Simon, only known from the male. Mopiopia Simon (Galiano 1963; Edwards et al. 2005), another genus from Brazil, shares with Tariona the palpal features mentioned in the diagnosis. Therefore, here I consider Tariona as a junior synonym of Mopiopia. However, whether the species from Cuba, T. maculata Franganillo also belongs to Mopiopia as the other species is uncertain due to the poor documentation. 72! Saphrys New Genus Type species: Euophrys tehuelche Galiano, 1968a. Etymology. The generic name consists of two parts: the first part sa refers to the South America, where the species were found; the second part phrys is from Euophrys, the genus in which the species were originally described; feminine in gender. Diagnosis. Spiders of Saphrys are relatively small and usually brownish or dark brown in color with foliate markings on the dorsal abdomen. The chelicera has two promarginal teeth and one retrolateral tooth. The fourth leg of the male is usually longest. The males of most species have a relatively hard area on the anterior part of the dorsum of the abdomen (ch. 42-1). The male palpal bulb is usually wide with a large proximal tegular lobe and a short and slightly curved embolus. The palpal tibia often has a ventral bump (ch. 48-1) and a long and finger-like RTA. The epigynal window is usually small. Saphrys differs from Euophrys by the wide palpal bulb, the much smaller embolic spiral, and the epigynum, which usually has the openings to the copulatory ducts further away from each other. Saphrys is similar to Mopiopia (Galiano 1963; Edwards et al. 2005) in the relatively small embolic spiral, but can be distinguished by the wide palpal bulb, the shorter embolus and the epigynal form. The following nine species are placed in this genus including the type species based on their similar genitalic and somatic morphology (Galiano 1968a; Richardson 2010). The other neotropical Euophrys species are also likely misplaced. However, their position is still uncertain. Saphrys a-notata (Mello-Leitão, 1940) (from Euophrys, New Combination) Saphrys flordellago (Richardson, 2010) (from Euophrys, New Combination) Saphrys laetata (Simon, 1904) (from Euophrys, New Combination) Saphrys mapuche (Galiano, 1968) (from Euophrys, New Combination, Fig. 3.17K-Q) Saphrys patagonica (Simon, 1905) (from Euophrys, New Combination) Saphrys rapida (C. L. Koch, 1846) (from Euophrys, New Combination) Saphrys rusticana (Nicolet, 1849) (from Euophrys, New Combination) Saphrys saitiformis (Simon, 1901) (from Euophrys, New Combination) Saphrys tehuelche (Galiano, 1968) (from Euophrys, New Combination, Fig. 3.17E-J) 73! Notes. Two South American species described as Euophrys, “Euophrys” a-notata Mello-Leitao and “Euophrys” cf. patagonica Simon were included in the phylogenetic study. The results show that they are not monophyletic with the Holarctic Euophrys species, but fall in a distinct clade. This confirms Logunov’s concern that the South American “Euophrys” species may belong to a distinct genus other than Euophrys C. L. Koch (see Logunov 1997). Hence, a new genus is erected here. 3.5.2.2.7 Chapoda-Maeota Clade: Chapoda, Maeota, Commoris. (Figs 3.18-3.19) Medium sized spiders. The chelicera usually has two promarginal teeth. The fourth leg of the male is usually longest. The male palpal bulb is usually relatively small with a short and slightly curved embolus. The RTA is finger-like. The epigynum has typical window structure with a median septum. A well-supported clade in the molecular phylogeny, including Chapoda and Maeota. Commoris Simon 1902 (see Galiano, 1963: 329, pl. XV, figs. 4-6) is also placed in the clade based on the similar male palpal structure. Chapoda Peckham & Peckham, 1896 Chapoda Peckham & Peckham, 1896: 26, type species: Chapoda festiva Peckham & Peckham, 1896. Diagnosis. Some species have a marble-white guanine deposit in the eye area. The chelicera has two promarginal teeth and one unident or bicuspid retromarginal tooth. The male palpal bulb is relatively small. The proximal tegular lobe is present or absent. The male palpal tibia has a ventral bump (ch. 48-1). The male palpal femur of some species has a ventral projection. The epigynal window is relatively small with a median septum. The beginning of the copulatory duct is swollen and becomes the secondary spermatheca (ch. 76-1). Chapoda are medium sized spiders usually found on tree trunks, branches or foliage. The following species are transferred to Chapoda here based on the features above and the molecular phylogeny: Chapoda recondita (Peckham & Peckham, 1896) (from Sidusa, New Combination) Chapoda maxillosa (F. O. P.-Cambridge, 1901) (from Compsodecta, New Combination) 74! Chapoda montana (Chickering, 1946) (from Compsodecta, New Combination) Notes. Typical Chapoda species, such as C. festiva Peckham & Peckham, have guanine deposits in the eye area and a swollen male palp, usually with a projection on the femur. Some species in this study lack some or all of the above characters, e.g. C. fortuna has no guanine deposit in the eye area and C. angusta lacks an obvious projection on the femur of the male palp (see Chapter 7). However, the results from the molecular phylogeny indicate that they are closely related to typical Chapoda species, and the other features of their genitalic organs are very similar to the typical Chapoda species. Rather than erecting new genera for these species and so adding to the glut of Salticidae genera, I extend the delimitation of Chapoda to comprise more species (see Chapter 7). The results show that “Sidusa” recondita Peckham & Peckham is not closely related with the other Sidusa species, and here I transfer it to Chapoda. Compsodecta maxillosa (F. O. P.- Cambridge) and C. montana Chickering are also transferred to Chapoda based on their similar genitalic structure. Maeota Simon, 1901 Maeota Simon, 1901a: 567, type species: Maeota dichrura Simon, 1901. Diagnosis. Almost all species currently placed in Maeota (see Chapter 7) share the character that the plane of the embolic spiral is perpendicular to the longitudinal axis of the bulb (ch. 64- 1), except in Maeota sp. [Manabi] (JXZ058). While these species are generally similar with a boxy carapace and somewhat delicate legs, a precise morphological delimitation is not yet clear; their placement in Maeota is based on the molecular phylogeny. The following species is transferred to Maeota here based on the features above and the molecular phylogeny: Maeota tuberculotibiata (Caporiacco, 1955) (from Pensacola, New Combination) Notes. The genus Maeota was monotypic with the type species, M. dichrura Simon, well known for the extremely long posterior lateral spinneret in the male. However, this study discovered quite a few species that are closely related to M. dichrura but only have normal posterior lateral 75! spinneret in the male (see Chapters 2 and 7). Hence, the abnormally long posterior lateral spinneret of male can be considered an autapomorphy of M. dichrura rather than the defining synapomorphy of the genus Maeota, and it makes more sense to extend the definition of Maeota to include more species. The results of molecular phylogeny reveal that Pensacola tuberculotibiata Caporiacco (Ruiz & Brescovit 2005: 756, figs. 11-12) is misplaced and here I transfer it to Maeota. 3.5.2.2.8 Amphidraus-Marma Clade (Fig 3.20) Spiders of this clade are small to medium sized. They usually have two promarginal teeth and one fissident retromarginal tooth of more than two cusps (ch. 21-3, ch. 23-4). The embolic disc has a large apophysis (\"PED\" in Fig. 3.20C, I) independent from the embolus on the male palp, and which might be mistaken for the embolus itself. The epigynum lacks a typical window structure (ch. 80-1). The morphological and molecular data confirm the suggestion by G. R. S. Ruiz (pers. comm.) that Marma and Amphidraus may be related. Although similar in the genitalic form to Amphidraus, Marma can be distinguished from it by the larger body; the absence of the tegular lobe, the shape of the apophysis of the embolic disc, the long embolus partly hidden beneath the bulb, the simple and finger-like RTA of the male palp. See following for discussion on Amphidraus. Amphidraus Simon, 1900 Amphidraus Simon, 1900: 60, type species: Amphidraus auriga Simon, 1900. Nebridia Simon, 1902: 373, type species: Nebridia semicana Simon, 1902. New Synonym Diagnosis. Some species are lighter in color with foliate markings on the dorsum of the abdomen; some species are dark with some small light colored spots and chevron-like markings. The male palpal bulb is large with a distinct proximal tegular lobe. The RTA is large and complex with multiple branches. Spiders of Amphidraus are small leaf litter or tree trunk dwelling spiders. According to the features above and the molecular phylogeny (see Chapter 2), the following species are transferred to Amphidraus here: 76! Amphidraus manni (Bryant, 1943) (from Nebridia, New Combination) Amphidraus mendicus (Bryant, 1943) (from Nebridia, New Combination) Amphidraus parvus (Mello-Leitão, 1945) (from Nebridia, New Combination) Amphidraus semicanus (Simon, 1902) (from Nebridia, New Combination) Notes. Here I consider Nebridia Simon (Galiano 1963: 401, pl. XXVII, figs. 15-18) as a junior synonym of Amphidraus because they share similar characters such as the complex RTA and the distinct tegular lobe over the tibia of the male palp. However, the two species described as Nebridia from Hispaniola, N. manni Bryant, 1943 and N. mendica Bryant, 1943, do not belong to Amphidraus; further study is needed to clarify the generic placement of these two species. 3.5.2.2.9 Coryphasia Clade: Coryphasia and possibly Semnolius. (Fig 3.21) This clade contains the genus Coryphasia. Semnolius may also belong to this clade based on its similar body form and male palp. Coryphasia Simon, 1902 Coryphasia Simon, 1902: 380, type species: Coryphasia albibarbis Simon, 1902. Asaphobelis Simon, 1902: 384, type species: Asaphobelis physonychus Simon, 1902. New Synonym Siloca Simon, 1902: 389, type species: Siloca sanguiniceps Simon, 1902. New Synonym Diagnosis. The chelicera has two promarginal teeth and one bicuspid retromarginal tooth (ch. 23-3). The male chelicera often has a longitudinal ridge on the front surface (ch. 25-1). The male palp is relatively large with a distinctive tegular lobe. The embolus is usually long and highly spiraled, except in C. physonycha (Simon). The retrolateral sperm duct loop is relatively wide. The RTA is finger-like. The epigynum has a typical window structure with a median septum. The vulva of many species has a pair of secondary spermathecae in addition to the primary spermathecae. Spiders of Coryphasia are medium sized, usually with foliate markings on the dorsum of the abdomen. The following species are transferred to Coryphasia here based on the characters above and the molecular phylogeny: Coryphasia physonycha (Simon, 1902) (from Asaphobelis, New Combination, Fig. 3.21H-O) 77! Coryphasia bulbosa (Tullgren, 1905) (from Siloca, New Combination) Coryphasia campestrata (Simon, 1902) (from Siloca, New Combination, Fig. 3.21P-V) Coryphasia monae (Petrunkevitch, 1930) (from Siloca, New Combination) Coryphasia sanguiniceps (Simon, 1902) (from Siloca, New Combination) Coryphasia septentrionalis (Caporiacco, 1954) (from Siloca, New Combination) Coryphasia viaria (Peckham & Peckham, 1901) (from Siloca, New Combination) Notes. Coryphasia, Asaphobelis and the South American species of Siloca including the type, S. sanguiniceps Simon, are similar in body form and cheliceral teeth pattern. The embolus of Coryphasia and Siloca is usually long with wide spiral, while that of Asaphobelis is short and stout. Hence, Edwards et al. (2005) concluded that Asaphobelis is a valid genus. However, this study suggests a relationship of (Coryphasia (Siloca, Asaphobelis)), and indicates the short and stout embolus is just an autapomorphy for the species Asaphobelis physonychus Simon (Galiano 1963: 298, pl. VIII, figs. 1-4). To eliminate unnecessary monotypic genera in Euophryinae, here I merge these three genera and consider both Asaphobelis and Siloca as junior synonyms of Coryphasia. However, transferring Siloca bulbosa Tullgren (Argentina) and S. septentrionalis Caporiacco (French Guiana) into Coryphasia is less certain because of the poor documentation of these two species. In addition, the two species described as Siloca from the Caribbean Islands, S. monae Petrunkevitch (Puerto Rico; Petrunkevitch 1930: 151, figs. 135-139) and S. viaria (Peckham & Peckham) (Jamaica; Peckham & Peckham 1901: 14, pl. 4, fig. 11, as Prostheclina) may not be congeneric with the type species of Siloca, but their taxonomic position still needs to be further investiagted. Semnolius Simon, 1902 Semnolius Simon, 1902: 369, type species: Semnolius chrysotrichus Simon, 1902. Ocnotelus Simon, 1902: 382, type species: Ocnotelus imberbis Simon, 1902. New Synonym Diagnosis. Semnolius Simon (Galiano 1963: 443, pl. XXXVII, figs. 10-11) is similar to Coryphasia in body form, cheliceral teeth pattern and male palpal structure, but differs in the presence of a lamella beside the embolus of the male palp. The following species are transferred to Semnolius based on the characters above: Semnolius imberbis (Simon, 1902) (from Ocnotelus, New Combination) 78! Semnolius lunatus (Mello-Leitão, 1947) (from Ocnotelus, New Combination) Semnolius rubrolunatus (Mello-Leitão, 1945) (from Ocnotelus, New Combination) Notes. Ocnotelus Simon (Galiano 1963: 415, pl. XXXI, figs. 1-4) is considered as a junior synonym of Semnolius because their type species have extremely similar male palpal structure. However, whether Ocnotelus lunatus Mello-Leitão and O. rubrolunatus Mello-Leitão also belong to Semnolius as the type species O. imberbis Simon is uncertain due to the poor documentation of these species. 3.5.2.2.10 Pensacola-Mexigonus Clade (Fig 3.22) Pensacola and Mexigonus are very different in both body form and genitalic structure. However, the molecular phylogeny suggests they are more closely related to each other than to other euophryine genera. They share certain characters such as the teeth pattern on the chelicera (two teeth on the promargin and one unident tooth on the retromargin), the finger-like RTA of the male palp and the large window of the epigynum. Nevertheless, no unambiguous morphological synapomorphy is resolved for this clade from the character optimization. 3.5.2.2.11 Neonella Clade: Neonella and Darwinneon (Fig 3.23) Of these two genera in this clade, only Neonella was included in the molecular analyses. However, the similar morphology (see below) of Darwinneon (see Cutler 1971: 511, figs. 1-3) with Neonella indicates they are closely related. Spiders of this clade are very small and usually found in leaf litter. The fovea on the carapace is absent. The male abdomen sometimes has a relatively hard area on the dorsum. The male palp has a distinctive proximal tegular lobe, but the retrolateral sperm duct loop is absent. The RTA is often short. The copulatory duct is usually short and wide. In Neonella the embolic disc is highly reduced and the embolus has an accompanying lamella of various shapes. 3.5.2.2.12 Belliena Clade: Belliena, Saitidops and possibly Stoidis (Fig 3.24) Spiders in this clade are also small. Belliena was sampled in the molecular analyses and the results of molecular phylogeny indicate it may be related to Ilargus and Neonella. Belliena species sometimes have a slightly sclerotized area on the dorsal abdomen of the male and a large prespiracular bump on the venter of the male abdomen. The genitalia are typical for euophryines: 79! the male palp has a distal and coiled embolus, a proximal tegular lobe and a retrolateral sperm duct loop; the epigynum has obvious window structure with a median septum. Inclusion of Saitidops in this clade is based on its similar male palpal shape with Belliena (Galiano 1963: 430, pl. XXXV, figs. 1-3). However, the placement of Stoidis in this clade is less certain. 3.5.2.2.13 Soesilarishius Clade: Soesilarishius and possibly Rhyphelia (Fig 3.25) Three Soesilarishius species, including the type species S. amrishi Makhan, 2007, were included in molecular phylogenetic analyses. The genus has most recently been reviewed by Ruiz (2011). They are small euophryine spiders known from leaf litter. The male abdomen sometimes has a relatively sclerotized area on the dorsum (ch. 42-1). The embolus of the male palp is usually not coiled (ch. 46-0) and the distal hematodocha is sometimes reduced (ch. 70-1). The epigynum usually lacks a distinct window but has a groove or a transverse roof (see Chapter 7). However, the species S. crispiventer Ruiz described from Brazil (Ruiz 2011: 26, figs. 8-14, 47-48, 64-65) has the embolus coiling anti-clockwise (left palp ventral view). Whether it is a derived condition within Soesilarishius or not needs further investigation. The male palpal shape of Rhyphelia Simon (Galiano, 1963: 429, pl. XXXIII, figs. 10-12) indicates it may be closely related to Soesilarishius. 3.5.2.2.14 Other genera from the New World: Ecuadattus, Ilargus, Popcornella, Tylogonus (Figs. 3.26-3.29) No clear groupings of these genera are indicated by the results. Ecuadattus is a newly discovered genus from Ecuador (see Chapter 7 for a detailed description). They have typical euophryine appearance, but the carapace usually has guanine deposits in the eye area. The abdomen usually has foliate markings. The female chelicera has two promarginal teeth and one retromarginal tooth. The embolic disc of the male palp is small or highly reduced. The embolus is short and slightly curved. The retrolateral sperm duct loop is narrow. The RTA is long and finger-like. The epigynum has the window with a median septum. The spermatheca is oval or round. Spiders of this genus are medium sized and usually found on foliage. 80! The results also reveal a lineage with a number of species related to Ilargus coccineus Simon, the type species of Ilargus. Some species are light colored with longitudinal stripes; some species have foliage-like markings on the dorsal abdomen; some species have the marble- appearance with guanine deposit in the eye area; some species are dark brown or brown with many light colored spots and chevron-like markings on the dorsal abdomen. The chelicera has two promarginal teeth and one retromarginal tooth. The male palpal bulb is relatively large with a big tegular lobe. The embolus is long with a wide spiral. The RTA is usually finger-like. The epigynal window is large (ch. 79-2) with a median septum. Spiders of Ilargus are medium sized and usually found on foliage (see Chapter 7 for detailed description). Popcornella is a newly discovered genus endemic to the Caribbean Islands (see Chapter 7 for a detailed description). The body is brown or dark brown in color without iridescent scales. Leg lacks fringe. The embolus of male palp is short and uncoiled with the embolic disc reduced or absent. The distal hematodocha is also reduced. The RTA is simple and small. The epigynum lacks distinct window structure. The copulatory duct of the vulva is usually very short. They are small euophryines usually found in leaf litter. Tylogonus is an unusual euophryine genus whose males have a body shape and longitudinal white stripes that make them appear, at first glance, to be dendryphantines. The genitalic structures are also different from the typical euophryine form: the embolus is fixed but not free, and the epigynal window is not obvious. The male palp has the retrolateral sperm duct loop on the tegulum like most of other euophryine genera. The male chelicera is usually modified with a projection on the front surface and a concaved mesal margin. The RTA is usually more complex but not finger-like. 3.5.2.3 Major euophryine clades from the Old World 3.5.2.3.1 Bathippus-Canama Clade: Bathippus, Canama and possibly Spilargis (Fig 3.30) This clade includes Bathippus, Canama and probably also Spilargis. The body is usually elongate with the female often having the third leg longest (ch. 31-0). Many species have elongate chelicerae in males. The promargin of the female chelicera usually has two teeth. The first tibia usually has three pairs of vental macrosetae. The male palp is usually elongate with a relatively small bulb and curved femur. The embolus coils with the plane of the spiral more or 81! less perpendicular to the longitudinal axis of the bulb. The epigynum has typical window structure with a median septum. They are usually medium sized foliage dwelling spiders. There has been a long debate on whether Bathippus and Canama should be merged or both of them are valid. Prószy!ski (1987) considered Canama as a junior synonym of Bathippus based on their similarities. However, Davies and Zabka (1989) disagreed and suggested that they differ in chelicerae and epigynum structure. The phylogenetic results indicate both Bathippus and Canama are monophyletic, and thus support the argument in Davies and Zabka (1989) that these two genera are both valid. They differ from each other as follows: Canama usually has a bicuspid retromarginal tooth on the female chelicera, while Bathippus usually has one unident retromarginal tooth (ch. 21-1); the embolus of Canama usually coils more than half a circle, while in Bathippus, the embolus coils no more than half a circle (ch. 46-1); the opening to the copulatory duct of Canama is usually posterior to the vulva (with exception of C. hinnulea, see Davies & Zabka, 1989: 220, pl. 29), while in Bathippus the opening is anterior or medial to the vulva; the primary spermatheca in Canama is usually narrow and coiled, not distinctive from the copulatory duct, while in Bathippus, the primary spermatheca is usually swollen and ovoid or kidney-shaped. 3.5.2.3.2 Omoedus Clade (Fig 3.31) Omoedus Thorell, 1881 Omoedus Thorell, 1881: 669, type species: Omoedus niger Thorell, 1881. Pystira Simon, 1901a: 656, type species: Pystira ephippigera (Simon, 1885). Zenodorus Peckham & Peckham, 1886: 297, type species: Zenodorus durvillei (Walckenaer, 1837). Diagnosis. Spiders of Omoedus are usually robust with various color patterns (see Chapter 6). The male palpal bulb usually lacks a proximal tegular lobe. The embolus of most species is highly spiraled and coils for more than one and a half circle. The retrolateral sperm duct loop is more than half of the bulb width. The RTA is finger-like. The epigynal window is present with a medium septum. The copulatory duct is usually extremely long and highly convoluted. The primary spermatheca is as small as the copulatory duct, and narrow and coiled. They are small to medium sized and can be found in various microhabitats, including leaf litter and foliage. 82! According to the characters above and the molecular phylogeny, the following species are transferred to Omoedus here: Omoedus insultans (Thorell, 1881) (from Margaromma, New Combination) Omoedus kochi (Simon, 1901) (from Margaromma, New Combination) Omoedus semirasus (Keyserling, 1882) (from Margaromma, New Combination) Omoedus sexualis (Strand, 1911) (from Margaromma, New Combination) Omoedus torquatus (Simon, 1902) (from Margaromma, New Combination) Notes. The monophyly of Omoedus is strongly supported by the molecular phylogeny. The morphological and molecular results support merging Pystira and Zenodorus with Omoedus (see Chapter 6). Although the carapace of Omoedus and Pystira is usually much higher and has a concavity at the posterior end, their genitalic organs share the same pattern as those of Zenodorus. Although the genus Margaromma is distinct from Omoedus based on the diagnostic drawings provided by Davies and Zabka (1989: 230, pl. 38) for the type species Margaromma funestum Keyserling, other species currently in Margaromma have genitalia matching Omoedus. Based on the genitalia, they are transferred to Omoedus here. 3.5.2.3.3 Bulolia-Coccorchestes Clade: Bulolia, Coccorchestes, Leptathamas, Variratina and possibly Athamas (Figs 3.32-3.33) Spiders in this clade are quite diverse in the body form. Leptathamas has a high carapace with a hump at the posterior slope bearing small projections, and when alive they usually hold strange poses and walk in jerky fashion, appearing to be piles of debris (Maddison & Zhang 2011). Coccorchestes is well-known as a beetle mimic. However, the male palps of these genera share the following characters: palpal femur with no or only one dorsal macroseta (ch. 51-0/1); retrolateral sperm duct loop absent or reduced with its width much narrower than half of bulb width; prolateral sperm duct loop present (ch. 57-1); embolus coiled with the plane of the spiral perpendicular to the longitudinal axis of bulb. Athamas is similar to Bulolia and Leptathamas in the four-row eye arrangement with ALEs posterior to the AMEs and forming their own row, and may also belong to this clade (see comment in Szüts 2003). However, their palpi lack the proximal or prolateral sperm duct loop like other members in this clade. The four–row eye arrangement also appears in Furculattus, a genus related to Diolenius based on genitalic structures (Wanless & Lubin 1986). This suggests 83! the four eye rows arrangement has evolved more than once independently within Euophryinae. The molecular data are ambiguous on the placement of Athamas on phylogeny because of its strange long branch in 28S analysis (see Chapter 2). Thus, the placement of Athamas in this clade is still uncertain. 3.5.2.3.4 Diolenius Clade: Chalcolecta, Chalcolemia, Diolenius, Efate, Furculattus, Ohilimia, Paraharmochirus, Sobasina, Tarodes, Udvardya (Figs 3.34-3.36) This clade well-supported in the molecular phylogeny includes genera with peculiar body forms such as Diolenius and Paraharmochirus. In addition to the genera sampled for phylogenetic analyses (Chalcolecta, Chalcolemia, Diolenius, Efate, Ohilimnia, Paraharmochirus and Sobasina), Furculattus, Tarodes and Udvardya are also included in this clade based on their body forms and genitalic structures. Spiders of this clade are elongate and relatively flattened (ch. 3-1). Many species lack a longitudinal fovea on the carapace (ch. 15-1). The first leg is usually very long with its tibia having no less than 5 pairs of ventral macrosetae (ch. 38-6). Some species have extremely long trochanters of the first leg (e.g. Diolenius and Ohilimia, see Gardzinska & Zabka 2006; Gardzinska 2006); some are more or less ant-like (e.g. species of the genera Efate, Paraharmochirus and Sobasina). They also show some abnormal palp forms, which are distinctive from the typical euophryines, such as the proximal position of the embolus in Diolenius (see Gardzinska & Zabka 2006) and Furculattus (see Szüts 2003), and the absence of the retrolateral sperm duct loop in Chalcolecta (see Gardzinska & Zabka 2005) and Sobasina (see Wanless 1978). The epigynal window is present or absent (e.g. Ohilimia and Sobasina). The primary spermatheca is swollen or as narrow as the copulatory duct (e.g. Sobasina). 3.5.2.3.5 Pristobaeus Clade: Pristobaeus, Opisthoncana, Ergane and Ascyltus (Fig 3.37) Of this clade, only Pristobaeus was sampled in the analyses. An additional three genera, Opisthoncana, Ergane and Ascyltus are grouped in this clade based on their similar body forms and genitalic structures. The genus Pristobaeus is further discussed below. Pristobaeus Simon, 1902 Pristobaeus Simon, 1902: 391, type species: Pristobaeus jocosus Simon, 1902. Palpelius Simon, 1903: 735, type species: Plexippus beccarii Thorell, 1881. New Synonym 84! Diagnosis. Pristobaeus are medium sized spiders. They are relatively elongate, usually with foliate markings on the dorsum of the abdomen. The cheliceral promargin usually has two teeth, while the retromargin varies with a unident tooth in some species and multiple teeth in others. The male palp is relatively long with the tibia often longer than the patella dorsally (ch. 53-1) and the femur often slightly curved (ch. 50-1). The male palpal bulb is relatively narrower with a proximal tegular lobe (ch. 47-1). The embolus sometimes has an accompanying lamella (ch. 65-1). The epigynal window is present with a median septum. Following species are transferred to Pristobaeus here based on the features above and the molecular phylogeny: Pristobaeus albofasciatus (Peckham & Peckham, 1907) (from Palpelius, New Combination) Pristobaeus arboreus (Peckham & Peckham, 1907) (from Palpelius, New Combination) Pristobaeus beccarii (Thorell, 1881) (from Palpelius, New Combination, Fig. 3.37H-N) Pristobaeus clarus (Roewer, 1938) (from Palpelius, New Combination) Pristobaeus dearmatus (Thorell, 1881) (from Palpelius, New Combination) Pristobaeus discedens (Kulczy!ski, 1910) (from Palpelius, New Combination) Pristobaeus fuscoannulatus (Strand, 1911) (from Palpelius, New Combination) Pristobaeus kuekenthali (Pocock, 1897) (from Palpelius, New Combination) Pristobaeus namosi (Berry, Beatty & Prószy!ski, 1996) (from Palpelius, New Combination) Pristobaeus nemoralis (Peckham & Peckham, 1907) (from Palpelius, New Combination) Pristobaeus taveuniensis (Patoleta, 2008) (from Palpelius, New Combination) Pristobaeus trigyrus (Berry, Beatty & Prószy!ski, 1996) (from Palpelius, New Combination) Pristobaeus vanuaensis (Patoleta, 2008) (from Palpelius, New Combination) Pristobaeus vitiensis (Patoleta, 2008) (from Palpelius, New Combination) Notes. Although Pristobaeus has multiple teeth on the cheliceral retromargin, while Palpelius usually has a unident tooth on the retromargin, they are very similar to each other in body form, color pattern and genitalic structures. Thus, I follow Dr. Jerzy Prószy!ski’s suggestion (pers. comm.) of merging these two genera, and consider Palpelius as a junior synonym of Pristobaeus. Species of Palpelius are therefore transferred to Pristobaeus. 85! 3.5.2.3.6 Phasmolia Clade: Bindax, Lakarobius, Phasmolia, Araneotanna (Fig 3.38) Of this clade only Phasmolia was sampled for phylogenetic analyses. The additional three genera, Bindax, Lakarobius and Araneotanna, are included in the clade because of their similarity in body form to Phasmolia. Spiders of this clade are small to medium sized. They are usually very delicate with light transverse bands against a dark background on the abdomen (see Berry et al. 1998 for Lakarobius). They all have the epigynal window with an obvious median septum, and only one pair of spermathecae, which are swollen and distinguishable from the copulatory duct. The male palp usually has a distal and coiled embolus, which is similar to most of the euophryine genera. In Bindax and Lakarobius, the male palp has the proximal tegular lobe and the retrolateral sperm duct loop, whereas both the tegular lobe and the retrolateral sperm duct loop are absent in Phasmolia. The male palp of Araneotanna is still unknown. 3.5.2.3.7 Parabathippus-Parvattus Clade (Fig 3.39) The close relationship of Parabathippus and Parvattus is well supported by molecular phylogeny but unexpected because they are very different in both body form and genitalia. No unambiguous synapomorphy for the pair was resolved from the optimization. Five species of Parabathippus and Bathippus each were sampled in phylogenetic studies including their type species, P. shelfordi (Peckham & Peckham) and B. macrognathus. The results indicate Parabathippus and Bathippus fall into different clades that are not even closely related, thus corroborating the separation of Southeast Asian species as Parabathippus (see Chapter 8 for detailed description). Parabathippus differs from Bathippus in the long and more coiled embolus with the plane of the spiral parallel to the longitudinal axis of the bulb (the plane of the embolic spiral is more perpendicular to the longitudinal axis of the bulb in Bathippus), the larger and oval embolic disc, and the longer and less sclerotized copulatory duct. The monophyly of Parabathippus is also strongly supported by the molecular phylogeny. 3.5.2.3.8 Euophrys Clade: Chalcoscirtus, Euophrys, Pseudeuophrys, Talavera and Featheroides (Figs 3.40-3.41) The close relationship of Euophrys (Holarctic species), Pseudeuophrys and Talavera has been proposed by Zabka and Prószy!ski (1998) based on morphological characters. The molecular phylogeny supports this proposal and indicates Chalcoscirtus is also closely related to them. In addition, Featheroides (Peng et al. 1994: 2-3, figs. 1-3, 4-6) is also included in this clade based 86! on the similarity of its male palpal structure with Euophrys. Spiders in this clade are small to medium sized, usually relatively robust and dark in color. The carapace usually lacks the longitudinal fovea (ch. 15-1). The genitalic structures vary. The male palp has a proximal tegular lobe. The embolus is thin or wide, coiled or uncoiled (e.g. Talavera). The retrolateral sperm duct loop is present and the prolateral sperm duct loop is absent except in Talavera. The RTA is usually finger-like, but is highly reduced in Talavera. The epigynal window is present or absent (e. g. Talavera). The spermatheca is usually swollen and oval or round. The molecular phylogeny also suggests that Chalcoscirtus diminutus (Banks) is misplaced; its generic placement needs to be further investigated. 3.5.2.3.9 Saitis Clade: Saitis and possibly Margaromma (Figs 3.42-3.43) The Saitis clade is well supported by the molecular phylogeny. Margaromma (see diagnostic drawings of the type species in Davies & Zabka 1989: 230, pl. 38) may be closely related to Saitis based on its similar morphology. Saitis Simon, 1876 Saitis Simon, 1876: 168, type species: Saitis barbipes (Simon, 1868). Lycidas Karsch, 1878: 26, type species: Lycidas anomalus Karsch, 1878. New Synonym Maratus Karsch, 1878: 27, type species: Maratus amabilis Karsch, 1878. New Synonym Jotus L. Koch, 1881: 1243, type species: Jotus auripes L. Koch, 1881. New Synonym Prostheclina Keyserling, 1882: 1368, type species: Prostheclina pallida Keyserling, 1882. Hypoblemum Peckham & Peckham, 1886: 326, type species: Hypoblemum villosum (Keyserling, 1883). New Synonym Maileus Peckham & Peckham, 1907: 612, type species: Maileus fuscus Peckham & Peckham, 1907. New Synonym Diagnosis. Spiders of Saitis are medium sized and usually relatively compact but not elongate. The male of some species (e.g. Saitis volans) has colorful abdominal flaps, which expand to display to the female during courtship. The male palpal bulb is relatively large with a retrolateral sperm duct loop and a tegular lobe. The RTA is finger-like. The ventral tibial bump of the male palp is usually present (ch. 48-1). The embolus usually coils for no more than half of a circle. 87! Saitis differs from other euophryine genera in the male palp, which has a lamella on the tegular shoulder (ch. 49-1). The epigynal window is present with a median septum. According to features above and the molecular phylogeny, the following species are transferred to Saitis here: Saitis anomaliformis (Zabka, 1987) (from Lycidas, New Combination) Saitis anomalus (Karsch, 1878) (from Lycidas, New Combination) Saitis bitaeniatus (Keyserling, 1882) (from Lycidas, New Combination) Saitis chlorophthalmus (Simon, 1909) (from Lycidas, New Combination) Saitis chrysomelas (Simon, 1909) (from Lycidas, New Combination) Saitis dialeucus (L. Koch, 1881) (from Lycidas, New Combination) Saitis furvus (Song & Chai, 1992) (from Lycidas, New Combination) Saitis griseus (Keyserling, 1882) (from Lycidas, New Combination) Saitis heteropogon (Simon, 1909) (from Lycidas, New Combination) Saitis karschi (Zabka, 1987) (from Lycidas, New Combination) Saitis kochi (Zabka, 1987) (from Lycidas, New Combination) Saitis michaelseni (Simon, 1909) (from Lycidas, New Combination) Saitis nigriceps (Keyserling, 1882) (from Lycidas, New Combination) Saitis nigromaculatus (Keyserling, 1883) (from Lycidas, New Combination) Saitis obscurior (Simon, 1909) (from Lycidas, New Combination) Saitis piliger (Keyserling, 1882) (from Lycidas, New Combination) Saitis pilosus (Keyserling, 1882) (from Lycidas, New Combination) Saitis scutulatus (L. Koch, 1881) (from Lycidas, New Combination) Saitis speculifer (Simon, 1909) (from Lycidas, New Combination) Saitis vittatus (Keyserling, 1881) (from Lycidas, New Combination) Saitis amabilis (Karsch, 1878) (from Maratus, New Combination) Saitis linnaei (Waldock, 2008) (from Maratus, New Combination) Saitis mungaich (Waldock, 1995) (from Maratus, New Combination) Saitis pavonis Dunn, 1947 (from Maratus) Saitis rainbowi (Roewer, 1951) (from Maratus) Saitis vespertilio Simon, 1901 (from Maratus) Saitis volans (O. P.-Cambridge, 1874) (from Maratus, Fig. 3.42G-L) Saitis auripes (L. Koch, 1881) (from Jotus, New Combination, Fig. 3.42A-F) 88! Saitis braccatus (L. Koch, 1881) (from Jotus, New Combination) Saitis debilis (L. Koch, 1881) (from Jotus, New Combination) Saitis frosti (Peckham & Peckham, 1901) (from Jotus, New Combination) Saitis insulanus (Rainbow, 1920) (from Jotus, New Combination) Saitis maculivertex (Strand, 1911) (from Jotus, New Combination) Saitis minutus (L. Koch, 1881) (from Jotus, New Combination) Saitis ravus (Urquhart, 1893) (from Jotus, New Combination) Saitis amplior (Richardson & Zabka, 2007) (from Prostheclina, New Combination) Saitis basilonesa (Richardson & Zabka, 2007) (from Prostheclina, New Combination) Saitis boreoaitha (Richardson & Zabka, 2007) (from Prostheclina, New Combination) Saitis boreoxantha (Richardson & Zabka, 2007) (from Prostheclina, New Combination) Saitis bulburin (Richardson & Zabka, 2007) (from Prostheclina, New Combination) Saitis eungella (Richardson & Zabka, 2007) (from Prostheclina, New Combination) Saitis pallidus (Keyserling, 1882) (from Prostheclina) Saitis albovittatus (Keyserling, 1882) (from Hypoblemum, New Combination) Saitis villosus (Keyserling, 1883) (from Hypoblemum, New Combination) Saitis fuscus (Peckham & Peckham, 1907) (from Maileus, New Combination, Fig. 3.43J-M) Notes. In spite of extensive efforts having been made on properly defining and distinguishing among the genera Lycidas, Hypoblemum, Saitis, Maratus and Jotus (e.g. Zabka 1987; Davies & Zabka 1989; Zabka & Pollard 2002; Richardson & Zabka 2007), it is still very difficult to tell them apart due to the high similarity in their genitalic organs. The molecular phylogeny (see Chapter 2) indicates that at least some of the above genera are not monophyletic (e.g. Lycidas and Hypoblemum). Study has shown that spiders of both Maratus and Saitis extend their third legs during courtship (Hill 2009a). Although the spectacular courtship ornaments of Maratus might seem to warrant a separate genus, in other salticid genera there are mixes of highly ornamented and unornamented species (e.g., Habronattus, Phidippus); separating Maratus would leave a paraphyletic group behind. Thus, I merge them into one single well-defined genus and consider Hypoblemum, Jotus, Lycidas, Maileus, Maratus and Prostheclina as junior synonyms of Saitis. Species of these genera are therefore transferred to Saitis. Saitis has been a problematic genus, with species from all over the world placed into it. However, the molecular phylogeny suggests that the two species from Africa (“Saitis” mundus 89! Peckham & Peckham and “Saitis” leighi Peckham & Peckham) are not closely related to Saitis barbipes (Simon), the type species of Saitis, but group with the African Thyenulla species. The Neotropical species also likely have been misplaced and need to be transferred when their proper placement is understood. 3.5.2.3.10 Laufeia Clade: Laufeia and possibly Magyarus (Fig 3.44) The monophyly of Laufeia is well supported. The monotypic genus Magyarus Zabka (1985: 237, figs. 268-271) from Vietnam may also belong to this clade. Laufeia Simon, 1889 Laufeia Simon, 1889: 248, type species: Laufeia aenea Simon, 1889. Orcevia Thorell, 1890: 166, type species: Orcevia keyserlingii Thorell, 1890. Junxattus Prószy!ski & Deeleeman-Reinhold, 2012: 40, type species: Junxattus daiqini Prószy!ski & Deeleeman-Reinhold, 2012. New Synonym Diagnosis. Body of Laufeia is usually relatively compact, dark brown or yellow brown in color covered with many hairs and setae. The male usually has a slightly sclerotized scutum on the dorsal abdomen. The chelicera usually has a bicuspid tooth on the retromargin (ch. 21-2). The genitalia show high interspecific variation. The male palp has a retrolateral sperm duct loop and a proximal tegular lobe. The embolus is long or short, coiled or uncoiled. The epigynal window is usually indistinct (ch. 80-1). Spiders of Laufeia are small, some dwelling on leaf litter. Following species are transferred to Laufeia based on the features above and the molecular phylogeny: Laufeia eucola (Thorell, 1890) (from Orcevia) Laufeia keyserlingii (Thorell, 1890) (from Orcevia, Fig. 3.44N-T) Laufeia kuloni (Prószy!ski & Deeleeman-Reinhold, 2012) (from Orcevia, New Combination) Laufeia perakensis (Simon, 1901) (from Orcevia) Laufeia proszynskii Song, Gu & Chen, 1988 (from Orcevia) Laufeia daiqini (Prószy!ski & Deeleeman-Reinhold, 2012) (from Junxattus, New Combination, Fig. 3.44G-M) 90! Notes. Orcevia Thorell (reinstated from Laufeia) and Junxattus Prószy!ski & Deeleeman- Reinhold are newly separated from Laufeia by Prószy!ski & Deeleeman-Reinhold (2012) based on their different genitalic structures. In the molecular phylogeny (see Chapter 2), I include the type species of Orcevia (O. keyserlingi Thorell), the type species of Junxattus (J. daiqini Prószy!ski & Deeleeman-Reinhold) and two Laufeia species newly discovered (L. eximia and L. concava, see Chapter 8). They all fall in a strongly supported clade. The type species of Laufeia, L. aenea Simon was not included because no material was available for the molecular work. However, the type specimens of L. aenea (examined) show similarity in genitalia with L. concava. For instance, they both have relatively wide bulb, small embolic disc and short embolus in the male palp, and a window structure in the epigynum. In spite of having genitalia more diverse than in other euophryine genera, the species included in phylogenetic study and L. aenea are very similar in body form and cheliceral teeth pattern. A high genitalic diversity could occur even in closely related species, if for instance strong sexual selection drives rapid divergence. Thus I am reluctant to follow Prószy!ski and Deeleeman-Reinhold’s classification (2012), which will result in at least four genera for this clade with each comprising very few or even single species. Instead, I treat all of them as one single genus, and consider Orcevia and Junxattus as junior synonyms of Laufeia. 3.5.2.3.11 Colyttus Clade (Fig 3.45) Colyttus Thorell, 1891 Colyttus Thorell, 1891: 132, type species: Colyttus bilineatus Thorell, 1891. Donoessus Simon, 1902: 376, type species: Hasarius nigriceps Simon, 1899. New Synonym Diagnosis. The male palpal femur is usually slightly curved. The chelicera usually has two promarginal teeth and a unident or bicuspid retromarginal tooth. The male palpal bulb has a proximal tegular lobe and a slightly coiled embolus. The plane of the embolic disc is ventral to the plane of the front tegulum (ch. 68-1). Some species have an additional projection on the embolic disc beside the embolus. The epigynal window is present with a median septum. The vulva has a pair of secondary spermathecae in addition to the primary spermathecae. Spiders of Colyttus are medium sized and relatively bright colored spiders usually found on foliage. The following species are transferred to Colyttus here based on the features above and the molecular phylogeny: 91! Colyttus kerinci (Prószy!ski & Deeleeman-Reinhold, 2012) (from Donoessus, New Combination) Colyttus nigriceps (Simon, 1899) (from Donoessus, New Combination) Colyttus striatus (Simon, 1902) (from Donoessus, New Combination, Fig. 3.45O-U) Notes. The molecular phylogeny indicates Colyttus is paraphyletic with Donoessus embedded. Based on their similar morphology, I consider Donoessus as a junior synonym of Colyttus, and the two species in Donoessus are therefore transferred to Colyttus. 3.5.2.3.12 Cytaea-Euryattus Clade: Cytaea, Euryattus, possibly Aruattus and Charippus (Fig 3.46) This well-supported clade in the molecular phylogeny includes Cytaea and Euryattus. Cytaea and Euryattus are medium sized spiders. The chelicera usually has multiple small teeth on the promargin and a bicuspid tooth on the retromargin. The male palpal bulb is wide or narrow, usually lacking a distinct proximal tegular lobe. The embolus is usually long and highly spiraled, and partly embedded in a distal furrow of the bulb. The retrolateral sperm duct loop is usually no more than half of the palpal bulb width (ch. 58-0/1). The epigynal window is present, with a median septum. The palpal bulb of Cytaea is usually much wider than that of Euryattus. Male Cytaea often have bright setae on the clypeus and the front surface of the chelicerae, which are absent in Euryattus. The vulva of Euryattus has a projection (secondary spermatheca) from the copulatory duct beside the primary spermatheca, while the vulva of Cytaea usually doesn’t have this projection. The results indicate Cytaea oreophila Simon (Fig. 3.46H-N) may belong to a distinct lineage from the other Cytaea species. Its genitalic structures show some difference from typical Cytaea species, and further study is needed to clarify its position. Charippus Thorell (Wanless 1988: 162, figs. 40A-H) and Aruattus Logunov & Azarkina (2008: 112, figs. 1-9) may also belong to this clade. Both of them have a bicuspid tooth on the retromargin like Cytaea and Euryattus, but have fewer teeth on the promargin (one tooth in Aruattus and two in Charippus). The wide palpal bulb of Charippus is similar to that of Cytaea, and the male palpal structure of Aruattus is similar to that of Euryattus. 92! 3.5.2.3.13 Thiania Clade (Fig 3.47) Thiania C. L. Koch, 1846 Thiania C. L. Koch, 1846: 171, type species: Thiania pulcherrima C. L. Koch, 1846. Nicylla Thorell, 1890: 171, type species: Nicylla sundevalli Thorell, 1890. New Synonym Pselcis Simon, 1903: 825, type species: Euophrys latefasciata Simon, 1877. New Synonym Thianitara Simon, 1903: 1054, type species: Thianitara spectrum Simon, 1903. New Synonym Diagnosis. Spiders of Thiania are medium sized with the body obviously flattened (ch. 3-1). Some species have many iridescent scales on the body. The longitudinal fovea is relatively far behind the PLEs (ch. 16-1). The male palpal bulb is large or small, with the tegular lobe present or absent. The embolus is usually wide and only slightly coiled. The plane of the embolic disc is often ventral to the plane of the front tegulum (ch. 68-1). The epigynal window is present with a median septum. According to above features and molecular phylogeny, the following species are transferred to Thiania here: Thiania spectrum (Simon, 1903) (from Thianitara, New Combination, Fig. 3.47H-L) Thiania thailandica (Prószy!ski & Deeleeman-Reinhold, 2012) (from Thianitara, New Combination) Thiania sundevalli (Thorell, 1890) (from Nicylla, New Combination) Thiania latefasciata (Simon, 1877) (from Pselcis, New Combination) Notes. Thianitara, Nicylla and Pselcis are monotypic genera, with Pselcis only known from juveniles. They all show a flattened body as in Thiania. The molecular phylogeny (see Chapter 2) indicates Thianitara is derived within Thiania. Based on the similar flattened body form and the genitalic structures, here I consider Thianitara, Nicylla and Pselcis as junior synonyms of Thiania. 3.5.2.3.14 Emathis-Lepidemathis Clade (Fig 3.48) Spiders of this clade are medium or large sized spiders. The chelicera usually has two promarginal teeth and a fissident retromarginal tooth of multiple cusps (ch. 21-3, ch. 23-4). The first metatarsus usually has four pairs of ventral macrosetae (ch. 40-3, ch. 41-4). The male 93! palpal bulb is usually small with a distal coiled embolus. The proximal tegular lobe is absent (ch. 47-0). The epigynal window is present. The copulatory duct is short or long and convoluted. Emathis is usually much smaller and less robust than Lepidemathis. The copulatory duct of Emathis is usually long and convoluted (except E. gombak, see Chapter 8), while the copulatory duct of Lepidemathis is much shorter. The male palpal tibia of Lepidemathis often has a ventral ridge, which is absent in Emathis. The following species is transferred to Emathis based on its genitalic structure (Wanless 1984b: 468, figs. 14A-I): Emathis armillata (Peckham & Peckham, 1907) (from Cyrba, New Combination) 3.5.2.3.15 Thyenula Clade: Thyenula, possibly Tanzania, Lophostica and Pseudemathis (Fig 3.49) Only Thyenula from Africa is included in the analysis. Another three genera, Tanzania, Lophostica and Pseudemathis, are also known to be endemic to the African region. Whether they fall in the same clade as Thyenula or represent independent radiations into Africa is still uncertain. The results from the ML analysis on the combined morphological and DNA data fail to recover Thyenula laxa and T. nelshoogte in the same clade as the other Thyenula species included. However, the monophyly of Thyenula including these two species is well supported by the Bayesian analysis on the combined all genes data with full taxa (posterior probability=1.0, see Chapter 2). Thus, I tentatively put them in Thyenula until more thorough study suggests otherwise. Thyenulla are medium sized spiders. The male palp has a proximal tegular lobe and a retrolateral sperm duct loop. The embolus is usually long and spiraled for more than half a circle. The RTA is finger-like. The epigynal window is present, with a median septum. The spermatheca is usually strongly swollen and not as narrow as the copulatory duct. The morphological and molecular data show that two species from Africa,“Saitis” mundus Peckham & Peckham and “Saitis” leighi Peckham & Peckham have been misplaced and I transfer them to Thyenula here. 94! Thyenula munda (Peckham & Peckham, 1903) (from Saitis, New Combination) Thyenula leighi (Peckham & Peckham, 1903) (from Saitis, New Combination) 3.5.2.3.16 Other genera from the Old World: Chalcotropis, Chinophrys, Foliabitus, Lagnus, Servaea, Sigytes, Thorelliola, Viribestus, Xenocytaea, Zabkattus (Figs 3.50-3.57) Proper groupings of these genera still need further studies. Chinophrys, Foliabitus, Viribestus and Zabkattus are recently discovered from Asia and Papua New Guinea (see Chapters 6 and 8 for detailed diagnoses and descriptions). This study recognizes a well-supported clade associated with the type species of Thorelliola, T. ensifera (Thorell). Typical Thorelliola species usually have the male clypeus armed with “horns” (Gardzinska & Patoleta 1997; Szüts & De Bakker 2004). However, some species in this clade from Papua New Guinea (see Chapter 6 for detailed descriptions) lack clypeal “horns”. Some Thorelliola species, e.g. T. mahunkai Szüts, has “horns” that aren’t as robust as other species but are more like ordinary macrosetae (Szüts 2002). To avoid erecting more euophryine genera for species closely related to typical Thorelliola but lacking horns, I expanded the delimitation of Thorelliola to enclose more species. All species now placed in Thorelliola share the synapomorphy of a female epigynum without median septum (ch. 74-1). Thorelliola can now be diagnosed as: the chelicera has a fissident retromarginal tooth, and many species have a process distally on the front surface of the male chelicera (ch. 27-1); the epigynum has a big window without the median septum (ch. 74-1); some species have a pair of secondary spermathecae in addition to the primary spermathecae; the male palp usually has macrosetae on the tibia and also on the femur in some species; the tegulum lacks the proximal lobe; the embolus is long or short (see Chapter 6). Three species of Xenocytaea were included in this study; they fall into a clade. Xenocytaea can be easily distinguished from other euophryine genera by the genitalic structures: the female epigynum usually has a large anterior arch instead of a distinct window typical of euophryines; the male palpal bulb is usually wide, sometimes with a large or small apophysis at the shoulder of the tegulum. Spiders of this genus are small to medium sized. The chelicera usually has two promarginal and one bicuspid or unident retromarginal tooth. The first tibia has two or three 95! pairs of ventral macrosetae and the first metatarsus has two pairs of ventral macrosetae. Spiders of Lagnus are medium-sized. The chelicera has two promarginal and one fissident retromarginal tooth of multiple cusps. They are distinctive in the extremely elongate and spiny legs, especially in males. The first tibia has more than seven pairs of ventral macrosetae and the first metatarsus has more than five pairs of ventral macrosetae. The embolus of the male palp is slender and coiled. The tegulum lacks the proximal lobe. The epigynal window is present with a median septum. Clear delimitation of the genera Chalcotropis, Servaea and Sigytes is still uncertain and more thorough comparative studies on described species are needed. 3.6 Conclusions Delimitations of some euophryine genera and phylogenetic placements of genera lacking molecular data are clarified, based on the extensive study on morphological traits of a broad range of euophryine genera and species. The results of formal phylogenetic analyses suggest that the morphological dataset does not perform as well as the molecular dataset in resolving the phylogeny of Euophryinae. However, morphology helps to extend the molecular phylogeny, allowing 36 genera without molecular data to be placed within Euophryinae. In addition, optimization of the morphological characters on the phylogeny provides a better understanding of the euophryine generic groups and the delimitation of many euophryine genera, and reveals certain characters, especially on the male palp, that are valuable in delimiting various euophryine groups. Some problems in the traditional taxonomy of euophryines are also corrected based on the revised generic delimitations. 96! Table 3.1. Substitution models selected by ModelTest for each molecular partition. Gene partition Sites included (bp) Substitution model 28SrDNA 1384 GTR+I+G Actin 5C 1 st codon position 239 GTR+I+G Actin 5C 2 nd codon position 239 HKY Actin 5C 3 rd codon position 239 GTR+I+G 16SrDNA 753 TrN+I+G ND1+COI 1 st codon position 456 GTR+I+G ND1+COI 2 nd codon position 456 GTR+I+G ND1+COI 3 rd codon position 456 GTR+G 97! Table 3.2. Optimization of morphological chatacters for major clades of euophryines on different phylogenies. Only the resulting character states (Character#-State#) from unambiguous changes of all most parsimonious reconstructions are shown here. Clade Optimization on DNA ML tree (Fig. 3.1) Optimization on combined DNA and morphology ML tree (Fig. 3.5) Euophryinae Clade 56-0 46-1; 56-0; 66-0; 79-1 Anasaitis- Corythalia Clade 20-2; 22-3; 35-1; 48-1 20-2; 22-3; 34-1; 35-1; 48-1 Anasaitis 70-1 46-0; 80-1 Corythalia 14-0 14-0 Antillattus Clade none none Antillattus none none Petemathis 21-3; 23-4 21-3; 23-4 Truncattus 21-2; 23-3; 48-1 21-2; 23-3; 48-1 Agobardus Clade 47-0 47-0 Agobardus 21-2; 40-2; 41-3; 58-2 21-2; 40-2; 41-3; 58-2 Bythocrotus 20-2; 22-3; 25-1; 26-1; 31-2; 43-1; 48-1; 51-1; 52-0; 61-1; 62-1; 76-1 20-2; 22-3; 25-1; 26-1; 31-2; 43-1; 48-1; 51-1; 61-1; 62-1; 76-1 Compsodecta none none Popcornella 14-0; 46-0; 55-2; 66-1; 70-1; 80-1 14-0; 46-0; 55-2; 66-1; 70-1; 80-1 Naphrys- Corticattus Clade 9-1; 20-2; 22-3; 38-1; 39-1; 43-1; 48- 1; 52-0 9-1; 20-2; 22-3; 32-2; 38-1; 39-1; 43-1; 48-1 Sidusa Clade 21-2; 23-3; 38-4; 39-4; 40-2; 41-3; 43-1; 47-0; 58-0 21-2; 23-3; 38-4; 39-4; 41-3; 43-1; 58-0; 79-2 Mopiopia- Saphrys Clade none none Saphrys 32-2; 42-1; 48-1; 64-1 32-2; 42-1; 48-1; 64-1 Chapoda- Maeota Clade none 32-2 Chapoda 48-1; 51-1; 76-1 48-1; 51-1; 76-1 Maeota none none Amphidraus- Marma Clade 21-3; 23-4; 52-1; 80-1 21-3; 23-4; 52-1; 67-1; 80-1 Coryphasia Clade 23-3; 25-1 23-3; 25-1 Pensacola- Mexigonus Clade none none Soesilarishius Clade 42-1; 43-0; 46-0; 70-1 42-1; 43-0; 46-0; 70-1 Ilargus 79-2 79-2 Bathippus- Canama Clade 31-0; 40-2; 41-3 31-0; 50-1; 64-1; 72-1 98! Clade Optimization on DNA ML tree (Fig. 3.1) Optimization on combined DNA and morphology ML tree (Fig. 3.5) Bathippus 21-1; 22-5; 40-3; 41-4; 46-1; 71-1 21-1; 40-3; 41-4; 71-1 Canama none 46-2 Omoedus Clade none 20-2; 22-3; 46-3; 75-1; 81-3 Bulolia- Coccorchestes Clade 47-1; 51-0/1; 57-1; 58-0 57-1; 58-0; 64-1 Diolenius Clade 3-1; 15-1; 38-6; 40-2 3-1; 15-1; 38-6; 40-2; 79-2 Pristobaeus Clade 47-1; 50-1; 51-3; 53-1; 65-1 47-1; 50-1; 51-3; 53-1; 65-1 Parabathippus- Parvattus Clade none none Parabathippus 17-1; 18-1; 22-5; 45-1; 46-2; 47-0; 50-1; 51-1/2; 53-1 17-1; 18-1; 22-5; 46-2 Euophrys Clade 15-1; 82-2 82-2 Saitis Clade 22-3; 48-1; 49-1 22-3; 48-1; 49-1; 72-1 Laufeia Clade 21-2; 80-1 21-2; 80-1 Colyttus Clade 68-1 50-1; 51-3; 53-1; 68-1; 76-1 Cytaea- Euryattus Clade 43-1; 51-4; 58-0 47-0; 51-4; 58-0 Thiania Clade 3-1; 16-1; 68-1 3-1; 16-1 Emathis- Lepidemathis Clade 21-3; 23-4; 40-3; 41-4; 47-0 31-0 Thyenula Clade none NA Thorelliola 27-1; 74-1 74-1 Xenocytaea none none 99! Table 3.3. Full list of euophryine genera. Genera Availability of DNA data If in Maddison and Hedin (2003a) Where in Simon's Classification (1901; 1903) Agobardus yes no NA Allodecta no no NA Amphidraus yes no Amycieae (Pluridentati) Anasaitis yes yes NA Antillattus yes no NA Araneotanna no no NA Aruattus no no NA Asaphobelis (=Coryphasia) yes no Hasarieae (Fissidentati) Ascyltus no yes Cytaeeae (Fissidentati) Athamas yes yes Athameae (Fissidentati) Bathippus yes yes Plexippeae (Unidentati) Belliena yes no Bellieneae (Fissidentati) Bindax no no Amycieae (Pluridentati) Bulolia yes no NA Bythocrotus yes no Bythocroteae (Unidentati) Canama yes yes Cytaeeae (Fissidentati) Caribattus no no NA Chalcolecta yes no Diolenieae (Pluridentati) Chalcolemia yes no NA Chalcoscirtus yes yes Chalcoscirteae (Unidentati) Chalcotropis yes yes Hasarieae (Fissidentati) Chapoda yes yes Hasarieae (Fissidentati) Charippus no no Astieae (Pluridentati) Chinophrys yes no NA Chloridusa (=Sidusa) no no Hasarieae (Fissidentati) Cobanus (=Sidusa) yes yes Amycieae (Pluridentati) Coccorchestes yes no Coccorchesteae (Unidentati) Colyttus yes no Plexippeae (Unidentati) Commoris no no Hasarieae (Fissidentati) Compsodecta yes no Pensacoleae (Unidentati) Corticattus yes no NA Coryphasia yes no Hasarieae (Fissidentati) Corythalia yes yes Zenodoreae (Unidentati) Cytaea yes yes Cytaeeae (Fissidentati) Darwinneon no no NA Dinattus (=Corythalia) yes no NA Diolenius yes no Diolenieae (Pluridentati) Donoessus (=Colyttus) yes no Hasarieae (Fissidentati) Ecuadattus yes no NA Efate yes no NA Emathis yes no Emathideae (Fissidentati) 100! Genera Availability of DNA data If in Maddison and Hedin (2003a) Where in Simon's Classification (1901; 1903) Euophrys yes yes Evophrydeae (Unidentati) Euryattus yes yes Cytaeeae (Fissidentati) Featheroides no no NA Foliabitus yes no NA Furculattus no no NA Hypoblemum (=Saitis) yes yes Hylleae (Unidentati) Ilargus yes no Saiteae (Unidentati) Jotus (=Saitis) yes yes Saiteae (Unidentati) Junxattus (=Laufeia) yes no NA Lagnus yes yes Astieae (Pluridentati) Lakarobius no no NA Laufeia yes no Laufeieae (Fissidentati) Lepidemathis yes yes Emathideae (Fissidentati) Leptathamas yes no NA Lophostica no no Emathideae (Fissidentati) Lycidas (=Saitis) yes no NA Maeota yes yes Saiteae (Unidentati) Maeotella (=Anasaitis) no no NA Magyarus no no NA Maileus (=Saitis) yes no NA Maratus (=Saitis) yes yes Saiteae (Unidentati) Margaromma no no Zenodoreae (Unidentati) Marma yes no Hasarieae (Fissidentati) Mexigonus yes yes NA Mopiopia no no Hasarieae (Fissidentati) Naphrys yes yes NA Nebridia (=Amphidraus) yes no Hasarieae (Fissidentati) Neonella yes no NA Nicylla (=Thiania) no no NA Ocnotelus (=Semnolius) no no Hasarieae (Fissidentati) Ohilimia yes no NA Omoedus yes yes Coccorchesteae (Unidentati) Opisthoncana no no NA Orcevia (=Laufeia) yes no NA Palpelius (=Pristobaeus) yes no Plexippeae (Unidentati) Parabathippus yes no NA Paradecta (=Compsodecta) no no NA Paraharmochirus yes no NA Parasaitis no no NA Parvattus yes no NA Pensacola yes yes Pensacoleae (Unidentati) Petemathis yes no NA Phasmolia yes no NA 101! Genera Availability of DNA data If in Maddison and Hedin (2003a) Where in Simon's Classification (1901; 1903) Popcornella yes no NA Pristobaeus yes no Emathideae (Fissidentati) Prostheclina (=Saitis) yes no Saiteae (Unidentati) Pselcis (=Thiania) no no Laufeieae (Fissidentati) Pseudemathis no no Emathideae (Fissidentati) Pseudeuophrys yes yes NA Pystira (=Omoedus) yes yes Zenodoreae (Unidentati) Rhyphelia no no Evophrydeae (Unidentati) Saitidops no no Aelurilleae (Unidentati) Saitis yes yes Saiteae (Unidentati) Saphrys yes no NA Semnolius no no Hasarieae (Fissidentati) Servaea yes yes Servaeeae (Fissidentati) Sidusa yes yes Hasarieae (Fissidentati) Sigytes no no Plexippeae (Unidentati) Siloca (=Coryphasia) yes yes Hasarieae (Fissidentati) Sobasina yes no Sobasineae (Pluridentati) Soesilarishius yes no NA Spilargis no yes Spilargeae (Fissidentati) Stoidis no no Zenodoreae (Unidentati) Talavera yes yes NA Tanzania no no NA Tariona (=Mopiopia) yes no Hasarieae (Fissidentati) Tarodes no no NA Thiania yes yes Thianieae (Unidentati) Thianitara (=Thiania) yes no Thianieae (Unidentati) Thorelliola yes yes Spilargeae (Fissidentati) Thyenula yes no Thyeneae (Unidentati) Truncattus yes no NA Tylogonus yes no Hasarieae (Fissidentati) Udvardya no no NA Variratolia yes no NA Viribestus yes no NA Wallaba (=Sidusa) no no NA Xenocytaea yes no NA Zabkattus yes no NA Zenodorus (=Omoedus) yes yes Zenodoreae (Unidentati) 102! Figure 3.1. Phylogeny of Euophryinae. Tree shown is modified from the best ML tree from the combined all genes dataset (see Chapter 2) with taxa that without morphological data trimmed off. Major euophryine clades are indicated along the branches. New World or Old World distribution of euophryine taxa is indicated in colored blocks in front of the taxon names. Ghelna canadensis 'Bathippus' pahang Chinattus parvulus Heliophanus cupreus Freya decorata Plexippus paykulli Salticus scenicus Corythalia porphyra Corythalia sulfurea Corythalia bicincta Corythalia electa Corythalia broccai Corythalia minor Corythalia peblique Corythalia bromelicola Corythalia coronai Corythalia decora Anasaitis canosa [USA] Anasaitis cf. canalis Anasaitis adorabilis Anasaitis brunnea Anasaitis hebetata Anasaitis banksi Anasaitis placida Anasaitis gloriae Anasaitis laxa Anasaitis elegantissima Anasaitis locuples Anasaitis sp. [Peblique] Euophrys frontalis Euophrys monodnock Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Parvattus zhui Parabathippus kiabau Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Sidusa sp. [CostaRica] Sidusa extensa Sidusa unicolor Pensacola signata Mexigonus arizonensis Mexigonus morosus Petemathis minuta Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis Antillattus cambridgei Antillattus darlingtoni Antillattus maxillosus Antillattus scutiformis Naphrys pulex Corticattus latus Popcornella nigromaculata Popcornella furcata Popcornella spiniformis Compsodecta haytiensis Compsodecta peckhami Bythocrotus cf. crypticus Bythocrotus crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo Chapoda recondita Chapoda gitae Chapoda angusta Chapoda montana Chapoda fortuna [Panama] Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota dichrura Maeota sp. [Napo] Maeota flava Maeota simoni Maeota sp. [MoronaSantiago] Maeota tuberculotibiata Amphidraus complexus Marma nigritarsis Tylogonus cf. auricapillus Tylogonus yanayacu Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Mopiopia cf. bruneti Saphrys a-notata Saphrys cf. patagonica cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Coryphasia physonycha Coryphasia cf. campestrata Neonella vinnula Ecuadattus napoensis Ecuadattus pichincha Belliena ecuadorica Ilargus galianoae Ilargus pilleolus Ilargus coccineus Ilargus serratus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Servaea vestita Saitis auripes Saitis cf. fuscus Saitis barbipes Saitis sp. [SouthAustralia] Saitis sp. [NewSouthWales] Saitis cf. griseus Emathis gombak Lepidemathis haemorrhoidalis Chalcotropis cf. caeruleus Chalcotropis luceroi Colyttus bilineatus Colyttus striatus Euophryine sp. [GentingHighlands] Lagnus edwardsi Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola cf. mahunkai Thorelliola tualapa Thorelliola ensifera Thorelliola tamasi Foliabitus longzhou Foliabitus sp. [Malaysia] Laufeia daiqini Laufeia eximia Laufeia keyserlingi Thiania spectrum Thiania latibola Thiania bhamoensis Thiania tenuis Chinophrys pengi Thyenula laxa Thyenula nelshoogte Thyenula leighi Thyenula sp. [SouthAfrica] Thyenula wesolowskae Thyenula cf. mundus Thyenula cf. aurantiaca Cytaea oreophila Cytaea mitellata Cytaea nimbata Euryattus bleekeri Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Phasmolia elegans Zabkattus furcatus Zabkattus richardsi Zabkattus brevis Zabkattus trapeziformis Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Bathippus macrognathus Bathippus directus [Tualapa] Bathippus gahavisuka Bathippus korei Bathippus madang Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Pristobaeus beccarii Pristobaeus cf. jocosus Xenocytaea agnarssoni Xenocytaea albomaculata Xenocytaea proszynskii Chalcolemia nakanai Chalcolecta prensitans Diolenius varicus Ohilimia scutellata Sobasina wanlessi Efate albobicinctus Paraharmochirus tualapaensis Omoedus orbiculatus Omoedus cf. piceus Omoedus ephippigera Omoedus darleyorum Omoedus papuanus Omoedus meyeri Omoedus omundseni Omoedus cf. torquatus Omoedus cf. danae Omoedus brevis Omoedus cf. semirasus Omoedus swiftorum Euophryinae Anasaitis-Corythalia Euophrys Clade Parabathippus-Parvattus Sidusa Clade Pensacola-Mexigonus Clade Clade Clade Antillattus Clade Naphrys-Corticattus Clade Agobardus Chapoda-Maeota Clade Amphidraus-Marma Clade Mopiopia-Saphrys Clade Coryphasia Ilargus Clade Saitis Clade Thorelliola Laufeia Clade Colyttus Clade Thiania Clade Thyenula Clade Cytaea-Euryattus Clade Emathis-Lepidemathis Clade Zabkattus Clade Bulolia-Coccorchestes Clade Clade Clade Bathippus-Canama Clade Omoedus Clade Diolenius Clade Pristobaeus Clade Xenocytaea Foliabitus Chalcotropis Ecuadattus Tylogonus Soesilarishius Popcornella Phasmolia Clade ******* * New World Old World Outgroup 103! Figure 3.2. Strict consensus of four equally parsimonious trees (score=1141) from the morphological dataset that found in TNT, without constraining euophryine taxa as a monophyletic group. Recovered euophryine clades and MP bootstrap support values are indicated along branches. New World or Old World distribution of euophryine taxa is indicated in colored blocks in front of the taxon names. Ghelna canadensis cf. Coryphasia sp. [Brazil] Zabkattus brevis Zabkattus furcatus Zabkattus richardsi Zabkattus trapeziformis Thyenula leighi Omoedus meyeri Omoedus papuanus Xenocytaea agnarssoni Omoedus orbiculatus Omoedus cf. piceus Omoedus ephippigera Omoedus darleyorum Servaea vestita Omoedus omundseni Omoedus cf. semirasus Omoedus swiftorum Omoedus cf. torquatus Omoedus cf. danae Coryphasia fasciiventris Thorelliola aliena Coryphasia physonycha Coryphasia cf. campestrata Mopiopia cf. bruneti Thyenula laxa Chinophrys pengi Ilargus moronatigus Saphrys a-notata Saphrys cf. patagonica Ecuadattus napoensis Ecuadattus pichincha Maeota sp. [Napo] 'Bathippus' pahang Bulolia excentrica Leptathamas paradoxus Cytaea oreophila Cytaea mitellata Cytaea nimbata Euryattus bleekeri Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Omoedus brevis Bythocrotus cf. crypticus Bythocrotus crypticus Thiania tenuis Thiania bhamoensis Thiania latibola Xenocytaea proszynskii Thorelliola crebra Thorelliola Joannae Thorelliola cf. mahunkai Thorelliola ensifera Thorelliola tualapa Thorelliola tamasi Emathis gombak Lagnus edwardsi Lepidemathis haemorrhoidalis Ilargus macrocornis Maeota sp. [JatunSacha] Thyenula wesolowskae Ilargus foliosus Ilargus galianoae Foliabitus longzhou Foliabitus sp. [Malaysia] Maeota tuberculotibiata Thyenula cf. mundus Thyenula cf. aurantiaca Parvattus zhui Mexigonus arizonensis Chapoda montana Chapoda fortuna [Panama] Thyenula sp. [SouthAfrica] Anasaitis banksi Anasaitis placida Anasaitis gloriae Corythalia broccai Corythalia bromelicola Corythalia minor Corythalia peblique Corythalia coronai Corythalia decora Chalcotropis cf. caeruleus Compsodecta haytiensis Pristobaeus cf. jocosus Pristobaeus beccarii Colyttus bilineatus Colyttus striatus Pseudeuophrys erratica Chalcoscirtus diminutus Talavera minuta Plexippus paykulli Freya decorata Thyenula nelshoogte Tylogonus cf. auricapillus Tylogonus yanayacu Heliophanus cupreus Salticus scenicus Popcornella spiniformis Popcornella furcata Popcornella nigromaculata Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Anasaitis brunnea Anasaitis hebetata Anasaitis adorabilis Laufeia daiqini Anasaitis canosa [USA] Anasaitis cf. canalis Corythalia sulfurea Corythalia porphyra Saitis barbipes Saitis sp. [NewSouthWales] Saitis cf. griseus Saitis auripes Saitis sp. [SouthAustralia] Corticattus latus Naphrys pulex Corythalia electa Corythalia bicincta Anasaitis elegantissima Anasaitis laxa Anasaitis locuples Anasaitis sp. [Peblique] Euophrys frontalis Euophrys monodnock Belliena ecuadorica Chalcoscirtus infimus Neonella vinnula Laufeia keyserlingi Laufeia eximia Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Coccorchestes cf. inermis Coccorchestes clavifemur Maeota sp. [Manabi] Chapoda gitae Chapoda angusta Chapoda recondita Antillattus scutiformis Antillattus darlingtoni Antillattus cf. applanatus Antillattus maxillosus Petemathis minuta Antillattus gracilis Antillattus cambridgei Pensacola signata Compsodecta peckhami Viribestus suyanensis Chalcotropis luceroi Canama hinnulea Canama cf. forceps Canama triramosa Canama extranea Canama fimoi Bathippus directus [Tualapa] Bathippus gahavisuka Bathippus korei Bathippus macrognathus Bathippus madang Parabathippus shelfordi Parabathippus kiabau Parabathippus cf. macilentus Parabathippus cuspidatus Parabathippus magnus Truncattus cachotensis Truncattus flavus Euophryine sp. [GentingHighlands] Truncattus dominicanus Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Amphidraus complexus Marma nigritarsis Ilargus coccineus Ilargus pilleolus Ilargus serratus cf. Maeota sp. [Panama] Xenocytaea albomaculata Chinattus parvulus Mexigonus morosus Maeota dorsalis Maeota sp. [MoronaSantiago] Variratina minuta Maeota flava Maeota dichrura Maeota simoni Chapoda cf. inermis [Panama] Chapoda cf. inermis [CostaRica] Chapoda peckhami Agobardus cordiformis Agobardus gramineus Agobardus phylladiphilus Saitis cf. fuscus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus bahoruco Agobardus oviedo Phasmolia elegans Sidusa sp.1 [FrenchGuiana] Sidusa sp.2 [FrenchGuiana] Sidusa sp. [CostaRica] Sidusa extensa Sidusa unicolor Thiania spectrum Efate albobicinctus Sobasina wanlessi Paraharmochirus tualapaensis Chalcolemia nakanai Diolenius varicus Chalcolecta prensitans Ohilimia scutellata * ** ** * * 0.54 0.57 0.72 0.92 0.52 0.59 0.76 0.93 0.69 0.53 0.68 0.6 0.82 0.59 0.93 0.63 0.75 0.570.52 Sidusa Clade Amphidraus-Marma Clade Tylogonus Soesilarishius Ecuadattus 1.0 Colyttus Clade Cytaea-Euryattus Clade Phasmolia Clade Diolenius Clade * New World Old World Outgroup 104! Figure 3.3. Strict consensus of seven equally parsimonious trees (score=1145) from the morphological dataset that found in TNT, with euophryine taxa being enforced as a monophyletic group. Recovered euophryine clades are indicated along branches. New World or Old World distribution of euophryine taxa is indicated in colored blocks in front of the taxon names. Ghelna canadensis Chinattus parvulus Freya decorata Plexippus paykulli Heliophanus cupreus Salticus scenicus 'Bathippus' pahang Xenocytaea albomaculata Chalcotropis cf. caeruleus Parvattus zhui Bythocrotus cf. crypticus Bythocrotus crypticus Ecuadattus napoensis Ecuadattus pichincha Thorelliola Joannae Xenocytaea proszynskii Ilargus pilleolus Ilargus serratus Maeota sp. [JatunSacha] Compsodecta haytiensis Maeota sp. [Napo] Pristobaeus cf. jocosus Pristobaeus beccarii Colyttus bilineatus Colyttus striatus Laufeia daiqini Laufeia keyserlingi Thiania bhamoensis Thiania latibola Thiania tenuis Cytaea mitellata Cytaea nimbata Cytaea oreophila Euryattus bleekeri Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Pseudeuophrys erratica Chalcoscirtus diminutus Talavera minuta Popcornella spiniformis Popcornella furcata Popcornella nigromaculata Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Anasaitis brunnea Anasaitis hebetata Anasaitis adorabilis Anasaitis canosa [USA] Anasaitis cf. canalis cf. Coryphasia sp. [Brazil] Bulolia excentrica Leptathamas paradoxus Zabkattus brevis Zabkattus trapeziformis Zabkattus richardsi Zabkattus furcatus Servaea vestita Thorelliola crebra Mopiopia cf. bruneti Coryphasia fasciiventris Coryphasia physonycha Coryphasia cf. campestrata Thorelliola aliena Lagnus edwardsi Emathis gombak Lepidemathis haemorrhoidalis Corythalia sulfurea Corythalia porphyra Saitis barbipes Maeota sp. [Manabi] Saitis sp. [NewSouthWales] Saitis cf. griseus Saitis auripes Saitis sp. [SouthAustralia] Naphrys pulex Corythalia electa Corythalia bicincta Anasaitis elegantissima Anasaitis laxa Anasaitis locuples Anasaitis sp. [Peblique] Corticattus latus Neonella vinnula Laufeia eximia Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Coccorchestes cf. inermis Coccorchestes clavifemur Mexigonus arizonensis Thyenula sp. [SouthAfrica] Anasaitis banksi Anasaitis placida Anasaitis gloriae Corythalia broccai Corythalia bromelicola Corythalia minor Corythalia peblique Corythalia coronai Corythalia decora Thyenula cf. aurantiaca Ilargus foliosus Ilargus macrocornis Thyenula wesolowskae Maeota tuberculotibiata Thyenula cf. mundus Foliabitus longzhou Foliabitus sp. [Malaysia] Thyenula leighi Compsodecta peckhami Tylogonus cf. auricapillus Tylogonus yanayacu Thorelliola tamasi Thorelliola tualapa Thorelliola cf. mahunkai Thorelliola ensifera Viribestus suyanensis Chalcotropis luceroi Canama hinnulea Canama cf. forceps Canama triramosa Canama extranea Canama fimoi Bathippus directus [Tualapa] Bathippus gahavisuka Bathippus macrognathus Bathippus korei Bathippus madang Parabathippus shelfordi Parabathippus kiabau Parabathippus cf. macilentus Parabathippus cuspidatus Parabathippus magnus Belliena ecuadorica Chalcoscirtus infimus Euophrys frontalis Euophrys monodnock Chapoda montana Saphrys a-notata Saphrys cf. patagonica Chinophrys pengi Ilargus moronatigus Thyenula laxa Ilargus galianoae Thyenula nelshoogte Xenocytaea agnarssoni Omoedus papuanus Omoedus darleyorum Omoedus meyeri Omoedus omundseni Omoedus cf. semirasus Omoedus cf. torquatus Omoedus cf. danae Omoedus swiftorum Omoedus brevis Omoedus orbiculatus Omoedus cf. piceus Omoedus ephippigera Chapoda fortuna [Panama] Ilargus coccineus Mexigonus morosus cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [MoronaSantiago] Variratina minuta Maeota flava Maeota dichrura Maeota simoni Chapoda angusta Chapoda gitae Chapoda recondita Antillattus scutiformis Antillattus cf. applanatus Antillattus darlingtoni Antillattus maxillosus Pensacola signata Petemathis minuta Antillattus gracilis Antillattus cambridgei Truncattus flavus Truncattus cachotensis Euophryine sp. [GentingHighlands] Truncattus dominicanus Amphidraus complexus Marma nigritarsis Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Chapoda cf. inermis [Panama] Chapoda cf. inermis [CostaRica] Chapoda peckhami Agobardus cordiformis Agobardus gramineus Agobardus phylladiphilus Saitis cf. fuscus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus bahoruco Agobardus oviedo Phasmolia elegans Sidusa sp.1 [FrenchGuiana] Sidusa sp.2 [FrenchGuiana] Sidusa sp. [CostaRica] Sidusa extensa Sidusa unicolor Thiania spectrum Efate albobicinctus Sobasina wanlessi Paraharmochirus tualapaensis Chalcolemia nakanai Diolenius varicus Chalcolecta prensitans Ohilimia scutellata * Euophryinae ** ** ** Amphidraus-MarmaClade Tylogonus Soesilarishius Ecuadattus Emathis-Lepidemathis Clade Colyttus Clade Foliabitus Cytaea-Euryattus Clade Phasmolia Clade Diolenius Clade * New World Old World Outgroup 105! Figure 3.4. The best tree from ML analysis on the DNA dataset (28S, Actin 5C, 16S-ND1 and COI) with lnL as -135203.637. Recovered euophryine clades and ML bootstrap support values are indicated along branches. New World or Old World distribution of euophryine taxa is indicated in colored blocks in front of the taxon names. Ghelna canadensis Freya decorata Salticus scenicus 'Bathippus' pahang Plexippus paykulli Chinattus parvulus Heliophanus cupreus Servaea vestita Phasmolia elegans Zabkattus furcatus Zabkattus richardsi Zabkattus brevis Zabkattus trapeziformis Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Bathippus macrognathus Bathippus directus [Tualapa] Bathippus gahavisuka Bathippus korei Bathippus madang Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Pristobaeus cf. jocosus Pristobaeus beccarii Xenocytaea agnarssoni Xenocytaea proszynskii Xenocytaea albomaculata Chalcolemia nakanai Sobasina wanlessi Paraharmochirus tualapaensis Efate albobicinctus Chalcolecta prensitans Diolenius varicus Ohilimia scutellata Omoedus cf. piceus Omoedus ephippigera Omoedus orbiculatus Omoedus meyeri Omoedus omundseni Omoedus darleyorum Omoedus papuanus Omoedus swiftorum Omoedus cf. semirasus Omoedus brevis Omoedus cf. torquatus Omoedus cf. danae Saitis auripes Saitis cf. fuscus Saitis barbipes Saitis sp. [SouthAustralia] Saitis sp. [NewSouthWales] Saitis cf. griseus Chalcotropis cf. caeruleus Chalcotropis luceroi Lagnus edwardsi Euophryine sp. [GentingHighlands] Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola tamasi Thorelliola ensifera Thorelliola tualapa Thorelliola cf. mahunkai Emathis gombak Lepidemathis haemorrhoidalis Colyttus bilineatus Colyttus striatus Foliabitus sp. [Malaysia] Foliabitus longzhou Laufeia daiqini Laufeia eximia Laufeia keyserlingi Thiania spectrum Thiania latibola Thiania tenuis Thiania bhamoensis Ecuadattus napoensis Ecuadattus pichincha Belliena ecuadorica Ilargus galianoae Ilargus pilleolus Ilargus serratus Ilargus coccineus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Tylogonus yanayacu Tylogonus cf. auricapillus Marma nigritarsis Amphidraus complexus Chinophrys pengi Soesilarishius cf. amrishi Soesilarishius ruizi Soesilarishius micaceus cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Coryphasia physonycha Coryphasia cf. campestrata Neonella vinnula Mopiopia cf. bruneti Saphrys a-notata Saphrys cf. patagonica Thyenula laxa Thyenula nelshoogte Thyenula wesolowskae Thyenula leighi Thyenula sp. [SouthAfrica] Thyenula cf. mundus Thyenula cf. aurantiaca Cytaea oreophila Cytaea nimbata Cytaea mitellata Euryattus bleekeri Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Parvattus zhui Parabathippus kiabau Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Euophrys frontalis Euophrys monodnock Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Sidusa sp. [CostaRica] Sidusa extensa Sidusa unicolor Corythalia porphyra Corythalia sulfurea Corythalia electa Corythalia bicincta Corythalia minor Corythalia broccai Corythalia peblique Corythalia bromelicola Corythalia coronai Corythalia decora Anasaitis canosa [USA] Anasaitis cf. canalis Anasaitis adorabilis Anasaitis hebetata Anasaitis brunnea Anasaitis banksi Anasaitis placida Anasaitis gloriae Anasaitis laxa Anasaitis elegantissima Anasaitis sp. [Peblique] Anasaitis locuples Pensacola signata Mexigonus arizonensis Mexigonus morosus Corticattus latus Naphrys pulex Popcornella nigromaculata Popcornella spiniformis Popcornella furcata Compsodecta haytiensis Compsodecta peckhami Bythocrotus crypticus Bythocrotus cf. crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo Petemathis minuta Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis Antillattus cambridgei Antillattus darlingtoni Antillattus scutiformis Antillattus maxillosus Chapoda recondita Chapoda gitae Chapoda angusta Chapoda fortuna [Panama] Chapoda montana Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota sp. [Napo] Maeota dichrura Maeota flava Maeota simoni Maeota sp. [MoronaSantiago] Maeota tuberculotibiata Euophryinae ** ** ** * Anasaitis-Corythalia Clade Euophrys Clade Parabathippus-Parvattus Clade Sidusa Clade Pensacola-Mexigonus Clade Antillattus Clade Naphrys-Corticattus Clade Popcornella Agobardus Clade Chapoda-Maeota Clade Tylogonus Soesilarishius Mopiopia-Saphrys Clade Coryphasia Clade Ecuadattus Ilargus Clade Saitis Clade Emathis-Lepidemathis Clade Colyttus Clade Thorelliola Foliabitus Laufeia Clade Thiania Clade Thyenula Clade Cytaea-Euryattus Clade Phasmolia Clade Xenocytaea Pristobaeus Clade Diolenius Clade Omoedus Clade Zabkattus Clade Bulolia-Coccorchestes Clade Bathippus-Canama Clade 0.94 0.73 0.97 0.91 1.0 1.0 1.0 0.98 0.66 0.99 0.99 0.74 1.0 0.85 1.0 1.0 0.87 0.96 1.0 0.95 1.0 1.0 0.98 1.0 0.76 0.94 1.0 1.0 1.0 0.89 0.99 0.90 0.59 1.0 1.0 0.89 0.87 0.86 0.99 1.0 1.0 0.96 0.96 1.0 1.0 0.98 1.0 0.86 0.99 * New World Old World Outgroup 106! Figure 3.5. The best tree from ML analysis on the combined morphology and DNA dataset (morphology, 28S, Actin 5C, 16S-ND1 and COI) with lnL as -140981.450. Recovered euophryine clades and ML bootstrap support values are indicated along branches. New World or Old World distribution of euophryine taxa is indicated in colored blocks in front of the taxon names. Ghelna canadensis Chinattus parvulus 'Bathippus' pahang Freya decorata Plexippus paykulli Heliophanus cupreus Salticus scenicus Saitis auripes Saitis cf. fuscus Saitis barbipes Saitis sp. [SouthAustralia] Saitis sp. [NewSouthWales] Saitis cf. griseus Colyttus bilineatus Colyttus striatus Foliabitus longzhou Foliabitus sp. [Malaysia] Laufeia daiqini Laufeia eximia Laufeia keyserlingi Thiania spectrum Thiania tenuis Thiania bhamoensis Thiania latibola Chalcotropis luceroi Lagnus edwardsi Emathis gombak Lepidemathis haemorrhoidalis Euophryine sp. [GentingHighlands] Thorelliola aliena Thorelliola Joannae Thorelliola crebra Thorelliola cf. mahunkai Thorelliola tualapa Thorelliola ensifera Thorelliola tamasi Chalcotropis cf. caeruleus Amphidraus complexus Marma nigritarsis Soesilarishius cf. amrishi Soesilarishius micaceus Soesilarishius ruizi Mopiopia cf. bruneti Saphrys a-notata Saphrys cf. patagonica cf. Coryphasia sp. [Brazil] Coryphasia fasciiventris Coryphasia physonycha Coryphasia cf. campestrata Neonella vinnula Ecuadattus napoensis Ecuadattus pichincha Belliena ecuadorica Ilargus galianoae Ilargus pilleolus Ilargus coccineus Ilargus serratus Ilargus moronatigus Ilargus foliosus Ilargus macrocornis Thyenula laxa Thyenula nelshoogte Servaea vestita Cytaea oreophila Cytaea mitellata Cytaea nimbata Euryattus bleekeri Euryattus sp.1 [Gahavisuka] Euryattus sp.2 [Gahavisuka] Chinophrys pengi Tylogonus cf. auricapillus Tylogonus yanayacu Thyenula sp. [SouthAfrica] Thyenula wesolowskae Thyenula leighi Thyenula cf. mundus Thyenula cf. aurantiaca Chalcoscirtus infimus Chalcoscirtus diminutus Talavera minuta Pseudeuophrys erratica Euophrys frontalis Euophrys monodnock Sidusa sp.2 [FrenchGuiana] Sidusa sp.1 [FrenchGuiana] Sidusa sp. [CostaRica] Sidusa extensa Sidusa unicolor Parvattus zhui Parabathippus kiabau Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Phasmolia elegans Bathippus macrognathus Bathippus directus [Tualapa] Bathippus gahavisuka Bathippus korei Bathippus madang Canama cf. forceps Canama hinnulea Canama fimoi Canama extranea Canama triramosa Zabkattus furcatus Zabkattus richardsi Zabkattus brevis Zabkattus trapeziformis Viribestus suyanensis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes clavifemur Coccorchestes cf. inermis Coccorchestes cf. aiyura Coccorchestes cf. ildikoae Pristobaeus beccarii Pristobaeus cf. jocosus Xenocytaea agnarssoni Xenocytaea albomaculata Xenocytaea proszynskii Chalcolemia nakanai Chalcolecta prensitans Diolenius varicus Ohilimia scutellata Sobasina wanlessi Efate albobicinctus Paraharmochirus tualapaensis Omoedus orbiculatus Omoedus cf. piceus Omoedus ephippigera Omoedus papuanus Omoedus darleyorum Omoedus meyeri Omoedus omundseni Omoedus cf. semirasus Omoedus swiftorum Omoedus brevis Omoedus cf. torquatus Omoedus cf. danae Corythalia porphyra Corythalia sulfurea Corythalia bicincta Corythalia electa Corythalia broccai Corythalia minor Corythalia peblique Corythalia bromelicola Corythalia coronai Corythalia decora Anasaitis canosa [USA] Anasaitis cf. canalis Anasaitis adorabilis Anasaitis brunnea Anasaitis hebetata Anasaitis banksi Anasaitis placida Anasaitis gloriae Anasaitis laxa Anasaitis elegantissima Anasaitis locuples Anasaitis sp. [Peblique] Pensacola signata Mexigonus arizonensis Mexigonus morosus Naphrys pulex Corticattus latus Popcornella nigromaculata Popcornella furcata Popcornella spiniformis Compsodecta haytiensis Compsodecta peckhami Bythocrotus cf. crypticus Bythocrotus crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo Petemathis minuta Petemathis tetuani Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis Antillattus cambridgei Antillattus darlingtoni Antillattus maxillosus Antillattus scutiformis Chapoda recondita Chapoda gitae Chapoda angusta Chapoda montana Chapoda fortuna [Panama] Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Chapoda peckhami cf. Maeota sp. [Panama] Maeota dorsalis Maeota sp. [JatunSacha] Maeota sp. [Manabi] Maeota dichrura Maeota sp. [Napo] Maeota flava Maeota simoni Maeota sp. [MoronaSantiago] Maeota tuberculotibiata ** ** ** * Anasaitis-Corythalia Clade Euophrys Clade Euophryinae Parabathippus-Parvattus Clade Sidusa Clade Pensacola-Mexigonus Clade Antillattus Clade Naphrys-Corticattus Clade Popcornella Agobardus Clade Chapoda-Maeota Clade Amphidraus-Marma Clade Tylogonus Soesilarishius Mopiopia-Saphrys Clade Coryphasia Clade Ecuadattus Ilargus Clade Saitis Clade Emathis-Lepidemathis Clade Colyttus Clade Thorelliola Foliabitus Laufeia Clade Thiania Clade Cytaea-Euryattus Clade Phasmolia Clade Zabkattus Clade Bulolia-Coccorchestes Clade Bathippus-Canama Clade Pristobaeus Clade Xenocytaea Diolenius Clade Omoedus Clade 0.95 0.54 0.88 0.97 0.99 1.0 1.0 1.0 1.0 0.91 1.0 0.70 0.90 0.70 1.0 0.95 1.0 1.0 0.85 1.0 0.98 1.0 1.0 0.99 0.99 1.0 1.0 1.0 0.98 1.0 1.0 1.0 1.0 1.0 0.96 0.65 1.0 0.92 1.0 1.0 0.93 0.68 1.0 0.94 1.0 0.99 0.67 1.0 1.0 0.93 0.96 * New World Old World Outgroup 107! Figure 3.6. Anasaitis-Corythalia Clade. A-G. Anasaitis canosa; H-N. Anasaitis cf. canalis; O- U. Anasaitis placida. A, H, O, male, dorsal view; B, I, P, female, dorsal view; C, J, Q, male left palp, ventral view; D, K, male left palp, retrolateral view; R, male left palp, prolateral view; E, S, male left chelicera, back view; L, female left chelicera, back view; F, M, T, epigynum, ventral view; G, N, U, cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; H-I, O-P, 0.5 mm; E, K, S, 0.2 mm; C-D, F-G, J-K, M-N, Q-R, T-U, 0.1 mm. 108! Figure 3.7. Anasaitis-Corythalia Clade. A-G. Corythalia electa; H-N. Corythalia minor; O-U. Corythalia decora. A, H, O, male, dorsal view; B, I, P, female, dorsal view; C, J, Q, male left palp, ventral view; D, K, R, male left chelicera, back view; E, female carapace, dorsal view; L, S, male carapace, dorsal view; F, M, T, epigynum, ventral view; G, N, U, cleared epigynum, dorsal view. Scale bars: A-B, H-I, O-P, 1.0 mm; D-E, K-L, R-S, 0.2 mm; C, F-G, J, M-N, Q, T- U, 0.1 mm. 109! Figure 3.8. Antillattus Clade. A-G. Antillattus gracilis; H-N. Antillattus cambridgei. A, H, male, dorsal view; B, I, female, dorsal view; C, J, male left palp, ventral view; D, male left chelicera, back view; E, male left chelicera, front view; K, female right chelicera, back view; L, male right chelicera, front view; F, M, epigynum, ventral view; G, N, cleared epigynum, dorsal view. Scale bars: A-B, H-I, 2.0 mm; C-G, J-L, 0.2 mm; M-N, 0.1 mm. 110! Figure 3.9. Antillattus Clade. A-H. Antillattus darlingtoni; I-P. Antillattus scutiformis. A, I, male, dorsal view; B, J, female, dorsal view; C, K, male left palp, ventral view; D, L, male left chelicera, front view; E, M, male left chelicera, back view; F, male endites and labium, ventral view; N, female left chelicera, back view; G, O, epigynum, ventral view; O, P, cleared epigynum, dorsal view. Scale bars: A-B, I-J, 1.0 mm; C-F, K-N, 0.2 mm; G-H, O-P, 0.1 mm. 111! Figure 3.10. Antillattus Clade. A-G. Petemathis minuta; H-M. Petemathis portoricensis; N-S, Truncattus flavus. A, H, N, male, dorsal view; B, I, O, female, dorsal view; C, J, P, male left palp, ventral view; D, K, male left chelicera, back view; E, male left chelicera, front view; Q, male right chelicera, back view; F, L, R, epigynum, ventral view; G, M, S, cleared epigynum, dorsal view. Scale bars: A-B, N-O, 0.5 mm; H-I, 2.0 mm; D-E, J-M, 0.2 mm; C, F-G, P-S, 0.1 mm. 112! Figure 3.11. Antillattus Clade. A-C. Allodecta maxillaris. A, male left palp, ventral view; B, male left chelicera, front view; C, male left chelicera, back view. Scale bars: A-C, 0.2 mm. 113! Figure 3.12. Agobardus Clade. A-F. Agobardus cf. anormalis montanus; G-M. Bythocrotus crypticus. A, G, male, dorsal view; B, H, female, dorsal view; C, I, male left palp, ventral view; J, male left palp, prolateral view; D, K, male left chelicera, back view; E, L, epigynum, ventral view; F, M, cleared epigynum, dorsal view. Scale bars: A-B, G-H, 1.0 mm; C-F, I-M, 0.2 mm. 114! Figure 3.13. Agobardus Clade. A-H. Compsodecta peckhami; I-P. Compsodecta haytiensis. A, I, male, dorsal view; B, J, female, dorsal view; C, K, male left palp, ventral view; D, male left palp, prolateral view; L, male left palpal tibia, retrolateral view; E, male left chelicera, front view; F, male endites and labium, ventral view; M, male left chelicera, back view; N, male right chelicera, front view; G, O, epigynum, ventral view; H, P, cleared epigynum, dorsal view. Scale bars: A-B, I-J, 1.0 mm; C-F, K-N, 0.2 mm; G-H, O-P, 0.1 mm. 115! Figure 3.14. Agobardus Clade. A-D. Compsodecta grisea; E-I. Compsodecta festiva. A, E, male left palp, ventral view; B, male left palp, retrolateal view; F, male left palp, prolateral view; C, male right chelicera, front view; G. male left chelicera, front view; H, male endites and labium, ventral view; D, I, epigynum, ventral view. Scale bars: A-I, 0.2 mm. 116! Figure 3.15. Naphrys-Corticattus Clade. A-E. Naphrys pulex; F-K. Corticattus latus. A, F, male, dorsal view; B, G, female, dorsal view; C, H, male left palp, ventral view; I, female right chelicera, back view; D, J, epigynum, ventral view; E, K, cleared epigynum, dorsal view. Scale bars: A-B, F-G, 0.5 mm; C-E, 0.2 mm; H-K, 0.1 mm. 117! Figure 3.16. Sidusa Clade. A-G. Sidusa cf. extensa; H-N. Sidusa sp.2 [French Guiana]; O-T. Sidusa sp.1 [French Guiana]. A, H, O, male, dorsal view; B, I, P. female, dorsal view; C, J, Q, male left palp, ventral view; D, K, R, male left palp, retrolateral view; E, male left chelicera, front view; L, female left chelicera, back view; F, M, S, epigynum, ventral view; G, N, T, cleared epigynum, dorsal view. Scale bars: A, 2.0 mm; B, H-I, O-P, 1.0 mm; C-G, J-N, Q-T, 0.2 mm. 118! Figure 3.17. Mopiopia-Saphrys Clade. A-D. Mopiopia cf. bruneti; E-J. Saphrys tehuelche; K- Q. Saphrys mapuche. A, E, K, male, dorsal view; F, L. female, dorsal view; B, G, M, male left palp, ventral view; H, N, male left palp, retrolateral view; C, male left chelicera, front view; O, female left chelicera, back view; D, male endites and labium, ventral view; I, P, epigynum, ventral view; J, Q, cleared epigynum, dorsal view. Scale bars: A, E, K, L, 1.0 mm; F, 2.0 mm; B-D, 0.2 mm; G-J, M-Q, 0.1 mm. 119! Figure 3.18. Chapoda-Maeota Clade. A-H. Chapoda cf. festiva; I-L. Chapoda peckhami; M-R. Chapoda recondita. A, I, M, male, dorsal view; B, N, female, dorsal view; C, J, O, male left palp, ventral view; D, K, male left palp, prolateral view; E, L, P, male left palp, retrolateral view; F, female left chelicera, back view; G, Q, epigynum, ventral view; H, R, cleared epigynum, dorsal view. Scale bars: A-B, I, M-N, 1.0 mm; C-F, J-L, O-P, 0.2 mm; G-H, Q-R, 0.1 mm. 120! Figure 3.19. Chapoda-Maeota Clade. A-D. Maeota dichrura. A, male, dorsal view; B, male left palp, ventral view; C, male left palp, retrolateral view; D, male right endite and right chelicera, ventral view; E, epigynum, ventral view. Scale bars: A, 0.5 mm; B-E, 0.2 mm. 121! Figure 3.20. Amphidraus-Marma Clade. A-F. Amphidraus complexus; G-L. Marma nigritarsis. A, G, male, dorsal view; B, H, female, dorsal view; C, I, male left palp, ventral view; D, male left palp, retrolateral view; J, male left chelicera, back view; E, K, epigynum, ventral view; F, L, cleared epigynum, dorsal view. Scale bars: A-B, G-H, 0.5 mm; C-F, I-L, 0.1 mm. 122! Figure 3.21. Coryphasia Clade. A-G. Coryphasia albibarbis; H-O. Coryphasia physonycha; P- V. Coryphasia cf. campestrata. A, H, P, male, dorsal view; B, I, Q, female, dorsal view; C, J, R, male left palp, ventral view; D, K, S, male left palp, retrolateral view; E, M, T, female left chelicera, back view; L, male left chelicera, back view; F, N, U, epigynum, ventral view; G, O, V, cleared epigynum, dorsal view. Scale bars: A-B, H-I, Q, 2.0 mm; P, 1.0 mm; C-G, J-O, R-V, 0.2 mm. 123! Figure 3.22. Pensacola-Mexigonus Clade. A-G. Pensacola signata; H-N. Mexigonus arizonensis. A, H, male, dorsal view; B, I, female, dorsal view; C, J, male left palp, ventral view; D, male right endite, ventral view; E, male right chelicera, back view; K, male left chelicera, front view; L. male left chelicera, back view; F, M, epigynum, ventral view; G, N, cleared epigynum, dorsal view. Scale bars: A, I, Q, 1.0 mm; B, H, 0.5 mm; C-G, J-N, 0.2 mm. 124! Figure 3.23. Neonella Clade. A-G. Neonella vinnula. A, male, dorsal view; B, female, dorsal view; C, male left palp, ventral view; D, male left palp, retrolateral view; E, male left chelicera, back view; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A, B, 0.2 mm; C-G, 0.05 mm. 125! Figure 3.24. Belliena Clade. A-G. Belliena ecuadorica. A. male, dorsal view; B. female, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. 126! Figure 3.25. Soesilarishius Clade. A-D. Soesilarishius cf. amrishi; E-J. Soesilarishius ruizi. A, E, male, dorsal view; F, female, dorsal view; B, G, male left palp, ventral view; C, H, male left palp, retrolateral view; D, male left chelicera, back view; I, epigynum, ventral view; J, cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B-D, 0.2 mm; E-F, 0.5 mm; G-J, 0.1 mm. 127! Figure 3.26. Ecuadattus. A-G. Ecuadattus typicus. A. male, dorsal view; B. female, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. female left chelicera, back view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-F, 0.2 mm; G-H, 0.1 mm. 128! Figure 3.27. Ilargus. A-G. Ilargus coccineus. A, male, dorsal view; B, female, dorsal view; C, male left palp, ventral view; D, male left palp, retrolateral view; E, female left chelicera, back view; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. 129! Figure 3.28. Popcornella. A-G. Popcornella spiniformis. A. male, dorsal view; B. female, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. 130! Figure 3.29. Tylogonus. A-C. Tylogonus cf. auricapillus; D-K. Tylogonus yanayacu. A, D, male, dorsal view; E, female, dorsal view; B, F, male left palp, ventral view; C, G, male left palp, retrolateral view; H, male right chelicera, front view; I. male right chelicera, back view; J, epigynum, ventral view; K, cleared epigynum, dorsal view. Scale bars: A, D-E, 0.5 mm; B-C, F- K, 0.2 mm. 131! Figure 3.30. Bathippus-Canama Clade. A-J. Bathippus macrognathus; K-Q. Canama forceps; R-Y. Canama hinnulea. A, K, R, male, dorsal view; B, female, dorsal view; C, L, S, male left palp, ventral view; D, M, T, male left palp, retrolateral view; E, N, male left palp with femur and patella, retrolateral view; U, male left palpal femur, retrolateral view; F, O, V, male left chelicera, dorsal view; G, P, W, male left chelicera, ventral view; H. female left chelicera, back view; X, male left chelicera, outer lateral view; Q, Y, male endites, labium, sternum, coxae of legs, ventral view; I, epigynum, ventral view; J, cleared epigynum, dorsal view. Scale bars: A-B, K, R, 2.0 mm; C-H, L-Q, S-Y, 0.2 mm; I-J, 0.1 mm. 132! Figure 3.31. Omoedus Clade. A-D. Omoedus cf. piceus; E-K. Omoedus ephippigerus; K-Q. Omoedus orbiculatus. A, E, L, male, dorsal view; F, M, female, dorsal view; B, G, N, male left palp, ventral view; C, H, O, male left palp, retrolateral view; D, I, male left chelicera, back view; J, P, epigynum, ventral view; K, Q, cleared epigynum, dorsal view. Scale bars: A, 0.5 mm; E-F, L-M, 1.0 mm; B-D, G-K, N-Q, 0.2 mm. 133! Figure 3.32. Bulolia-Coccorchestes Clade. A-H. Bulolia excentrica; I-P. Leptathamas paradoxus. A, I, male, dorsal view; B, J, female, dorsal view; C, K, male left palp, ventral view; E, L, male left palp, retrolateral view; D, male left palp, prolateral view; F, female left chelicera, backl view; M, male right chelicera, back view; N, male right chelicera, front view; G, O, epigynum, ventral view; H, P, cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B, I-J, 0.5 mm; C-F, K-N, 0.2 mm; G-H, O-P, 0.1 mm. 134! Figure 3.33. Bulolia-Coccorchestes Clade. A-G. Coccorchestes cf. aiyura; H-M. Variratolia minuta. A, H, male, dorsal view; B, I, female, dorsal view; C, J, male left palp, ventral view; D, K, male left palp, retrolateral view; E, female left chelicera, back view; F, L, epigynum, ventral view; G, M, cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B, H, 0.5 mm; I, 0.2 mm; C- G, J-M, 0.1 mm. 135! Figure 3.34. Diolenius Clade. A-G. Diolenius varicus; H-N. Ohilimia scutellata. A, H, male, dorsal view; B, I, female, dorsal view; C, J, male left palp, ventral view; D, K, male left palp, retrolateral view; E, L, female right chelicera, back view; F, M, epigynum, ventral view; G, N, cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-G, J-N, 0.2 mm. 136! Figure 3.35. Diolenius Clade. A-E. Chalcolemia nakanai; F-H. Chalcolecta prensitans. I-K. Efate albobicinctus. A, F, female, dorsal view; I, male, dorsal view; B, G, epigynum, ventral view; D, H, cleared epigynum, dorsal view; C, cleared epigynum, ventral view; E, female right chelicera, back view; J, male left palp, ventral view; K, male left palp, retrolateral view. Scale bars: A, F, 1.0 mm; I, 0.5 mm; E, 0.2 mm; B-D, G-H, J-K, 0.1 mm. 137! Figure 3.36. Diolenius Clade. A-G. Paraharmochirus tualapaensis; H-N. Sobasina wanlessi. A, H, male, dorsal view; B, I, female, dorsal view; C, J, male left palp, ventral view; D, male left palp, retrolateral view; E, female left chelicera, back view; L, male left chelicera, back view; K, male right leg I, retrolateral view; F, M, epigynum, ventral view; G, N, cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; H-I, 0.5 mm; K, 0.2 mm; C-G, L-N, 0.1 mm 138! Figure 3.37. Pristobaeus Clade. A-G. Pristobaeus cf. jocosus; H-N. Pristobaeus beccarii. A, H, male, dorsal view; B, I, female, dorsal view; C, J, male left palp, ventral view; K, male left palp, retrolateral view; D, male left chelicera, back view; E, female right chelicera, back view; L, female left chelicera, back view; F, M, epigynum, ventral view; G, N, cleared epigynum, dorsal view. Scale bars: A-B, H-I, 2.0 mm; C-G, J-N, 0.2 mm. 139! Figure 3.38. Phasmolia Clade. A-G. Phasmolia elegans. A. male, dorsal view; B. female, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female right chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. 140! Figure 3.39. Parabathippus-Parvattus Clade. A-H. Parabathippus shelfordi; I-M. Parvattus zhui. A, I, male, dorsal view; B, female, dorsal view; C, J, male left palp, ventral view; D, K, male left palp, retrolateral view; E, male left chelicera, inner lateral view; F, female left chelicera, back view; L, male endites, labium and chelicerae, ventral view; M, male carapace, lateral view; G, epigynum, ventral view; H, cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; I, 0.5 mm; C-H, L-M, 0.2 mm; J-K, 0.1 mm. 141! Figure 3.40. Euophrys Clade. A-E. Chalcoscirtus infimus; F-K. Euophrys frontalis. A, F, male, dorsal view; B, G, female, dorsal view; C, H, male left palp, ventral view; I, female right chelicera, back view; D, J, epigynum, ventral view; E, K, cleared epigynum, dorsal view. Scale bars: A-B, F-G, 0.5 mm; C-E, H-K, 0.1 mm. 142! Figure 3.41. Euophrys Clade. A-E. Pseudeuophrys erratica; F-J. Talavera minuta. A, F, male, dorsal view; B, G, female, dorsal view; C, H, male left palp, ventral view; D, I, epigynum, ventral view; E, J, cleared epigynum, dorsal view. Scale bars: A-B, F-G, 0.5 mm; C-E, H-J, 0.1 mm. 143! Figure 3.42. Saitis Clade. A-F. Saitis barbipes; G-L. Saitis volans. A, G, male, dorsal view; B, H, female, dorsal view; C, I, male left palp, ventral view; D, J, male left palp, retrolateral view; E, K, epigynum, ventral view; F, L, cleared epigynum, dorsal view. Scale bars: A, G, 0.5 mm; B, H, 1.0 mm; C-F, I-L, 0.2 mm. 144! Figure 3.43. Saitis Clade. A-F. Saitis auripes; G-I. Saitis sp.; J-M. Saitis cf. fuscus. A, G, male, dorsal view; B, J, female, dorsal view; C, H, male left palp, ventral view; D, I, male left palp, retrolateral view; K, female right chelicera, back view; E, L, epigynum, ventral view; F, M, cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; G, J, 1.0 mm; C-F, H-I, K-M, 0.2 mm. 145! Figure 3.44. Laufeia Clade. A-F. Laufeia aenea; G-M. Laufeia daiqini; N-T. Laufeia keyserlingi. A, G, N, male, dorsal view; B, H, O, female, dorsal view; C, I, P, male left palp, ventral view; D, J, Q, male left palp, retrolateral view; E, K, R, female left chelicera, back view; F, L, S, epigynum, ventral view; M, T, cleared epigynum, dorsal view. Scale bars: A-B, N-O, 1.0 mm; G-H, 0.5 mm; C-F, I-M, P-T, 0.2 mm. 146! Figure 3.45. Colyttus Clade. A-G. Colyttus bilineatus; H-N. Colyttus cf. lehtineni; O-U. Colyttus striatus. A, H, O, male, dorsal view; B, I, P, female, dorsal view; C, J, Q, male left palp, ventral view; D, K, R, male left palp, retrolateral view; E, L, S, female left chelicera, back view; F, M, T, epigynum, ventral view; G, N, U, cleared epigynum, dorsal view. Scale bars: A- B, H-I, O-P, 2.0 mm; C-G, J-N, Q-U, 0.2 mm. 147! Figure 3.46. Cytaea-Euryattus Clade. A-G. Cytaea nimbata; H-N. Cytaea oreophila; O-U. Euryattus bleekeri. A, H, O, male, dorsal view; B, I, P, female, dorsal view; C, J, Q, male left palp, ventral view; D, K, R, male left palp, retrolateral view; E, L, female left chelicera, back view; S, male left chelicera, back view; F, M, T, epigynum, ventral view; G, N, U, cleared epigynum, dorsal view. Scale bars: A-B, H-I, O-P, 2.0 mm; C-G, J-N, Q-U, 0.2 mm. 148! Figure 3.47. Thiania Clade. A-G. Thiania bhamoensis; H-L. Thiania spectrum. A, H, male, dorsal view; B, female, dorsal view; C, I, male left palp, ventral view; D, J, male left palp, retrolateral view; E, female left chelicera, back view; K, male left chelicera, back view; L, male right first leg; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; H, 1.0 mm; C-G, I-L, 0.2 mm. 149! Figure 3.48. Emathis-Lepidemathis Clade. A-F. Emathis gombak; G-L. Lepidemathis sericea. A, G, male, dorsal view; B, H, female, dorsal view; C, I, male left palp, ventral view; D, J, male left chelicera, back view; E, F, epigynum, ventral view; K, L, cleared epigynum, dorsal view. Scale bars: A, G-H, 2.0 mm; B, 1.0 mm; C-F, I-L, 0.2 mm. 150! Figure 3.49. Thyenula Clade. A-G. Thyenula wesolowskae. A, male, dorsal view; B, female, dorsal view; C, male left palp, ventral view; D, male left palp, retrolateral view; E, female left chelicera, back view; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.2 mm. 151! Figure 3.50. Chalcotropis. A-E. Chalcotropis acutefrenata. A, male, dorsal view; B, male left palp, ventral view; C, male left palp, retrolateral view; D, male right chelicera, back view; E, male right chelicera, front view. Scale bars: A, 2.0 mm; B-E, 0.2 mm. 152! Figure 3.51. Chinophrys. A-G. Chinophrys pengi. A, male, dorsal view; B, female paratype, dorsal view; C, male left palp, ventral view; D, male left palp, retrolateral view; E, male left chelicera, back view; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. 153! Figure 3.52. Foliabitus. A-G. Foliabitus longzhou. A, male, dorsal view; B, female, dorsal view; C, male left palp, ventral view; D, male left palp, retrolateral view; E, male left chelicera, back view; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. 154! Figure 3.53. Lagnus. A-F. Lagnus edwardsi. A, male, dorsal view; B, female, dorsal view; C, male left palp, ventral view; D, male left chelicera, back view; E, epigynum, ventral view; F, cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-F, 0.2 mm. 155! Figure 3.54. Thorelliola. A-H. Thorelliola ensifera. A, male, dorsal view; B, female, dorsal view; C, male left palp, ventral view; D, male left palp, retrolateral view; E, male left chelicera, front view; F, female left chelicera, back view; G, epigynum, ventral view; H, cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-F, 0.2 mm; G-H, 0.1 mm. 156! Figure 3.55. Viribestus. A-D. Viribestus suyanensis. A, male, dorsal view; B, male cephalothorax, front view; C, male left palp, ventral view; D, male left palp, retrolateral view. Scale bars: A, 1.0 mm; B-D, 0.2 mm. 157! Figure 3.56. Xenocytaea. A-C. Xenocytaea albomaculata; D-E. Xenocytaea proszynskii. A, male, dorsal view; B, male left palp, ventral view; C, male left palp, retrolateral view; D, female, dorsal view; E, epigynum, ventral view; F, cleared epigynum, dorsal view. Scale bars: A, D, 0.5 mm; B-C, E-F, 0.1 mm. 158! Figure 3.57. Zabkattus. A-G. Zabkattus brevis. A, male, dorsal view; B, female, dorsal view; C, male left chelicera, front view; D, male left palp, ventral view; E, male left palp, retrolateral view; F, epigynum, ventral view; G, cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C, 0.2 mm; D-G, 0.1 mm. 159! Figure 3.58. Expanded male left palp. A. Chapoda angusta; B. Colyttus striatus; C. Corythalia electa; D. Ilargus pilleolus; E. Laufeia keyserlingi; F. Parabathippus shelfordi; G. Parvattus zhui; H. Sidusa sp.1 [French Guiana]; I. Tylogonus yanayacu. Scale bars: A-I, 0.2 mm. 160! 4 Intersexual correlated evolution of genitalia in euophryine jumping spiders (Araneae: Salticidae): sexual selection or “lock- and-key”? 4.1 Synopsis A long-standing question in spider evolution is the extent to which genitalia are sexually selected or serve as species recognition mechanisms (“lock-and-key”). The first step to investigate this question is to see if female and male genitalic traits are coevolved, which is predicted by either mechanism. In this study, I investigated the lengths of the male embolus and female copulatory duct of euophryine jumping spiders, and found they are positively correlated among the 142 species studied. This correlation confirms an interaction between them, but it does not indicate the selective mechanism involved. In an attempt to narrow down the possible mechanisms, I investigated intra-specific variation of traits in two euophryine species that show considerable difference in sexual dimorphism: Chapoda recondita and Antillattus cambridgei. The \"lock-and-key\" mechanism predicted genitalia would show lower intra-specific variation relative to somatic body parts; whereas, literature suggested if post-copulatory sexual selection were acting on genitalia, they would show higher intra-specific variation than somatic body parts. Intra-specific variation of embolus and copulatory duct in these two species shows negative allometry — females or males tend to retain a common copulatory duct or embolus size even as their bodies vary. However, while conservation of size within species may appear to favor the “lock-and-key” scenario, it could also occur via post-copulatory sexual selection. The size-corrected intra-specific variation is high for genitalia, which has been argued to indicate sexual selection, but could also arise through developmental mechanisms to achieve negative allometry. Nevertheless, if high intra-specific variation is indicative of post-copulatory sexual selection, there remains the question of the mechanism: cryptic female choice or sexually antagonistic coevolution. It is difficult to determine the selection mechanism for the evolution of genitalia through the inter-specific and intra-specific phenotypic variation patterns of genitalic traits, because a certain pattern can usually be explained by more than one hypothesis. The mechanisms underlying the variability of euophryine genitalia remain unclear, and in fact sexual selection and species isolation mechanisms in principle could coexist. Unlike the genitalic traits directly associated with copulation, the pre-copulatory sexually-selected traits (e.g. male chelicerae) tend to show positive intra-specific allometry, and thus may have evolved under 161! strong directional selection. The fact that the species with stronger somatic sexual dimorphism (Antillattus cambridgei) has the lower intra-specific variation in genitalia may imply a trade-off between pre- and post-copulatory sexual selection. 4.2 Introduction Strongly correlated evolution of female and male reproductive structures has been revealed in birds (Brennan et al. 2007), mammals (Gomendio & Roldan 1993), insects (Presgraves et al. 1999; Ilango & Lane 2000; Rodriguez et al. 2004; Rönn et al. 2007; McPeek et al. 2008; Joly & Schiffer 2010; Tatarnic & Cassis 2010) and web-building spiders (Ramos et al. 2005; Kuntner et al. 2009). The covaried pattern of inter-sexual genitalic traits can be explained by different hypotheses. The classic non-sexual selection explanation is the “lock-and-key” hypothesis, which suggests that the divergence of genitalic evolution has resulted from hybridization avoidance (Hosken & Stockley 2004). As a matter of a fact, genitalic traits have long been used as criteria to distinguish species in taxonomic studies across various animal groups (Eberhard 1985; 2010), especially in spiders (Foelix 1996), because their morphology usually diverge relatively rapidly, and therefore are often species-specific and show more variation in closely related species than other somatic traits such as legs and eyes. However, more and more evidence suggests that sexual selection mechanisms, such as cryptic female choice that affect male paternity success (Eberhard 1996) and sexual conflict that propel sexually antagonistic coevolution (Arnqvist & Rowe 2002a), may have played a more important role in the evolution of genitalia (Arnqvist 1998; Hosken & Stockley 2004). Nevertheless, which hypothesis better explains the evolution of genitalia is still an ongoing debate (Arnqvist & Rowe 2002a; Mikkola 2008; Eberhard 2010). In spiders, the female copulatory duct and male embolus are the key components of genitalia that interact directly during copulation: the male spider inserts the embolus into the female copulatory duct through the genitalic opening and deposits sperms into the spermathecae (Foelix 1996). However, the intersexual correlated variation between the lengths of female copulatory duct and male embolus has never been examined in spiders. Jumping spiders are well-known for their acute vision and complex courtship behavior (Richman & Jackson 1992; Foelix 1996). Sexual selection has been suggested to drive the 162! divergence of phenotypes potentially crucial to the speciation process in some jumping spider species (Masta & Maddison 2002). However, previous studies of sexual selection of jumping spiders have been almost exclusively focused on pre-copulatory selection (Clark & Biesiadecki 2002; Hebets & Maddison 2005; Elias et al. 2006; Sivalinghem et al. 2010). In comparison, little attention has been placed on post-copulatory selection mechanisms, which are usually associated with the evolution of genitalia. Euophryine jumping spiders are an ideal system in which to study intersexual correlation of genitalic traits, as their genitalia are relatively simple in shape, and the lengths of the female copulatory duct and male embolus account for most of the inter-specific variation (Fig. 4.1). Here, I gather data on the lengths of female copulatory duct and male embolus of 142 euophryine species and use the phylogeny obtained from molecular data (see Chapter 2) to investigate their relationship for the first time. I expected a positive correlation between the lengths of male embolus and female copulatory duct. An association is expected because an embolus too short compared to the female copulatory duct may result in failure of fertilization, whereas an extremely long embolus may not only be developmentally expensive but also potentially harmful to the female reproductive structures. Intra-specific variation of genitalic structures could be informative about evolutionary processes (Eberhard et al. 1998; Eberhard et al. 2009). Although intra-specific genitalic variation has been widely investigated in insects and other spiders (e.g. Palestrini et al. 2000; Uhl & Vollrath 2000; Hosken et al. 2005; Mutanen et al. 2006; Tatsuta et al. 2007; Eberhard et al.1998; Eberhard 2009), it has been rarely studied in jumping spiders. Here I investigated intra-specific variation of multiple non-genitalic and genitalic traits in two euophryine species that show considerable difference in the degree of sexual dimorphism: Chapoda recondita and Antillattus cambridgei (Fig. 4.2). The “lock-and-key” hypothesis predicts relatively low intra-specific phenotypic variation since it implies a strict fit between female and male genitalia, whereas high intra- specific phenotypic variation may indicate sexual selection (Eberhard et al. 1998; Eberhard et al. 2009). 163! 4.3 Material and methods 4.3.1 Intersexual correlated evolution The lengths of the copulatory duct and embolus were measured for 142 euophryine species. Additionally, two somatic parts (carapace length and the 2 nd leg patella plus tibia length for both female and male) and two genitalic parts (female vulva width and male palpal bulb width) were also measured for the purpose of standardization (see below). A medium-body-sized specimen was chosen to represent the species when multiple specimens were available. Somatic parts were measured under a dissecting microscope with ocular micrometer. Genitalic parts were measured using ImageJ version 1.40 (Rasband 1997-2005; Abramoff et al. 2004) either from drawings made with a drawing tube on a Nikon ME600L compound microscope, or from photos taken under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0. The length of the carapace was measured from the base of the anterior median eyes (not including the lens) to the posterior end of the carapace. The length of the copulatory duct in females was measured from the genitalic opening to the beginning of spermatheca. In a few cases where a secondary spermatheca is present, such as in Antillattus cambridgei and Coryphasia physonycha, the length of copulatory duct was measured from the genitalic opening to the beginning of the primary spermatheca instead of the secondary spermatheca, with the assumption that males all deposit sperms in the primary spermatheca. The absolute lengths of copulatory duct and embolus may be influenced by developmental constraints associated with factors such as body size or genitalia size. Thus, the observed covariation in the absolute lengths of these two traits may simply represent the variation in body size or genitalia size among euophryine species, and not the result of selection from intersexual interaction during copulation. To exclude possible size-mediated effects, we tried different methods of standardization. The copulatory duct length and embolus length were standardized alternatively with the carapace length, the 2 nd leg patella plus tibia length, and the genitalia size (vulva width in female and bulb width in male). I did not simply standardize against a single overall size measurement (e.g. from a principal components analysis) because such standardization would be dominated by somatic size, which does not seem appropriate. For instance, we suspect there is a negative allometry in genitalia size — small species may have disproportionately larger genitalia. Such a pattern may not be driven by sexual selection, but perhaps by functional constraints on genitalic function, that is, small-bodied species have similar 164! sized genitalia as large-bodied species in order to fulfill copulation and sperm transfer. Then standardizing only against overall size could generate a false correlation, as small-bodied species would appear to have relatively large embolus and copulatory duct while large-bodied species would appear to have relatively small embolus and copulatory duct after standardization. Thus, I used both a simple somatic size indicator as well as genitalic size for standardization, to explore whether they give the same results. The evolutionary histories of the lengths of the embolus and copulatory duct were inferred on the phylogeny in Mesquite version 2.74 (Maddison & Maddison 2010b) using the squared- change parsimony model. We used the species-level phylogeny of euophryines and outgroups from molecular data (see Chapter 2). Taxa for which measurements of somatic and genitalic parts unavailable were trimmed from the phylogeny. I conducted phylogenetically independent contrast (PIC) analyses using PDAP version 1.15 (Felsenstein 1985a; Midford et al. 2010) as implemented for Mesquite version 2.74 (Maddison & Maddison 2010b). PIC analyses were conducted using the topology of the ML best tree directly from GARLI analysis (Chapter 2). Two kinds of branch length were tried for the PIC analyses with one as the expected number of DNA sequence changes along the branch and the other calibrated to represent the divergence time. The divergence time was estimated with two sets of calibrations respectively from the fossil records of jumping spiders using r8s v1.70 (Sanderson 2003; 2004; see Chapter 2 for details on dating the divergence of euophryines). The PIC analyses were conducted on the untransformed embolus length and copulatory duct length (both with no standardization and all three standardizations), as well as the log10-transformed embolus length and copulatory duct length (both with no standardization and all three standardizations). The relationship between the absolute lengths of embolus and copulatory duct was also examined using linear ordinary least squares regression performed in JMP 8.0 (SAS Institute, USA). 4.3.2 Intra-specific variation I studied intra-specific variation in two euophryine species, Chapoda recondita and Antillattus cambridgei. They show considerable difference in the degree of sexual dimorphism: in Chapoda recondita the males are very similar to females, whereas males of Antillattus cambridgei are quite different from females and usually armed with strongly elongate chelicerae (Fig. 4.2). 165! Specimens of Chapoda recondita used in this study were collected from Costa Rica: Heredia: Est. Biol. La Selva, and specimens of Antillattus cambridgei were from Dominican Republic: La Vega: P. N. Armando Bermúdez. I measured five non-genitalic traits (Fig. 4.3) in both females and males of each species: carapace length, carapace width, 2 nd leg femur length, sternum length and chelicera length. Among them, the carapace length was chosen as an indicator of overall body size for the allometric regression analyses since it has been commonly used in similar studies of spiders (Greenstone et al. 1985; Eberhard et al. 1998). The male chelicera length was treated differently from the other non-genitalic traits and not included when computing the mean or median for the somatic traits, given that male chelicerae, especially in Antillattus cambridgei, may be under direct sexual selection. Genitalic traits measured in males and females are shown in Fig. 4.4. For males, I measured the embolus length, bulb width, bulb length and palpal cymbium length. For females of Chapoda recondita, I measured the copulatory duct (CD) length, vulva width, vulva length, primary spermatheca (PS) length, primary spermatheca width, window length, window width and epigynum length. Females of Antillattus cambridgei have secondary spermathecae (SS) in addition to the primary spermathecae, and two sets of copulatory ducts, with one connecting the genitalic opening and the secondary spermatheca (CD1), and the other connecting the secondary spermatheca and the primary spermatheca (CD2). Thus, four additional traits (length of CD1, length of CD2, length of SS, width of SS) were measured in this species. Because the CD2 is right beside the CD1, the combined length of CD1 and CD2 was considered as the total length of copulatory duct. All non-genitalic and genitalic traits were measured from photos taken under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0 using the software ImageJ 1.40 (Rasband 1997-2005; Abramoff et al. 2004). To reduce measurement error, three independent measurements were taken for each trait and the mean was used for analysis. All variables were log10-transformed to evaluate allometric slopes using linear ordinary least squares regressions. I determined the positive (higher slope) or negative (lower slope) allometry by comparing the allometric slope of the trait with the median and mean allometric slope of the somatic traits (excluding the male chelicerae length), rather than by comparing them to the 166! absolute value of “1.0” (see discussion in Eberhard et al. 2009). A positive allometry of a trait means it is disproportionally larger to the body size of the organism, whereas a negative allometry indicates the trait is disproportionally larger for organisms of smaller body size, but disproportionally smaller for those of larger body size. Both the allometric slope and the degree of dispersion around the regression line were considered to evaluate phenotypic variation of a given trait (Eberhard et al. 1998; Eberhard et al. 2009). I used two indices to evaluate the dispersion of points around the regression line: CV' (the coefficient of variation that y would have if x were held constant) and standard error of the estimate (SE: the square root of the residuals from the regression line). CV' was calculated as CV (y) ! (1-r 2 ) 1/2 (Eberhard et al. 1998; Eberhard et al. 2009), where CV (y) is the coefficient variation of the variable y and r is the correlation coefficient (both from untransformed data). When r was not statistically significant, CV' was not used in any comparison or computation of medians or means, and only the standard error of the estimate was used. Linear ordinary least squares regression analyses were performed using JMP 8.0 (SAS Institute, USA). 4.4 Results 4.4.1 Intersexual correlation of embolus length and copulatory duct length Tracing the evolutionary history of the standardized lengths of embolus and copulatory duct on the phylogeny shows apparent covariation of the two traits (Fig. 4.5). Tracing the absolute lengths of embolus and copulatory duct on the phylogeny shows similar pattern as the standardized lengths. The PDAP diagnostic chart was checked to see whether branch lengths of the phylogenetic tree adequately fit the tip data (Midford et al. 2010). Neither the untransformed embolus length nor the copulatory duct length (both with no standardization and all three standardizations) showed significant correlation with its standard deviations under all “real” branch length assumptions (two-tailed P = 0.136-0.530), which indicated the data meet PIC statistical assumptions. In contrast, some log10-transformed variables (e.g. embolus length standardized with carapace length) showed significant correlation with its standard deviations under “real” branch length 167! assumptions (two-tailed P <0.05), which suggested the log10-transformed data do not meet the branch lengths and assumed Brownian motion model of character evolution (Midford et al. 2010). Thus, even though the PIC analyses on the log10-transformed data were still carried out to see if they show similar results as the untransformed data, only the results from the untransformed data were shown in Fig. 4.6. The phylogenetic independent contrast results (PIC) show significant positive correlation between the untransformed lengths of male embolus and female copulatory duct regardless of standardizations and branch length assumptions (Fig. 4.6, n = 141, r 2 = 0.738-0.881, df = 140, one-tailed P < 0.0001). Similarly, the log10-transformed lengths of male embolus and female copulatory duct are also positively correlated regardless of standardizations and branch length assumptions (n = 141, r 2 = 0.676-0.759, df = 140, one-tailed P < 0.0001). Regression of the absolute length of copulatory duct with that of embolus recovers a relationship of “CD length = 0.039 + 0.748!Embolus length” (Fig. 4.7). This indicates in general the embolus is slightly longer than the copulatory duct, as measured, in euophryine jumping spiders. 4.4.2 Intra-specific variation Table 4.1 lists details of intra-specific variation and statistics of the genitalic and non-genitalic traits in the two species, Chapoda recondita and Antillattus cambridgei. For both Chapoda recondita (Fig. 4.8) and Antillattus cambridgei (Fig. 4.9), the genitalic traits tend to show lower allometric slopes than somatic traits, when regressed against carapace length (used as the general body size indicator). In Chapoda recondita, the median of the seven somatic allometric slope values (both female and male, not including the male chelicera length) is 0.790, and the mean is 0.832. Of the 12 genitalic traits, 11 have allometric slopes lower than the median (or mean) slope of the somatic traits, but the regressions of CD length and window length are not statistically significant. The primary spermatheca length, on the other hand, shows higher slope. For Antillattus cambridgei, the median allometric slope of the seven somatic traits is 0.918, and the mean is 0.942. In total, 16 genitalic traits were examined for this species. All of them show lower allometric slopes than the median or mean slope of the somatic traits. However, four of them (CD2 length, secondary spermatheca width, window length and window 168! width) have regressions that are not statistically significant. In contrast, the male chelicera length of both species tends to have higher slope than the median (or mean) slope of the somatic traits, especially in Antillattus cambridgei, of which the slope of male chelicera length is more than threefold the median or mean slope of the somatic traits, and thus shows positive allometry. The CV' and standard error of the estimate (SE) tend to be higher for the genitalic traits than the somatic traits in both species. In Chapoda recondita, 10 of 12 genitalic traits (with the exception of CD length and window length whose linear regressions are not statistically significant) have the CV' higher than the median (2.905) or mean (2.980) CV' of the seven somatic traits (not including the male chelicera length); all of them have the SE higher than the median (0.076) or mean (0.077) SE of the seven somatic traits. The median and mean CV' for the 10 genitalic traits are 6.108 and 6.307 respectively; the median and mean SE for the 12 genitalic traits are 0.179 and 0.174 respectively. All of them are much higher than the corresponding values for the somatic traits. In Antillattus cambridgei, 12 of 16 genitalic traits (not including the four traits whose linear regressions are not statistically significant) have the CV' higher than the median (1.908) or mean (2.378) CV' of the seven somatic traits; 15 (with the only exception of male bulb width) have the SE higher than the median (0.060) or mean (0.062) SE of the seven somatic traits. The median and mean CV' for the 12 genitalic traits are 4.570 and 4.767 respectively; the median and mean SE for the 16 genitalic traits are 0.145 and 0.146 respectively. All of them are much higher than the corresponding values for the somatic traits. In both species, the male chelicera length has higher CV' and SE than the somatic traits. In addition, the CV' of the genitalic traits in Chapoda recondita (median=6.108 and mean=6.307) tends to be higher than those in Antillattus cambridgei (median=4.570 and mean=4.767), whereas the CV' of the male chelicera length in Chapoda recondita (5.685) tends to be much lower than that in Antillattus cambridgei (11.910). ! 4.5 Discussion 4.5.1 Intersexual correlated evolution of embolus and copulatory duct The results strongly support the positive correlation of the lengths of male embolus and female copulatory duct in euophryine jumping spiders, and confirm the interaction of these two traits between males and females during copulation. This is the first study to show quantitatively that 169! the embolus and copulatory duct, as important copulatory organs in spiders, are likely to have coevolved. The male embolus is usually slightly longer than the copulatory duct of conspecific females. This may indicate that in euophryine jumping spiders, males need to insert the embolus into the spermatheca of females in order to get sperms in place, or males insert only part of the embolus for sperm deposit. The covaried pattern of genitalia could be due to the “lock-and-key” mechanism in order to avoid hybridization. The general shape of genitalic traits may be more crucial to function as “lock” and “key” than their size. However, in euophryines the difference in lengths of embolus and copulatory duct accounts for most of the inter-specific variation of genitalia, especially among closely related species. Therefore, in euophryines the length of the male embolus may function as the “key” while the length of female copulatory duct function as the “lock”, and only males of the same species have the right “key” to fit the female “lock”. Alternatively, this pattern could be explained by post-copulatory sexual selection mechanisms. Females may increase the length of copulatory duct to retain better control over fertilization, for instance to screen for more viable (“stronger”) sperm. In response, males would gain an advantage by evolving a long enough embolus to reach the spermatheca. Thus, the male embolus and female copulatory duct could coevolve through the evolutionary arms race caused by intersexual conflict (sexual antagonistic coevolution, Arnqvist & Rowe 2002a). An alternative sexual selection scenario would involve female choice for males with the embolus that has the best stimulatory or mechanical fit to the copulatory tract (cryptic female choice, Eberhard 1996). Intersexual covariance in genitalic structures with similar functions as the spider embolus and copulatory duct has also been found in sand flies (Ilango & Lane 2000), tortoise beetles (Rodriguez et al. 2004\"!#$%!waterfowls (Brennan et al. 2007). Post-copulatory sexual selection has been suggested to be the most likely mechanism to explain their correlated evolution. 4.5.2 “Lock-and-key” or sexual selection Jumping spiders have elaborate species-specific courtship movements before copulation, enabled by their acute vision (Richman & Jackson 1992; Foelix 1996). The complex courtship may serve in species-recognition and/or sexual selection between the conspecific female and male. However, we cannot rule out the possibility that species-recognition and sexual selection may continue during the copulation. 170! Although the “lock-and-key” theory involves an interaction among species, it does not have a clear prediction about whether the interspecific variation of genitalic traits should be higher or lower than somatic traits. “Lock-and-key” might predict strong differences between species and thus high interspecific variation. However, the variation of genitalic traits among species could still be lower than that of somatic traits, if ecological factors select for high somatic variation among species while genitalic variation is just enough for species to be distinguished. Thus, “lock-and-key” could result in either high or low interspecific variation of genitalic traits as compared to somatic traits. Both the male embolus and female copulatory duct show negative intra-specific allometries when compared to somatic traits (Table 4.1; Figs 4.8-4.9). This is compatible with the traditional “lock-and-key” hypothesis (Eberhard 2009), which supposes that the embolus and copulatory duct are under strong stabilizing selection and thus a standard sized embolus and copulatory duct would be favored. However, post-copulatory sexual selection mechanisms — sexual conflict or female cryptic choice — could also result in negative intra-specific allometry (see below). While the low allometric slope is consistent with the “lock-and-key” hypothesis, a low slope does not guarantee genitalia will be of consistent absolute size, as that also requires low dispersion around the regression line. The “lock-and-key” hypothesis implies a strict “fit” between the female and male genitalic traits that interact and thus the “lock-and-key” hypothesis also predicts low size-corrected intra-specific phenotypic variation (CV'). The rather high size- corrected intra-specific phenotypic variation of embolus and copulatory duct found in Chapoda recondita and Antillattus cambridgei would therefore appear to be against the “lock and key” hypothesis. High phenotypic variation has been proposed to be an indicator of the opportunity for selection to act, and a higher degree of phenotypic variation than ordinary somatic traits is usually associated with sexual selection (Eberhard et al. 1998; Eberhard et al. 2009). However, a high size-corrected variation (CV') for genitalic traits could still be consistent with “lock-and-key”, explained by decoupled developmental mechanisms of somatic and genitalic traits. If somatic traits, such as carapace width and leg length, are linked to common developmental mechanisms, when regressed to each other they may show low dispersion. 171! However, the low allometric slopes for genitalic traits when regressed to the carapace length indicates that genitalia fail to respond fully to changes in body size, perhaps suggesting developmental mechanisms for genitalic and somatic traits are somewhat decoupled. If so, the genitalic traits could have more dispersion than the somatic traits when regressed to one somatic trait (carapace length) simply because of the decoupled variation, even if the genitalic traits actually have similar variability as the somatic traits. Thus the dispersion in genitalic traits may be an artifact of regressing against a decoupled trait, and overestimate the real intra-specific variation. Another possible cause for the high dispersion of points around the line is the elevated error in measuring genitalic traits, since they are usually small. Nevertheless, discriminating these sources from the observed variation is empirically difficult. Thus, whether or what proportion of the observed variation in the genitalic traits is indeed subject to sexual selection is hard to determine. Even if high phenotypic variation of a trait provides the opportunity for sexual selection to act, whether the trait is actually sexually selected is still an open question, needing experimental data to confirm. Also, although the high phenotypic variation may be indicative of sexual selection, the relatively low phenotypic variation of a trait does not necessarily mean it is not sexually- selected, because sexual selection could result in either increased or decreased intra-specific variation. Bertin and Fairbairn (2007) have found, for example that some traits are under sexual selection but have similar size-corrected intra-specific variation as somatic traits. 4.5.3 Sexual selection mechanisms If high intra-specific phenotypic variation in genitalia is indeed a valid indicator of post- copulatory sexual selection, there remains the question of which sexual selection mechanism is more likely, sexually antagonistic coevolution or cryptic female choice? The sexually antagonistic coevolution theory supposes that female and male have different interests in the events associated with copulation, insemination and fertilization, and thus genitalic divergence occurs as a result of the evolutionary arms race for control over these processes (Arnqvist & Rowe 2002a; Eberhard 2010). In contrast, the female cryptic choice hypothesis proposes that the divergence of genitalic evolution is driven by female choice for males with genitalia that best stimulate them or have best mechanical fit during copulation via biases in post-copulatory processes such as sperm transport, oviposition and remating (Eberhard 172! 1996; 2010). The sexual antagonistic coevolution hypothesis has been suggested to be the most likely mechanism to explain the covaried female and male genitalia traits in various animals such as waterfowl (Brennan et al. 2007), sand fly (Ilango & Lane 2000) and nephilid spiders (Kuntner et al. 2009). However, the two hypotheses are not mutually exclusive and discriminating between them is sometimes difficult (Eberhard 2010). It has been suggested that the sexual antagonistic coevolution hypothesis predicts at least a trend toward positive intra-specific allometry in genitalia (Eberhard 2009). Individuals should be selected to have as long an embolus or copulatory duct as possible, in order to gain control over reproduction, but resource limitations combined with developmental processes may enable larger individuals to develop disproportionately longer structures, resulting in positive allometry (Faber 1984; Nijhout & Wheeler 1996). This could be true if selection intensity is constant across the body size spectrum. However, if for whatever reason the intensity of selection for larger trait is greater for individuals of small body size than for individuals of large body size, then a negative intra-specific allometry will result (Eberhard et al. 2009). On the contrary, the “cryptic female choice” hypothesis usually predicts negative intra-specific allometry. Unless there is size-assortative mating, selection on males may favor males to have intermediate-and- standard-sized genitalia (and thus low allometric slopes) so that they can fit appropriately with majority of females in the population; this will in turn result in selection favoring females that possess intermediate-sized genitalia themselves (Eberhard et al. 1998; Eberhard 2009). Consequently, the negative intra-specific allometry of embolus and copulatory duct as well as other genitalic traits of these two euophryine species may result either from stabilizing sexual selection for standardized sizes of traits or from directional selection with greater intensity of selection favoring larger traits for small individuals than for large individuals (Eberhard et al. 2009). The scaling and variation pattern within the population may be ineffective to discern the ultimate causes of genitalic evolution (Bertin & Fairbairn 2007). In contrast to the genitalic traits directly associated with copulation, the male chelicerae presumably evolving under pre-copulatory sexual selection have obvious positive intra-specific allometry. They may have evolved under strong directional selection that favors disproportionally larger chelicerae in large individuals (Eberhard et al. 2009). The positive allometry of male chelicera length has been previously found in other jumping spider species, such as Zygoballus rufipes Peckham & Peckham (Faber 1984). Elongate male chelicerae could 173! act in male-male combat (Jackson 1989) or as a visual display character. Either way, elongate chelicerae may indicate “good viability” of males. Sexual selection may be present both before and after copulation. However, few studies have been conducted on the relationship between pre- and post-copulatory sexual selection mechanisms (see Simmons & Emlen 2006; Parzer & Moczek 2008). The extremely elongate chelicerae in some male jumping spiders are likely associated with strong pre-copulatory sexual selection. The high phenotypic variation of genitalia in Antillattus cambridgei suggests that post-copulatory sexual selection may still exist in spite of the presence of strong pre-copulatory sexual selection (elongate male chelicerae). However, the higher phenotypic variation (CV') of male chelicera length but lower variation (median or mean of CV') of genitalic traits in Antillattus cambridgei when compared to Chapoda recondita hints that there may be trade-offs between pre- and post-copulatory sexual selection. Because investment in pre- and post- copulatory sexual selection is costly to a spider, it is probably beneficial to invest more in one of the two selections but less in the other. 4.6 Conclusions and future directions Investigation on the correlated evolutionary pattern of the lengths of male embolus and female copulatory duct under the phylogenetic context shows that they are positively correlated among euophryine species. Study of intra-specific variation of two euophryine species indicates that genitalic traits including the copulatory duct and embolus length usually show negative allometry but high size-corrected variation. However, I find it is difficult to determine the mechanism for genitalic evolution (“lock-and-key” or post-copulatory sexual selection, sexually antagonistic coevolution or cryptic female choice) by examining only inter-specific and intra- specific patterns of phenotypic variation. A given pattern can usually be explained by more than one hypothesis. The available data on intra-specific variation in euophryine spiders did not allow us to distinguish whether genitalic divergence is due to post-copulatory sexual selection or functions as a species-recognition mechanism among species. In principle, the sexual selection and non-sexual selection mechanisms may coexist and the divergent genitalia across euophryines species may be the result of a balance between the sexual and non-sexual selection forces (House & Simmon 2003). In contrast, the pre-copulatory sexually-selected traits (e.g. 174! male chelicerae) tend to show positive allometry, and thus may have evolved under strong directional selection. More data, especially from experimental and behavioral studies, are needed to better understand the underlying mechanisms of genitalic evolution. A comparative study on the genitalia morphology of closely related sympatric species is essential to undercover the presence of species-specific “lock-and-key”. Evidence from experiments on the relationship between paternity and the size of a male’s genitalic structures (as in the Tortoise beetle Chelymorpha alternans, Rodriguez et al. 2004) would be particularly useful to test sexual selection hypothesis for genitalic evolution in euophryines. In addition, a better understanding of the courtship and copulation mechanisms in a wide range of jumping spider species is also important to detect intersexual conflict and reveal the relationship between the pre- and post-copulatory selection. 175! Table 4.1. Intra-specific variation of somatic and genitalic traits in Chapoda recondita and Antillattus cambridgei. 1 Mean (SD) mm CV (%) Slope r 2 SE CV’ Chapoda recondita Female Body Part (n=50) Carapace length 1.319 (0.058) 4.424 - - - - Carapace width 1.140 (0.051) 4.514 0.791*** 0.586*** 0.093 2.905 2 nd femur length 0.768 (0.036) 4.630 0.903*** 0.748*** 0.076 2.324 Sternum length 0.581 (0.021) 3.563 0.500*** 0.399*** 0.089 2.763 Chelicera length 0.442 (0.020) 4.484 0.734*** 0.526*** 0.100 3.087 Female Genitalic Part (n=50) CD length 0.107 (0.005) 5.106 0.288 ns 0.060 ns 0.160 4.951 ns Vulva width 0.211 (0.016) 7.745 0.725** 0.162** 0.232 7.090 Vulva length 0.141 (0.009) 6.743 0.567** 0.126* 0.203 6.306 Primary spermatheca length 0.067 (0.006) 8.498 0.960*** 0.253*** 0.238 7.345 Primary spermatheca width 0.107 (0.010) 9.561 0.663* 0.086* 0.294 9.141 Window length 0.073 (0.004) 5.696 0.076 ns 0.002 ns 0.183 5.691 ns Window width 0.169 (0.011) 6.447 0.424* 0.085* 0.203 6.166 Epigynum length 0.171 (0.010) 5.710 0.478** 0.119* 0.176 5.360 Male Body Part (n=38) !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 1 Mean (SD): mean and standard deviation of each trait (untransformed data); CV: coefficient of variation of each trait (untransformed data); Slope: slope when regressed on carapace length which is used as the indicator of body size (both variables log10-transformed); r 2 : squared coefficient of correlation of the regression on carapace length (untransformed data); SE: standard error of estimate of the regression on carapace length (log10-transformed data); CV’: size-corrected intra-specific variation, calculated as CV’ = CV (y) ! (1-r 2 ) 1/2 with both CV (y) and r 2 from untransformed data (*** P < 0.001; ** P < 0.01; * P < 0.05; “ns” not statistically significant at a = 0.05). 176! Mean (SD) mm CV (%) Slope r 2 SE CV’ Carapace length 1.353 (0.123) 9.120 - - - - Carapace width 1.173 (0.091) 7.793 0.789*** 0.866*** 0.053 2.851 2 nd femur length 0.891 (0.104) 11.656 1.181*** 0.888*** 0.070 3.903 Sternum length 0.614 (0.051) 8.380 0.864*** 0.870*** 0.054 3.030 Chelicera length 0.505 (0.072) 14.330 1.387*** 0.843*** 0.101 5.685 Male Genitalic Part (n=38) Embolus length 0.095 (0.007) 7.332 0.511*** 0.416*** 0.104 5.601 Bulb width 0.187 (0.015) 7.851 0.570*** 0.406*** 0.108 6.051 Bulb length 0.247 (0.019) 7.876 0.606*** 0.498*** 0.103 5.579 Palpal cymbium length 0.431 (0.034) 7.828 0.680*** 0.680*** 0.084 4.426 Antillattus cambridgei Female Body Part (n=34) Carapace length 1.567 (0.087) 5.574 - - - - Carapace width 1.253 (0.051) 4.105 0.658*** 0.820*** 0.056 1.740 2 nd femur length 0.897 (0.047) 5.241 0.880*** 0.882*** 0.058 1.803 Sternum length 0.725 (0.040) 5.516 0.918*** 0.880*** 0.061 1.908 Chelicera length 0.610 (0.047) 7.659 1.276*** 0.881*** 0.083 2.638 Female Genitalic Part (n=34) CD 1 length 0.032 (0.002) 5.502 0.493** 0.211** 0.152 4.888 CD 2 length 0.034 (0.002) 6.484 0.247 ns 0.070 ns 0.130 6.252 ns CD 1+2 length 0.066 (0.003) 4.905 0.368* 0.191** 0.138 4.412 Vulva width 0.166 (0.009) 5.279 0.375* 0.160* 0.152 4.839 Vulva length 0.139 (0.010) 6.890 0.700*** 0.322*** 0.186 5.672 177! Mean (SD) mm CV (%) Slope r 2 SE CV’ Primary spermatheca length 0.070 (0.006) 7.934 0.676** 0.264** 0.215 6.808 Primary spermatheca width 0.077 (0.005) 6.623 0.480* 0.144* 0.191 6.127 Secondary spermatheca length 0.066 (0.004) 6.180 0.491* 0.185* 0.181 5.580 Secondary spermatheca width 0.067 (0.005) 7.380 0.312 ns 0.079 ns 0.229 7.083 ns Window length 0.132 ( 0.005) 3.988 0.243 ns 0.097 ns 0.120 3.789 ns Window width 0.243 (0.011) 4.371 0.142 ns 0.029 ns 0.139 4.309 ns Epigynum length 0.187 (0.010) 5.176 0.557*** 0.340*** 0.135 4.205 Male Body Part (n=32) Carapace length 1.713 (0.157) 9.148 - - - - Carapace width 1.303 (0.086) 6.606 0.710*** 0.933*** 0.035 1.710 2 nd femur length 1.091 (0.117) 10.716 1.086*** 0.875*** 0.078 3.790 Sternum length 0.855 (0.087) 10.134 1.064*** 0.909*** 0.060 3.057 Chelicera length 2.106 (0.609) 28.931 2.897*** 0.831*** 0.261 11.910 Male Genitalic Part (n=32) Embolus length 0.096 (0.004) 3.768 0.192** 0.249** 0.069 3.265 Bulb width 0.213 (0.010) 4.537 0.405*** 0.625*** 0.056 2.778 Bulb length 0.230 (0.013) 5.839 0.396*** 0.387*** 0.091 4.570 Palpal cymbium length 0.467 (0.033) 7.034 0.627*** 0.666*** 0.083 4.067 178! Figure 4.1. Variation in the lengths of embolus and copulatory duct among euophryines. A-D, male left palpi; E-H, female vulvae. A and E, Anasaitis canosa (Walckenaer); B and F, Corythalia decora Bryant; C and G, Coryphasia campestrata Simon; D and H, Omoedus darleyorum Zhang & Maddison. Scales = 0.2mm. 179! Figure 4.2. Comparison of sexual dimorphism in Chapoda recondita and Antillattus cambridgei. A-B, Chapoda recondita: A, female; B, male. C-D, Antillattus cambridgei: C, female; D, male. 180! Figure 4.3. Non-genitalic traits measured for Antillattus cambridgei (same for Chapoda recondita). A, carapace, dorsal view; B, sternum, ventral view; C, 2 nd leg, retrolateral view; D, female chelicera, back view; E, male chelicera, back view. Scales: A, C, E = 0.5mm; B, D = 0.2mm. Abbreviations: CL, carapace length; CW, carapace width; SL, sternum length; FL, femur length; ChL, chelicera length. 181! Figure 4.4. Genitalic traits measured. A-C, G-H, Chapoda recondita; D-F, I-J, Antillattus cambridgei. A, D, epigynum not cleared, ventral view; B, C, E, A, epigynum cleared, dorsal view; F, epigynum cleared, ventral view; G, I, male palp, ventral view; H, J, male palp, dorsal view. Scales: A-F = 0.1 mm; G-J = 0.2 mm. Abbreviations: WL, window length; WW, window width; EPL, epigynum length; PSL, primary spermatheca length; PSW, primary spermatheca width; SSL, secondary spermatheca length; SSW, secondary spermatheca width; VL, vulva length; VW, vulva width; CDL, copulatory duct length; CD1L, length of copulatory duct connecting the genitalic opening and the secondary spermatheca; CD2L, length of copulatory duct connecting the secondary spermatheca and the primary spermatheca; EL, embolus length; BL, palpal bulb length; BW, palpal bulb width; PCL, palpal cymbium length. 182! Figure 4.5. Correlated evolution of the lengths of male embolus (left) and female copulatory duct (right) in euophryine jumping spiders, both standardized by carapace length. Anasaitis adorabilis Anasaitis brunnea Anasaitis cf. canalis Anasaitis canosa [USA] Anasaitis banksi Anasaitis placida Anasaitis gloriae Anasaitis laxa Anasaitis elegantissima Anasaitis sp. [Peblique] Anasaitis locuples Corythalia porphyra Corythalia sulfurea Corythalia minor Corythalia broccai Corythalia peblique Corythalia bromelicola Corythalia coronai Corythalia decora Corythalia bicincta Corythalia electa Parabathippus shelfordi Parabathippus magnus Parabathippus cf. macilentus Parabathippus cuspidatus Euophrys frontalis Chalcoscirtus infimus Talavera minuta Chalcoscirtus diminutus Pseudeuophrys erratica Sidusa sp.2 [FrenchGuiana] Sidusa silvae Sidusa sp. [CostaRica] Sidusa unicolor Sidusa extensa Pensacola signata Mexigonus morosus Mexigonus arizonensis Petemathis minuta Petemathis portoricensis [Maricao] Petemathis portoricensis [Adjuntas] Truncattus flavus Truncattus cachotensis Truncattus dominicanus Antillattus cf. applanatus Antillattus gracilis Antillattus cambridgei Antillattus darlingtoni Antillattus scutiformis Antillattus maxillosus Corticattus latus Naphrys pulex Popcornella spiniformis Popcornella furcata Popcornella nigromaculata Compsodecta haytiensis Compsodecta peckhami Bythocrotus crypticus Bythocrotus cf. crypticus Agobardus bahoruco Agobardus phylladiphilus Agobardus cf. anormalis montanus Agobardus cf. brevitarsus Agobardus cordiformis Agobardus gramineus Agobardus oviedo Chapoda recondita Chapoda gitae Chapoda angusta Chapoda cf. inermis [CostaRica] Chapoda cf. inermis [Panama] Marma nigritarsis Amphidraus complexus Tylogonus yanayacu Soesilarishius ruizi Soesilarishius micaceus Saphrys a-notata Saphrys cf. patagonica Coryphasia fasciiventris Coryphasia physonycha Coryphasia cf. campestrata Belliena ecuadorica Ilargus galianoae Ilargus pilleolus Ilargus serratus Ilargus coccineus Ilargus foliosus Saitis auripes Saitis barbipes Saitis sp. [SouthAustralia] Saitis cf. griseus Emathis gombak Lepidemathis haemorrhoidalis Colyttus bilineatus Colyttus striatus Lagnus edwardsi Thorelliola aliena Thorelliola crebra Thorelliola ensifera Thorelliola tualapa Thorelliola cf. mahunkai Foliabitus longzhou Laufeia daiqini Laufeia keyserlingi Laufeia eximia Thiania bhamoensis Chinophrys pengi Thyenula wesolowskae Cytaea oreophila Cytaea nimbata Euryattus bleekeri Phasmolia elegans Zabkattus brevis Variratina minuta Bulolia excentrica Leptathamas paradoxus Coccorchestes cf. inermis Coccorchestes cf. aiyura Canama fimoi Canama extranea Canama triramosa Bathippus macrognathus Bathippus korei Bathippus gahavisuka Bathippus directus [Tualapa] Pristobaeus cf. jocosus Pristobaeus beccarii Xenocytaea agnarssoni Sobasina wanlessi Paraharmochirus tualapaensis Ohilimia scutellata Diolenius varicus Omoedus ephippigera Omoedus orbiculatus Omoedus swiftorum Omoedus cf. semirasus Omoedus cf. torquatus Omoedus cf. danae Omoedus darleyorum Omoedus papuanus Omoedus tortuosus Omoedus omundseni 0.35 to 0.45 0.45 to 0.55 0.55 to 0.65 0.65 to 0.75 0.75 to 0.85 0.85 to 1.4 Embolus length / Carapace length 0.01 to 0.05 0.05 to 0.15 0.15 to 0.25 0.25 to 0.35 Copulatory duct length / Carapace length 0.01 to 0.05 0.05 to 0.15 0.15 to 0.25 0.25 to 0.35 0.35 to 0.45 0.45 to 0.55 0.55 to 0.65 0.65 to 0.75 0.75 to 0.85 0.85 to 1.0 183! Figure 4.6. Independent contrasts of male embolus length (x) vs. female copulatory duct length (y) are positively correlated in euophryine species (n = 141, df = 140, one-tailed P < 0.0001, branch length as the number of DNA sequence changes along the branch): A, not standardized; B, standardized by carapace length; C, standardized by 2nd leg patella plus tibia length; D, standardized by genitalia size (vulva width in female and bulb width in male). 184! Figure 4.7. Linear regression of absolute lengths of male embolus (x) vs. female copulatory duct (y) in euophryine species, not corrected for phylogeny (n = 142, df = 141, r 2 = 0.909, one- tailed P < 0.0001). 185! Figure 4.8. Allometric relationships of genitalic and non-genitalic traits in Chapoda recondita, using carapace length as the indicator of body size: A-C, female; D-F, male. A and D, carapace length; B and E, chelicera length; C, copulatory duct length; F, embolus length. 186! Figure 4.9. Allometric relationships of genitalic and non-genitalic traits in Antillattus cambridgei, using carapace length as the indicator of body size: A-C, female; D-F, male. A and D, carapace length; B and E, chelicera length; C, copulatory duct length; F, embolus length. ! 187! 5 New euophryine jumping spiders from the Dominican Republic and Puerto Rico (Araneae: Salticidae: Euophryinae) 1 5.1 Synopsis Twenty four new species and three new genera of euophryine jumping spiders from two Caribbean Islands, Hispaniola and Puerto Rico, are described. The new genera are Corticattus (C. guajataca sp. nov. and the type species C. latus sp. nov.), Popcornella (P. furcata sp. nov., P. nigromaculata sp. nov., P. yunque sp. nov. and the type species P. spiniformis sp. nov.) and Truncattus (T. cachotensis sp. nov., T. dominicanus sp. nov. and the type species T. flavus sp. nov.). The other new species belong to the genera Agobardus (A. bahoruco sp. nov., A. cordiformis sp. nov., A. gramineus sp. nov., A. oviedo sp. nov., A. phylladiphilus sp. nov.), Anasaitis (A. adorabilis sp. nov., A. brunnea sp. nov., A. hebetata sp. nov., A. laxa sp. nov.), Antillattus (A. applanatus sp. nov.), Bythocrotus (B. crypticus sp. nov.) and Corythalia (C. broccai sp. nov., C. bromelicola sp. nov., C. coronai sp. nov., C. peblique sp. nov.). Photographs of living spiders and diagnostic illustrations are provided for all of the new species. 5.2 Introduction As one of the most diverse groups in the Salticidae, the subfamily Euophryinae has about 900 described species, with the majority found in the tropics of both the Old and the New World (Prószy!ski 1976; Maddison & Hedin 2003a; Platnick 2011). Euophryine jumping spiders are abundant and diverse in the Caribbean Islands (e.g. Bryant 1940, 1943, 1947a, b, 1950; Galiano 1988; Peckham & Peckham 1901; Petrunkevitch 1930), with a total of 84 species in 27 genera reported (Platnick 2011). However, much of the euophryine jumping spider fauna in the Caribbean Islands remains undiscovered. In expeditions to two of the larger islands, Hispaniola and Puerto Rico in 2009, we collected about 66 euophryine jumping spider species, of which many are new to science. Of these, 24 species were chosen for description here in order to give names for the taxa included in the molecular phylogenetic study (Chapter 2) on the subfamily Euophryinae. These molecular data also provide evidence for the generic placement of species described below. Thus, I report here !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! \"!A version of this chapter has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from the Dominican Republic and Puerto Rico (Araneae: Salticidae: Euophryinae). Names of new taxa are not finalized for nomenclature. ! 188! three new genera, Corticattus (two species), Popcornella (four species) and Truncattus (three species). An additional 15 new euophryine species are described and included in the genera Agobardus (five species), Anasaitis (four species), Antillattus (one species), Bythocrotus (one species) and Corythalia (four species). Although I do not describe all 66 species, this expedition revealed how much Caribbean euophryine diversity remains to be explored. The collected species in total belong to the described genera Agobardus (ca. 16 species), Anasaitis (ca. 15 species), Antillattus (ca. seven species), Bythocrotus (one species), Compsodecta (two species), Corythalia (six species), Dinattus (one species), Caribbean “Emathis” (three species), Wallaba (one species), and the newly described genera Corticattus gen. nov. (two species), Popcornella gen. nov. (five species) and Truncattus gen. nov. (ca. seven species). The Agobardus species have radiated into various habitats ranging from the rain forest to the desert dry forest, and can be found in different microhabitats such as foliage, on ground (leaf litter, grass clumps or on rocks) and tree trunks. Typical Antillattus species (e.g. A. gracilis Bryant) are foliage dwellers, but a few species are found on tree trunks (e.g. A. applanatus sp. nov.). Most species of Anasaitis, Corythalia, Dinattus and Wallaba are ground dwellers, but a few species are associated with lower foliage. Both Bythocrotus and Compsodecta are foliage-dwelling, but the Bythocrotus species described here is found in the relatively dry areas while the Compsodecta species are found in moist forests. The Caribbean “Emathis” species are collected from foliage or tree trunks. Species of Corticattus and Truncattus are exclusively tree-trunk dwellers, while the Popcornella species are found in leaf-litter in forest. 5.3 Material and methods During an expedition to the Dominican Republic and Puerto Rico in 2009, we explored various habitats ranging from the highland pine forest and cloud forest to the lowland humid forest and desert dry forest. Multiple collecting techniques, including beating foliage, brushing tree trunks and searching by eye on the ground and leaf-litter, were used. Photographs of living specimens were taken with a Pentax Optio 33WR digital camera. For macro capability, a small lens was glued to it. Photographs of preserved specimens were taken under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0. ! 189! Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Terminology is standard for Araneae. All measurements are given in millimeters. Descriptions of color pattern are based on the alcohol-preserved specimens. Carapace length was measured from the base of the anterior median eyes, not including the lenses, to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; AME, anterior median eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the \"small eyes\"). Specimens are deposited in the Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM). 5.4 Taxonomy 5.4.1 Genus Agobardus Keyserling, 1885 Small to medium sized spiders. Body is usually relatively robust, not elongate. Male cheek is strongly swollen in some species. Chelicera usually has two promarginal teeth and one bicuspid retromarginal tooth. Male chelicera of some species is relatively enlarged with modifications. Tibia and metatarsus of first leg usually have three pairs of ventral macrosetae each. Embolus usually coils for no more than one circle. Tegulum lacks proximal lobe. Window of epigynum is large or relatively small, with a median septum. Spermatheca is strongly swollen. Agobardus shows similar body form as Bythocrotus, Compsodecta and some species of Antillattus, but differs from them by the bicuspid retromarginal tooth on chelicera. It also differs from Bythocrotus by the absence of stout macrosetae on the male palpal tibia, and from most species of Compsodecta by the absence of additional apophysis on the male palpal tibia or patella besides the retrolateral tibial apophysis. Eleven species and one subspecies have been reported from the Caribbean Islands (Platnick 2011). However, some species described from Cuba (Bryant 1940) appear to belong to Antillattus Bryant, 1943 based on their diagnostic drawings. Also, some species described as Siloca Simon, 1902 (Galiano 1963) from Cuba may actually belong to Agobardus. Five new species from the Dominican Republic are described here. Placement of these new species in ! 190! Agobardus is supported by molecular data (see Chapter 2). 5.4.1.1 Agobardus bahoruco sp. nov. Figs 5.1-5.2 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: P. N. Sierra de Bahoruco, 18.128° N, 71.558° W, elev. 1340 m, 15 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-033. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 female, DOMINICAN REPUBLIC: Pedernales: P. N. Sierra de Bahoruco, 18.15° N, 71.60-71.62° W, elev. 1400 m, 15 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-034. Etymology. A noun in apposition taken from the type locality. Diagnosis. A. bahoruco can be distinguished from other Agobardus by the flattened body. Similar in epigynum to Agobardus phylladiphilus, but differs by abdominal markings, the modified male chelicera and the longer tibia of the male palp. The flattened body is similar to Antillattus applanatus and Commoris modesta Bryant, 1943. It can be distinguished from Antillattus applanatus by the bicuspid retromarginal tooth on the female chelicera (one unident tooth in Antillattus applanatus), the male cheliceral shape, the absence of a proximal tegular lobe and the wide retrolateral sperm duct loop of the male palp. It differs from Commoris modesta Bryant, 1943 in the shorter embolus and the longer tibia of the male palp. Description. Male (holotype, UBC-SEM AR00030). Carapace length 2.2; abdomen length 2.4. Body dorsal-ventrally slightly flattened. Chelicera (Fig. 5.2D): dark brown; with two promarginal teeth and one retromarginal tooth of six cusps; retromargin also with a blunt spur between the retromarginal tooth and the base of fang; fang with a small cusp near the middle. Palp (Fig. 5.2C): yellow brown. Embolus relatively short and curved. Retrolateral tibial apophysis finger-like. Palpal femur and patella dorsally with white hairs. Measurements of legs: I 4.9, II 4.0, III 4.4, IV 4.7. Color in alcohol (Fig. 5.2A): carapace dark brown, but lateral margins covered with white setae; abdomen gray brown with a light brownish yellow marking and some small light colored speckles; venter gray brown with two light brownish yellow longitudinal stripes; legs gray brown with light yellow markings. ! 191! Female (paratype, UBC-SEM AR00031). Carapace length 2.3 (variation 1.8-2.3, n=3); abdomen length 2.8. Measurements of legs: I 4.1, II 3.9, III 4.5, IV 5.0. Epigynum (Figs 5.2E- F): window occupying less than half of the epigynal plate, with relatively wide median septum; opening of copulatory duct at the posterior margin of the window. Copulatory ducts short and with accessory gland; spermathecae pear-shaped. Color in alcohol (Fig. 5.2B): similar to that of the male. Natural history. Specimens were found on pine tree trunks in a forest. 5.4.1.2 Agobardus cordiformis sp. nov. Figs 5.3-5.4 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: east of Pedernales, 17.965° N, 71.635° W, elev. 30 m, 17 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-040. Paratypes: 1 female, same data as holotype; 1 male and 1 juvenile, same data as holotype. Etymology. The specific epithet is an adjective from the combination of the Latin cordis (heart) and formis (shaped), and refers to the dark heart-like marking on the front face of male chelicerae. Diagnosis. Resembles Agobardus oviedo in markings, but differs from it by the heart-like marking on the front face of the male chelicerae, the non-modified male chelicera, and the spermathecae which are further away from each other. It resembles A. gramineus in color pattern and markings, but differs from that species by the narrower median septum of the epigynum and the wider embolic disc of the male palp. Description. Male (holotype, UBC-SEM AR00032). Carapace length 1.3 (variation 1.3-1.4, n=2); abdomen length 1.2. Chelicera: dark; proximal part covered with yellow scales; not elaborate. Palp (Figs 5.4C-D): light yellow. Embolus curved for less than half a circle. Retrolateral tibial apophysis finger-like. Measurements of legs: I 2.9, II 2.8, III 3.4, IV 3.6. Color in alcohol (Fig. 5.4A): carapace red brown, covered with orange scales, with margins light ! 192! yellow brown and a light yellow brown stripe behind fovea; abdomen light yellow, with gray brown patches and markings and many light colored speckles; tibiae, metatarsi and tarsi of legs brownish; other segments light yellow. Female (paratype, UBC-SEM AR00033). Carapace length 1.6; abdomen length 1.5. Measurements of legs: I 3.6, II 3.4, III 4.3, IV 4.5. Epigynum (Figs 5.4E-F): window occupying about half of the epigynal plate, median septum narrow. Copulatory ducts short, without obvious accessory gland; spermathecae oval and apart from each other. Color in alcohol (Fig. 5.4B): similar to that of the male except markings on abdomen darker. Natural history. Specimens were found on rocks in a desert. 5.4.1.3 Agobardus gramineus sp. nov. Figs 5.5-5.6 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: near Pedernales, 17.970° N, 71.651° W, elev. 8 m, 17 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-042. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 female and 10 males, same data as holotype; 2 males and 3 females, same data as holotype; 1 male and 1 female, DOMINICAN REPUBLIC: Pedernales: east of Pedernales, 17.965° N, 71.635° W, elev. 30 m, 17 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-040; 1 male, same data as previous; 2 females and 2 males, DOMINICAN REPUBLIC: Pedernales: near Pedernales, 17.964° N, 71.652° W, elev. 13 m, 18 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-044. Etymology. Latin adjective gramineus (of grass), referring to the habitat (grass clumps). Diagnosis. Distinguished from other Agobardus by the two pairs of distinct dark patches on the dorsum of the abdomen and the wide median septum of the epigynum. Description. Male (holotype, UBC-SEM AR00034). Carapace length 1.1 (variation 1.1-1.2, n=8); abdomen length 1.1. Chelicera: not elaborate; yellowish with gray pigments. Palp (Figs ! 193! 5.6C-D): tibia and tarsi gray brown, other segments light yellow. Embolus slightly curved. Retrolateral tibial apophysis finger-like. Measurements of legs: I 2.0, II 1.8, III 2.3, IV 2.5. Color in alcohol (Fig. 5.6A): carapace gray brown, darker in eye area, with wide lateral marginal stripes and a central stripe behind fovea composed of white setae; abdomen light yellow, with light brown markings and two pairs of dark patches, the anterior pair smaller than the posterior pair; venter with a few dark brown irregular markings and a big dark brown patch in front of spiracle; legs light brown. Female (paratype, UBC-SEM AR00035). Carapace length 1.3 (variation 1.2-1.4, n=18); abdomen length 1.5. Measurements of legs: I 2.5, II 2.2, III 2.8, IV 3.1. Epigynum (Figs 5.6E- F): window relatively big, occupying more than half of the epigynal plate; median septum wide, with opening of copulatory duct anterior. Copulatory ducts short, without obvious accessory gland; spermathecae spherical. Color in alcohol (Fig. 5.6B): similar to that of the male, but lacking lateral marginal and distinct central stripes on carapace. Natural history. Specimens were found in grass clumps in a desert. 5.4.1.4 Agobardus oviedo sp. nov. Figs 5.7-5.8 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo, 17.802° N, 71.349° W, elev. 5 m, 14 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-032. Paratype: 1 female, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Similar to Agobardus brevitarsus Bryant, 1943 in the shape of male chelicera, and to A. perpilosus Bryant, 1943 in the shape of male palp. It differs from A. brevitarsus in the shorter embolus, the dentition on the male chelicera, and the smaller epigynal window. It can be distinguished from A. perpilosus by the teeth pattern and the presence of a cusp on the fang of the male chelicera. Description. Male (holotype, UBC-SEM AR00036). Carapace length 1.5; abdomen length 1.2. ! 194! Chelicera (Fig. 5.8D): yellow brown; fang with a small cusp near the middle. Palp (Fig. 5.8C): light brown. Embolus short but curved for about half a circle. Retrolateral tibial apophysis finger-like. Palpal femur and patella with white hairs. Measurements of legs: I 3.4, II 2.9, III 3.4, IV 3.6. Color in alcohol (Fig. 5.8A): carapace dark brown, covered with dark hairs, and with wide lateral margins and a central stripe composed of white setae; abdomen light yellow, with a pair of dark patches near the center and some orange markings; venter with a central grayish marking; legs light yellow. Female (paratype, UBC-SEM AR00037). Carapace length 1.4; abdomen length 1.4. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 2.7, II 2.5, III 3.1, IV 3.5. Epigynum (Figs 5.8E-F): window small, with opening of copulatory duct at the posterior margin of the window. Copulatory ducts short and with big accessory gland; spermathecae spherical. Color in alcohol (Fig. 5.8B): similar to that of male. Natural history. Specimens were found on rocks in a dry forest. 5.4.1.5 Agobardus phylladiphilus sp. nov. Figs 5.9-5.10 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: P. N. Sierra de Bahoruco, 18.128° N, 71.558° W, elev.1340 m, 15 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-033. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 1 female 1 male, same data as holotype; 1 male, same data as holotype; 1 female 1 male, same data as holotype; 1 male, same data as holotype. Etymology. The specific epithet means \"leaf litter loving\", based on the Greek \"phyllas\" for leaf litter, and refers to its habitat. Diagnosis. Male chelicerae are not modified, as in Agobardus cordiformis and A. gramineus. Differs from A. cordiformis by the wider retrolateral sperm duct loop, and the pear-shaped spermathecae, which are closer to each other. Differs from A. gramineus by the narrower median septum of the epigynum, and the pear-shaped spermathecae, which are closer to the epigynal groove. ! 195! Description. Male (holotype, UBC-SEM AR00038). Carapace length 1.6 (variation 1.4-1.7, n=6); abdomen length 1.6. Chelicera (Fig. 5.10E): yellow brown; not elaborate. Palp (Figs 5.10C-D): light brown. Embolus slightly curved. Retrolateral tibial apophysis finger-like. Palpal femur and patella with long white hairs. Measurements of legs: I 3.7, II 3.4, III 3.7, IV 4.4. Color in alcohol (Fig. 5.10A): carapace dark brown, with wide lateral margins and a central stripe behind fovea covered with white setae; abdomen light brown, with a pair of dark brown irregular stripes; venter with dark brown speckles; legs I and II dark brown, legs III and IV light brown. Female (paratype, UBC-SEM AR00039). Carapace length 1.6 (variation 1.6-1.9, n=3); abdomen length 1.7. Measurements of legs: I 3.6, II 3.4, III 3.9, IV 4.7. Epigynum (Figs 5.10F- G): window occupying about half of the epigynal plate; opening of copulatory duct at the posterior margin of the window. Copulatory ducts short and with accessory gland; spermathecae pear-shaped and close to each other. Color in alcohol (Fig. 5.10B): similar to that of the male, but the white lateral margins on carapace narrower, and the dark colored stripes on abdomen more obvious. Natural history. Specimens were found in leaf litter in a pine forest, usually beneath the litter. 5.4.2 Genus Anasaitis Bryant, 1950 Small to medium sized, usually with iridescent scales. They are mainly ground dwellers and can be found in leaf litter, on rocks or in grass clumps. Chelicera usually has one bicuspid promarginal tooth and one retromarginal tooth. First tibia often has three pairs of ventral macrosetae; first metatarsus has two pairs of ventral macrosetae. Embolus usually is very short and not coiled. Tegulum usually has distinct proximal lobe. Epigynum usually lacks obvious window. Copulatory duct is short. Spermatheca is swollen. As indicated by Bryant (1950: 169), this genus is probably close to Corythalia C. L. Koch and can be distinguished from it mainly by the genitalic structure: the embolus of the male palp is usually short with the spiral highly reduced, and the female epigynum usually lacks a distinct window. The reduced embolic spiral is also seen in Soesilarishius Makhan 2007 (see Ruiz 2011) and Popcornella, both of which are small leaf-litter dwelling euophryines, with Soesilarishius found in South America and Popcornella endemic to Caribbean. Anasaitis can be distinguished from Soesilarishius by the ! 196! presence of iridescent scales on the body and the highly reduced copulatory duct. It differs from Popcornella in the hairy appearance, the presence of iridescent scales on the body, and the extremely short embolus of the male palp. Molecular data (see Chapter 2) also suggest that Anasaitis, Soesilarishius and Popcornella fall into three distinct clades and the reduced embolic spiral may have evolved independently in these genera. Five species have been recorded from the Caribbean Islands and USA (Platnick 2011). Some other Caribbean species presently placed in Corythalia undoubtedly belong in Anasaitis instead based on their genitalic structures. Four new species from the Dominican Republic are described here. 5.4.2.1 Anasaitis adorabilis sp. nov. Figs 5.11-5.12 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo, 17.802° N, 71.349° W, elev. 5 m, 14 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-032. Paratypes: 1 female, same data as holotype; 7 males and 10 females, same data as holotype; 1 male and 1 female, same data as holotype; 1 male, DOMINICAN REPUBLIC: Pedernales: near Cabo Rojo, 17.915° N, 71.657° W, elev. 2 m, 16 July 2009, coll. G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-038; 6 males and 1 female, DOMINICAN REPUBLIC: Pedernales: near Pedernales, N 18.021° W 71.730°, 16 July 2009, coll. G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-039. Etymology. Latin adjective adorabilis (adorable), referring to the cute nature of the species. Diagnosis. Similar to Anasaitis brunnea in the color pattern and markings, but its carapace has a ‘U’-shaped marking behind PLEs, unlike A. brunnea. It can also be distinguished from A. brunnea by the shape of the genitalia, in which the proximal tegular lobe of the male palp is larger and the palpal bulb is narrower; the copulatory ducts of vulva are shorter. Description. Male (holotype, UBC-SEM AR00040). Carapace length 1.7 (variation 1.2-1.8, n=16); abdomen length 1.6. Chelicera dark brown. Palp (Fig. 5.12C): yellow brown. Proximal tegular lobe wide, retrolateral sperm duct loop narrower. Embolus very short. Retrolateral tibial ! 197! apophysis finger-like, tibia without ventral bump. Tibia of first leg with three ventral macrosetae retrolaterally and two ventral macroseta prolaterally. Measurements of legs: I 3.3, II 3.0, III 3.2, IV 2.7. Color in alcohol (Fig. 5.12A): carapace dark brown to brown, with a U-shaped marking behind PLEs composed of yellow brown iridescent scales; abdomen dark brown, light brownish yellow laterally, with a middle earthy yellow stripe extending to posterior end of abdomen; venter of abdomen brown with light yellow brown speckles; legs light yellow brown to brown, with dorsal and ventral fringes on first two pairs of legs. Face iridescent blue in life (Fig. 5.11C). Female (paratype, UBC-SEM AR00041). Carapace length 1.4 (variation 1.4-1.6, n=13); abdomen length 1.8. Tibia of first leg with three pairs of ventral macrosetae but two of them more prolateral. Measurements of legs: I 2.2, II 2.0, III 2.5, IV 2.3. Epigynum (Figs 5.12D-E): no obvious window or median septum. Copulatory ducts very short, with small accessory gland; spermathecae oval. Color in alcohol (Fig. 5.12B): similar to that of male, but the stripes on abdomen discontinuous and less distinct. Natural history. Specimens were found on shaded leaf litter in dry forest. 5.4.2.2 Anasaitis brunnea sp. nov. Figs 5.13-5.14 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Highway 44 south of Barahona, 18.138° N, 71.070° W, elev. 50 m, 14 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-031. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 1 female, same data as holotype; 2 females, same data as holotype; 1 female, same data as holotype; 1 female and 1 male, same data as holotype. Etymology. Latin adjective brunnea (brown), referring to the brown markings on the male dorsal abdomen. Diagnosis. Resembles Anasaitis adorabilis in color pattern, but differs by the lack of the ‘U’- shaped marking on the carapace, the shape of the embolus and the proximal tegular lobe, and the longer copulatory ducts. ! 198! Description. Male (holotype, UBC-SEM AR00042). Carapace length 1.4 (variation 1.1-1.4, n=3); abdomen length 1.3. Chelicera: dark brown. Palp (Figs 5.14C-D): light yellow brown to gray brown. Proximal tegular lobe obvious, retrolateral sperm duct loop wide. Embolus short. Retrolateral tibial apophysis finger-like, tibia without ventral bump. Measurements of legs: I 3.4, II 2.2, III 2.5, IV 2.3. Color in alcohol (Fig. 5.14A): carapace dark brown, with a pair of markings extending to posterior end of carapace composed of yellowish scales; abdomen dark brown, with a middle yellow brown stripe and yellow brown posterior end; ventral abdomen light yellow laterally and gray brown in the middle; legs light yellow brown to yellow brown. Female (paratype, UBC-SEM AR00043). Carapace length 1.3 (variation 1.1-1.3, n=6); abdomen length 1.5. Measurements of legs: I 2.1, II 1.8, III 2.4, IV 2.2. Epigynum (Figs 5.14E- F): no obvious window or median septum. Copulatory ducts narrow and short, with accessory gland; spermathecae spherical. Color in alcohol (Fig. 5.14B): similar to that of male, but the middle stripe on abdomen less distinct. Natural history. Specimens were found on leaf litter in a humid forest. 5.4.2.3 Anasaitis hebetata sp. nov. Fig 5.15 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García, 18.424° N, 71.112° W, elev.170 m, 21 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-049. Paratype: 1 male, same data as holotype. Etymology. Latin adjective hebetata (dull, not shining), referring to the non-colorful body of the species. Diagnosis. Palpal structure is very similar to that of Anasaitis brunnea, from which A. hebetata can be distinguished by the absence of distinct markings on the carapace and abdomen. Description. Male (holotype, UBC-SEM AR00044). Carapace length 1.4 (variation 1.4-1.5, n=2); abdomen length 1.3. Chelicera: dark brown. Palp (Fig. 5.15E): femur, patella and tibia ! 199! light yellow, tarsus yellow brown. Proximal tegular lobe big; embolus short; retrolateral sperm duct loop wide. Retrolateral tibial apophysis finger-like; tibia without ventral bump. Tibia of first leg with three ventral macrosetae retrolaterally and two macrosetae prolaterally. Measurements of legs: I 3.4, II 2.1, III 2.5, IV 2.1. Color in alcohol (Fig. 5.15D): carapace dark brown, abdomen gray brown, without distinct markings; legs yellow brown to dark yellow brown. Female. Unknown. Natural history. Specimens were found in grass clumps in dry desert forest. 5.4.2.4 Anasaitis laxa sp. nov. Figs 5.16-5.17 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García, 18.424° N, 71.112° W, elev.170 m, 21 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-049. Paratype: 1 female, same data as holotype. Etymology. Latin adjective laxa (wide), referring to the wide retrolateral tibial apophysis of the male palp. Diagnosis. Can be easily distinguished from other Anasaitis species by the V-shaped marking on the carapace, the wide retrolateral tibial apophysis, the more posteriorly located opening to the copulatory duct, and the longer and wider copulatory ducts. Description. Male (holotype, UBC-SEM AR00045). Carapace length 1.3; abdomen length 1.2. Chelicera: dark brown; with white scales on front face. Palp (Figs 5.17C-D): femur, patella and tibia light yellow, tarsus yellow brown. Proximal tegular lobe relatively small; retrolateral sperm duct loop wide; embolus very short. Retrolateral tibial apophysis wide and axe-like; tibia with an obvious ventral bump. Tibia of first leg with three ventral macrosetae retrolaterally and two ventral macrosetae prolaterally. Measurements of legs: I 2.1, II 2.0, III 2.7, IV 2.7. Color in alcohol (Fig. 5.17A): carapace dark brown, with a V-shaped marking behind PLEs and lateral marginal stripes at posterior half part composed of white or yellow iridescent scales; abdomen ! 200! dark brown, with a brown stripe and distinct earthy yellow markings; venter earthy yellow; legs light yellowish brown, femur of first leg brown. Female (paratype, UBC-SEM AR00046). Carapace length 1.4; abdomen length 1.5. Tibia of first leg with three ventral macrosetae retrolaterally and two prolaterally. Measurements of legs: I 2.2, II 2.1, III 2.8, IV 2.8. Epigynum (Figs 5.17E-F): no obvious window or median septum; opening to copulatory duct at posterior end of epigynum. Copulatory ducts short; spermathecae spherical. Color in alcohol (Fig. 5.17B): similar to that of male, but light colored markings on carapace and abdomen more distinct. Natural history. Specimens were found in grass clumps in dry desert forest. 5.4.3 Genus Antillattus Bryant, 1943 Medium sized spiders. Male chelicerae are usually elongate or enlarged with modifications such as projections on the front and/or ectal surface. Endites are usually enlarged distally. Chelicera usually has a fissident retromarginal tooth with more than two cusps or a unident retromarginal tooth (e.g. Antillattus applanatus). Embolus of the male palp is usually short with a simple curve. Epigynum has an obvious window with a median septum. Vulva usually has a pair of secondary spermathecae in addition to the primary spermathecae. Antillattus differs from Agobardus in the dentition of the female chelicera (Agobardus has one bicuspid retromarginal tooth in female) and the presence of the secondary spermatheca in vulva (Agobardus lacks secondary spermatheca). Although some species of Agobardus also have enlarged chelicerae in male, they usually lack a projection on the front or ectal surface of the chelicera as seen in Antillattus. Only two species have been reported from Hispaniola (Bryant 1943; Platnick 2011). However, some species placed in Agobardus, Pensacola and Siloca may actually belong to Antillattus. The new species is placed in Antillattus mainly because molecular data (see Chapter 2) indicates that it is closely related to the type species of Antillattus, A. gracilis Bryant, 1943. ! 201! 5.4.3.1 Antillattus applanatus sp. nov. Figs 5.18-5.19 Type material. Holotype: male, DOMINICAN REPUBLIC: La Vega: road Constanza to Ocoa, Valle Nuevo, 18.829° N, 70.691° W, elev. 2070 m, 12 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-023. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype. Etymology. Latin adjective applanatus (flattened), referring to the flattened body of the species. Diagnosis. Distinguished from other known Antillatus by the flattened body, the less elongated male chelicera, the absence of a spur on the front surface of the male chelicera, the unident retromarginal tooth and the vulva shape. Also, see the diagnosis of Agobardus bahoruco. Description. Male (holotype, UBC-SEM AR00047). Body relatively flattened. Carapace length 2.7; abdomen length 2.8. Chelicera (Fig. 5.19E): brown; internal margin concave and with two small processes; promargin with two teeth and retromargin with one tooth. Palp (Fig. 5.19C): tibia and tarsus yellow brown, other segments light yellow. Embolus short, retrolateral sperm duct loop narrow. Retrolateral tibial apophysis finger-like. Tibia of first leg with three pairs of ventral macrosetae and metatarsus with two pairs. Measurements of legs: I 6.6, II 5.1, III 5.7, IV 6.1. Color in alcohol (Fig. 5.19A): carapace gray brown, paler around fovea, lateral margins white; abdomen light yellow brown, with a pair of elongated gray brown medial markings and paired white spots at posterior part, venter with gray patches; first pair of legs brown to light brown, other legs light yellow with brown annuli. Female (paratype, UBC-SEM AR00048). Carapace length 2.5 (variation 2.1-2.5, n=2); abdomen length 3.5. Chelicera: with two promarginal teeth and one retromarginal tooth. First leg with three pairs of ventral macrosetae on tibia and two pairs on metatarsus. Measurements of legs: I 4.5, II 4.2, III 5.1, IV 5.5. Epigynum (Figs 5.19F-G): window relatively small, with opening to copulatory duct at its posterior margin. Copulatory ducts short, without accessory gland; secondary spermathecae not obvious, primary spermatheca oval. Color in alcohol (Fig. 5.19B): similar to that of the male. ! 202! Natural history. Specimens were collected on pine tree trunks in forest. 5.4.4 Genus Bythocrotus Simon, 1903 Medium sized spiders. Dull in color. Carapace is broad and high. Tibia of the male palp is widened and armed with multiple short and stout macrosetae. Palpal bulb is usually small with short and slightly curved embolus. Tegulum lacks distinct tegular lobe. Epigynum has window with a narrow median septum. Vulva has a pair of secondary spermathecae in addition to the primary spermathecae. Bythocrotus can be easily distinguished from other Caribbean euophryine genera by the unique male palpal structure. Only the type species, Bythocrotus cephalotes (Simon) (Bryant 1943) has been reported from Hispaniola. A new species from the Dominican Republic is described here. 5.4.4.1 Bythocrotus crypticus sp. nov. Figs 5.20-5.21 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García, 18.424° N, 71.112° W, elev. 170 m, 21 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-049. Paratypes: 1 female, same data as holotype; 2 females and 3 males, same data as holotype; 1 female, same data as holotype; 1 male, DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García, 18.417° N, 71.109° W, elev. 200 m, 13 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-030. Etymology. Latin adjective crypticus, referring to the camouflaging color pattern. Diagnosis. Differs from Bythocrotus cephalotes (Simon) (Bryant 1943; Galiano 1963) in the presence of a process at the distal end of the cymbium, the narrower palpal bulb and the smaller window of the epigynum. Description. Male (holotype, UBC-SEM AR00049). Carapace length 2.9 (variation 2.3-2.9, n=4); abdomen length 3.0. Chelicera (Fig. 5.21E): dark red brown; promargin with one bicuspid tooth and retromargin with one tooth. Palp (Figs 5.21C-D): dark yellow brown; cymbium ! 203! narrow, with a process at tip. Retrolateral tibial apophysis finger-like. Tibia with a prolateral spur and ten stout macrosetae (variation 8-11 stout macrosetae, n=4). First leg with three pairs of ventral macrosetae on tibia and two pairs on metatarsus. Measurements of legs: I 6.8, II 5.4, III 5.8, IV 5.7. Color in alcohol (Fig. 5.21A): carapace dark brown, with white scales at lateral margins; abdomen gray brown, with light brown speckles; legs dark brown to light yellow brown. Female (paratype, UBC-SEM AR00050). Carapace length 2.8 (variation 2.0-2.8, n=4); abdomen length 3.0. Chelicera: red brown; promargin with one bicuspid tooth and retromargin with one tooth. First leg with three pairs of ventral macrosetae on tibia and two pairs on metatarsus. Measurements of legs: I 5.8, II 5.1, III 5.7, IV 5.5. Epigynum (Figs 5.21F-G): window with very narrow median septum; opening to copulatory duct at posterior end of window. Copulatory ducts very short; secondary spermathecae smaller, primary spermathecae larger and spherical. Color in alcohol (Fig. 5.21B): carapace with no distinct lateral marginal white stripes; abdomen brown with light brown and dark brown markings; legs light yellow brown to dark brown. Natural history. Specimens were collected by beating tree branches with many small Tillandsia-like bromeliads, in desert savannah. 5.4.5 Genus Corticattus new genus Type species: Corticattus latus Zhang & Maddison, sp. nov. Etymology. The generic name is from the combination of cortex (bark) and attus (a common ending for salticid genera), referring to the habitat of the spiders; masculine in gender. Diagnosis. Small tree trunk dwelling spiders. Body is flattened. Chelicera has one bicuspid promarginal tooth and one unident retromarginal tooth. First pair of legs in male sometimes are widened. Embolus is long and coiled. Tegulum has distinctive proximal lobe and small distal triangular projection. Epigynum has windows. Corticattus is similar to the marpissoids Balmaceda (see Edwards 2006) and Metacyrba (see Edwards 2006) in the body form, but can be easily distinguished by the genitalia. Corticattus differs from other euophryine genera by the flattened body, the absence of retrolateral sperm duct loop and the presence of prolateral sperm ! 204! duct loop on the bulb, the presence of a triangular projection at the distal end of the tegulum, and the presence of a large accessory gland on the copulatory duct. 5.4.5.1 Corticattus guajataca sp. nov. Figs 5.22-5.23 Type material. Holotype: male, PUERTO RICO: Isabela: Bosque de Guajataca, 18.421° N, 66.966° W, elev. 217-250m, wet forest, 25 July 2009, coll. J. Zhang & G. B. Edwards, WPM#09-063. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 1 female, PUERTO RICO: Maricao: Bosque de Maricao, 18.150° N, 66.994° W, elev. 790m, wet forest, 26 Jul. 2009, Zhang & Edwards, WPM#09-064; 1 female, PUERTO RICO: Maricao: Bosque de Maricao at HWY105, 18.1768° N, 67.0105° W, elev. 400m, 26 July 2009, coll. J. Zhang & G. B. Edwards, WPM#09-065; 1 female, same data as previous. Etymology. A noun in apposition taken from the holotype locality. Diagnosis. See the diagnosis of Corticattus latus. Description. Male (holotype, UBC-SEM AR00075). Carapace length 1.1; abdomen length 0.9. Fovea far behind PLEs. Chelicera: dark brown. Palp (Figs 5.23C-D): yellowish brown. Embolus widened near the tip. Retrolateral tibial apophysis long and finger-like. First pair of legs robust, tibia with one ventral macroseta prolaterally and two ventral macrosetae retrolaterally; metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 1.9, II 1.5, III 1.8, IV 1.8. Color in alcohol (Fig. 5.23A): carapace dark, covered with many white setae; abdomen brownish, with some pale markings and a medial dark brown stripe; venter brown; first pair of legs yellow brown to dark yellow brown, other legs light yellow with dark annuli, anterior surface of femur of second leg dark. Female (paratype, UBC-SEM AR00076). Carapace length 1.0 (variation 1.0-1.3, n=4); abdomen length 1.1. Tibia and metatarsus of first leg with two pairs of ventral macrosetae each. Measurements of legs: I 1.5, II 1.3, III 1.7, IV 2.0. Epigynum (Figs 5.23E-F): window large, with a very narrow median septum; opening to copulatory duct along the median septum. Copulatory ducts short, with relatively large accessory gland; spermathecae small and oval. ! 205! Color in alcohol (Fig. 5.23B): similar to that of male except first pair of legs lighter in color. Natural history. Specimens were found on tree trunks in humid forest. 5.4.5.2 Corticattus latus sp. nov. Figs 5.24-5.25 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo, 17.802° N, 71.349° W, elev. 5 m, 18 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-045. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 female, same data as holotype; 1 female, DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo, 17.802° N, 71.349° W, elev.5 m, 14 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, WPM#09-032; 2 females, same data as previous. Etymology. Latin adjective latus (wide), referring to the wide first pair of legs in male. Diagnosis. Differs from Corticattus guajataca by the widened first pair of legs in male, the thinner embolus, the longer copulatory ducts, the smaller epigynal window and the more laterally located opening to copulatory duct. Description. Male (holotype, UBC-SEM AR00051). Carapace length 1.3; abdomen length 1.4. Fovea far behind PLEs. Chelicera: red brown. Palp (Figs 5.25C-D): gray brown to yellow brown. Embolus relatively thin. Retrolateral tibial apophysis long and finger-like; ventral tibial bump distinct. First pair of legs robust, with no macrosetae on tibia and metatarsus. Measurements of legs: I 1.7, II 1.6, III 1.8, IV 1.9. Color in alcohol (Fig. 5.25A): carapace eye area dark brown, remainder yellowish brown; dorsal area of carapace covered with white setae; abdomen light gray brown, with a medial gray brown stripe; venter gray brown; first pair of legs yellowish brown to dark yellowish brown, other legs light yellow with gray annuli. Female (paratype, UBC-SEM AR00052). Carapace length 1.3 (variation 1.2-1.3, n=6); abdomen length 1.6. Tibia of first leg with three ventral macrosetae; first metatarsus with two pairs of macrosetae. Measurements of legs: I 1.6, II 1.5, III 2.0, IV 2.1. Epigynum (Figs 5.25F- G): window with very narrow median septum; opening to copulatory duct almost at its center. ! 206! Copulatory ducts long, with large accessory gland; spermathecae spherical. Color in alcohol (Fig. 5.25B): similar to that of male except first pairs of legs lighter in color. Natural history. Specimens were found on or under bark or by beating branches in dry forest. 5.4.6 Genus Corythalia C. L. Koch, 1850 Small to large sized spiders. Corythalia can usually be distinguished from the closely related genus Anasaitis by the long and coiled embolus, the presence of a distinct epigynal window with median septum, and the longer copulatory duct. Corythalia is one of the most diverse although poorly studied euophryine genera in the New World, with 73 species described (Platnick 2011). The monophyly of this genus has never been tested and some species reported from the Caribbean Islands may be misplaced. The new species described here are placed in Corythalia because molecular data (see Chapter 2) indicate that they fall into a clade with the other Corythalia species from the South and Central America. 5.4.6.1 Corythalia broccai sp. nov. Figs 5.26-5.27 Type material. Holotype: male, DOMINICAN REPUBLIC: El Seibo: Pedro Sanchez - Miches Road, 18.958° N, 69.069° W, elev. 157 m, 22 July 2009, coll. W. Maddison, G. Ruiz, J. Brocca, WPM#09-053. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 male and 1 female, same data as holotype; 2 females, same data as holotype. Etymology. The specific epithet is a patronym in honor of Mr. J. Brocca, who provided great help in organizing the field work in the Dominican Republic in 2009 during which this species was discovered. Diagnosis. Can be distinguished from Dinattus minor Bryant, 1943 by the absence of distinct lateral extensions on the male carapace and the detailed structure of the male palp. Also, see the diagnosis of Corythalia bromelicola. Description. Male (holotype, UBC-SEM AR00053). Carapace length 1.8; abdomen length 1.6. ! 207! Chelicera (Fig. 5.27E): red brown; with one bicuspid promarginal tooth and one retromarginal tooth. Palp (Fig. 5.27C): dark brown. Embolus short and slightly curved; retrolateral sperm duct loop reduced. Retrolateral tibial apophysis long and finger-like. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. First pair of legs with dorsal fringe on femur and ventral fringes on femur, patella and tibia; second pair of legs with ventral fringe on femur. Measurements of legs: I 3.9, II 3.2, III 3.8, IV 3.8. Color in alcohol (Fig. 5.27A): carapace dark brown, with narrow lateral margins and markings around posterior eyes and behind fovea composed of iridescent scales; abdomen brown with light yellow brown markings; first two pairs of legs dark brown, last two pairs of legs yellow brown with dark brown markings. Female (paratype, UBC-SEM AR00054). Carapace length 2.0 (variation 1.7-2.0, n=5); abdomen length 2.2. Chelicera: with one bicuspid promarginal and one retromarginal tooth. First leg with three pairs of ventral macrosetae on tibia and metatarsus each. Measurements of legs: I 3.9, II 3.5, III 4.3, IV 4.2. Epigynum (Figs 5.27F-G): window occupying about half of epigynal plate, with opening to copulatory duct at its posterior margin. Spermathecae oval, located within the window. Color in alcohol (Fig. 5.27B): similar to that of the male except legs lighter in color. Natural history. Specimens were collected on leaf litter in a disturbed forest with planted cacao trees. 5.4.6.2 Corythalia bromelicola sp. nov. Figs 5.28-5.29 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Cachote-La Cienega Road, 18.0562 N, 71.1416 W, elev. 730 m, 20 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-048. Paratype: 1 female, same data as holotype. Etymology. Refers to the habitat, bromeliads. Diagnosis. Differs from other species by the larger and flatter body, the long legs, and the markings on the abdomen. Similar in palp to Corythalia broccai, but differs in the wider median septum of the epigynum and the shape of the vulva. ! 208! Description. Male (holotype, UBC-SEM AR00055). Carapace length 2.9; abdomen length 2.9. Chelicera (Fig. 5.29E): red brown; with one bicuspid promarginal tooth and one retromarginal tooth. Palp (Fig. 5.29C): yellow brown. Embolus short and slightly curved; retrolateral sperm duct loop reduced. Retrolateral tibial apophysis long and finger-like. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 8.7, II 6.2, III 6.6, IV 7.3. Abdomen with a mound in front of the spiracle opening. Color in alcohol (Fig. 5.29A): carapace dark brown, with narrow white lateral margins and markings around PLEs composed of white scales; abdomen dark brown with a white medial marking and white margins; venter gray; legs gray brown. Female (paratype, UBC-SEM AR00056). Carapace length 2.7; abdomen length 3.4. Chelicera: with one bicuspid promarginal tooth and one retromarginal tooth. First leg with three pairs of ventral macrosetae on tibia and metatarsus each. Measurements of legs: I 5.9, II 5.5, III 6.2, IV 6.8. Epigynum (Figs 5.29F-G): window almost round, occupying about half of epigynal plate, with opening to copulatory duct at its posterior margin. Copulatory ducts with accessory gland near the opening; spermathecae small and oval, located right anterior to the window. Color in alcohol (Fig. 5.29B): similar to that of the male except legs lighter in color. Natural history. Specimens were found in bromeliads in cloud forest. 5.4.6.3 Corythalia coronai sp. nov. Figs 5.30-5.31 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: Rio Mulito, 18.155° N, 71.758° W, elev. 270 m, 16 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-036. Paratypes: 1 female, same data as holotype; 7 females, same data as holotype; 12 males, same data as holotype; 1 female and 1 male, same data as holotype. Etymology. The specific epithet is a patronym in honor of Mr. Nicolas Corona, who helped in collecting specimens during the expedition to the Dominican Republic in 2009. Diagnosis. Unlike Corythalia broccai and C. bromelicola, the carapace is not swollen behind ! 209! the ALEs, and the copulatory duct openings are more anterior. It can be distinguished from Wallaba decora Bryant, 1943 by the shorter embolus and the narrower retrolateral sperm duct loop of the male palp, and the more widely spread iridescent scales on the male carapace. See also the diagnosis of C. peblique for differences with that species. Description. Male (holotype, UBC-SEM AR00057). Carapace length 2.0 (variation 1.5-2.1, n=14); abdomen length 1.8. Chelicera (Figs 5.31D-E): dark brown; with one bicuspid promarginal tooth and one retromarginal tooth; front surface with a straight ridge. Palp (Fig. 5.31C): tarsus brown, other segments light yellow. Embolus short and slightly curved; retrolateral sperm duct loop less than half of bulb width. Retrolateral tibial apophysis long and finger-like. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. First two pairs of legs with dorsal and ventral fringes. Measurements of legs: I 4.2, II 3.5, III 4.1, IV 4.4. Color in alcohol (Fig. 5.31A): carapace dark brown, with markings behind PLEs composed of iridescent scales; abdomen dark brown with a medial brown stripe; venter with two lines of yellow brown small spots behind genital groove; first two pairs of legs dark brown and last two pairs of legs yellow brown. Female (paratype, UBC-SEM AR00058). Carapace length 1.9 (variation 1.9-2.1, n=9); abdomen length 2.5. Chelicera: with one bicuspid promarginal and one retromarginal tooth. First leg with three pairs of ventral macrosetae on tibia and metatarsus each. Measurements of legs: I 3.3, II 3.1, III 3.8, IV 4.2. Epigynum (Figs 5.31F-G): window occupying about two thirds of epigynal plate, with opening to copulatory duct close to the margin. Copulatory ducts with accessory gland near the opening; spermathecae oval and located within the window. Color in alcohol (Fig. 5.31B): similar to that of the male but legs brown. Natural history. Specimens were found on leaf litter in broad-leaf forest. 5.4.6.4 Corythalia peblique sp. nov. Figs 5.32-5.33 Type material. Holotype: male, DOMINICAN REPUBLIC: Pedernales: Peblique, 18.059° N, 71.638° W, elev. 270 m, 17 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-041. Paratypes: 1 female, same data as holotype; 1 female and 1 male, same ! 210! data as holotype; 9 females, same data as holotype; 6 males, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Similar to Corythalia coronai, but can be distinguished by the markings on the carapace and abdomen, the longer male legs, the curved ridge on the front surface of the male chelicera, and the wider embolus. Description. Male (holotype, UBC-SEM AR00059). Carapace length 2.5 (variation 2.2-2.7, n=8); abdomen length 2.4. Chelicera (Figs 5.33D-E): dark brown; with one bicuspid promarginal tooth and one retromarginal tooth; anterior surface with a ridge. Palp (Fig. 5.33C): dark brown. Embolus wide with a simple and short curve. Retrolateral tibial apophysis long and finger-like. Tibia of first leg with three pairs of ventral macrosetae and a retrolateral macroseta partially ventral; metatarsus with three pairs of ventral macrosetae. Fringes present on ventral tibiae and patellae of first two pairs of legs, but relatively sparse on leg II. Measurements of legs: I 8.0, II 5.0, III 5.7, IV 5.5. Color in alcohol (Fig. 5.33A): carapace dark brown, without distinct markings; abdomen gray brown scattered with yellow brown speckles, with a light yellow brown band at anterior margin and two yellow brown stripes along the lateral margins of the middle brown patch; venter with two lines of yellow brown speckles; first three pairs of legs dark brown, fourth leg slightly lighter in color with proximal end of femur light yellow. Female (paratype, UBC-SEM AR00060). Carapace length 2.5 (variation 2.1-2.5, n=11); abdomen length 3.0. Chelicera: with one bicuspid promarginal and one retromarginal tooth. First leg with three pairs of ventral macrosetae on tibia and metatarsus each. Measurements of legs: I 4.9, II 4.5, III 5.2, IV 5.4. Epigynum (Figs 5.33F-G): window occupying more than half of epigynal plate, with opening to copulatory duct close to the anterior margin. Copulatory ducts with accessory gland near the opening; spermathecae oval and located within the window. Color in alcohol (Fig. 5.33B): similar to that of the male but legs yellow brown. Natural history. Specimens were found on leaf litter or on rocks in a relatively dry forest. This species seems to become active in the late afternoon. Several hours of collecting in the forest yielded many salticids, including many from leaf litter and rocks, but none (or very few) of this species until approximately 6:00 pm, when over the span of about 15 minutes they became ! 211! commonly seen. 5.4.7 Genus Popcornella new genus Type species: Popcornella spiniformis Zhang & Maddison, sp. nov. Etymology. The generic name is derived from the English “popcorn”, inspired by the nature of the spiders to jump away quickly in apparently random directions when disturbed; feminine in gender. Diagnosis. Small leaf-litter dwelling spiders. They are usually brown or dark brown in color. Chelicera has two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae; first metatarsus with two pairs. Embolus of the male palp is short and uncoiled. Epigynum has no distinct window structure, which is similar to Anasaitis Bryant, 1950, another leaf-litter associated genus mainly from the Caribbean. However, Popcornella differs from Anasaitis in lacking iridescent scales on the carapace and abdomen, and fringes on male legs; the retrolateral tibial apophysis of male palp is usually short. Popcornella is also distinguishable from Soesilarishius Makhan 2007 (see Ruiz 2011) by the short retrolateral tibial apophysis of the male palp and the usually highly shortened copulatory duct of the female vulva. Molecular data (see Chapter 2) indicate that Popcornella falls into a clade distinct from both Anasaitis and Soesilarishius. 5.4.7.1 Popcornella furcata sp. nov. Figs 5.34-5.35 Type material. Holotype: male, DOMINICAN REPUBLIC: La Vega: Reserva Científica Ébano Verde (antenna), 19.04° N, 70.518° W, elev. 1460m, cloud forest, 11 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, WPM#09-022. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 3 females, same data as holotype; 1 male, same data as holotype; 1 female, same data as holotype; 5 females, same data as holotype. Etymology. Latin adjective furcata, referring to the forked retrolateral tibial apophysis of male palp. ! 212! Diagnosis. Similar in color pattern to the other species in the Dominican Republic, Popcornella spiniformis, but easily distinguished by the forked retrolateral tibial apophysis, the distinct proximal tegular lobe of the male palp, and the shape of the epigynum and vulva. Description. Male (holotype, UBC-SEM AR00061). Carapace length 1.2 (variation 1.2-1.3, n=3); abdomen length 1.1. Chelicera: yellow brown. Palp (Figs 5.35C-D): yellow brown to gray brown; dorsal femur and patella covered with white setae. Retrolateral sperm duct loop almost half as wide as the tegulum width; tegular lobe distinct. Retrolateral tibial apophysis short and forked. Measurements of legs: I 2.6, II 2, III 2.2, IV 2.3. Color in alcohol (Fig. 5.35A): carapace dark to dark brown, without distinct markings except with some white setae behind fovea; abdomen light yellow brown, with parallel chevron-shaped dark markings; venter of abdomen gray brown; legs yellow brown. Female (paratype, UBC-SEM AR00062). Carapace length 1.2 (variation 1.0-1.2, n=10); abdomen length 1.3Measurements of legs: I 2.3, II 1.7, III 1.8, IV 2.2. Epigynum (Figs 5.35E- F): without distinct window structure; opening to copulatory duct close to epigynal groove. Copulatory ducts very short; spermathecae kidney-shaped. Color in alcohol (Fig. 5.35B): carapace dark brown to yellow brown; abdomen dark brown with light yellow brown speckles and markings; legs light yellow proximally and gray brown distally. Natural history. Specimens were found in leaf litter in a cloud forest, especially by taking loose well-drained litter onto a beating sheet and sorting through it. 5.4.7.2 Popcornella nigromaculata sp. nov. Figs 5.36-5.37 Type material. Holotype: male, PUERTO RICO: Maricao: Bosque de Maricao, 18.150° N, 66.994° W, elev. 790m, wet forest, 24 July 2009, J. Zhang & G. B. Edwards, WPM#09-062. Paratypes: 1 female, same data as holotype; 1 female and 1 male, same data as holotype; 1 male and 2 females, same data as holotype; 2 females, PUERTO RICO: Maricao: Bosque de Maricao, 18.150° N, 66.994° W, elev. 790m, wet forest, 26 July 2009, coll. J. Zhang & G. B. Edwards, WPM#09-064. ! 213! Etymology. The specific epithet (an adjective) refers to the dark markings at the posterior end of dorsal abdomen of male. Diagnosis. Differs from the other Puerto Rican species, Popcornella yunque, by the markings on the abdomen, the shorter embolus and the shorter copulatory ducts. Description. Male (holotype, UBC-SEM AR00063). Carapace length 0.8; abdomen length 0.7. Chelicera: dark brown. Palp (Figs 5.37C-D): dark yellow brown. Retrolateral sperm duct loop invisible. Retrolateral tibial apophysis short, shaped like a bird beak. Measurements of legs: I 1.8, II 1.2, III 1.5, IV 1.6. Color in alcohol (Fig. 5.37A): carapace dark to dark brown, without distinct markings; abdomen light gray brown to light brown, with three parallel chevron-shaped dark markings at posterior part, posterior end of abdomen dark; venter of abdomen light brown; legs light yellow. Female (paratype, UBC-SEM AR00064). Carapace length 0.8; abdomen length 0.7. Measurements of legs: I 1.5, II 1.2, III 1.4, IV 1.5. Epigynum (Figs 5.37E-F): without distinct window structure, posterior part with a transverse margin; opening to copulatory duct anterior to spermatheca. Copulatory ducts short; spermathecae oval. Color in alcohol (Fig. 5.37B): similar to that of male. Natural history. Specimens were found in leaf litter in wet forest. 5.4.7.3 Popcornella spiniformis sp. nov. Figs 5.38-5.39 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Cachote, 18.098° N, 71.187° W, elev. 1220 m, 19-20 July 2009, coll. coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-047. Paratypes: 1 female, same data as holotype; 1 male and 1 female, same data as holotype; 1 male and 2 females, same data as holotype; 1 male 5 females, DOMINICAN REPUBLIC: Barahona: Cachote, 18.101° N, 71.194° W, elev. 1200 m, 18-19 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-046. Etymology. The specific epithet, an adjective, combines the Latin spinus (spine) and formis ! 214! (shaped), referring to the spine-like embolus of the male palp. Diagnosis. Differs from the other species in the Dominican Republic, Popcornella furcata, by the hooked retrolateral tibial apophysis, the palpal bulb shape, and the shape of the epigynum and spermatheca. Description. Male (holotype, UBC-SEM AR00065). Carapace length 1.0 (variation 0.9-1.1, n=4); abdomen length 0.9. Chelicera: yellow brown. Palp (Figs 5.39C-D): coxa, trochanter and femur dark brown, patella, tibia and cymbium light yellow. Retrolateral sperm duct loop almost half as wide as tegulum width. Retrolateral tibial apophysis short and hooked at the tip, with a hump at the base. Measurements of legs: I 2.2, II 1.6, III 1.9, IV 1.9. Color in alcohol (Fig. 5.39A): carapace dark to dark brown, without distinct markings; abdomen gray brown, with light yellow brown speckles and markings; legs light yellow to brownish. Female (paratype, UBC-SEM AR00066). Carapace length 0.9 (variation 0.8-0.9, n=9); abdomen length 1.0. Measurements of legs: I 1.8, II 1.5, III 1.8, IV 1.9. Epigynum (Figs 5.39F- G): without distinct window structure; opening to copulatory duct far anterior to the genital groove. Copulatory ducts very short; spermathecae almost oval. Color in alcohol (Figs 5.39B): similar to that of male. Natural history. Specimens were found in leaf litter in cloud forest. 5.4.7.4 Popcornella yunque sp. nov. Figs 5.40-5.41 Type material. Holotype: male, PUERTO RICO: Río Grande: El Yunque National Forest: trail from HWY186 S of El Verde station, 18.3174° N, 65.8314° W, elev. 470m, 29 July 2009, coll. J. Zhang & G. B. Edwards, WPM#09-071. Paratypes: 1 female, same data as holotype; 2 males and 1 female, same data as holotype; 4 males and 1 female, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. See the diagnosis of Popcornella nigromaculata. ! 215! Description. Male (holotype, UBC-SEM AR00073). Carapace length 0.8 (variation 0.8-0.9, n=7); abdomen length 0.5. Chelicera: yellow brown. Palp (Figs 5.41C-D): pale yellow with cymbium brown. Embolus long spur-like, not spiral, retrolateral sperm duct loop not obvious. Retrolateral tibial apophysis short. Measurements of legs: I 1.5, II 1.1, III 1.3, IV 1.4. Color in alcohol (Fig. 5.41A): carapace dark brown, without distinct markings; abdomen dark brown, with large brownish patches; ventral abdomen brownish, with two dark markings at posterior end; legs grayish brown to brownish. Female (paratype, UBC-SEM AR00074). Carapace length 0.8 (variation 0.8-0.9, n=3); abdomen length 0.8. Measurements of legs: I 1.3, II 1.1, III 1.2, IV 1.4. Epigynum (Figs 5.41E- F): without distinct window structure. Copulatory ducts short; spermathecae kidney-shaped. Color in alcohol (Fig. 5.41B): similar to that of male. Natural history. Specimens were found in leaf litter in cloud forest. 5.4.8 Genus Truncattus new genus Type species: Truncattus flavus Zhang & Maddison, sp. nov. Etymology. The generic name is from the combination of truncus (tree trunk, where the species is usually found) and -attus (a common ending for salticid genera); masculine in gender. Diagnosis. Small tree trunk dwelling spiders. Chelicera has two promarginal teeth and one bicuspid retromarginal tooth. First tibia has three pairs of ventral macrosetae; first metatarsus has two pairs. Male chelicerae are not modified. Embolus is usually not very long but curved. Tegulum has an obvious proximal lobe, and palpal tibia has a ventral bump. Epigynum has a window with median septum. Some species have obvious secondary spermathecae in addition to the primary spermathecae. Genitalia are similar to those of Antillattus Bryant, 1943 and Petemathis Prószy!ski & Deeleeman-Reinhold, 2012. It differs from Antillattus by the non- modified male chelicerae and endites; and from Petemathis by the bicuspid retromarginal tooth on the chelicera (three or more cusps in Petemathis). This genus is also similar to Caribattus (see Peckham & Peckham 1901) in color pattern, but can be distinguished by the bicuspid retromarginal tooth (Caribattus has one unident retromarginal tooth), and the presence of a ! 216! proximal tegular lobe on the male palp (the tegular lobe is absent in Caribattus). 5.4.8.1 Truncattus cachotensis sp. nov. Figs 5.42-5.43 Type material. Holotype: male, DOMINICAN REPUBLIC: Barahona: Cachote, 18.098° N, 71.187° W, elev. 1220 m, 19-20 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-047. Paratypes: 1 female, same data as holotype; 1 male, DOMINICAN REPUBLIC: Barahona: Cachote, 18.101° N, 71.194° W, elev. 1200 m, 18-19 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-046; 1 female and 1 male, same data as previous; 1 male and 3 females, DOMINICAN REPUBLIC: Barahona: Cachote-La Cienega Road, 18.0562° N, 71.1416° W, elev. 730 m, 20 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, G. Ruiz, N. Corona, WPM#09-048. Etymology. The specific epithet, to be treated as a Latin adjective, refers to the type locality. Diagnosis. Differs from other Truncattus species by the wider median septum of the epigynum, the shape of the vulva and the shape of the retrolateral tibial apophysis. Description. Male (holotype, UBC-SEM AR00067). Carapace length 1.1 (variation 1.1-1.3, n=4); abdomen length 1.1. Chelicera: dark brown to yellow brown. Palp (Fig. 5.43C): light yellow. Retrolateral sperm duct loop very narrow. Retrolateral tibial apophysis short, with tip pointed. Measurements of legs: I 2.1, II 2.0, III 2.2, IV 2.4. Color in alcohol (Fig. 5.43A): carapace dark brown with narrow lateral white margins; abdomen gray brown with light yellow markings in the middle and many white yellow speckles, and with a pair of dark spots in the center; venter sandy yellow with brown patches; legs light yellow to dark brown, although in life the femora are greenish yellow (Figs 5.42A-B). Female (paratype, UBC-SEM AR00068). Carapace length 1.2 (variation 1.1-1.3, n=5); abdomen length 1.6. Measurements of legs: I 2.1, II 2.0, III 2.3, IV 2.7. Epigynum (Figs 5.43D- E): window small, with opening to copulatory duct at posterior end. Copulatory ducts short, with accessory gland; secondary spermathecae not obvious, primary spermathecae large and spherical. Color in alcohol (Fig. 5.43B): similar to that of male, but first pair of legs lighter in ! 217! color. Natural history. Specimens were found on tree trunks in a cloud forest or along its edge. 5.4.8.2 Truncattus dominicanus sp. nov. Figs 5.44-5.45 Type material. Holotype: male, DOMINICAN REPUBLIC: La Vega: P. N. Armando Bermúdez, 19.06-19.07° N, 70.86-70.88° W, elev. 1120-1250 m, 8-9 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, J. Brocca, WPM#09-018. Paratypes: 1 female, same data as holotype; 10 males and 7 females, same data as holotype; 1 female and 1 male, DOMINICAN REPUBLIC: La Vega: near Manabao, 19.076° N, 70.827° W, elev. 980 m, 8-10 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, J. Brocca, WPM#09-017; 9 males and 7 females, same data as previous; 1 male, DOMINICAN REPUBLIC: La Vega: Reserva Científica Ébano Verde (station), 19.033° N, 70.543° W, elev. 1080 m, 10-11 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, WPM#09-021; 1 female, same data as previous. Etymology. The specific epithet, to be treated as a Latin adjective, refers to the country where the species was found. Diagnosis. Easily distinguished from the other two species by the large window and the narrow median septum of the epigynum, the shape of the vulva and the larger embolic disc. Description. Male (holotype, UBC-SEM AR00069). Carapace length 1.4 (variation 1.2-1.5, n=22); abdomen length 1.5. Chelicera (Fig. 5.45D): dark yellow brown. Palp (Fig. 5.45C): femur, patella and tibia light yellow, cymbium yellow brown. Embolus relatively longer, retrolateral sperm duct loop very narrow. Retrolateral tibial apophysis long and finger-like; ventral tibial bump small. Measurements of legs: I 2.7, II 2.5, III 2.6, IV 3.0. Color in alcohol (Fig. 5.45A): carapace dark red brown without lateral white margins, posterior part with a medial yellow brown stripe; abdomen brown with a medial light yellow elongated marking and many light yellow speckles; venter light brown, with a wide medial gray brown stripe behind genital groove; legs yellow brown, with indistinct gray brown annuli. ! 218! Female (paratype, UBC-SEM AR00070). Carapace length 1.6 (variation 1.2-1.6, n=17); abdomen length 2.5. Tibia of first leg with three pairs of ventral macrosetae; first metatarsus with two pairs. Measurements of legs: I 2.7, II 2.5, III 3.1, IV 3.4. Epigynum (Figs 5.45E-F): window large, with opening to copulatory duct close to the middle along its outer margin. Secondary spermatheca close to the opening, primary spermatheca small and oval. Color in alcohol (Fig. 5.45B): similar to that of male. Natural history. Specimens were found on tree trunks. 5.4.8.3 Truncattus flavus sp. nov. Figs 5.46-5.47 Type material. Holotype: male, DOMINICAN REPUBLIC: La Vega: P. N. Armando Bermúdez, 19.06-19.07° N, 70.86-70.88° W, elev. 1120-1250 m, 8-9 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, J. Brocca, WPM#09-018. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 1 female and 1 male, DOMINICAN REPUBLIC: La Vega: near Manabao, 19.076° N, 70.827° W, elev. 980 m, 8-10 July 2009, coll. W. Maddison, G. B. Edwards, J. Zhang, J. Brocca, WPM#09-017; 12 females and 8 males, same data as previous. Etymology. Latin adjective flavus (yellow), referring to the yellow palp of the male. Diagnosis. Similar to Truncattus cachotensis in shape of the epigynum and palp, but differs in the narrower median septum of the epigynum, the large secondary spermatheca and the long retrolateral tibial apophysis. Description. Male (holotype, UBC-SEM AR00071). Carapace length 1.3 (variation 1.1-1.4, n=11); abdomen length 1.4. Chelicera (Fig. 5.47D): yellow brown. Palp (Fig. 5.47C): femur, patella and tibia light yellow; cymbium brown. Retrolateral sperm duct loop narrow. Retrolateral tibial apophysis long and thin; ventral tibial bump obvious. First leg with ventral fringes on tibia and patella. Measurements of legs: I 2.5, II 1.9, III 2.1, IV 2.4. Color in alcohol (Fig. 5.47A): carapace dark brown to red brown with narrow lateral marginal stripes composed of white setae, posterior part with white setae medially; abdomen gray brown with light yellow ! 219! markings in the middle, posterior end dark brown; venter light yellow; first pair of legs dark brown with tarsus light yellow, other legs light yellow to brown. Female (paratype, UBC-SEM AR00072). Carapace length 1.1 (variation 1.0-1.2, n=14); abdomen length 1.7. Measurements of legs: I 1.7, II 1.6, III 1.7, IV 2.1. Epigynum (Figs 5.47E- F): window small, with opening to copulatory duct at posterior end. Copulatory ducts short, with accessory gland; secondary spermathecae obvious; primary spermathecae oval. Color in alcohol (Fig. 5.47B): similar to that of male, but first pair of legs lighter in color. Natural history. Specimens were found on tree trunks. ! 220! Figure 5.1. Agobardus bahoruco sp. nov. A-C. male holotype; D. female paratype. ! 221! Figure 5.2. Agobardus bahoruco sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, back view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; D, 0.2 mm; C, E-F, 0.1 mm. ! 222! Figure 5.3. Agobardus cordiformis sp. nov. A-B. male holotype; C-D. female paratype. ! 223! Figure 5.4. Agobardus cordiformis sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 224! Figure 5.5. Agobardus gramineus sp. nov. A-B. male paratype; C-D. female paratype. ! 225! Figure 5.6. Agobardus gramineus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 226! Figure 5.7. Agobardus oviedo sp. nov. A-D. male holotype; E-F. female paratype. ! 227! Figure 5.8. Agobardus oviedo sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, back view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; D, 0.2 mm; C, E-F, 0.1 mm. ! 228! Figure 5.9. Agobardus phylladiphilus sp. nov. A-C. male paratype; D. female paratype. ! 229! Figure 5.10. Agobardus phylladiphilus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B, 2.0 mm; E, 0.2 mm; C-D, F-G, 0.1 mm. ! 230! Figure 5.11. Anasaitis adorabilis sp. nov. A-C. male paratype; D. female paratype. ! 231! Figure 5.12. Anasaitis adorabilis sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. epigynum, ventral view; E. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-E, 0.1 mm. ! 232! Figure 5.13. Anasaitis brunnea sp. nov. A-B. male holotype; C-D. male paratype; E. one female paratype; F. another female paratype. ! 233! Figure 5.14. Anasaitis brunnea sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 234! Figure 5.15. Anasaitis hebetata sp. nov. A-C. male paratype; D. male paratype, dorsal view; E. male left palp, ventral view. Scale bars: D, 0.5 mm; E, 0.1 mm. ! 235! Figure 5.16. Anasaitis laxa sp. nov. A-C. male holotype; D. female paratype. ! 236! Figure 5.17. Anasaitis laxa sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 237! Figure 5.18. Antillatus applanatus sp. nov. A-C. male holotype; D. female paratype. ! 238! Figure 5.19. Antillatus applanatus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male endites and labium, ventral view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A, 2.0 mm; B, 1.0 mm; D-E, 0.2 mm; C, F-G, 0.1 mm. ! 239! Figure 5.20. Bythocrotus crypticus sp. nov. A-D. male paratype; E-F. female paratype. ! 240! Figure 5.21. Bythocrotus crypticus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, prolateral view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. ! 241! Figure 5.22. Corticattus guajataca sp. nov. A-C. male holotype; D. one female paratype from Marico; E-F. another female paratype from Guajataca. ! 242! Figure 5.23. Corticattus guajataca sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 243! Figure 5.24. Corticattus latus sp. nov. A-D. male holotype; E-F. female paratype. ! 244! Figure 5.25. Corticattus latus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female right chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. ! 245! Figure 5.26. Corythalia broccai sp. nov. A-D. male paratype; E-F. female paratype. ! 246! Figure 5.27. Corythalia broccai sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male carapace, dorsal view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A- B, 1.0 mm; D, 0.2 mm; C, E-G, 0.1 mm. ! 247! Figure 5.28. Corythalia bromelicola sp. nov. A-D. male holotype; E-F. female paratype. ! 248! Figure 5.29. Corythalia bromelicola sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male carapace, dorsal view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A, 2.0 mm; B, 1.0 mm; C-G, 0.2 mm. ! 249! Figure 5.30. Corythalia coronai sp. nov. A-D. male paratype; E-F. female paratype. ! 250! Figure 5.31. Corythalia coronai sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, front view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; D-E, 0.2 mm; C, F-G, 0.1 mm. ! 251! Figure 5.32. Corythalia peblique sp. nov. A-D. male paratype; E-F. female paratype. ! 252! Figure 5.33. Corythalia peblique sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, front view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; D-E, 0.2 mm; C, F-G, 0.1 mm. ! 253! Figure 5.34. Popcornella furcata sp. nov. A-C. male paratype; D. female paratype. ! 254! Figure 5.35. Popcornella furcata sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 255! Figure 5.36. Popcornella nigromaculata sp. nov. A-B. male paratype; C-D. female paratype. ! 256! Figure 5.37. Popcornella nigromaculata sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 257! Figure 5.38. Popcornella spiniformis sp. nov. A-D. male paratype; E-F. female paratype. ! 258! Figure 5.39. Popcornella spiniformis sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. ! 259! Figure 5.40. Popcornella yunque sp. nov. A-C. male paratype; D. female paratype. ! 260! Figure 5.41. Popcornella yunque sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 261! Figure 5.42. Truncattus cachotensis sp. nov. A-B. male paratype; C-D. female paratype. ! 262! Figure 5.43. Truncattus cachotensis sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. epigynum, ventral view; E. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-E, 0.1 mm. ! 263! Figure 5.44. Truncattus dominicanus sp. nov. A-B. male paratype; C-D. female paratype. ! 264! Figure 5.45. Truncattus dominicanus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, back view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 265! Figure 5.46. Truncattus flavus sp. nov. A-B. male paratype; C-D. female paratype. ! 266! Figure 5.47. Truncattus flavus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male right chelicera, back view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! ! 267! 6 New euophryine jumping spiders from Papua New Guinea (Araneae: Salticidae: Euophryinae) 1 6.1 Synopsis Thirty-four new species and five new genera of euophryine jumping spiders from Papua New Guinea are described. The new genera are Chalcolemia (type species C. nakanai sp. nov.), Phasmolia (type species P. elegans sp. nov.), Variratina (type species V. minuta sp. nov.), Viribestus (type species V. suyanensis sp. nov.) and Zabkattus (type species Z. brevis sp. nov., plus new species Z. furcatus sp. nov., Z. richardsi sp. nov. and Z. trapeziformis sp. nov.). The other new species belong to the genera Bathippus (B. directus sp. nov., B. gahavisuka sp. nov., B. korei sp. nov., B. madang sp. nov.), Canama (C. extranea sp. nov., C. fimoi sp. nov., C. triramosa sp. nov.), Omoedus (O. brevis sp. nov., O. darleyorum sp. nov., O. meyeri sp. nov., O. omundseni sp. nov., O. papuanus sp. nov., O. swiftorum sp. nov., O. tortuosus sp. nov.), Paraharmochirus (P. tualapaensis sp. nov.), Sobasina (S. wanlessi sp. nov.), Thorelliola (T. aliena sp. nov., T. crebra sp. nov., T. joannae sp. nov., T. squamosa sp. nov., T. tamasi sp. nov., T. tualapa sp. nov., T. zabkai sp. nov.) and Xenocytaea (X. agnarssoni sp. nov., X. albomaculata sp. nov., X. proszynskii sp. nov.). The genera Pystira and Zenodorus are both considered as junior synonyms of Omoedus because of their similar genitalic structure. Species of these two genera are therefore transferred to Omoedus. Diagnostic illustrations are provided for all of the described species, and photographs of living spiders are also added when available. 6.2 Introduction Euophryine jumping spiders are unusually abundant and diverse in New Guinea (Maddison & Zhang 2011). In expeditions to Papua New Guinea in 2008 and 2009 organized by Conservation International, the majority of salticid species collected were euophryines, about 80 species. In addition to the high species diversity, the fauna of euophryines in Papua New Guinea is also remarkable for the presence of some unique body forms not seen in euophryines elsewhere. For instance, Coccorchestes closely resemble curculionid beetles; Leptathamas and Athamas have a very high carapace, and males of Leptathamas resemble piles of debris, holding legs in strange !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! \"!A version of this chapter has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from Papua New Guinea (Araneae: Salticidae: Euophryinae). Names of new taxa are not finalized for nomenclature. ! ! 268! poses and walking in a jerky fashion (Maddison & Zhang 2011); spiders of Sobasina and Paraharmochirus are ant-like. Euophryines have also adapted to various microhabitats in New Guinea. On leaf litter can be found Zabkattus, Sobasina and some species of Omoedus and Thorelliola, on tree trunks can be found Variratina, Paraharmochirus and other Thorelliola, while on foliage are many genera including Bathippus, Bulolia, Canama, Chalcolemia, Coccorchestes, Cytaea, Euryattus, Palpelius, Phasmolia, Xenocytaea and other species of Omoedus. In this Chapter, I report five new genera: Chalcolemia (one species), Phasmolia (one species), Variratina (one species), Viribestus (one species) and Zabkattus (four species). In addition, 26 species of the genera Bathippus (four species), Canama (three species), Omoedus (seven species), Paraharmochirus (one species), Sobasina (one species), Thorelliola (seven species) and Xenocytaea (three species) are also described. Here I consider the genera Pystira and Zenodorus as junior synonyms of Omoedus because of their similar genitalic structure. Species of these two genera are transferred to Omoedus. Most of the species described here were chosen in order to give names for the taxa included in molecular phylogenetic study on the subfamily Euophryinae (see Chapter 2). 6.3 Material and methods During expeditions to Papua New Guina in 2008 and 2009, jumping spiders from five areas were sampled (Maddison & Zhang 2011): Wanakipa (Southern Highlands Province), near Porgera (Enga Province), Mt. Gahavisuka (Eastern Highlands Province), Varirata National Park (Central Province), and Mts. Nakanai (New Britain). Multiple habitats ranging from the highland wet forest to the lowland rainforest and disturbed forest were explored. Collecting mainly involved beating foliage and visual inspection on ground and on tree trunks. Photographs of living specimens were mostly taken with a Pentax Optio 33WR digital camera. For the macro capability, a small lens was glued to it. Living spider photographs of Xenocytaea albomaculata were taken by Piotr Naskrecki with a Canon 40D camera with a 100 mm macrolens. Photographs of preserved specimens were taken under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0. Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings ! 269! were made with a drawing tube on a Nikon ME600L compound microscope. Terminology is standard for Araneae. All measurements are given in millimeters. Descriptions of color pattern are based on the alcohol-preserved specimens. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; AME, anterior median eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the \"small eyes\"). Specimens are deposited in the Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM). 6.4 Taxonomy 6.4.1 Genus Bathippus Thorell, 1892 Medium to large sized spiders. Males have elongate chelicerae. Female chelicera has a unident retromarginal tooth. Male palpal bulb is oval, without proximal tegular lobe. Retrolateral tibial apophysis is long or short, and usually finger-like. Epigynum has a window, with the opening to the copulatory duct at the anterior or median of the vulva. Thirty-one species have been described from Southeast Asia, Papua New Guinea, Pacific islands and Australia (Platnick 2011). The four new species are placed within this genus because of their similar morphology to the type species Bathippus macrognathus (see Prószy!ski 1976) and their close relationship with B. macrognathus as indicated by molecular data (see Chapter 2). 6.4.1.1 Bathippus directus sp. nov. Figs 6.1-6.2 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1000-1100 m a.s.l., 11-22 July 2008, forest interior and river side, coll. W. Maddison & Luc Fimo Tuki, WPM#08-008. Paratypes: 1 female, same data as holotype; 4 males and 2 females, same data as holotype; 1 male, same data as holotype; 1 female, same data as holotype; 1 female, PAPUA NEW GUINEA: Southern ! 270! Highlands Province: Umgé, 5.304-5.305° S, 142.510-142.512° E, elev. 1400-1450 m a.s.l., 15- 19 July 2008, coll. W. Maddison & Aislan Tama Wanakipa Indiaf, WPM#08-012. Etymology. Latin directus, referring to the male chelicerae projecting almost straight forward. Diagnosis. Differs from other species by the presence of two cusps near the base of the male cheliceral fang, the long embolus and the abruptly narrowed distal end of the retrolateral tibial apophysis of the male palp, the large window of the epigynum, and the large and spherical spermatheca of the vulva. Description. Male (holotype, UBC-SEM AR00077). Carapace length 3.6 (variation 3.5-3.8, n=6); abdomen length 5.0. Chelicera (Figs 6.2F-G): dark red brown; elongate and extending forward; promargin with four teeth and retromargin with one big tooth; fang with two cusps near the base. Palp (Figs 6.2C-E): sandy yellow. Retrolateral loop of sperm duct wide, occuping about three quaters of the bulb width. Embolus coiled for about half a circle. Retrolateral tibial apophysis thick proximally and narrowed abruptly near distal end. Femur of palp long and curved. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with four pairs. Measurements of legs: I 13.7, II 9.0, III 10.7, IV 9.3. Color in alcohol (Fig. 6.2A): carapace yellow orange with indistinct gray markings; abdomen gray brown, with yellow orange speckles and yellowish stripes; legs light yellow to brown. Female (paratype, UBC-SEM AR00078). Carapace length 2.8 (variation 2.8-3.1, n=5); abdomen length 4.9. Chelicera (Fig. 6.2H): with two promarginal teeth and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with four pairs. Measurements of legs: I 6.3, II 5.7, III 7.5, IV 6.7. Epigynum (Figs 6.2I-J): window large with a narrow median septum; opening to copulatory duct at anterior lateral edge of the window. Copulatory duct short and slightly curved; spermatheca large and spherical. Color in alcohol (Fig. 6.2B): similar but lighter than that of male. Natural history. Specimens were collected by beating forest understory vegetation. ! 271! 6.4.1.2 Bathippus gahavisuka sp. nov. Figs 6.3-6.4 Type material. Holotype: male, PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park, 6.015° S, 145.412° E, elev. 2320 m a.s.l., 1-2 August 2008, coll. W. Maddison, WPM#08-025. Paratype: 1 female, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Similar in color pattern and markings to B. korei, but differs in the presence of a curved spur on the prolateral surface of male chelicera, the thinner embolus and the shorter retrolateral tibial apophysis of male palp, and the wider median septum of epigynum. Description. Male (holotype, UBC-SEM AR00079). Carapace length 2.9; abdomen length 3.8. Chelicera (Figs 6.4C-D): dark red brown; elongate; promargin with three teeth and retromargin with one tooth; fang with a small cusp in the middle; prolateral surface with a curved spur. Palp (Figs 6.4E-G): sandy yellow to red brown. Tegulum narrow. Embolus short and slightly curved. Retrolateral tibial apophysis thin and finger-like. Femur of palp very long and curved. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with four pairs. Measurements of legs: I 8.9, II 7.2, III 7.8, IV 7.1. Color in alcohol (Fig. 6.4A): carapace brown with two lateral cream stripes; abdomen grayish brown, with light yellow markings; legs yellowish to dark red brown. Female (paratype, UBC-SEM AR00080). Carapace length 2.7; abdomen length 2.8. Chelicera: with two promarginal and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with four pairs. Measurements of legs: I 6.0, II 5.4, III 6.7, IV 6.2. Epigynum (Figs 6.4H-I): window occupying about half of the epigynal plate; median septum wide almost reaching the anterior margin of window. Copulatory duct short with an accessory gland; spermatheca relatively small and oval. Color in alcohol (Fig. 6.4B): carapace light orange, with two lateral yellowish stripes; abdomen brownish, with yellowish speckles and a large leaf-like marking in the middle. Natural history. Specimens were collected by beating forest understory vegetation. ! 272! 6.4.1.3 Bathippus korei sp. nov. Figs 6.5-6.6 Type material. Holotype: male, PAPUA NEW GUINEA: Central Province: Varirata National Park, 9.436° S, 147.364° E, elev. 740 m a.s.l., 4 August 2008, coll. W. Maddison, A. Kore & J. Kore, WPM#08-029. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 1 female, same data as holotype; 1 male, same data as holotype. Etymology. The specific epithet is a patronym in honor of Mr. Agustus Kore, who participated in collecting specimens in Varirata National Park. Diagnosis. Differs from other species by the presence of seven teeth on the promargin of female and male chelicerae, and the wider embolus of male palp. Description. Male (holotype, UBC-SEM AR00081). Carapace length 3.5 (variation 3.5-3.9, n=3); abdomen length 4.9. Chelicera (Figs 6.6F-G): dark red brown; elongate; promargin with seven teeth and a small cusp near the fang base, retromargin without distinct tooth but with a big process at the base of the fang. Palp (Figs 6.6C-E): light yellow. Tegulum narrow. Embolus relatively wide and slightly curved. Retrolateral tibial apophysis finger-like with a small cusp near the tip. Femur of palp long and curved. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with four pairs. Measurements of legs: I 12.2, II 9.5, III 10.8, IV 9.3. Color in alcohol (Fig. 6.6A): carapace orange; abdomen grayish brown, with light yellow speckles and markings; legs sandy yellow to yellow brown. Female (paratype, UBC-SEM AR00082). Carapace length 2.9; abdomen length 4.5. Chelicera (Fig. 6.6H): with seven promarginal and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with four pairs. Measurements of legs: I 6.2, II 5.5, III 7.2, IV 6.5. Epigynum (Figs 6.6I-J): window occupying almost three quaters of the epigynal plate; median septum relatively wide; opening to the copulatory duct at anterior edge of median septum. Copulatory duct short; spermatheca long oval. Color in alcohol (Fig. 6.6B): carapace orange, with a light yellow stripe in thoracic area; abdomen brown, with a yellowish leaf-like marking in the middle; legs sandy yellow. ! 273! 6.4.1.4 Bathippus madang sp. nov. Fig 6.7 Type material. Holotype: male, PAPUA NEW GUINEA: Madang, Adalbert Mts., Sewan-Keki, 4.704° S, 145.419° E, elev. 700m, 4 May 2006, coll. Balke & Manaono (PNG 51). Etymology. A noun in apposition taken from the type locality. Diagnosis. Differs from the other species by the unique teeth pattern on the male chelicerae. Similar in the male palpal structure to B. macrognathus (see Prószy!ski 1976) and B. dilanians (see Prószy!ski 1984). It differs from B. macrognathus in the longer retrolateral tibial apophysis, and from B. dilanians in the straight and finger-like retrolateral tibial apophysis (retrolateral tibial apophysis hooked in B. dilanians). Description. Male (holotype, UBC-SEM AR00083). Carapace length 3.0; abdomen length 4.4. Chelicera (Figs 6.7E-F): light orange to brown; elongate; promargin with two teeth, one of them really big with five cusps on one side; retromargin with one tooth; fang with a central cusp. Palp (Figs 6.7B-D): light yellow to yellowish brown. Tegulum narrow. Embolus short and spiral, with the plane of spiral perpendicular to the longitudinal axis of the palpal bulb. Retrolateral tibial apophysis long and finger-like. Femur of palp long and curved. Tibia of first leg with four pairs of ventral macrosetae; metatarsus with three pairs. Measurements of legs: I 9.7, II 8.3, III 9.4, IV 8.3. Color in alcohol (Fig. 6.7A): carapace orange; abdomen grayish brown, with light yellow speckles and streaks; legs sandy yellow to brown. Female. Unknown. 6.4.2 Genus Canama Simon, 1903 Males of the described species usually have elongate chelicera, similar to Bathippus Thorell. Prószy!ski (1987) considered it as a junior synonym of Bathippus based on their similarities. But Davies and Zabka (1989) rejected this synonym, and suggested that it differed from Bathippus in the cheliceral and epigynal structure. We agree with Davies and Zabka (1989) that they are distinct genera. Females of Canama have one bicuspid tooth on the retromargin of the ! 274! chelicera; the spermatheca usually is not highly swollen, but rather coiled and continuous with the copulatory duct; males of Canama usually have a longer embolus and wider embolic spiral, with the plane of the embolic spiral usually perpendicular to the longitudinal axis of the tegulum. Six species have been included in this genus (Platnick 2011). An additional three new species are described here. The placement of these species in Canama is based both on their similarities to the type species Canama forceps (see Prószy!ski 1984, 1987) and their close relationship with C. forceps indicated by molecular data (see Chapter 2). 6.4.2.1 Canama extranea sp. nov. Figs 6.8-6.9 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1100 m a.s.l., 13-22 July 2008, forest edge, coll. W. Maddison & Luc Fimo Tuki, WPM#08-010. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 male, PAPUA NEW GUINEA: Southern Highlands Province: Putuwé, junction of Lagaip & Uruwabwa Rivers, 5.231° S, 142.532° E, elev. 570 m a.s.l., 23-26 July 2008, coll. W. Maddison & Luc Fimo Tuki, WPM#08-019; 1 female, same data as previous. Etymology. Latin extranea (strange), refering to the peculiar male palpal structure. Diagnosis. Male chelicerae are not as elongate as those of C. forceps (see Prószy!ski 1984, 1987) and C. hinnulea (see Davies & Zabka 1989). Also differs from other species by the presence of dark markings in the eye area of male and female, the big and bowl-like embolic disc and the presence of proximal tegular lobe of the male palp, and the narrow and convoluted copulatory duct of the vulva. Description. Male (holotype, UBC-SEM AR00084). Carapace length 2.1 (variation 2.1-2.7, n=2); abdomen length 2.2. Chelicera (Figs 6.9F-G): yellow brown; not very elongate; with two promarginal teeth and one long bicuspid tooth on retromargin (the paratype male with one big dagger-like retromarginal tooth). Palp (Figs 6.9C-E): yellow brown. Cymbial distal groove more ! 275! retrolateral. Proximal tegular lobe present. Embolic disc large and bowl-like. Retrolateral tibial apophysis finger-like with distal end hooked from ventral view; tibial ventral bump absent. Femur of palp slightly curved. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 4.2, II 3.7, III 4.6, IV 4.1. Color in alcohol (Fig. 6.9A): carapace light orange, with dark markings in eye area; abdomen anterior end light yellow, other part dark; legs light red brown to light yellow. Female (paratype, UBC-SEM AR00085). Carapace length 2.3 (variation 2.2-2.3, n=3); abdomen length 2.9. Chelicera (Fig. 6.9H): yellowish; with two promarginal and one bicuspid retromarginal tooth. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 3.4, II 3.5, III 4.1, IV 3.7. Epigynum (Figs 6.9I-J): median septum relatively narrow; opening to the copulatory duct at posterior margin of window. Copulatory duct long and convoluted; spermatheca small and oval. Color in alcohol (Fig. 6.9B): carapace light yellow, also with dark markings within eye area; abdomen yellowish, without distinct dark markings, legs light yellow. Body green when alive (Figs 6.8C-D) Natural history. Specimens were found beating vegetation in forest edge and disturbed forest. 6.4.2.2 Canama fimoi sp. nov. Figs 6.10-6.11 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1000-1100 m a.s.l., 11-22 July 2008, forest interior and river side, coll. W. Maddison & Luc Fimo Tuki, WPM#08-008; Paratypes: 1 female, same data as holotype; 3 females and 6 males, same data as holotype; 1male, same data as holotype; 1 male, same data as holotype; 1 female, same data as holotype; 1 male and 1 female, PAPUA NEW GUINEA: Southern Highlands Province: Putuwé, junction of Lagaip & Uruwabwa Rivers, 5.231° S, 142.532° E, elev. 570 m a.s.l., 23-26 July 2008, coll. W. Maddison & Luc Fimo Tuki, WPM#08-019; 1 female, PAPUA NEW GUINEA: Southern Highlands Province: Umgé, 5.304-5.305° S, 142.510-142.512° E, elev. 1400-1450 m a.s.l., 15-19 July 2008, coll. W. Maddison & Aislan Tama Wanakipa Indiaf, WPM#08-012; 1 male and 2 females, PAPUA NEW GUINEA: Southern Highlands Province: trail from Tualapa to Umgé, ! 276! 5.2912° S, 142.5006° E to 5.2918° S, 142.5001° E, elev. 1170 m a.s.l., 21 July 2008, coll. W. Maddison & Luc Fimo Tuki, WPM#08-018. Etymology. The specific epithet is a patronym in honor of Mr. Luc Fimo Tuki, who helped in collecting specimens of this species. Diagnosis. Male chelicerae similar to those of C. extranea in being only slightly elongate, but unlike that species having additional projections on the front and back surfaces. Female differs from other species in the short median septum and the wide copulatory duct; male can be distinguished by the wide bulb and the less spiraled embolus of the palp. Description. Male (holotype, UBC-SEM AR00086). Carapace length 3.3 (variation 2.9-4.1, n=11); abdomen length 3.6. Chelicera (Figs 6.11F-H): light orange; robust but not very elongate; promargin with two teeth, retromargin with one large dagger-like tooth; front surface with a spur at the base of fang, back surface with a distal process and a median process. Palp (Figs 6.11C-E): yellowish to yellow brown. Cymbial distal groove more retrolateral. Tegulum wide; proximal tegular lobe absent. Embolus long and coiled for less than a circle. Retrolateral tibial apophysis finger-like with distal end hooked; tibial ventral bump absent. Femur of palp slightly curved. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 11.9, II 8.4, III 9.1, IV 8.2. Color in alcohol (Fig. 6.11A): carapace orange, eye area slightly grayish; abdomen gray brown, with a medial yellowish leaf-like marking; legs light orange. Female (paratype, UBC-SEM AR00087). Carapace length 3.2 (variation 3.1-3.5, n=9); abdomen length 3.5. Chelicera (Fig. 6.11I): with two promarginal and one bicuspid retromarginal tooth. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 7.0, II 6.5, III 8.1, IV 7.6. Leg formula: 3412. Epigynum (Figs 6.11J- K): window large; median septum short and not even reaching the center of window; opening to copulatory duct at posterior end of window. Copulatory duct wide and convoluted; spermatheca relatively large. Color in alcohol (Fig. 6.11B): similar to that male. Natural history. Specimens were collected by beating in forest understory, especially suspended litter. ! 277! 6.4.2.3 Canama triramosa sp. nov. Figs 6.12-6.13 Type material. Holotype: male, PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park, 6.016° S, 145.417° E to 6.017° E, 145.416° E, elev. 2450 - 2490 m a.s.l., 2 August 2008, coll. W. Maddison, WPM#08-027. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 female, PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park, 6.015° S, 145.412° E, elev. 2320 m a.s.l., 1-2 August 2008, coll. W. Maddison, WPM#08-025. Etymology. The specific epithet is from the combination of the prefix tri- (three) and the Latin ramosa (branched), and refers to the large three-forked tooth on the promargin of male chelicerae. Diagnosis. Differs from other species by the unique three-branched promarginal tooth and the one large and wide retromarginal tooth on the male chelicera. Similar in male palpal structure to Canama forceps (see Prószy!ski 1984, 1987), but can be easily distinguished by the shorter and wider male chelicera, the tooth pattern on the male chelicera, the wider palpal bulb and the shape of the retrolateral tibial apophysis of the male palp. Description. Male (holotype, UBC-SEM AR00088). Carapace length 2.1; abdomen length 2.5. Chelicera (Figs 6.13F-G): yellow brown; robust with a tri-forked big tooth on promargin and an axe-like tooth on retromargin. Palp (Figs 6.13C-E): yellowish to brownish. Cymbial distal groove more retrolateral. Proximal tegular lobe absent. Embolus long and coiled for.about two circles. Retrolateral tibial apophysis finger-like; tibial ventral bump absent. Femur of palp curved. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 6.5, II 5.1, III 5.6, IV 4.8. Color in alcohol (Fig. 6.13A): carapace yellowish, eye area light orange; abdomen grayish without distinct markings; legs light yellow with yellow brown annuli. Body more greenish when alive (Figs 6.12A-C). Female (paratype, UBC-SEM AR00089). Carapace length 2.0 (variation 2.0-2.2, n=3); abdomen length 3.2. Chelicera (Fig. 6.13H): with two promarginal and one long bicuspid ! 278! retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.2, II 3.7, III 4.7, IV 4.4. Epigynum (Figs 6.13I-J): window with a narrow median septum; opening at posterior end of window close to median septum. Copulatory duct long and convoluted with a gland-like process near the beginning. Color in alcohol (Fig. 6.13B): similar to that of male, but eye area of carapace yellow, abdomen orange with symmetrical gray markings. Body more greenish when alive (Fig. 6.12D). 6.4.3 Genus Chalcolemia new genus Type species: Chalcolemia nakanai Zhang & Maddison, sp. nov. Etymology. Chosen to resemble the name Chalcolecta; feminine in gender. Diagnosis. Medium-sized spiders with narrow body and slender legs. First pair of legs has many ventral macrosetae. Similar in general body form to Chalcolecta (see Gardzinska & Zabka 2005), but differs in the lateral margins of carapace which is almost parallel instead of convex in Chalcolecta; the thin femur, patella and tibia of first pair of legs which are not much thicker than metatarsus and tarsus; and the presence of distinct median septum in the epigynum. Chalcolemia also resembles Gambaquezonia Barrion & Litsinger 1995 (Edwards 2009) in the delicate body and the presence of a fissident retromarginal tooth on the chelicera, but differs in the much slender legs and the narrower body; the PLEs closer to the lateral margins of the carapace; and the presence of large secondary spermatheca in addition to the primary spermatheca. 6.4.3.1 Chalcolemia nakanai sp. nov. Fig 6.14 Type material. Holotype: female, PAPUA NEW GUINEA: New Britain, Nakanai Mts, Camp 1, Lamas, 3-8 April 2009, 5.614° S, 151.408° E, elev. 200 m, coll. I. Agnarsson. Etymology. A noun in apposition taken from the type locality. Diagnosis. The retromargin of chelicera has one fissident tooth of four cusps; the opening to the copulatory duct is near the center of the window; the primary spermatheca is smaller than the ! 279! secondary spermatheca. Description. Female (holotype, UBC-SEM AR00090). Carapace length 1.7; abdomen length 2.9. Chelicera (Fig. 6.14B): light yellow; with two promarginal teeth and one retromarginal tooth of four cusps. Epigynum (Figs 6.14C-E): median septum of window relatively wide; opening to copulatory duct close to the center of window. Copulatory duct without accessory gland; secondary spermatheca large and oval, primary spermatheca smaller. First pair of legs very long; tibia with nine ventral macrosetae on prolateral side and eight ventral macrosetae on retrolateral side; metatarsus with four pairs of ventral macrosetae. Measurements of legs: I 7.4, II 3.9, III 4.7, IV 5.8. Color in alcohol (Fig. 6.14A): carapace yellow brown, eye area gray brown with indistinct guanine deposit; posterior part of carapace with a “U”-shaped gray brown marking; abdomen light sandy brown, with brown to dark brown irregular markings in the middle; venter of abdomen and legs light sandy yellow. Male. Unknown. Natural history. The specimen was collected by beating foliage in forest. 6.4.4 Genus Omoedus Thorell, 1881 Omoedus Thorell, 1881: 668, type species: Omoedus niger Thorell, 1881. Zenodorus Peckham & Peckham, 1886: 287, type species: Attus d'urvillii Walckenaer, 1837. New Synonym Pystira Simon, 1901a: 656, type species: Hadrosoma ephippigerum Simon, 1885. New Synonym Small to large spiders with various color patterns. Male usually has long and highly coiled embolus. Epigynum has a large window with a median septum of various shapes. Vulva is posterior to the window. Copulatory duct is usually long and convoluted. Spermatheca is not strongly swollen, but small and tubular, and not very distinctive from the copulatory duct. Remarks. Molecular data (see Chapter 2) indicate Omoedus, Pystira and Zenodorus form a clade, with Omoedus and Pystira embedded within Zenodorus. Although the carapace of Omoedus and Pystira is higher than that of typical Zenodorus and has a shallow concavity at the ! 280! posterior end, their genitalic organs do share the same pattern with those of Zenodorus. Hence, we merge them as one genus and consider Pystira and Zenodorus as junior synonyms of Omoedus. Five species once placed in Pystira and 23 species in Zenodorus (Platnick 2011) are therefore transferred into Omoedus: Omoedus cyanothorax (Thorell, 1881) (from Pystira, New Combination) Omoedus ephippigerus (Simon, 1885) (from Pystira, New Combination) Omoedus karschi (Thorell, 1881) (from Pystira, New Combination) Omoedus nigripalpis (Thorell, 1877) (from Pystira, New Combination) Omoedus versicolor (Dyal, 1935) (from Pystira, New Combination) Omoedus albertisi (Thorell, 1881) (from Zenodorus, New Combination) Omoedus arcipluvii (Peckham & Peckham, 1901) (from Zenodorus, New Combination) Omoedus asper (Karsch, 1878) (from Zenodorus, New Combination) Omoedus danae (Hogg, 1915) (from Zenodorus, New Combination) Omoedus durvillei (Walckenaer, 1837) (from Zenodorus, New Combination) Omoedus formosus (Rainbow, 1899) (from Zenodorus, New Combination) Omoedus jucundus (Rainbow, 1912) (from Zenodorus, New Combination) Omoedus juliae (Thorell, 1881) (from Zenodorus, New Combination) Omoedus lepidus (Guérin, 1834) (from Zenodorus, New Combination) Omoedus marginatus (Simon, 1902) (from Zenodorus, New Combination) Omoedus metallescens (L. Koch, 1879) (from Zenodorus, New Combination) Omoedus microphthalmus (L. Koch, 1881) (from Zenodorus, New Combination) Omoedus niger (Karsch, 1878) (from Zenodorus, New Combination) Omoedus obscurofemoratus (Keyserling, 1883) (from Zenodorus, New Combination) Omoedus orbiculatus (Keyserling, 1881) (from Zenodorus, New Combination) Omoedus ponapensis (Berry, Beatty & Prószy!ski, 1996) (from Zenodorus, New Combination) Omoedus pupulus (Thorell, 1881) (from Zenodorus, New Combination) Omoedus pusillus (Strand, 1913) (from Zenodorus, New Combination) Omoedus rhodopae (Hogg, 1915) (from Zenodorus, New Combination) Omoedus syrinx (Hogg, 1915) (from Zenodorus, New Combination) Omoedus variatus (Pocock, 1899) (from Zenodorus, New Combination) Omoedus varicans (Thorell, 1881) (from Zenodorus, New Combination) ! 281! Omoedus wangillus (Strand, 1911) (from Zenodorus, New Combination) Twelve species have been reported from Papua New Guinea (Platnick 2011). Seven new species from Papua New Guinea are described here to provide names for the taxa included in the upcoming molecular phylogeny of euophryines. 6.4.4.1 Omoedus brevis sp. nov. Fig 6.15 Type material. Holotype: male, PAPUA NEW GUINEA: Central Province: Varirata National Park, 9.436° S, 147.364° E, elev. 740 m a.s.l., 4 August 2008, coll. W. Maddison, A. Kore & J. Kore, WPM#08-029. Paratype: 1 male, same data as holotype. Etymology. Latin brevis (short), referring to the short embolus of male palp. Diagnosis. Can be easily distinguished from other species by the short embolus of the male palp. Description. Male (holotype, UBC-SEM AR00124). Carapace length 1.6 (variation 1.6-1.7, n=2); abdomen length 1.4. Chelicera: dark red brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.15D-E): dark yellow brown with gray pigments. Proximal tegular lobe absent; retrolateral sperm duct loop about three quaters of tegulum width. Embolus very short and not coiled. Retrolateral tibial apophysis relatively short and finger-like. Ventral tibial bump big. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.8, II 2.6, III 3.1, IV 3.0. Color in alcohol (Fig. 6.15C): carapace dark brown, with iridescent scales behind PLEs and AMEs; dorsal abdomen dark brown, anterior margin with a wide band composed of golden iridescent scales, posterior part with a few transverse light yellow bands; ventral abdomen gray brown with brownish speckles; first pair of legs cream on tarsi, other segments dark brown; other legs dark brown on coxae, trochanters and femora, other segments cream. Female. Unknown. ! 282! Natural history. Specimens were found on leaf litter in forest. 6.4.4.2 Omoedus darleyorum sp. nov. Figs 6.16-6.17 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1100 m a.s.l., 13-22 July 2008, Forest edge, coll. W. Maddison & Luc Fimo Tuki, WPM#08-010. Paratypes: 1 female, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1100 m a.s.l., 11-22 July 2008, old clearing (former garden), coll. W. Maddison, WPM#08-009; 1 male, same data as previous. Etymology. Named in honour of Merrick and Lorraine Darley, important supporters of Conservation International's efforts to document biodiversity. Diagnosis. Similar in color pattern and markings to Omoedus brevis, but can be easily distinguished by the long and highly coiled embolus and the paler legs. Median septum of the epigynum is relatively wide, distinct from that in other species. Description. Male (holotype, UBC-SEM AR00127). Carapace length 1.6 (variation 1.6-1.8, n=2); abdomen length 1.7. Chelicera (Fig. 6.17E): dark; with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.17C-D): pale yellow. Proximal tegular lobe absent; retrolateral sperm duct loop about three quarters of bulb width. Embolus long and coiled for more than five circles. Retrolateral tibial apophysis long and gradually narrowed towards the tip. Tibia of first leg with six ventral macrosetae prolaterally and three ventral macrosetae retrolaterally; metatarsus with five ventral macrosetae prolaterally and three ventral macrosetae retrolaterally. Measurements of legs: I 5.8, II 3.6, III 4.1, IV 3.9. Color in alcohol (Fig. 6.17A): carapace dark brown to red brown, with iridescent scales in front of and behind PLEs; abdomen brown, anterior end light brown, covered with iridescent scales, posterior part with a pair of brownish markings; legs pale yellow without distinct markings. Female (paratype, UBC-SEM AR00128). Carapace length 1.4; abdomen length 1.8. Chelicera: with two promarginal teeth and one retromarginal teeth. Tibia of first leg with four ventral ! 283! macrosetae prolaterally and three ventral macrosetae retrolaterally; metatarsus with three pairs of ventral macrosetae. Measurements of legs: I 2.9, II 2.5, III 2.9, IV 3.0. Epigynum (Figs 6.17F-G): window large; anterior end of median septum relatively wide. Copulatory duct long and highly convoluted; spermatheca irregular in shape. Color in alcohol (Fig. 6.17B): similar to that of male. Natural history. Specimens were found beating understory vegetation. 6.4.4.3 Omoedus meyeri sp. nov. Figs 6.18-6.19 Type material. Holotype: male, PAPUA NEW GUINEA: Enga Province: Kai-ingri, 5.574° S, 143.048° E, elev. 3315 m a.s.l., 5-8 July 2008, coll. W. Maddison,WPM#08-004. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 1 male, same data as holotype; 1 male, same data as holotype; 1 male, PAPUA NEW GUINEA: Enga Province: Kai-ingri, 5.579° S, 143.053° E, elev. 3240 m a.s.l., 7-9 July 2008, coll. W. Maddison & Manisé Kulé, WPM#08- 005; 1 male, same data as previous; 1 male, same data as previous. Etymology. Named for George Meyer, an important supporter of Conservation International's efforts in Papua New Guinea. Diagnosis. Similar in body form to Omoedus omundseni, but differs in the presence of dark patches on the dorsum of abdomen; the narrower bulb and embolic spiral of the male palp; and the narrower window of the epigynum. This species can be distinguished from O. papuanus by the presence of guanine deposits in the eye area; the markings on the abdomen; the thinner and more coiled embolus (four circles in O. meyeri; three circles in O. papuanus); and the wider median septum of the epigynum Description. Male (holotype, UBC-SEM AR00125). Carapace length 1.8 (variation 1.8-2.2, n=7); abdomen length 2.3. Chelicera (Fig. 6.19E): dark; with one bicuspid promarginal tooth and one retromarginal tooth. Palp (Figs 6.19C-D): tibia and cymbium dark brown, other segments pale yellow. Proximal tegular lobe absent; retrolateral sperm duct loop about two thirds of tegulum width. Embolus long, coiled for more than four circles. Retrolateral tibial ! 284! apophysis relatively thin. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.8, II 3.9, III 4.1, IV 4.1. Color in alcohol (Fig. 6.19A): eye area with guanine deposit, carapace dark brown, with a medial light yellow brown stripe behind fovea, lateral margins light yellow brown; abdomen brown, with brownish markings and a pair of large dark patches near the center; legs light brown with dark borwn annuli. Some specimens darker in Color in alcohol (Figs 6.18D-E). Female (paratype, UBC-SEM AR00126). Carapace length 1.7; abdomen length 2.5. Chelicera: with one tooth on promargin and retromargin each. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.5, II 3.2, III 3.8, IV 4.1. Epigynum (Figs 6.19F-G): window large, with opening to copulatory duct close to its posterior end. Copulatory duct long and highly convoluted; spermatheca not very distinctive from copulatory duct in shape. Color in alcohol (Fig. 6.19B): similar to that of male except carapace and abdomen a bit darker in color. Natural history. Specimens were found beating trees at forest edge and grassland bushes, at high elevation. 6.4.4.4 Omoedus omundseni sp. nov. Figs 6.20-6.21 Type material. Holotype: male, PAPUA NEW GUINEA: Enga Province: Kai-ingri, 5.579° S, 143.053° E, elev. 3240 m a.s.l., 7-9 July 2008, coll. W. Maddison & Manisé Kulé, WPM#08- 005. Paratypes: 1 female, same data as holotype; 2 females, same data as holotype; 1 female, same data as holotype; 7 males and 2 females, PAPUA NEW GUINEA: Enga Province: Kai- ingri, 5.574° S, 143.048° E, elev. 3315 m a.s.l., 5-8 July 2008, coll. W. Maddison, WPM#08- 004. Etymology. Named after Tim Omundsen, for his assistance facilitating the 2008 expedition to Papua New Guinea. Diagnosis. Similar to Omoedus meyeri in the presence of guanine deposits in the eye area and the genitalic structures, but differs in the markings on the abdomen, the wider male palpal bulb ! 285! and the wider window of the epigynum. Description. Male (holotype, UBC-SEM AR00122). Carapace length 2.1 (variation 1.9-2.7, n=8); abdomen length 2.5. Chelicera (Fig. 6.21E): yellow brown; with one bicuspid promarginal tooth and one retromarginal tooth; front surface with a longitudinal ridge and a small process. Palp (Figs 6.21C-D): light brown. Proximal tegular lobe absent; retrolateral sperm duct loop about three quarters of bulb width. Embolus long and coiled for four circles. Palpal bulb wide. Retrolateral tibial apophysis long and thin. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 6.3, II 4.8, III 5.1, IV 4.9. Color in alcohol (Fig. 6.21A): carapace yellow brown, eye area with guanine deposit, lateral margins and around fovea light yellow brown; dorsal abdomen dark brown, with sandy yellow medial markings, ventral abdomen sandy yellow, with grayish brown markings; legs light yellow to yellow brown. Female (paratype, UBC-SEM AR00123). Carapace length 2.1 (variation 2.0-2.3, n=6); abdomen length 3.0. Chelicera: with one bicuspid promarginal tooth and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.3, II 3.8, III 4.3, IV 4.4. Epigynum (Figs 6.21F-G): window large; median septum almost triangular with anterior part very narrow; opening to copulatory duct close to posterior end of window. Copulatory duct long and highly convoluted; spermatheca not distinctive from copulatory duct in shape. Color in alcohol (Figs 6.21B): similar to that of male, except the light colored markings on abdomen more distinctive. Natural history. Specimens were found beating trees at forest edge, especially those with dark lichen-covered branches. 6.4.4.5 Omoedus papuanus sp. nov. Figs 6.22-6.23 Type material. Holotype: male, PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park, 6.015° S, 145.412° E, elev. 2320 m a.s.l., 1-2 August 2008, coll. W. Maddison, WPM#08-025. Paratypes: 1 female, same data as holotype; 1 female and 1 male, same data as holotype; 1 male, same data as holotype; 1 male and 3 females, PAPUA NEW ! 286! GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park, 6.016° S, 145.417° E to 6.017° S, 145.416° E, elev. 2450-2490 m a.s.l., 2 August 2008, coll. W. Maddison, WPM#08- 027. Etymology. The specific epithet refers to the country where the species is found. Diagnosis. Similar in markings on the abdomen to Omoedus microphthalmus (see Berry et al. 1996), but can be distinguished by the wider spirals of the embolus, the wider palpal bulb, and the shape of the epigynum. Description. Male (holotype, UBC-SEM AR00129). Carapace length 1.9 (variation 1.8-2.0, n=4); abdomen length 1.8. Chelicera (Fig. 6.23E): dark brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.23C-D): cymbium yellow brown, other segments cream. Proximal tegular lobe absent; retrolateral sperm duct loop about two thirds of bulb width. Embolus long and coiled for three circles. Retrolateral tibial apophysis long and thin. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.2, II 3.6, III 3.9, IV 4.3. Color in alcohol (Fig. 6.23A): carapace dark yellow brown, with many white scales, lateral margins light yellow brown, with a light yellow brown stripe medially behind fovea; dorsal abdomen dark brown, anterior end and most of lateral margins light yellow, posterior end with a few paralleled chevron-like markings; ventral abdomen light yellow, with an indistinct middle stripe; legs light yellow with yellow brown annuli. Female (paratype, UBC-SEM AR00130). Carapace length 1.9 (variation 1.8-1.9, n=5); abdomen length 1.9. Chelicera: also with two promarginal teeth and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.5, II 3.3, III 3.7, IV 4.2. Epigynum (Figs 6.23F-G): window large; median septum almost triangular with anterior part very narrow. Copulatory duct long and highly convoluted; spermatheca not very distinctive from copulatory duct. Color in alcohol (Fig. 6.23B): similar to that of male. Natural history. Specimens were found beating forest understory. ! 287! 6.4.4.6 Omoedus swiftorum sp. nov. Figs 6.24-6.25 Type material. Holotype: male, PAPUA NEW GUINEA: National Capital District: Port Moresby, 9.443° S, 147.179° E, elev. 75 m a.s.l., 3-4 July 2008, coll. W. Maddison WPM#08- 002. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 female, same data as holotype; 1 male, same data as holotype. Etymology. In honor of Kristin Swift and John Swift, who have generously supported Conservation International's work in New Guinea. Diagnosis. Differs from other species by the light colored longitudinal stripes on the body and the genitalic shape. Description. Male (holotype, UBC-SEM AR00120). Carapace length 1.8 (variation 1.8-2.0, n=2); abdomen length 2.1. Chelicera (Figs 6.25E-F): red brown; with one bicuspid promarginal tooth and one retromarginal tooth; inner margin concaved; front surface with a longitudinal ridge and a small process. Palp (Figs 6.25C-D): brown. Proximal tegular lobe absent; retrolateral sperm duct loop almost as wide as bulb. Embolus long and coiled for more than two circles. Retrolateral tibial apophysis very thin. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.0, II 3.5, III 3.8, IV 4.2. Color in alcohol (Fig. 6.25A): eye area dark, other region of carapace red brown, with a medial stripe behind PLEs and two lateral stripes composed of white scales; abdomen dark brown, margins sandy yellow, with a wide medial light yellow longitudinal stripe containing a brown marking in the middle; ventral abdomen pale yellow with gray brown longitudinal markings; first pair of legs reddish brown to gray brown, other legs cream to gray brown. Female (paratype, UBC-SEM AR00121). Carapace length 2.0 (variation 1.8-2.0, n=3); abdomen length 3.0. Chelicera: also with one bicuspid promarginal tooth and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.7, II 3.3, III 3.8, IV 4.3. Epigynum (Figs 6.25G-H): window large; median septum almost triangular; opening to the copulatory duct at posterior end of window. Copulatory duct long and convoluted; spermatheca not distinctive from copulatory duct. Color ! 288! in alcohol (Fig. 6.25B): similar to that of male except the brownish marking in the middle of light yellow stripe on dorsal abdomen not obvious and legs cream lacking obvious markings. Natural history. Specimens were found on low bushes in an open disturbed area. 6.4.4.7 Omoedus tortuosus sp. nov. Figs 6.26-6.27 Type material. Holotype: male, PAPUA NEW GUINEA: Enga Province: Paiam Forest, near Suyan Village, 5.495° S, 143.144° E, elev. 2400 m a.s.l., 10 July 2008, coll. W. Maddison, Pingisa Saiké, Yainé Ribson, S. Soté, & N. Soté, WPM#08-007. Paratypes: 1 female, same data as holotype; 1 male and 2 females, same data as holotype; 2 females, PAPUA NEW GUINEA: Enga Province: Kai-ingri, 5.574° S, 143.048° E, elev. 3315 m a.s.l., 5-8 July 2008, coll. W. Maddison, WPM#08-004; 1 male, PAPUA NEW GUINEA: Enga Province: Kai-ingri, 5.579° S, 143.053° E, elev. 3240 m a.s.l., 7-9 July 2008, coll. W. Maddison & Manisé Kulé, WPM#08- 005. Etymology. Latin tortuosus (twisting), referring to the convoluted copulatory duct of vulva. Diagnosis. Resembles Omoedus papuanus in body form, but differs from it by the color pattern, the shape of the median septum of epigynum, the thicker retrolateral tibial apophysis and the narrower but more spiraled embolus of the male palp. Description. Male (holotype, UBC-SEM AR00131). Carapace length 2.5 (variation 2.2-2.5, n=3); abdomen length 2.7. Chelicera (Fig. 6.27E): dark brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.27C-D): yellow brown. Proximal tegular lobe absent; retrolateral sperm duct loop occupying about three quarters of bulb width. Embolus long and coiled for more than seven circles. Retrolateral tibial apophysis long and finger-like. Tibia of first leg with four ventral macrosetae prolaterally and three ventral macrosetae retrolaterally; metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 7.2, II 5.2, III 5.4, IV 6.1. Color in alcohol (Fig. 6.27A): eye area dark, other regions of carapace red brown; abdomen dark brown, with some brownish spots; legs light yellow proximally and dark brown distally. ! 289! Female (paratype, UBC-SEM AR00132). Carapace length 2.2 (variation 2.1-2.3, n=5); abdomen length 2.6. Chelicera: with two promarginal teeth and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.1, II 3.8, III 4.5, IV 5.1. Epigynum (Figs 6.27F-G): window large; margins of median septum slightly convex. Copulatory duct long and convoluted; spermatheca not distinctive from copulatory duct. Color in alcohol (Fig. 6.27B): similar to that of male except the brownish spots on dorsal abdomen more irregular. Natural history. Specimens were found in suspended leaf litter in forest. 6.4.5 Genus Paraharmochirus Szombathy, 1915 Ant-like jumping spiders. Tibia, patella and femur of first pair of legs are robust; tibia has ventral fringes; metatarsus and tarsus are much thinner. Lateral margins of carapace are convex at PLEs and carapace has many punctures. Abdomen has a constriction dorsally. Epigynum has window, which is typical in Euophryinae. However, the male palp is quite distinctive from other euophryines in that embolus coils clockwise (left palp ventral view) instead of anti-clockwise as seen in most other euophryines. The inclusion of this genus within the subfamily Euophryinae is mainly based on molecular data (see Chapter 2). Only one species, Paraharmochirus monstrosus Szombathy, 1915 (from New Guinea), has been reported, of which the type specimen may have been lost (Tamas Szüts, pers. comm.). The species recently collected from Papua New Guinea is similar to P. monstrosus in the shape of carapace and first pair of legs, and the presence of punctures on the carapace. However, it is not congruent with the original description of P. monstrosus Szombathy in the male palpal structure and the tooth pattern of chelicera, and therefore is described here as a new species. 6.4.5.1 Paraharmochirus tualapaensis sp. nov. Figs 6.28-6.29 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1100 m a.s.l., 13-22 July 2008, forest edge, coll. W. Maddison & Luc Fimo Tuki, WPM#08-010. Paratypes: 1 female, same data as holotype; 1 female, same data as holotype; 1 male, PAPUA NEW GUINEA: Southern ! 290! Highlands Province: Putuwé, junction of Lagaip & Uruwabwa Rivers, 5.231° S, 142.532° E, elev. 570 m a.s.l., 23-26 July 2008, coll. W. Maddison & Luc Fimo Tuki, WPM#08-019; 1 female, same data as previous; 1 female, same data as previous. Etymology. The specific epithet refers to the type locality. Diagnosis. Resembles P. monstrosus Szombathy, 1915 in general body form and color pattern, but differs in the presence of two promarginal teeth on the male chelicera (no promarginal tooth in P. monstrosus), and the embolus of the male palp coiling no more than one circle (1.5 circles in P. monstrosus). Epigynum has a large window with a median septum, and the spermathecae are round and relatively small. Description. Male (holotype, UBC-SEM AR00091). Carapace length 1.8; abdomen length 1.9. Carapace with many small punctures, and widest at PLEs. Chelicera: yellow brown; with two promarginal teeth and one bicuspid retromarginal tooth. Palp (Figs 6.29C-D): light yellow brown. Embolus long and wide, coiled clockwise (left palp ventral view); embolic disc at prolateral side of palpal bulb; retrolateral sperm duct loop at distal end of tegulum; tegular lobe absent. Retrolateral tibial apophysis robust and pointed at tip. First pair of legs long and robust; with fringe on ventral tibia; tibia with seven ventral macrosetae on prolateral side and six ventral macrosetae on retrolateral side; metatarsus with three pairs of ventral macrosetae. Measurements of legs: I 4.8, II 2.8, III 2.8, IV 3.5. Color in alcohol (Fig. 6.29A): carapace orange, with some white setae; dorsal abdomen constricted at 1/3 from anterior end, with white markings within the constriction; abdomen light brown anterior to the constriction and dark brown posterior to the constriction; first pair of legs light orange, other legs light yellow to gray brown. Some specimens darker in color. Female (paratype, UBC-SEM AR00092). Carapace length 1.5 (variation 1.5-1.8, n=4); abdomen length 1.9. Chelicera (Fig. 6.29E): with two promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with six pairs of ventral macrosetae; metatarsus of first leg with three pairs of ventral macrosetae. Measurements of legs: I 2.7, II 2.0, III 2.1, IV 2.6. Epigynum (Figs 6.29F-G): window relatively big, with a median septum. Copulatory duct long and sac-like, without accessory gland; spermatheca small and spherical. Color in alcohol (Fig. 6.29B): similar to that of male. ! 291! Natural history. Specimens were collected from tree trunks at forest edge. 6.4.6 Genus Phasmolia new genus Type species: Phasmolia elegans Zhang & Maddison, sp. nov. Etymology. The generic name is derived from the Latin phasma (ghost), referring to the ghost- like appearance of the species; feminine in gender. Diagnosis. Resembles Lakarobius Berry, Beatty and Prószy!ski, 1998 and Bindax (see Prószy!ski 1984) in body form and color pattern, but differs from them by the absence of a proximal tegular lobe and retrolateral sperm duct loop in the male palp. This genus can also be distinguished from Lakarobius by the chelicera with three promarginal teeth (two in Lakarobius) and one bicuspid retromarginal tooth (four cusps in Lakarobius), and the median septum of female epigynum, which is not continuous with the anterior rim of the window. Also similar to Athamas (see Jendrzejewska 1995), Bulolia Zabka, 1996 and Leptathamas Balogh, 1980a (Szüts 2003) in the ALEs, which are posterior to the AMEs, but can be easily distinguished from them by the shape of the genitalic organs. 6.4.6.1 Phasmolia elegans sp. nov. Figs 6.30-6.31 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1100 m a.s.l., 13-22 July 2008, forest edge, coll. W. Maddison & Luc Fimo Tuki, WPM#08-010. Paratypes: 1 female, same data as holotype; 5 males and 1 female, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1000-1100 m a.s.l., 11-22 July 2008, forest interior and river side, coll. W. Maddison & Luc Fimo Tuki, WPM#08-008; 1 female, same data as previous; 1 male, same data as previous; 1 female, PAPUA NEW GUINEA: Southern Highlands Province: trail from Tualapa to Umgé, 5.2933° S, 142.4999° E, elev. 1210 m a.s.l., 19 July 2008, coll. W. Maddison, WPM#08-015. Etymology. Latin elegans, referring to the elegant body form. ! 292! Diagnosis. See the diagnosis of the genus. Description. Male (holotype, UBC-SEM AR00093). Carapace length 1.5 (variation 1.5-1.7, n=7); abdomen length 1.8. ALEs relatively posterior to AMEs. Chelicera: gray brown; with three promarginal teeth and one bicuspid retromarginal tooth. Palp (Figs 6.31C-D): cymbium light yellow, and other segments gray brown. Proximal tegular lobe absent; sperm duct not forming a loop at retrolateral side of bulb. Embolic disc almost round; embolus slender and curved for about half a circle. Retrolateral tibial apophysis long and finger-like. Tibia of first leg with four pairs of ventral macrosetae; metatarsus with three pairs. Measurements of legs: I 3.4, II 3.3, III 3.8, IV 3.4. Color in alcohol (Fig. 6.31A): carapace dark brown with a large light brown marking behind PLEs; eye area covered with gray white long scales between ALEs and PMEs; abdomen dark gray, with one anterior light yellow band and two lateral round light yellow patches near the middle; legs light yellow, all femora with distinct dark gray markings. Female (paratype, UBC-SEM AR00094). Carapace length 1.5 (variation 1.5-1.7, n=4); abdomen length 1.9. Chelicera (Fig. 6.31E): light yellow; with three promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with five pairs of ventral macrosetae; metatarsus with three pairs. Measurements of legs: I 3.2, II 3.0, III 3.4, IV 3.5. Epigynum (Figs 6.31F-G): window relatively large; opening to copulatory duct near anterior edge of the window; median septum narrow. Copulatory duct relatively narrow and not convoluted; spermatheca almost spherical. Color in alcohol (Figs 6.31B): eye area dark, other regions light yellow; abdomen dark with a light yellow transverse band near the middle and two lateral light yellow patches at posterior end; legs light yellow without dark markings. Natural history. Specimens were collected by beating forest understory vegetation. 6.4.7 Genus Sobasina Simon, 1898 Ant-like jumping spiders. Wanless (1978) revised the genus and indicated that it is distinctive from the other Oriental ant-like salticids by the structure of the genitalia, the strongly recurved anterior eye row in frontal view, the scalloped sternum, and the elongate coxa and trochanter of first leg. Another ant-like salticid genus, Paraharmochirus Szombathy, 1915 from Papua New Guinea shares some similarities to Sobasina, such as the scalloped sternum, the elongate coxa ! 293! and trochanter of first leg, and the presence of punctures on the carapace. But Sobasina can be distinguished from it by the elongate carapace which is not strongly convex at PLEs, the short embolus, the absence of the retrolateral sperm duct loop in the male palp, the absence of a window in the epigynum, and the indistinctive spermatheca. Placement of Sobasina in the subfamily Euophryinae is based on molecular data (see Chapter 2). Fourteen species of this genus have been described from the Pacific Islands and Malaysia (Wanless 1978; Berry et al. 1998; Edmunds & Prószy!ski 2001; Platnick 2011). One new species from Papua New Guinea is described here. 6.4.7.1 Sobasina wanlessi sp. nov. Fig 6.32 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev.1000-1100 m a.s.l., 11-22 July 2008, forest interior and river side, coll. W. Maddison & Luc Fimo Tuki, WPM#08-008. Paratypes: 5 males, same data as holotype; 1 female, same data as holotype; 2 males, same data as holotype. Etymology. The specific epithet is a patronym in honor of Dr. F. R. Wanless, who made great contributions in the study of jumping spider systematics, and revised this genus. Diagnosis. Similar to Sobasina cutleri Berry, Beatty & Prószy!ski, 1998 and S. platypoda Berry, Beatty & Prószy!ski, 1998, but differs from S. cutleri in the presence of ventral fringes on male first leg, the relatively wider palpal bulb, and the shape of vulva; and from S. platypoda in the shape of epigynum and vulva. Description. Male (holotype, UBC-SEM AR00095). Carapace length 1.2 (variation 1.2-1.5, n=8); abdomen length 1.1. Carapace with many small punctures, and a hump at posterior part. Chelicera (Fig. 6.32E): yellow brown; with two promarginal teeth and one bicuspid retromarginal tooth. Palp (Figs 6.32G): light yellow brown. Embolus short and not coiled; retrolateral sperm duct loop absent; tegular lobe absent. Retrolateral tibial apophysis finger-like. First pair of legs robust, with fringes on ventral tibia; tibia with four ventral macrosetae on prolateral side and five ventral macrosetae on retrolateral side (some specimens with five pairs ! 294! of ventral macrosetae on first tibia); metatarsus with three ventral macrosetae on prolateral side and two ventral macrosetae on retrolateral side (some specimens with three pairs of ventral macrosetae on first metatarsus). Measurements of legs: I 2.0, II 1.5, III 1.5, IV 1.9. Color in alcohol (Fig. 6.32C): carapace dark brown to red brown; dorsal abdomen slightly constricted at 1/3 from anterior end; abdomen brown with a sandy yellow band at constriction; first pair of legs yellow brown, with tarsus and metatarsus light yellow; other legs light yellow to light brown. Female (paratype, UBC-SEM AR00096). Carapace length 1.5; abdomen length 1.5. Chelicera: with two promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with four ventral macrosetae on prolateral side and five ventral macrosetae on retrolateral side; metatarsus of first leg with three pairs of ventral macrosetae. Measurements of legs: I 2.7, II 2.1, III 2.2, IV 3.1. Epigynum (Figs 6.32H-I): without window. Copulatory duct long and tubular, without accessory gland; spermatheca not distinctively larger than copulatory duct. Color in alcohol (Fig. 6.32D): darker than that of male. Natural history. Specimens were collected by beating forest understory, possibly with suspended litter. 6.4.8 Genus Thorelliola Strand, 1942 Small to medium sized spiders. Carapace is usually high. Males sometimes have setae enlarged into “horns” on the clypeus and some of them have a truncus for the “horns” on the clypeus (Gardzinska & Patoleta 1997; Szüts & De Bakker 2004). Chelicera has a fissident retromarginal tooth. Many species have a process distally on the front surface of the male chelicera. Epigynum has a big window without median septum. Some species have a pair of secondary spermathecae in addition to the primary spermathecae. Male palp usually has prominent macrosetae on the tibia and also on the femur in some species; tegulum lacks proximal lobe; embolus is long or short. In total, 12 species have been reported from Southeast Asia, Papua New Guinea and the Pacific Islands (Platnick 2011). Some of the new species described here are not congruent with the described species in the appearance and in that males only have ordinary setae rather than robust “horns” on the clypeus. However, molecular data (see Chapter 2) indicate that they fall into a ! 295! clade with the typical Thorelliola species including the type species Thorelliola ensifera. Thorelliola mahunkai Szüts has “horns” not robust but more like ordinary setae (Szüts 2002). Thus, using the “horns” on the male clypeus to define the genus Thorelliola may be too restricting. Here we expand the delimitation of the genus Thorelliola to comprise more species to avoid erecting more new genera for jumping spiders. 6.4.8.1 Thorelliola aliena sp. nov. Figs 6.33-6.34 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Umgé, 5.304° S, 142.512° E, elev. 1450 m a.s.l., 15-19 July 2008, coll. W. Maddison & Aislan Tama Wanakipa Indiaf, WPM#08-013. Paratypes: 1 female, same data as holotype; 2 females and 2 males, same data as holotype. Etymology. Latin aliena (foreign or incongruous), referring to the atypical male palpal structure and body form. Diagnosis. This new species resembles other species of Thorelliola in the absence of a median septum in the epigynum. It can be easily distinguished by the presence of longitudinal stripes on the dorsum of abdomen (red in male and yellow in female), and the unusually long tibia and patella of the male palp. It also differs in the long embolus of the male palp, the sperm duct loop located at the proximal side instead of retrolateral side of the bulb, the absence of an apophysis on the femur of male palp, the presence of a pair of secondary spermathecae in addition to the primary spermathecae in the vulva, and the absence of guanine deposits in the carapace. Description. Male (holotype, UBC-SEM AR00097). Carapace length 1.9 (variation 1.8-2.2, n=3); abdomen length 2.0. Chelicera (Fig. 6.34C): with two promarginal teeth and one bicuspid retromarginal tooth; with a ectal protrusion at distal end. Palp (Figs 6.34D-F): dark brown. Embolus long and spiral; sperm duct loop proximal and pointing towards center of bulb. Tibia of palp with one prolateral and one ventral macrosetae. Retrolateral tibial apophysis finger-like. Tibia and metatarsus of first leg with four pairs of ventral macrosetae each. Measurements of legs: I 4.6, II 3.4, III 4.4, IV 4.1. Color in alcohol (Fig. 6.34A): carapace dark brown, scattered with orange scales; abdomen yellowish brown, with gray markings and two orange stripes ! 296! laterally; first pair of legs dark brown, other legs yellowish brown. Female (paratype, UBC-SEM AR00098). Carapace length 1.8 (variation 1.8-2.0, n=3); abdomen length 2.0. Chelicera: with two promarginal teeth and one bicuspid retromarginal tooth. Tibia and metatarsus of first leg with four pairs of ventral macrosetae each. Measurements of legs: I 3.5, II 2.7, III 3.5, IV 3.6. Epigynum (Figs 6.34G-H): with a big window lacking median septum. Copulatory duct thick at the beginning and then divided, with one leading to secondary spermatheca and the other to the kidney-shaped primary spermatheca with rather thin connecting duct. Color in alcohol (Fig. 6.34B): similar to that of male, but carapace a bit lighter in color, stripes on dorsal abdomen yellow, and the first pair of legs yellowish brown. Natural history. Specimens were found on leaf litter in a mid-elevation forest. 6.4.8.2 Thorelliola crebra sp. nov. Figs 6.35-6.36 Type material. Holotype: male, PAPUA NEW GUINEA: Enga Province: Suyan Camp, Porgera, 5.4833° S, 143.1337° E, elev. 2300 m a.s.l., 28-29 July 2008, coll. W. Maddison, WPM#08-022. Paratypes: 1 female, same data as holotype; 1 male and 2 females, same data as holotype. Etymology. Latin crebra (thick), referring to the thick patella and tibia of male palp. Diagnosis. Can be easily distinguished from Thorelliola aliena and previously reported species by the rough mottled apprearance of bark, the presence of guanine deposits in the eye area of carapace, and the femoral protuberance of the male palp. Similar in male palp structure to T. zabkai, but differs in the male chelicera with a triangular process near the fang base on the front surface; the male endite with two lateral bulges; the shorter embolus, the much wider retrolateral sperm duct loop and the oval embolic disc of the male palp. The new species has similar color pattern and epigynal shape as T. squamosa, but can be distinguished by the large dark patches at the posterior part of the dorsal abdomen and the spherical secondary spermatheca which is further behind the posterior end of the window. ! 297! Description. Male (holotype, UBC-SEM AR00099). Carapace length 1.9 (variation 1.9-2.2, n=2); abdomen length 2.2. Chelicera (Fig. 6.36F): dark brown; with two promarginal teeth and one bicuspid retromarginal tooth; with a triangular process near the base of the fang on front surface. Endite (Fig. 6.36F): grey brown; with two lateral bulges, one at distal end and the other near middle. Palp (Figs 6.36C-E): yellow brown. Embolic disc oval; embolus long with a tiny cusp near the end; loop of sperm duct wide almost occupying the whole width of bulb. Tibia, patella and femur of palp with multiple macrosetae; femur with a big distal protuberance prolaterally; patella with a retrolateral lump and a thick macroseta at its top. Retrolateral tibial apophysis finger-like reaching the proximal edge of embolic disc. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 5.1, II 4.1, III 4.5, IV 4.6. Color in alcohol (Fig. 6.36A): carapace yellow brown, eye area with white guanine deposit, with white stripes laterally behind eye area composed of white scales; abdomen light yellow with a few symmetrical dark gray markings, heart mark also dark gray, almost rhomboid; legs light yellow to yellowish brown, with dark gray markings. Female (paratype, UBC-SEM AR00100). Carapace length 1.8 (variation 1.8-1.9, n=3); abdomen length 2.5. Chelicera (Fig. 6.36G): with two promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.4, II 3.3, III 3.5, IV 4.0. Epigynum (Figs 6.36H-I): window almost semicircular with opening to copulatory duct at its posterior end. Copulatory duct short; secondary and primary spermatheca oval. Color in alcohol (Fig. 6.36B): similar to that of male. Natural history. Specimens were found on bark of Casuarina tree trunks. 6.4.8.3 Thorelliola joannae sp. nov. Fig 6.37 Type material. Holotype: female, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1000-1100 m a.s.l. 11-22 July 2008, forest interior and river side, coll. W. Maddison & Luc Fimo Tuki, WPM#08-008. Paratypes: 1 female, PAPUA NEW GUINEA: Southern Highlands Province: Putuwé, junction of Lagaip & Uruwabwa Rivers, 5.231° S, 142.532° E, elev. 570 m a.s.l. 23-26 July 2008, coll. W. Maddison & Luc Fimo Tuki, WPM#08-019; 1 female, PAPUA NEW GUINEA: Southern Highlands ! 298! Province: Umgé, 5.304-5.305° S, 142.510-142.512° E, elev. 1400-1450 m a.s.l., 15-19 July 2008, coll. W. Maddison & Aislan Tama Wanakipa Indiaf, WPM#08-012. Etymology. The specific epithet is a patronym in honor of Dr. Joannae Gardzinska, who has contributed much to the taxonomy of this genus. Diagnosis. Differs from other described species (Gardzinska & Patoleta 1997; Gardzinska 2009) and T. tualapa by the vulva located posterior to the window. The absence of guanine deposits in the eye area can distinguish the species from T. creba, T. zabkai and T. squamosa. Differs from T. aliena by the absence of bright stripes on the abdomen and the epigynal shape. Description. Female (holotype, UBC-SEM AR00101). Carapace length 1.8 (variation 1.8-1.9, n=3); abdomen length 2.0. Chelicera: red brown; with two promarginal teeth and one retromarginal tooth of five cusps. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.5, II 3.4, III 4.4, IV 4.7. Epigynum (Figs 6.37C-D): window big; vulva posterior to window. Copulatory duct long and coiled; spermatheca small. Color in alcohol (Fig. 6.37B): carapace dark brown with a yellow brown marking behind PLEs; abdomen dark brown, scattered with small brownish speckles, with a wide yellowish brown band at anterior end and in the middle each; venter of abdomen sandy yellow with gray patches; legs yellow brown with gray brown annuli. Male. Unknown. 6.4.8.4 Thorelliola squamosa sp. nov. Fig 6.38 Type material. Holotype: female, PAPUA NEW GUINEA: Central Province: Varirata National Park, 9.436° S, 147.364° E, elev. 740 m a.s.l., 4 August 2008, coll. W. Maddison, A. Kore & J. Kore, WPM#08-029. Paratype: 1 female, same data as holotype. Etymology. Latin squamosa, referring to the scale-like markings on the dorsal abdomen. Diagnosis. Differs from T. crebra and T. zabkai by the posterior end of lateral rims of the ! 299! epigynal window curving anteriorly; and also from T. zabkai in the presence of guanine deposits in the eye area. Description. Female (holotype, UBC-SEM AR00102). Carapace length 1.9 (variation 1.8-1.9, n=2); abdomen length 2.1. Chelicera: red brown; with two promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.3, II 3.2, III 3.5, IV 3.8. Epigynum (Figs 6.38D-F): lateral rims of window curved anteriorly. Copulatory duct short and divided, with one leading to the long and oval secondary spermatheca and the other leading to the spherical primary spermatheca. Color in alcohol (Fig. 6.38C): carapace yellow brown, eye area with distinct white guanine deposits, eyes with dark surroundings; abdomen covered with squamous markings and a few grayish markings, heart mark also grayish; legs light yellow with gray markings. Male. Unknown. Natural history. Specimens were found on tree trunks in a clearing forest. 6.4.8.5 Thorelliola tamasi sp. nov. Figs 6.39-6.40 Type material. Holotype: male, PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park, 6.016° S, 145.417° E to 6.017° S, 145.416° E, elev. 2450 - 2490 m a.s.l., 2 August 2008, coll. W. Maddison, WPM#08-027. Paratypes: 2 males, same data as holotype; 1 male, same data as holotype; 1 male, same data as holotype. Etymology. The specific epithet is a patronym in honor of Dr. Tamas Szüts, who has contributed much to the taxonomy of this genus. Diagnosis. Resembles Thorelliola dumicola Berry, Beatty & Prószy!ski, 1997 in the absence of macrosetae on the tibia of male palp, but differs by the presence of a truncus on the male clypeus equipped with two ”horns”, and the short embolus. Can also be distinguished from other species that have the clypeal truncus by the combination of following chatacters: embolus is small; retrolateral sperm duct loop occupies about half of the bulb width; tibia of male palp ! 300! lacks prolateral macroseta. Description. Male (holotype, UBC-SEM AR00103). Carapace length 2.2 (variation 1.8-2.3, n=5); abdomen length 2.3. Truncus in the middle of clypeus and armed with two horn-like macrosetae with bases fused together. Chelicera (Figs 6.40D-E): dark brown to red brown; with two promarginal teeth and one retromarginal tooth of six or seven cusps; with a small projection on the front surface. Palp (Figs 6.40B-C): dark brown. Embolus very short; sperm duct loop occupying about half of the bulb width. Macroseta absent on the prolateral side of palpal tibia. Tibia of palp with a clump of long setae retrolaterally in front of retrolateral tibial apophysis. Retrolateral tibial apophysis relatively thick. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 6.3, II 4.6, III 4.9, IV 5.2. Color in alcohol (Fig. 6.40A): carapace dark brown, area around fovea yellow brown; abdomen gray brown, with many irregular sandy yellow speckles, posterior part with 3-4 parallel yellowish bands; first two pairs of legs dark brown proximally and sandy yellow distally, last two pairs of legs gray brown proximally and sandy yellow distally. Female. Unknown. Natural history. Specimens were collected from suspended litter. 6.4.8.6 Thorelliola tualapa sp. nov. Figs 6.41-6.42 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa, 5.283° S, 142.498° E, elev. 1000-1100 m a.s.l., 11-22 July 2008, forest interior and river side, coll. W. Maddison & Luc Fimo Tuki, WPM#08-008. Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 4 males and 1 female, same data as holotype; 1 female, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Differs from other species with a truncus on the clypeus of male by the wide retrolateral sperm duct loop, the shape of the embolus and the embolic disc of male palp. ! 301! Resembles Thorelliola pallidula Gardzinska, 2009 in female epigynum, but can be distinguished by the opening to the copulatory duct which is far anterior to the epigynal groove. Description. Male (holotype, UBC-SEM AR00104). Carapace length 2.4 (variation 1.8-2.4, n=6); abdomen length 2.5. Clypeus with a long truncus in the middle armed with two relatively short macrosetae at the tip. Chelicera (Figs 6.42E-F): dark brown, distal end red brown; with two promarginal teeth and one retromarginal tooth of six or seven cusps; with a distal process on front surface. Palp (Figs 6.42C-D): dark red brown. Bulb and embolic disc oval; embolus short and not curved; retrolateral sperm duct loop about three quarters of bulb width. Tibia of palp with multiple macrosetae, the prolateral distal macroseta of tibia on top of a protuberance. Palpal tibia with a clump of long and curved stiff setae retrolaterally in front of retrolateral tibial apophysis. Retrolateral tibial apophysis relatively wide and flap-like with a curved longitudinal ridge. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 7.8, II 5.3, III 5.8, IV 6.1. Color in alcohol (Fig. 6.42A): carapace dark red brown; abdomen sandy yellow, with gray and dark brown markings, anterior part with long and stiff setae; ventral abdomen pale yellow with irregular gray brown markings; first pair of legs dark red brown, other legs dark brown to light yellow brown. Female (paratype, UBC-SEM AR00105). Carapace length 1.8 (variation 1.7-1.8, n=3); abdomen length 2.6. Chelicera: with two promarginal teeth and one retromarginal tooth of four or five cusps. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.6, II 3.3, III 3.9, IV 4.2. Epigynum (Figs 6.42G-H): window not obvious; opening to copulatory duct far away from the epigynal groove. Copulatory duct short and thin; spermatheca almost spherical. Color in alcohol (Figs 6.42B): similar to that of male, but dorsal abdomen with two pairs of dark brown patches, carapace and legs paler. Natural history. Specimens were collected by beating forest understory. 6.4.8.7 Thorelliola zabkai sp. nov. Figs 6.43-6.44 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: Umgé, 5.304° S, 142.512° E, elev. 1450 m a.s.l., 15-19 July 2008, coll. W. Maddison & Aislan Tama ! 302! Wanakipa Indiaf, WPM#08-013. Paratype: 1 female, same data as holotype. Etymology. The specific epithet is a patronym in honor of Dr. Marek Zabka, who has made great contributions to the study of jumping spider systematics and biodiversity. Diagnosis. Female of this species can be distinguished from T. crebra and T. squamosa by the more obscure guanine deposits in the eye area and the indistinctive secondary spermatheca. Male of this species differs from T. crebra in the longer embolus, the shape of the embolic disc, the narrower sperm duct loop, the thinner retrolateral tibial apophysis and the bump-like protuberance on the palpal femur. Description. Male (holotype, UBC-SEM AR00106). Carapace length 2.0; abdomen length 2.1. Chelicera: red brown; with two promarginal teeth and one bicuspid retromarginal tooth; without distal process on front surface. Endite: grayish brown; without lateral bulge. Palp (Figs 6.44C- E): yellow brown. Embolic disc almost rectangular with the retrolateral margin almost straight; embolus long and curved; loop of sperm duct almost half as wide as bulb. Tibia, patella and femur of palp with multiple macrosetae; femur with a hump near distal end pro-ventrally. Retrolateral tibial apophysis long and finger-like. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.8, II 4.1, III 4.8, IV 5.0. Color in alcohol (Fig. 6.44A): carapace gray brown, with lateral margins and posterior eye area light yellow, guanine deposit in eye area not obvious; abdomen light yellow, markings not distinct in the preserved specimen; legs light yellow with dark gray markings, first pair of legs darker than others. Female (paratype, UBC-SEM AR00107). Carapace length 1.8; abdomen length 2.6. Chelicera: with two promarginal and one bicuspid retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.5, II 3.4, III 3.8, IV 4.4. Epigynum (Figs 6.44F-G): window large; opening to copulatory duct at posterior end of window laterally. Copulatory duct long and convoluted; secondary spermatheca not obvious, primary spermatheca small and spherical. Color in alcohol (Fig. 6.44B): similar to that of male, but lighter. Natural history. Specimens were collected on tree trunks in forest. ! 303! 6.4.9 Genus Variratina new genus Type species. Variratina minuta Zhang & Maddison, sp. nov. Etymology. The generic name is a reminiscent of “Varirata”, the name of the National Park where the species was first found. Diagnosis. Small tree-trunk dwelling jumping spiders. Resembles Bulolia Zabka, 1996, Leptathamas Balogh, 1980a (Szüts 2003) and Coccorchestes (see Balogh 1980b) in the sperm duct of male palp forming a loop at the prolateral side of the bulb, but can be easily distinguished by the body form and color pattern. ALEs and AMEs are in typical salticid configuration, unlike Bulolia and Leptathamas whose ALEs are posterior to the AMEs. Also differs from Coccorchestes in its carapace lacking crenellated posterior margin overlying the abdomen. 6.4.9.1 Variratina minuta sp. nov. Figs 6.45-6.46 Type material. Holotype: male, PAPUA NEW GUINEA: Central Province: Varirata National Park, 9.436° S, 147.364° E, elev. 740 m a.s.l., 4 August 2008, coll. W. Maddison, A. Kore & J. Kore, WPM#08–029. Paratype: 1 female, same data as holotype. Etymology. The specific epithet refers to the small size of the spider. Diagnosis. See diagnosis of the genus. Embolus of the male palp is slender with the plane of its spiral almost perpendicular to the longitudinal axis of the bulb. Epigynal window is large, and copulatory duct is convoluted. Description. Male (holotype, UBC-SEM AR00108). Carapace length 0.9; abdomen length 0.8. Chelicera: with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.46C-D): cream. Embolus long and spiral. Retrolateral tibial apophysis finger-like. Tibia of first leg with three ventral macrosetae; first metatarsus with two pairs. Measurements of legs: I 1.4, II 1.4, III 1.7, IV 1.9. Color in alcohol (Fig. 6.46A): carapace covered with pale yellow setae, eye area dark, ! 304! with a medial stripe composed of white setae extending to posterior part of carapace; behind eye area grey brown, with medial and lateral cream stripes; abdomen with a medial white stripe and some symmetrical dark markings; legs pale yellow, with indistinct grey marking at distal end of each segment. Female (paratype, UBC-SEM AR00109). Carapace length 1.0; abdomen length 1.4. Chelicera: with two promarginal teeth and one retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; first metatarsus with two pairs. Measurements of legs: I 1.6, II 1.6, III 2.0, IV 2.2. Epigynum (Figs 6.46E-F): window wide. Copulatory duct convoluted; spermatheca small. Color in alcohol (Fig. 6.46B): similar to that of male. Natural history. Specimens were found on tree trunks. 6.4.10 Genus Viribestus new genus Type species: Viribestus suyanensis Zhang & Maddison, sp. nov. Etymology. The generic name is from the combination of virilis (virile) and bestia (animal), referring to the masculine nature the male spider; masculine in gender. Diagnosis. Similar in body form and the presence of a lamella accompanying the embolus to Colyttus, but differs in the flat protrusion on the front surface of the male chelicera, the absence of proximal tegular lobe and the forked retrolateral tibial apophysis. Also differs from Canama and Bathippus in the wide carapace, the wide palpal bulb, the presence of a flat protrusion on the front surface of male chelicera, the presence of a lamella right beside the embolus and the forked retrolateral tibial apophysis of the male palp. 6.4.10.1 Viribestus suyanensis sp. nov. Figs 6.47-6.48 Type material. Holotype: male, PAPUA NEW GUINEA: Enga Province: Suyan Camp, Porgera, 5.4833° S, 143.1337° E, elev. 2300 m a.s.l., 28-29 July 2008, coll. W. Maddison, WPM#08-022. ! 305! Etymology. The specific epithet refers to the type locality. Diagnosis. Male has unique flat lateral extension on the front surface of chelicera. Palpal bulb is wide; embolus is short, thick and slightly curved, with a lamella beside it; retrolateral tibial apophysis is large and forked. Description. Male (holotype, UBC-SEM AR00110). Carapace length 2.0; abdomen length 3.0. Chelicera (Fig. 6.48B): red brown; with two promarginal teeth and one bicuspid retromarginal tooth; front surface with a flat extension laterally. Palp (Figs 6.48C-D): light yellow to reddish brown. Bulb wide with retrolateral sperm duct loop almost occupying its whole width. Embolus short and lightly curved, with a distinct lamella beside it. Retrolateral tibial apophysis with two branches, tip of ventral branch round and tip of dorsal branch pointed from retrolateral view. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 4.8, II 3.9, III 4.1, IV 4.2. Color in alcohol (Fig. 6.48A): carapace pale yellow, area between PLEs light orange, with white scales in eye area; abdomen pale yellow without distinct markings; first legs yellow brown, other legs pale yellow, coxa, trochanter, femur and patella of first leg and femur of second leg with dark markings prolaterally. Body greenish when alive (Figs 6.47A-D). Female. Unknown. Natural history. The holotype was found beating vegetation in a distrubed area. 6.4.11 Genus Xenocytaea Berry, Beatty & Prószy!ski, 1998 Berry et al. (1998) erected this genus to comprise five species from the Pacific Islands. Spiders of this genus are small to medium sized. Chelicera usually has two promarginal teeth and one bicuspid retromarginal tooth. First tibia has two or three pairs of ventral macrosetae and first metatarsus has two pairs of ventral macrosetae. Epigynum usually has a large anterior arch and lacks distinct window. Embolus is coiled and palpal bulb is usually wide. Retrolateral tibial apophysis is long and finger-like or wide with multiple branches. Some species have a protrusion on the distal end of the tegulum. Species described here have a unident rather than a bicuspid retromarginal tooth on the ! 306! chelicera, which is not congruent with previously reporated species. However, we place them in Xenocytaea because of their similar genitalic shape as the type species X. triramosa Berry, Beatty & Prószy!ski, 1998. 6.4.11.1 Xenocytaea agnarssoni sp. nov. Fig 6.49 Type material. Holotype: male, PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 1, Lamas, 5.614° S, 151.408° E, elev. 200 m, 3-8 April 2009, coll. I. Agnarsson. Paratypes: 1 female, same data as holotype; 2 female, same data as holotype; 1 male, PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 2, Vouvou, 5.446° S, 151.464° E, elev. 859 m, 10- 18 April 2009, coll. I. Agnarsson. Etymology. The specific epithet is a patronym in honor of Dr. I. Agnarsson, who collected specimens of this species and provided other material for this study. Diagnosis. Differs from other species by the absence of obvious anterior arch of the epigynum; the presence of a large protrusion at the distal end of the tegulum, the narrower embolic spiral, and the shape of the retrolateral tibial apophysis of the male palp. Description. Male (holotype, UBC-SEM AR00111). Carapace length 1.6 (variation 1.6-1.8, n=2); abdomen length 1.5. Chelicera: dark brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.49C-D): cream to light brown, with dark pigments. Embolus relatively wide and slightly coiled; retrolateral sperm duct loop present and wide; proximal tegular lobe absent. Retrolateral tibial apophysis with with two branches near the tip. First pair of legs with three pairs of ventral macrosetae on tibia and two pairs of ventral macrosetae on metatarsus. Measurements of legs: I 2.9, II 2.4, III 3.9, IV 3.8. Color in alcohol (Fig. 6.49A): carapace dark brown, with two white patches within eye area and one white patch right behind fovea composed of white scales; lateral margins with white stripes also composed of white scales; abdomen pale yellow, with two wide lateral stripes and one narrow medial stripe, all dark brown in color, ventral abdomen pale yellow with a wide dark stripe medially; legs cream without distinct markings. ! 307! Female (paratype, UBC-SEM AR00112). Carapace length 1.8 (variation 1.6-1.8, n=3); abdomen length 2.2. Chelicera (Fig. 6.49E): with two promarginal teeth and one retromarginal tooth. Tibia of first leg with five ventral macrosetae; metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.9, II 2.8, III 3.9, IV 3.7. Epigynum (Figs 6.49F-G): window relatively small, at anterior part of epigynal plate; opening to copulatory duct at posterior rim of the window. Copulatory duct wide and not coiled; spermatheca long and kidney-like. Color in alcohol (Fig. 6.49B): similar to that of male except without white stripes at lateral margins of carapace. 6.4.11.2 Xenocytaea albomaculata sp. nov. Fig 6.50 Type material. Holotype: male, PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 1, Lamas, 5.614° S, 151.408° E, elev. 200 m, 3-8 April 2009, coll. I. Agnarsson. Paratypes: 2 males, same data as holotype; 1 male, same as holotype; 1 male, same as holotype. Etymology. The specific epithet refers to the white markings on the carapace and abdomen. Diagnosis. Can be easily distinguished from other species by the short and thick embolus and the shape of the retrolateral tibial apophysis of male palp. Description. Male (holotype, UBC-SEM AR00113). Carapace length 1.4 (variation 1.4-1.5, n=5); abdomen length 1.2. Chelicera (Fig. 6.50F): dark yellow brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 6.50D-E): light yellow. Embolus wide and short, slightly coiled; sperm duct with a loop at the proximal side of bulb, retrolateral sperm duct loop absent; proximal tegular lobe absent. Retrolateral tibial apophysis finger-like, with tip beak-like. First pair of legs with three pairs of ventral macrosetae on tibia and two pairs of ventral macrosetae on metatarsus. Measurements of legs: I 2.8, II 2.6, III 3.5, IV 3.3. Color in alcohol (Fig. 6.50C): carapace dark brown with two white patches before PLEs and a white patch around fovea composed of white scales; lateral margins with white stripes composed of white scales too; abdomen dark, anterior margin covered with white scales, and with a pair of posterior markings composed of white scales; first pair of legs light yellow on metatarsi and tarsi, other segments brown or dark brown, other legs light yellow to gray brown with dark ! 308! stripes. Female. Unknown. Natural history. Specimens were collected by beating foliage in forest. 6.4.11.3 Xenocytaea proszynskii sp. nov. Fig 6.51 Type material. Holotype: female, PAPUA NEW GUINEA: New Britain: Nakanai Mts., near Camp 3, Tompoi, 5.343° S, 151.315° E, elev. 1590 m, 20 April 2009, coll. D. Gassmann & K. Sagatasame. Etymology. The specific epithet is a patronym in honor of Dr. J. Prószy!ski, who has made great contribution to the study of jumping spider systematics and biodiversity. Diagnosis. Resembles Xenocytaea triramosa Berry, Beatty & Prószy!ski, 1998 in lacking the posterior pocket of epigynum, but differs in the spermatheca closer to the epigynal groove and the shape of the anterior arch of the epigynum. Also similar in the epigynal shape to X. zabakai Berry, Beatty & Prószy!ski, 1998, but can be distinguished by the spherical spermatheca. Description. Female (holotype, UBC-SEM AR00114). Carapace length 1.6; abdomen length 1.7. Chelicera: red brown; with two promarginal teeth and one retromarginal tooth. Epigynum (Figs 6.51B-C): with a convex arch anteriorly; without median septum; opening to copulatory duct at posterior end of the arch. Copulatory duct relatively short, without accessory gland; spermatheca spherical. First pair of legs with three pairs of ventral macrosetae on tibia and two pairs of ventral macrosetae on metatarsus. Measurements of legs: I 2.9, II 2.8, III 3.7, IV 3.8. Color in alcohol (Fig. 6.51A): carapace dark red brown, with some white scales; abdomen light yellow with brown to brownish markings; venter with two brownish stripes behind genital groove; legs light yellow. Male. Unknown. ! 309! 6.4.12 Genus Zabkattus new genus Type species: Zabkattus brevis Zhang & Maddison, sp. nov. Etymology. The generic name combines “zabk” in honor of Dr. Marek Zabka, who has made great contributions in the study of jumping spider systematics and biodiversity, with “attus”, often used as an ending for salticid genera; masculine in gender. Diagnosis. Small to medium sized leaf litter dwelling spiders. Body is usually dark brown or brown. Chelicera has two promarginal teeth and one bicuspid retromarginal tooth. Male chelicera usually has a protrusion of varied shape on the front surface. Epigynum has a window with median septum. Spermatheca is usually relatively small. Resembles another leaf litter dwelling genus, Laufeia (see Bohdanowicz & Prószy!ski 1987) in body form, but differs in the shape of genitalia and the presence of a protrusion on the front surface of the male chelicera. 6.4.12.1 Zabkattus brevis sp. nov. Figs 6.52-6.53 Type material. Holotype: male, PAPUA NEW GUINEA: Southern Highlands Province: trail from Tualapa to Umgé, 5.2933° S, 142.4999° E, elev. 1210 m a.s.l., 19 July 2008, coll. W. Maddison, WPM#08-015. Paratypes: 1 female, same data as holotype; 1 male, PAPUA NEW GUINEA: Southern Highlands Province: Umgé, 5.304-5.305° S, 142.510-142.512° E, elev. 1400-1450 m a.s.l., 15-19 July 2008, coll. W. Maddison & Aislan Tama Wanakipa Indiaf, WPM#08-012; 1 female, same data as previous; 1 female and 1 male, same data as previous. Etymology. Latin brevis (short), referring to the short embolus of male palp. Diagnosis. Male can be easily distinguished from other species by the very short embolus and the long protrusion on the front surface of the chelicera. Female differs from Zabkattus trapeziformis in the median septum of the epigynum, which is not continuous with the edge of the window anteriorly, and the thicker and shorter copulatory duct. Description. Male (holotype, UBC-SEM AR00115). Carapace length 1.4 (variation 1.3-1.4, n=3); abdomen length 1.1. Chelicera (Fig. 6.53C): red brown; with two promarginal teeth and ! 310! one bicuspid retromarginal tooth; front surface with a long protrusion at the distal half part laterally. Palp (Figs 6.53D-E): light yellow. Proximal tegular lobe present. Embolus short and slightly curved. Retrolateral tibial apophysis short with distal end hawk-beak shaped. Tibia of first leg with three pairs of ventral macrosetae; first metatarsus with two pairs. Measurements of legs: I 3.0, II 2.4, III 2.7, IV 2.8. Color in alcohol (Fig. 6.53A): carapace dark red brown with orange and white scales; abdomen gray brown with irregular light yellow markings, covered with orange and white scales; legs light yellow, with distinct gray brown markings. Female (paratype, UBC-SEM AR00116). Carapace length 1.2 (variation 1.2-1.5, n=3); abdomen length 1.5. Chelicera: with two promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; first metatarsus with two pairs. Measurements of legs: I 2.3, II 2.2, III 2.5, IV 2.7. Epigynum (Figs 6.53F-G): window long and oval; median septum not reaching anterior edge of the window. Copulatory duct well sclerotized and thick; spermatheca small. Color in alcohol (Fig. 6.53B): similar to that of male. Natural history. Specimens were found on leaf litter in forest. 6.4.12.2 Zabkattus furcatus sp. nov. Fig 6.54 Type material. Holotype: male, PAPUA NEW GUINEA: Central Province: Varirata National Park, 9.436° S, 147.364° E, elev. 740 m a.s.l., 4 August 2008, coll. W. Maddison, A. Kore & J. Kore, WPM#08-029. Etymology. Latin furcatus (forked), referring to the forked protrusion on the front surface of male chelicera. Diagnosis. Differs from other species of the genus in the presence of two lateral white stripes on the carapace, the small and forked cheliceral protrusion, and the round tip of the retrolateral tibial apophysis from retrolateral view. Description. Male (holotype, UBC-SEM AR00117). Carapace length 2.5; abdomen length 2.2. Clypeus and cheeks covered with white scales. Chelicera (Fig. 6.54D): red brown; with two ! 311! promarginal teeth and one bicuspid retromarginal tooth; front protrusion near distal end, small and forked. Palp (Figs 6.54E-F): yellow brown. Proximal tegular lobe big. Embolus long and spiral. Retrolateral tibial apophysis relatively long, with distal end round from retrolateral view. Tibia and metatarsus of first leg with three pairs of ventral macrosetae each. Measurements of legs: I 5.1, II 4.2, III 4.9, IV 4.5. Color in alcohol (Fig. 6.54C): carapace red brown, with orange and white scales, with lateral stripes composed of white scales behind eye area; abdomen grayish brown, with irregular light yellow markings; femur of first leg red brown, other segments and legs yellowish with gray brown markings. Female. Unknown. Natural history. Specimens were found on leaf litter in forest. 6.4.12.3 Zabkattus richardsi sp. nov. Fig 6.55 Type material. Holotype: male, PAPUA NEW GUINEA: Eastern Highlands Province: Goroka, 6.07° S, 145.40° E, elev. 1650 m a.s.l., 31 July-1 August 2008, coll. W. Maddison, WPM#08- 024. Paratype: 1 male, same data as holotype. Etymology. The specific epithet is a patronym in honor of Dr. S. Richards, who designed and executed the successful expeditions to Papua New Guinea, during which this species was first found. Diagnosis. Resembles Zabkattus furcatus in male palpal shape, but can be easily distinguished by the body form and the color pattern, the big and hooked cheliceral protrusion on the front surface, and the pointed distal end of the retrolateral tibial apophysis. Description. Male (holotype, UBC-SEM AR00118). Carapace length 2.0 (variation 1.8-2.0, n=2); abdomen length 1.7. Chelicera (Fig. 6.55D): red brown; with two promarginal teeth and one bicuspid retromarginal tooth; front surface with a hooked protrusion at the distal end laterally. Palp (Figs 6.55E-F): light yellow with gray markings. Proximal tegular lobe big and round. Embolus long and coiled for about one circle. Retrolateral tibial apophysis short with ! 312! distal end pointed. Tibia of first leg with three pairs of ventral macrosetae; first metatarsus with two pairs. Measurements of legs: I 4.4, II 3.7, III 4.0, IV 3.8. Color in alcohol (Fig. 6.55C): carapace dark brown, covered with orange and white scales; abdomen gray brown, without distinct markings; legs dark brown, with light yellow annuli. Female. Unknown. Natural history. Specimens were found on leaf litter in forest. 6.4.12.4 Zabkattus trapeziformis sp. nov. Fig 6.56 Type material. Holotype: female, PAPUA NEW GUINEA: Central Province: Varirata National Park, 9.436° S, 147.364° E, elev. 740 m a.s.l., 4 August 2008, coll. W. Maddison, A. Kore & J. Kore, WPM#08-029. Etymology. The specific epithet refers to the trapezoid-shaped median septum of the epigynum. Diagnosis. This species differs from Zabkattus brevis by the median septum, which is continuous with the anterior margin of the window, the long and convoluted copulatory duct, and the presence of small accessory gland near the beginning of copulatory duct. Description. Female (holotype, UBC-SEM AR00119). Carapace length 1.5; abdomen length 1.5. Chelicera: red brown; with two promarginal teeth and one bicuspid retromarginal tooth. Tibia of first leg with three pairs of ventral macrosetae; metatarsus with two pairs. Measurements of legs: I 3.2, II 2.6, III 3.1, IV 3.1. Epigynum (Figs 6.56D-E): median septum continuous with the anterior edge of the window, with posterior end wider than anterior end. Copulatory duct long and convoluted, with an accessary gland near the beginning; spermatheca small and oval. Color in alcohol (Fig. 6.56C): carapace dark brown, covered with many white and a few orange scales; abdomen grayish brown, scattered with light yellow dots; legs grayish brown, with some yellowish markings. Male. Unknown. ! 313! Natural history. Specimens were found on leaf litter in forest. ! 314! Figure 6.1. Bathippus directus sp. nov. A-B. male paratype; C-D. female paratype. ! 315! Figure 6.2. Bathippus directus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), retrolateral view; F. male left chelicera, retrolateral view; G. male left chelicera, prolateral view; H. female left chelicera, back view; I. epigynum, ventral view; J. cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-H, 0.2 mm; I-J, 0.1 mm. ! 316! Figure 6.3. Bathippus gahavisuka sp. nov. A-C. male holotype; D. female paratype. ! 317! Figure 6.4. Bathippus gahavisuka sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left chelicera, prolateral view; D. male left chelicera, retrolateral view; E. male left palp, ventral view; F. male left palp, retrolateral view; G. male left palp (with patella and femur), retrolateral view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A, 2.0 mm; B, 1.0 mm; C-G, 0.2 mm; H-I, 0.1 mm. ! 318! Figure 6.5. Bathippus korei sp. nov. A-C. male paratype; D. female paratype. ! 319! Figure 6.6. Bathippus korei sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), ventral view; F. male left chelicera, prolateral view; G. male left chelicera, retrolateral view; H. female left chelicera, back view; I. epigynum, ventral view; J. cleared epigynum, dorsal view. Scale bars: A, 2.0 mm; B, 1.0 mm; C-H, 0.2 mm; I-J, 0.1 mm. ! 320! Figure 6.7. Bathippus madang sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view; D. male left palp (with patella and femur), ventral view; E. male left chelicera, retrolateral view; F. male left chelicera, prolateral view. Scale bars: A, 2.0 mm; B-F, 0.2 mm. ! 321! Figure 6.8. Canama extranea sp. nov. A-B. male paratype; C-D. female paratype. ! 322! Figure 6.9. Canama extranea sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), retrolateral view; F. male left chelicera, prolateral view; G. male left chelicera, retrolateral view; H. female left chelicera, back view; I. epigynum, ventral view; J. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-H, 0.2 mm; I-J, 0.1 mm. ! 323! Figure 6.10. Canama fimoi sp. nov. A-B. male paratype; C-D. female paratype. ! 324! Figure 6.11. Canama fimoi sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), retrolateral view; F. male left chelicera, front view; G. male left chelicera, medial view; H. male left chelicera, back view; I. female left chelicera, back view; J. epigynum, ventral view; K. cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-K, 0.2 mm. ! 325! Figure 6.12. Canama triramosa sp. nov. A-C. male holotype; D. female paratype. ! 326! Figure 6.13. Canama triramosa sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), retrolateral view; F. male left chelicera, front view; G. male left chelicera, back view; H. female left chelicera, back view; I. epigynum, ventral view; J. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-H, 0.2 mm; I-J, 0.1 mm. ! 327! Figure 6.14. Chalcolemia nakanai sp. nov. A. female holotype, dorsal view; B. female right chelicera, back view; C. epigynum, ventral view; D. cleared epigynum, ventral view; E. cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B, 0.2 mm; C-E. 0.1 mm. ! 328! Figure 6.15. Omoedus brevis sp. nov. A-B. male holotype; C. male paratype, dorsal view; D. male left palp, ventral view; E. male left palp, retrolateral view. Scale bars: C, 0.5 mm; D-E, 0.1 mm. ! 329! Figure 6.16. Omoedus darleyorum sp. nov. A-B. male paratype; C-D. female paratype. ! 330! Figure 6.17. Omoedus darleyorum sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.2 mm. ! 331! Figure 6.18. Omoedus meyeri sp. nov. A. male holotype; B-C. male paratype (lighter color form as holotype); D-E. male paratype (darker color form); F. female paratype. ! 332! Figure 6.19. Omoedus meyeri sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. ! 333! Figure 6.20. Omoedus omundseni sp. nov. A-B. male paratype; C-D. female paratype. ! 334! Figure 6.21. Omoedus omundseni sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, front view; F epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. ! 335! Figure 6.22. Omoedus papuanus sp. nov. A-C. male paratype; D. female paratype. ! 336! Figure 6.23. Omoedus papuanus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.2 mm. ! 337! Figure 6.24. Omoedus swiftorum sp. nov. A-B. male paratype; C-D. female paratype. ! 338! Figure 6.25. Omoedus swiftorum sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, front view; F. male left chelicera, back view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B, 0.5 mm; C-H, 0.2 mm. ! 339! Figure 6.26. Omoedus tortuosus sp. nov. A-C. male holotype; D. female paratype. ! 340! Figure 6.27. Omoedus tortuosus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.2 mm. ! 341! Figure 6.28. Paraharmochirus tualapaensis sp. nov. A-D. male holotype; E-F. female paratype. ! 342! Figure 6.29. Paraharmochirus tualapaensis sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-G, 0.1 mm. ! 343! Figure 6.30. Phasmolia elegans sp. nov. A-C. male paratype; D-F. female paratype. ! 344! Figure 6.31. Phasmolia elegans sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female right chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. ! 345! Figure 6.32. Sobasina wanlessi sp. nov. A. male; B. female paratype; C. male holotype, dorsal view; D. female paratype, dorsal view; E. male left chelicera, back view; F. male right leg I, retrolateral view; G. male left palp, ventral view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: C-D, 0.5 mm; F, 0.2 mm; E, G-I, 0.1 mm. ! 346! Figure 6.33. Thorelliola aliena sp. nov. A-C. male holotype; D. female paratype. ! 347! Figure 6.34. Thorelliola aliena sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left chelicera, back view; D. male left palp, ventral view; E. male left palp, retrolateral view; F. male left palp (with patella and femur), retrolateral view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-H, 0.2 mm. ! 348! Figure 6.35. Thorelliola crebra sp. nov. A-C. male holotype; D. female paratype. ! 349! Figure 6.36. Thorelliola crebra sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), ventral view; F. male chelicerae, endites and labium, ventral view; G. female left chelicera, back view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B, 0.5 mm; C-I, 0.2 mm. ! 350! Figure 6.37. Thorelliola joannae sp. nov. A. female holotype; B. female paratype, dorsal view; C. epigynum, ventral view; D. cleared epigynum, dorsal view. Scale bars: B, 0.5 mm; C-D, 0.1 mm. ! 351! Figure 6.38. Thorelliola squamosa sp. nov. A-B. female holotype; C. female paratype, dorsal view; D. epigynum, ventral view; E. cleared epigynum, dorsal view; F. cleared epigynum, ventral view. Scale bars: C, 0.5 mm; D-F, 0.1 mm. ! 352! Figure 6.39. Thorelliola tamasi sp. nov. A-D. male holotype. ! 353! Figure 6.40. Thorelliola tamasi sp. nov. A. male paratype, dorsal view; B. male left palp, ventral view; C. male left palp (with patella and femur), retrolateral view; D. male left chelicera, front view; E. male left chelicera, back view. Scale bars: A, 0.5 mm; B-E, 0.2 mm. ! 354! Figure 6.41. Thorelliola tualapa sp. nov. A-C. male holotype; D. female paratype. ! 355! Figure 6.42. Thorelliola tualapa sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp (with patella and femur), retrolateral view; E. male right chelicera, back view; F. male right chelicera, front view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-F, 0.2 mm; G-H, 0.1 mm. ! 356! Figure 6.43. Thorelliola zabkai sp. nov. A-C. male holotype; D. female paratype. ! 357! Figure 6.44. Thorelliola zabkai sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp (with patella and femur), ventral view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-E, 0.2 mm; F-G, 0.1 mm. ! 358! Figure 6.45. Variratina minuta sp. nov. A-B. male holotype; C-D. female paratype. ! 359! Figure 6.46. Variratina minuta sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A, 0.5 mm; B, 0.2 mm; C-F, 0.1 mm. ! 360! Figure 6.47. Viribestus suyanensis sp. nov. A-D. male holotype. ! 361! Figure 6.48. Viribestus suyanensis sp. nov. A. male holotype, dorsal view; B. male cephalothorax, front view; C. male left palp, ventral view; D. male left palp, retrolateral view. Scale bars: A, 1.0 mm; B-D, 0.2 mm. ! 362! Figure 6.49. Xenocytaea agnarssoni sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. ! 363! Figure 6.50. Xenocytaea albomaculata sp. nov. A-B. male paratype (copyright to P. Naskrecki, with permission); C. male paratype, dorsal view; D. male left palp, ventral view; E. male left palp, retrolateral view; F. male right chelicera, back view. Scale bars: C, 0.5 mm; D-F, 0.1 mm. ! 364! Figure 6.51. Xenocytaea proszynskii sp. nov. A. female holotype, dorsal view; B. epigynum, ventral view; C. cleared epigynum, dorsal view. Scale bars: A, 0.5 mm; B-C, 0.1 mm. ! 365! Figure 6.52. Zabkattus brevis sp. nov. A-D. male holotype; E-F. female paratype. ! 366! Figure 6.53. Zabkattus brevis sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left chelicera, front view; D. male left palp, ventral view; E. male left palp, retrolateral view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A- B, 0.5 mm; C, 0.2 mm; D-G, 0.1 mm. ! 367! Figure 6.54. Zabkattus furcatus sp. nov. A-B. male holotype; C. male holotype, dorsal view; D. male left chelicerae, front view; E. male left palp, ventral view; F. male left palp, retrolateral view. Scale bars: C, 1.0 mm; D-F, 0.2 mm. ! 368! Figure 6.55. Zabkattus richardsi sp. nov. A-B. male holotype; C. male paratype, dorsal view; D. male left chelicera, front view; E. male left palp, ventral view; F. male left palp, retrolateral view. Scale bars: C, 0.5 mm; D-F, 0.2 mm. ! 369! Figure 6.56. Zabkattus trapeziformis sp. nov. A-B. female holotype; C. female holotype, dorsal view; D. epigynum, ventral view; E. cleared epigynum, dorsal view. Scale bars: C, 0.5 mm; D- E, 0.1 mm. ! ! 370! 7 New euophryine jumping spiders from Central America and South America (Araneae: Salticidae: Euophryinae) 1 7.1 Synopsis Twenty-two new species and one new genus of euophryine jumping spiders from the Central America and South America are described. The new genus is Ecuadattus (E. elongatus sp. nov., E. napoensis sp. nov., E. pichincha sp. nov. and the type species E. typicus sp. n). The other new species belong to the genera Amphidraus (A. complexus sp. nov.), Belliena (B. ecuadorica sp. nov.), Chapoda (C. angusta sp. nov., C. fortuna sp. nov. and C. gitae sp. nov.), Ilargus (I. foliosus sp. nov., I. galianoae sp. nov., I. macrocornis sp. nov., I. moronatigus sp. nov., I. pilleolus sp. nov. and I. serratus sp. nov.), Maeota (M. dorsalis sp. nov., M. flava sp. nov. and M. simoni sp. nov.), Soesilarishius (S. micaceus sp. nov. and S. ruizi sp. nov.) and Tylogonus (T. parvus sp. nov. and T. yanayacu sp. nov.). Diagnostic illustrations are provided for all of the new species. Photographs of living spiders are also added for some new species. 7.2 Introduction The subfamily Euophryinae is one of the most diverse groups in Salticidae and has about 900 described species, with the majority found in the tropics of both the Old and the New World (Prószy!ski 1976; Maddison & Hedin 2003a; Platnick 2011). As one of the major jumping spider groups in the Central and South America (the others are Amycoid, Freyinae and Dendryphantinae), Euophryinae has 218 species of 31 genera reported in these areas (Platnick 2011). Although many taxonomic studies have been conducted (e.g. Chickering 1946; Edwards et al. 2005; Galiano 1960, 1962, 1963, 1967, 1976, 1985, 1988; Ruiz 2011), lots of euophryine jumping spider fauna in the Central and South America remains undiscovered. In this Chapter, I describe 22 new species and one new genus of euophryine jumping spiders from Ecuador, Brazil and Panama in order to give names for the taxa included in the molecular phylogenetic study on the subfamily Euophryinae (See Chapter 2). These molecular data also provide evidence for the generic placement of species described below. The new genus is !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! \"!A version of this chapter has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from Central America and South America (Araneae: Salticidae: Euophryinae). Names of new taxa are not finalized for nomenclature. ! ! 371! Ecuadattus (four species). The other new species belong to the genera Amphidraus (one species), Belliena (one species), Chapoda (three species), Ilargus (six new species), Maeota (three species), Soesilarishius (two species) and Tylogonus (two species). Diagnostic illustrations are provided for all of the new species. Photographs of living spiders are also added for some new species. 7.3 Material and methods Photographs of living specimens were taken with a Pentax Optio 33WR digital camera. For macro capability, a small lens was attached to it. Photographs of preserved specimens were taken under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0. Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Terminology is standard for Araneae. All measurements are given in millimeters. Descriptions of color pattern are based on the alcohol-preserved specimens. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; AME, anterior median eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the \"small eyes\"). Depository of type specimens are as following: Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM); Instituto Butantan, São Paulo, Brazil (IBSP); Museum of Zoology, Pontificia Universidad Católica, Quito, Ecuador (QCAZ). Specimens are deposited in UBC-SEM unless it is indicated otherwise. 7.4 Taxonomy 7.4.1 Genus Amphidraus Simon, 1900 Small sized spiders. Chelicera with two promarginal teeth and one fissident retromarginal tooth of more than two cusps. Embolic disc usually with an additional process besides embolus. Tegulum with an obvious proximal lobe. Retrolateral sperm duct loop not obvious. Retrolateral tibial apophysis usually large and complex in shape. Epigynum without window but with a pair ! 372! of copulatory openings anteriorly. Copulatory duct short or long and convoluted. Amphidraus differs from other Neotropical euophryine genera by the unique male palpal structures. Amphidraus is similar to Marma Simon, 1902 (Galiano 1962; 1963) in the cheliceral teeth pattern, but differs in the more delicate body form and the genitalic structures. Four species have been reported from the South America (Platnick 2011). A new species from Ecuador is described here. 7.4.1.1 Amphidraus complexus sp. nov. Fig 7.1 Type material. Holotype: male, ECUADOR: Napo: Estación Biológica Jatun Sacha, 1.067° S, 77.617° W, elev. 400 m, forest and nearby disturbed areas, 21-24 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-055 (QCAZ). Paratypes: 1 female, same data as holotype; 1 female, same data as holotype. Etymology. Latin complexus (an aggregate of parts, a complex), referring to the elaborate retrolateral tibial apophysis of the male palp. Diagnosis. Differs from other Amphidraus species in the shape of the retrolateral tibial apophysis and the process on the embolic disc of the male palp, and the extremely short copulatory duct and the large and swollen spermatheca of the vulva. Description. Male (holotype, UBC-SEM AR00133). Carapace length 1.1; abdomen length 1.1. Chelicera: sandy yellow, with two promarginal teeth and one fissident retromarginal tooth of four cusps. Palp (Figs 7.1C-D): light yellow brown to dark brown. Embolus slender with tip branched, embolic disc with a big process; retrolateral sperm duct loop very narrow. Retrolateral tibial apophysis complex, with a dorsal branch long and pointed at the tip, and a ventral branch wide and short. First pair of legs with three pairs of ventral macrosetae on tibia and two pairs of macrosetae on metatarsus. Measurements of legs: I 1.7, II 1.7, III 2.0, IV 2.1. Color in alcohol (Fig. 7.1A): carapace sandy yellow, with two gray stripes behind PLEs, eye area dark; dorsal abdomen light yellow, with a pair of dark brown longitudinal markings, posterior end of abdomen dark; anterior part of dorsum of abdomen with a relatively hard area; legs light yellow, ! 373! with a few dark annuli. Female (paratype, UBC-SEM AR00134). Carapace length 1.0; abdomen length 1.4. Chelicera: with two promarginal teeth and one retromarginal tooth of four cusps. First tibia with three pairs of ventral macrosetae and first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 1.4, II 1.3, III 1.6, IV 1.9. Epigynum (Fig. 7.1E): without window, opening to copulatory duct relatively anterior. Vulva (Fig. 7.1F): copulatory duct very short; spermatheca almost spherical. Color in alcohol (Fig. 7.1B): similar to that of male, but markings on dorsum of abdomen more distinct. 7.4.2 Genus Belliena Simon, 1902 Small sized spiders with carapace relatively smooth. Some species with prespiracular bump on the venter of abdomen in male. Genitalic organs typical euophryine-like: embolus coiled anti- clockwise (left palp, ventral view), tegulum with a proximal tegular lobe, retrolateral sperm duct loop present; epigynum with a window separated by the median septum. Similar to Neonella (see Galiano 1988) in the body form, but differs in the absence of a lamella along the embolus and the prolateral sperm duct loop, the presence of the retrolateral sperm duct loop, and the epigynal form. Four species have been reported from Venezuela and Trinidad (Platnick 2011). A new species from Ecuador is described here. 7.4.2.1 Belliena ecuadorica sp. nov. Fig 7.2 Type material. Holotype: male, ECUADOR: Morona Santiago: km 13 from Limón towards Gualaceo, 3.0093° S, 78.4939° W, elev. 1750 m, 12 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-034 (QCAZ); Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 3 females, ECUADOR: Morona Santiago: km 32 from Limón towards Gualaceo, 3.0221° S, 78.5833° W, elev. 2250 m. cloud forest, 11 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-028; 1 male, ECUADOR: Morona Santiago: km 38 from Limón towards Gualaceo, 3.0108° S, 78.6150° W, elev. 2725 m, cloud forest, 11-15 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-026. ! 374! Etymology. The specific epithet refers to the country where the species was found. Diagnosis. The male can be distinguished from other Belliena species (Galiano 1963) by the short embolus and the narrower embolic spiral. The female differs from that of B. biocellosa (see Galiano 1963) in the smaller window and the wider median septum of the epigynum. Description. Male (holotype, UBC-SEM AR00135). Carapace length 1.0 (variation 0.9-1.2, n=3); abdomen length 0.9. Chelicera: gray brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 7.2C-D): gray brown, with white hairs on dorsal femur. Embolus short and slightly curved; retrolateral sperm duct loop about half as wide as palpal bulb, tegular lobe large; retrolateral tibial apophysis long, tibial ventral bump small. First pair of legs with three pairs of ventral macrosetae on tibia and two pairs of macrosetae on metatarsus. Measurements of legs: I 1.8, II 1.7, III 2.1, IV 2.2. Color in alcohol (Fig. 7.2A): carapace gray brown; dorsal abdomen sandy yellow with dark brown markings, anterior and median part with a relatively hard area; legs light yellow. Some male specimens with a small prespiracular bump on the venter of abdomen. Female (paratype, UBC-SEM AR00136). Carapace length 1.0 (variation 1.0-1.1, n=4); abdomen length 1.6. Chelicera (Fig. 7.2E): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 1.8, II 1.7, III 2.0, IV 2.3. Epigynum (Fig. 7.2F): window relatively small, with wide median septum, opening to copulatory duct at posterior end of window. Vulva (Fig. 7.2G): copulatory duct swollen at beginning; spermatheca almost spherical. Color in alcohol (Fig. 7.2B): carapace sandy yellow; abdomen brownish with light yellow speckles and markings. 7.4.3 Genus Chapoda Peckham & Peckham, 1896 Medium sized spiders living on tree trunks, on branches or on foliage. Carapace sometimes with guanine deposit within eye area. Male palp sometimes with process on femur or/and patella; embolus slightly curved and coiled for less than one circle; proximal tegular lobe present or absent; ventral tibial bump usually present; retrolateral tibial apophysis finger-like. Epigynal window relatively small with a median septum. Beginning of copulatory duct swollen becoming ! 375! secondary spermatheca. Chapoda is one of the poorly studied genus with four species have been reported from the Central and South America (Platnick 2011). Three more species are described here. Unlike the known species, some species described here have no guanine deposit in the eye area and the male palp lacks modification on femur or patella. However, the molecular data suggest that they fall into the same clade as the typical Chapoda species (see Chapter 2). Thus, here we expand the delimitation of Chapoda to enclose these species. 7.4.3.1 Chapoda angusta sp. nov. Fig 7.3 Type material. Holotype: male, ECUADOR: Pichincha: near Puerto Quito, km 113 on road from Quito, ENDESA Campamento Maderero, 0.083° N, 79.117° W, 9-12 July 1988, coll. W. Maddison, WPM#88-014 (QCAZ). Paratypes: 1 female, same data as holotype; 6 males and 4 females, same data as holotype. Etymology. Latin angusta (narrow), referring to the narrow retrolateral sperm duct loop of the male palp. Diagnosis. Differs from the species once reported by the presence of an obvious proximal tegular lobe and the absence of modification on the femur or/and patella of the male palp. Similar to Chapoda gitae and C. fortuna in the presence of a proximal tegular lobe on the male palp, but differs from the in the scale-like markings on the abdomen and the narrower retrolateral sperm duct loop of the male palp. This species can also be distinguished from C. gitae by the wider embolic spiral of the male palp; and from C. fortuna by the absence of a process on the femur of the male palp ventrally. Description. Male (holotype, UBC-SEM AR00137). Carapace length 1.4 (variation 1.3-1.4, n=7); abdomen length 1.3. Chelicera: yellow brown; promargin with two teeth, retromargin with one tooth. Palp (Figs 7.3C-D): yellowish brown. Proximal tegular lobe distinct; embolus short and slightly curved; retrolateral tibial apophysis finger-like; ventral tibial bump small. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. ! 376! Measurements of legs: I 2.6, II 2.4, III 2.8, IV 3.1. Color in alcohol (Fig. 7.3A): carapace red brown, lateral and posterior margins covered with white scales, PMEs and PLEs with dark surroundings; abdomen sandy yellow with white scale-like markings, anterior end gray brown; legs yellowish to reddish brown. Female (paratype, UBC-SEM AR00138). Carapace length 1.5 (1.4-1.6, n=5); abdomen length 2.4. Chelicera (Fig. 7.3F): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.7, II 2.6, III 3.2, IV 3.2. Epigynum (Fig. 7.3G): epigynal window relatively small and median septum wide; opening to copulatory duct at posterior end of window. Vulva (Fig. 7.3H): copulatory duct swollen near the beginning forming a secondary spermatheca; primary spermatheca relatively long and oval. Color in alcohol (Fig. 7.3B): similar to that of male, but carapace and legs lighter in color. 7.4.3.2 Chapoda fortuna sp. nov. Fig 7.4 Type material. Holotype: male, PANAMA: Chiriqui: Fortuna, Quebrada Samudio, 8.73464° N, 82.24839° W, elev. 1209-1245 m, 21 September 2008, coll. J. X. Zhang & G. B. Edwards, JXZ08#002. Paratype: 1 male, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Similar to Chapoda panamana Chickering, 1946 in the shape of the ventral process on the palpal femur, but differs in the presence of a proximal tegular lobe and the wide retrolateral sperm duct loop of the male palp, the one unident retromarginal tooth on the male chelicera (C. panamana has one bicuspid retromarginal tooth on the chelicera). Description. Male (holotype, UBC-SEM AR00141). Carapace length 1.4 (variation 1.4-1.6, n=2); abdomen length 1.4. Carapace with two tufts behind AMEs. Chelicera: promargin with two teeth, retromargin with one tooth. Palp (Figs 7.4B-C): yellow brown. Proximal tegular lobe present; retrolateral sperm duct loop almost as wide as palpal bulb; embolus short and slightly curved; retrolateral tibial apophysis narrow and finger-like; ventral tibial bump distinct; palpal ! 377! patella not wide, femur of palp with a ventral process near the center. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.9, II 2.5, III 3.0, IV 3.4. Color in alcohol (Fig. 7.4A): carapace dark brown, lateral margins with sparse white scales, without guanine deposit in eye area; abdomen dark brown with many brownish speckles, anterior part with a large middle stripe followed with four paralleled streaks, all brownish in color; legs dark yellow brown, last two pairs of legs with yellowish brown annuli. Female. Unknown. Natural history. Specimens were collected by beating mossy branches in wet forest. 7.4.3.3 Chapoda gitae sp. nov. Figs 7.5, 7.23 Type material. Holotype: male, ECUADOR: Esmeraldas: Reserva Canandé, Grd.-Cuckoo Trail, 0.5214-0.5216° N, 79.2049-79.2045° W, elev. ca. 550 m, 22 August 2011, coll. Piascik & Vega, WPM#11-177 (QCAZ). Paratype: 1 female, ECUADOR: ESMERALDAS: Reserva Canandé, Choco Tapa Culo Trail, 0.524-0.526° N, 79.212-79.213° W, elev. 350 m, 21-23 August 2011, coll. Maddison & Vega, WPM#11-173. Etymology. The specific epithet is a patronym in honor of Dr. Gita S. S. Bodner, who studied the phylogeny of Euophryinae in her PhD thesis. Diagnosis. Similar to Chapoda angusta in the epigynal form, but differs in the color pattern of the body, the narrower embolic spiral and the wider retrolateral sperm duct loop of the male palpal bulb. Description. Male (holotype, UBC-SEM AR00139). Carapace length 1.5; abdomen length 1.4. Chelicera: yellow brown to gray brown; promargin with two teeth, retromargin with one tooth. Palp (Figs 7.5C-D): femur dark, other segments yellow brown to light yellow. Proximal tegular lobe distinct; retrolateral sperm duct loop wide; embolus short and slightly curved with the spiral relatively narrow; retrolateral tibial apophysis finger-like; ventral tibial bump obvious. ! 378! First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.0, II 2.8, III 3.4, IV 3.5. Color in alcohol (Fig. 7.5A): carapace dark brown, middle area and lateral margins with white scales, clypeus also covered with white scales; abdomen anterior margin and lateral margins dark, other area pale yellow with some gray markings; legs yellowish to dark brown. Female (paratype, UBC-SEM AR00140). Carapace length 1.6; abdomen length 2.1. Chelicera: with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macroseate. Measurements of legs: I 3.0, II 2.9, III 3.4, IV 3.9. Leg formula: 4312. Epigynum (Fig. 7.5E): window relatively small with a median septum. Vulva (Fig. 7.5F): copulatory duct very short; with a pair of spherical secondary spermathecae; primary spermatheca long and oval. Color in alcohol (Fig. 7.5B): similar to that of male, but carapace with a stripe behind PLEs composed of white scales; abdomen with two lateral dark stripes; legs paler than those of male. 7.4.4 Genus Ecuadattus new genus Type species: Ecuadattus typicus Zhang & Maddison, sp. nov. Etymology. The first part of the generic name, “Ecuad” is derived from Ecuador (where the species were found), and the second part “attus” is a commonly used ending for salticid genera; masculine in gender. Diagnosis. Medium sized spiders usually found on foliage. Carapace usually with guanine deposit in eye area and abdomen usually with foliage-like light colored markings. Female chelicera with two promarginal teeth and one retromarginal tooth, but male chelicera usually with a bicuspid tooth on promargin. Epigynal window present with a median septum. Spermatheca oval or round. Differs from other Neotropical foliage-dwelling euophryine genera by the unique male palpal structures: embolic disc small or highly reduced; embolus short and slightly curved; retrolateral sperm duct loop narrow; retrolateral tibial apophysis long and finger-like. ! 379! 7.4.4.1 Ecuadattus elongatus sp. nov. Fig 7.6 Type material. Holotype: male, ECUADOR: Morona Santiago: km 16 from Limón towards Gualaceo, 3.0060° S, 78.4997° W, elev. 2000 m, 12-15 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-035 (QCAZ). Etymology. Latin elongatus (elongated), referring to the elongated male palpal tibia. Diagnosis. Similar to Ecuadattus napoensis in the color pattern and markings, but differs in the darker colored second, third and fourth legs, the relatively large dark markings on dorsum of the abdomen; the long male palpal tibia, the highly reduced embolic disc, and the narrower palpal bulb. This species also differs from E. typicus in the indistinctive proximal tegular lobe and embolic disc of the male palp. Description. Male (holotype, UBC-SEM AR00142). Carapace length 2.7; abdomen length 3.1. Chelicera: red brown, back surface with a depression. Palp (Figs 7.6B-C): yellow brown. Embolic disc highly reduced; embolus short and slightly curved; palpal tibia long; retrolateral tibial apophysis long and finger-like; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 5.7, II 4.8, III 5.7, IV 5.7. Color in alcohol (Fig. 7.6A): carapace brown, eye area with guanine deposit, posterior eye area with a narrow yellowish stripe, lateral margins behind eye area with yellowish stripes; abdomen light sandy yellow, with a pair of dark patches and some obscure markings; first leg brown, the other legs sandy yellow to brown. Female. Unknown. 7.4.4.2 Ecuadattus napoensis sp. nov. Fig 7.7 Type material. Holotype: male, ECUADOR: Napo: Cocodrilo, 0.6490° S, 77.7927° W, elev. 2040 m, 20-24 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-054 (QCAZ). ! 380! Etymology. The specific epithet refers to the type locality. Diagnosis. Differs from Ecuadattus elongatus by the shorter tibia of the male palp, the presence of an obvious embolic disc, and the wider palpal bulb. This species can also be distinguished from E. typicus and E. pichincha in the indistinctive proximal tegular lobe of the male palp. The palpal bulb of this species is wider than that of E. typicus. The retrolateral sperm duct loop of this species is narrower than that of E. pichincha. Description. Male (holotype, UBC-SEM AR00143). Carapace length 2.9; abdomen length 3.2. Chelicera (Fig. 7.7B): red brown; back surface with a depression; promargin with one bicuspid tooth and retromargin with one tooth. Palp (Figs 7.7C-D): red brown to yellow brown. Embolic disc small; embolus slightly curved. Retrolateral tibial apophysis long and finger-like; palpal tibia with a ventral ridge. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 7.2, II 5.4, III 6.0, IV 6.3. Color in alcohol (Fig. 7.7A): carapace brown, eye area with guanine deposit, posterior eye area with a narrow yellowish stripe; abdomen brownish, with a longitudinal pale yellow stripe; first leg brown, other legs pale yellow. Female. Unknown. 7.4.4.3 Ecuadattus pichincha sp. nov. Figs 7.8, 7.24 Type material. Holotype: male, ECUADOR: Pichincha: Bellavista, Cloud Forest Reserve, 0.012-016° S, 78.682° W, elev. 2050-2240 m, 9-10 November 2010, coll. W. Maddison, D. Maddison, M. Vega & M. Reyes, WPM#10-065 (QCAZ). Paratypes: 1 female, same data as holotype; 1 male, same data as holotype; 3 males and 1 female, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. This species resembles Ecuadattus typicus in the coloration and markings, but it can be distinguished f by the wider palpal bulb, the shorter male palpal tibia, the presence of a ! 381! ventral ridge on the palpal tibia, and the wider median septum of the epigynum. Also see the diagnosis of E. napoensis. Description. Male (holotype, UBC-SEM AR00144). Carapace length 2.3 (variation 1.9-2.3, n=5); abdomen length 2.3 (variation 2.0-2.2, n=2). Chelicera (Fig. 7.8E): dark red brown; promargin with one bicuspid tooth and retromargin with one tooth. Palp (Figs 7.8C-D): yellow brown. Bulb wide; tegular lobe present; embolic disc obvious; embolus slightly curved; retrolateral tibial apophysis long and finger-like; palpal tibia with a ventral ridge. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 4.8, II 4.0, III 4.2, IV 4.9. Color in alcohol (Fig. 7.8A): carapace dark brown, eye area with guanine deposit, behind eye area and lateral margins with yellowish stripes; abdomen brownish, with numerous light yellow speckles and a longitudinal wide stripe; first leg dark brown, other legs pale yellow with brownish annuli. Female (paratype, UBC-SEM AR00145). Carapace length 2.2; abdomen length 3.5. Chelicera: with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.8, II 3.6, III 4.3, IV 4.9. Epigynum (Fig. 7.8F): median septum present and wide, with opening to the copulatory duct anteriorly. Vulva (Fig. 7.8G): copulatory duct relatively long but not coiled; spermatheca oval. Color in alcohol (Fig. 7.8B): similar to that of male, but first leg paler in color and dorsum of abdomen obviously with a pair of white spots near posterior end. 7.4.4.4 Ecuadattus typicus sp. nov. Figs 7.9, 7.25 Type material. Holotype: male, ECUADOR: Napo: Vinillos, near Cosanga, 0.6025° S, 77.8508° W, elev. 2080 m, 29-30 October 2010, coll. W. Maddison, D. Maddison, M. Vega & M. Reyes, WPM#10-036 (QCAZ). Paratypes: 1 female, same data as holotype; 6 males and 4 females, same data as holotype; 3 males, ECUADOR: Napo: Reserva Ecologica Antisana, Sendero Jumandy, 0.624-0.625° S, 77.840-77.842° W, elev. 2260 m, 29 October 2010, coll. W. Maddison, D. Maddison & M. Reyes, WPM#10-035; 1 male, ECUADOR: Napo: Cosanga, Yanayacu Biological Station, forest, 0.600-0.601° S, 77.888-77.890° W, elev. 2100 m, 7 November 2010, coll. W. Maddison, D. Maddison, M. Vega & M. Reyes, WPM#10-058. ! 382! Etymology. Latin typicus (typical), referring that this is the type species of the genus. Diagnosis. Differs from Ecuadattus napoensis by the long palpal tibia, the distinct tegular lobe and the absence of a ventral ridge on the palpal tibia. This species can be distinguished from E. pichincha by the narrower palpal bulb, the longer palpal tibia, the absence of a ventral ridge on the palpal tibia, and the narrower median septum of the epigynum. Description. Male (holotype, UBC-SEM AR00146). Carapace length 2.5 (variation 1.9-2.8, n=11); abdomen length 2.8. Chelicera (Fig. 7.9E): dark red brown; promargin with one bicuspid tooth and retromargin with one tooth. Palp (Figs 7.9C-D): dark yellow brown. Palpal bulb narrow; tegular lobe distinct; embolic disc obvious; embolus slightly curved; retrolateral tibial apophysis long and finger-like. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 6.4, II 4.8, III 5.5, IV 5.9. Color in alcohol (Fig. 7.9A): carapace dark brown, eye area with guanine deposit, behind eye area with a longitudinal yellowish stripe; abdomen brown, with numerous light yellow speckles and a longitudinal stripe; first leg dark brown, other legs pale yellow to brown. Some male specimens with yellowish margins on dorsal abdomen. Female (paratype, UBC-SEM AR00147). Carapace length 2.0 (variation 2.0-2.3, n=5); abdomen length 2.0. Chelicera: with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.8, II 3.4, III 4.3, IV 4.7. Epigynum (Fig. 7.9G): median septum wide, with opening to the copulatory duct anteriorly. Vulva (Fig. 7.9H): copulatory duct not coiled; spermatheca round. Color in alcohol (Fig. 7.9B): paler than that of male, and markings on dorsal abdomen more obvious, first leg light yellow. Some female specimens as dark in color as that of holotype male. 7.4.5 Genus Ilargus Simon, 1901 Medium sized spiders with various color and marking patterns. Chelicera with two promarginal teeth and one unident retromarginal tooth. Male palpal bulb usually large and wide, with obvious proximal tegular lobe; embolus usually long and coiled; retrolateral tibial apophysis finger-like. Epigynal window present with a median septum. Spermatheca swollen. ! 383! Three species have been reported from the South America (Platnick 2011), of which only the type species, Ilargus coccineus Simon, 1901b has been well documented (Galiano 1963; Edwards et al. 2005). However, Ilargus is much more diverse than that. During the expeditions to Ecuador we discovered six new species, which will be described here. The placement of these species in Ilargus is partly based on unpublished molecular data. 7.4.5.1 Ilargus foliosus sp. nov. Fig 7.10 Type material. Holotype: male, ECUADOR: Morona Santiago: km 15 from Limón towards Gualaceo, 3.0090° S, 78.5024° W, elev. 1940 m, 15 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-043 (QCAZ). Paratype: 1 female, ECUADOR: Morona Santiago: km 7 from Limón towards Gualaceo, 2.9962° S, 78.4558° W, elev. 1415 m, 12-15 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-032. Etymology. Latin foliosus (leafy), referring to the leaf-like marking on the dorsum of the abdomen. Diagnosis. Similar to Ilargus macrocornis in the color and marking patterns, and the presence of guanine deposit in the eye area, but differs in the absence of a spur on the front surface of the male chelicera. Description. Male (holotype, UBC-SEM AR00148). Carapace length 1.7; abdomen length 1.7. Chelicera: red brown. Palp (Figs 7.10C-D): tibia and cymbium dark brown, other segments cream. Embolus long with a relatively wide spiral; proximal tegular lobe large; retrolateral sperm duct loop occupying more than half of bulb width; retrolateral tibial apophysis finger- like; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.3, II 2.9, III 3.4, IV 3.6. Color in alcohol (Fig. 7.10A): carapace dark brown, eye area with guanine deposit, behind eye area with a medial and two marginal yellowish stripes; abdomen dark brown, with many silvery white speckles and a large leaf-like mark medially; legs yellowish to dark brown. ! 384! Female (paratype, UBC-SEM AR00149). Carapace length 1.4; abdomen length 1.8. Chelicera (Fig. 7.10E): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.4, II 2.2, III 2.6, IV 3.2. Epigynum (Fig. 7.10F): window large; median septum relatively narrow; opening to copulatory duct at each side of median septum anteriorly. Vulva (Fig. 7.10G): copulatory duct slightly convoluted; spermatheca almost stomach-shaped. Color in alcohol (Fig. 7.10B): similar to that of male, but first two pairs of legs lighter in color than those of male. 7.4.5.2 Ilargus galianoae sp. nov. Fig 7.11 Type material. Holotype: male, ECUADOR: Morona Santiago: km 16 from Limón towards Gualaceo, 3.0060° S, 78.4997° W, elev. 2000 m, 12-15 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-035 (QCAZ). Paratypes: 1 female, same data as holotype; 1 female, same data as holotype. Etymology. The specific epithet is a patronym in honor of Dr. M. E. Galiano, who made great contributions in taxonomic studies of jumping spiders from Neotropical region. Diagnosis. Similar to Ilargus moronatigus in the color and marking patterns, but can be distinguished by the large epigynal window, the presence of the wave-shaped ridges in the window, and the long and convoluted copulatory duct. Also differs from other Ilargus species by the male palpal structures. Description. Male (holotype, UBC-SEM AR00150). Carapace length 1.5; abdomen length 1.7. Chelicera: yellow brown; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.11C-D): yellowish to brown. Embolus long and coiled for about a circle; retrolateral sperm duct occupying about two thirds of bulb width; retrolateral tibial apophysis finger-like; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.1, II 2.6, III 3.0, IV 3.5 Color in alcohol (Fig. 7.11A): carapace gray brown with indistinctive yellowish markings, eye area dark brown, behind eye area with a narrow sandy yellow stripe; abdomen burlywood in color, with brownish ! 385! markings and speckles; legs bisque, with gray brown annuli. Female (paratype, UBC-SEM AR00151). Carapace length 1.6; abdomen length 2.0. Chelicera (Fig. 7.11E): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.8, II 2.6, III 3.1, IV 3.8. Epigynum (Fig. 7.11F): window large, with transverse wave- shaped ridges in the window; median septum relatively narrow; opening to copulatory duct at anterior margin of window. Vulva (Fig. 7.11G): copulatory duct long and convoluted, with accessory gland near the beginning; spermatheca almost kidney-shaped. Color in alcohol (Fig. 7.11B): similar to that of male, but abdomen dark brown with burlywood markings and speckles. 7.4.5.3 Ilargus macrocornis sp. nov. Figs 7.12, 7.26C Type material. Holotype: male, ECUADOR: Morona Santiago: Cordillera de Cutucú; road from Patuca to Santiago, 2.8058° S, 78.2462° W, elev.1000 m, 13 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-040 (QCAZ). Etymology. The specific epithet is from the combination of macro- (long, large) and cornis (horned), referring to the large spur on the front surface of the male chelicera. Diagnosis. It can be easily distinguished from the other species by the presence of spurs on the front surface of male chelicera. Description. Male (holotype, UBC-SEM AR00152). Carapace length 1.7; abdomen length 1.7. Chelicera (Fig. 7.12D): red brown; promargin with two teeth, retromargin with one tooth; front surface with a long curved spur proximally. Palp (Fig. 7.12B-C): tibia and cymbium yellow brown, other segments cream. Embolus long and coiled for less a circle, with a short and thin lamella; proximal tegular lobe large; retrolateral tibial apophysis finger-like; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 4.2, II 2.9, III 3.3, IV 3.8. Color in alcohol (Fig. 7.12A): carapace dark brown, eye area with guanine deposit, behind eye area with a medial and ! 386! two marginal yellowish stripes; abdomen dark brown, with many silvery white speckles and a large leaf-like marking medially; legs yellowish to dark brown. Female. Unknown. 7.4.5.4 Ilargus moronatigus sp. nov. Fig 7.13 Type material. Holotype: female, ECUADOR: Morona Santiago: km 44 from Limón towards Gualaceo, 3.0069° S, 78.6425° W, elev. 3000 m, cloud forest, 11 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-025 (QCAZ). Paratypes: 1 female, same data as holotype; 2 females, ECUADOR: Morona Santiago: km 38 from Limón towards Gualaceo, 3.0108° S, 78.6150° W, elev. 2725 m, cloud forest, 11-15 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-026. Etymology. The specific epithet is derived from the Province name where the species was discovered. Diagnosis. Similar to Ilargus galianoae in the marking patterns, but differs in the smaller window, the absence of wave-like ridge in the window of the epigynum; and the shorter and less convoluted copulatory duct of the vulva. Description. Female (holotype, UBC-SEM AR00153). Carapace length 1.6 (variation 1.4-1.6, n=4); abdomen length 1.9. Chelicera (Fig. 7.13B): yellow brown; with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.4, II 2.1, III 2.6, IV 3.4. Epigynum (Fig. 7.13C): windows small; opening at posterior margin of window. Vulva (Fig. 13D): copulatory duct thin and slightly convoluted, with accessory gland near the beginning; spermatheca almost kidney-shaped. Color in alcohol (Fig. 7.13A): carapace gray brown with indistinct yellowish markings, eye area dark brown, thoracic area with a narrow yellowish brown stripe; abdomen dark brown, with light yellow speckles and markings; legs yellowish brown. ! 387! Male. Unknown. 7.4.5.5 Ilargus pilleolus sp. nov. Figs 7.14, 7.26A-B Type material. Holotype: male, ECUADOR: Pichincha: Crater del Pululahua, 0.033° S, 78.500° W, 19 June 1988, coll. W. Maddison, WPM#88-001 (QCAZ). Paratypes: 1 female, same data as holotype; 5 males and 3 females, same data as holotype. Etymology. Latin pilleolus (little cap), referring to the tuft behind the anterior eye row on the male carapace. Diagnosis. Differs from other Ilargus species by the presence of a row of tuft on the male carapace and the foliage-like marking on the abdomen. This species can also be distinguished from Ilargus serratus by the shorter embolus, the smooth margin of the retrolateral tibial apophysis of the male palp; and the wider median septum of the epigynal window. Description. Male (holotype, UBC-SEM AR00154). Carapace length 1.5 (variation 1.4-1.6, n=6); abdomen length 1.5. Chelicera: yellow brown; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.14C-D): reddish brown, tibia and patella dark ventrally. Palpal bulb wide; embolus wide; proximal tegular lobe large; retrolateral sperm duct loop almost occupying half of the bulb width; retrolateral tibial apophysis finger-like; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.3, II 2.2, III 2.4, IV 2.8. Color in alcohol (Fig. 7.14A): carapace red brown, with a row of dense sandy yellow tuft behind the first eye row, eye area yellowish, thoracic area with a narrow yellowish stripe medially; abdomen grayish brown, with many yellowish speckles and a medial yellowish leaf-like marking, posterior end of abdomen with a dark triangular marking; legs yellowish. Female (paratype, UBC-SEM AR00155). Carapace length 1.5 (variation 1.5-1.6, n=4); abdomen length 2.1. Chelicera (Fig. 7.14G): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of macrosetae. Measurements of legs: I 2.6, II 2.5, III 2.8, IV 3.1. Epigynum (Fig. 7.14H): window ! 388! large, median septum wide. Vulva (Fig. 7.14I): copulatory duct swollen at the beginning; spermatheca almost round. Color in alcohol (Fig. 7.14B): similar to that of male, thoracic area of carapace with yellowish lateral margins. 7.4.5.6 Ilargus serratus sp. nov. Fig 7.15 Type material. Holotype: male, ECUADOR: Morona Santiago: km 13 from Limón towards Gualaceo, 3.0093° S, 78.4939° W, elev. 1750 m, 12 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-034 (QCAZ). Paratypes: 1 female, same data as holotype; 1 female, same data as holotype. Etymology. Latin serratus (serrated), referring to the serrated edge of the retrolateral tibial apophysis of the male palp. Diagnosis. Similar in the markings to Ilargus coccineus (see Edwards et al. 2005) in the markings, but can be distinguished by the less elongated body; the swollen part at the beginning of the copulatory duct; the wider and branched embolus, and the serrated retrolateral tibial apophysis of the male palp. Description. Male (holotype, UBC-SEM AR00156). Carapace length 1.7; abdomen length 2.1. Chelicera: pale yellow; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.15C-D): yellowish to yellow brown with gray pigment. Palpal bulb wide; embolus long and coiled for less than a circle, branched from the base; retrolateral tibial apophysis finger-like with dorsal margin serrated; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 4.6, II 3.7, III 4.4, IV 4.3. Color in alcohol (Fig. 7.15A): carapace yellowish, with two dark brown wide stripes from anterior eyes to posterior end of carapace; abdomen yellowish, with two lateral dark brown stripes from anterior margin to posterior margin; first pairs of legs yellow brown, other legs yellowish. Female (paratype, UBC-SEM AR00157). Carapace length 2.2 (variation 2.0-2.2, n=2); abdomen length 2.2. Chelicera (Fig. 7.15F): with two promarginal teeth and one retromarginal ! 389! tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.6, II 3.3, III 4.2, IV 4.5. Epigynum (Fig. 7.15G): window large; median septum narrow; opening at anterior margin of window. Vulva (Fig. 7.15H): copulatory duct thick, swollen at the beginning; spermatheca oval. Color in alcohol (Fig. 7.15B): paler than that of male, stripes on carapace light orange, stripes on abdomen gray brown, all legs pale yellow. 7.4.6 Genus Maeota Simon, 1901 The genus was described for Maeota dichrura Simon, 1901a, which is quite unique for the abnormally elongate posterior lateral spinneret in the male. However, the preliminary unpublished molecular data discovered a group of species that are closely related to M. dichrura, but have no elongate posterior lateral spinneret in the male. Thus, the elongate spinneret might be an automorphy just for M. dichrura, and the delimitation of Maeota is extended here to include the closely related species: medium sized spiders usually live on foliage; the chelicera has two promarginal teeth and one retromarginal tooth, without obvious modifications; the embolus is coiled usually with the plane of the spiral more or less perpendicular to the longitudinal axis of the bulb; the tegulum has a proximal lobe. 7.4.6.1 Maeota dorsalis sp. nov. Fig 7.16 Type material. Holotype: male, BRAZIL: São Paulo: São Paulo, 2008, coll. G. Ruiz (IBSP, #96121). Etymology. Latin dorsalis (dorsal), and refers to the dorsal apophysis on tibia of male palp. Diagnosis. Differs from other Maeota species by the presence of a dorsal tibial apophysis of the male palp. This species can also be distinguished from Maeota flava by the marking on the dorsum of the male abdomen; the shape of the embolic disc and the longer retrolateral tibial apophysis of the male palp. Description. Male (holotype, UBC-SEM AR00158). Carapace length 1.4; abdomen length 1.3. Chelicera: gray brown; promargin with two teeth and retromargin with one tooth. Palp (Figs ! 390! 7.16B-D): brown to yellowish, with gray pigment. Proximal tegular lobe large; retrolateral sperm duct loop very narrow; embolus relatively short and slightly coiled; retrolateral tibial apophysis finger-like; dorsal tibial apophysis present and slightly curved distally; ventral tibial bump present. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.6, II 2.5, III 3.0, IV 3.4. Color in alcohol (Fig. 7.16A): carapace dark brown and yellow brown around fovea region, with a narrow medial stripe behind fovea and two lateral stripes composed of white scales; abdomen gray brown with many brownish speckles, and a wide sandy yellow stripe medially, ventral abdomen and legs sandy yellow without distinct markings. Female. Unknown. Natural history. Specimen collected on low vegetations in synanthropic habitat. 7.4.6.2 Maeota flava sp. nov. Fig 7.17 Type material. Holotype: male, BRAZIL: São Paulo: São Paulo, 2008, coll. G. Ruiz (IBSP, #96118). Etymology. Latin flava (yellow), referring to the yellowish color of the species. Diagnosis. See the diagnoses of Maeota dorsalis and M. simoni. Differs from M. dichrura in the short spinnerets of the male and the narrower embolic spiral of the male palp. Description. Male (holotype, UBC-SEM AR00159). Carapace length 1.6; abdomen length 1.7. Chelicera: gray brown; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.17B-C): yellowish. Retrolateral sperm duct loop less than half of the bulb width; embolus relatively short and slightly coiled; proximal tegular lobe obvious; retrolateral tibial apophysis finger-like; ventral tibial bump present; palpal patella slightly swollen. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.1, II 3.0, III 3.4, IV 3.8. Color in alcohol (Fig. 7.17A): carapace dark brown to dark yellow brown, covered with brownish scales; abdomen gray brown with many brownish ! 391! speckles and a medial brownish stripe, posterior part of the stripe with a few unobvious yellow brown bands, ventral abdomen grayish; legs sandy yellow. Female. Unknown. Natural history. Specimen collected on low vegetations in synanthropic habitat. 7.4.6.3 Maeota simoni sp. nov. Figs 7.18, 7.26D Type material. Holotype: male, PANAMA: Panamá: Panama City, Parque Metropolitano, 8.99442° N, 79.54301° W, elev. 96 m, 29 September 2008, beating at ground level, coll. J. X. Zhang & G. B. Edwards, JXZ08#013. Paratype: 1 male, same data as holotype. Etymology. The specific epithet is a patronym in honor of Mr. E. Simon, who made great contribution to the systematics of jumping spiders. Diagnosis. Similar to Maeota flava in the color pattern, but differs in the narrower embolic spiral, the smaller embolic disc, the narrower palpal bulb, the shape of the retrolateral tibial apophysis and the absence of the ventral tibial bump of the male palp. Description. Male (holotype, UBC-SEM AR00160). Carapace length 1.2 (variation 1.2-1.3, n=2); abdomen length 1.2. Palp (Figs 7.18B-C): dark yellow brown. Spiral of embolus relatively narrow; proximal tegular lobe present; retrolateral tibial apophysis finger-like, abruptly narrowed at the tip; ventral tibial bump absent. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.5, II 2.2, III 2.7, IV 3.0. Color in alcohol (Fig. 7.18A): carapace dark brown, covered with white scales, eye area also with orange scales, with a narrow and short brownish stripe behind fovea; abdomen yellow brown to gray brown, with a wide medial sandy yellow stripe and many small brownish speckles; first pair of legs yellow brown, with dark markings, other legs pale yellow. Female. Unknown. ! 392! Natural history. Specimens were collected by beating foliage in wet forest. 7.4.7 Genus Soesilarishius Makhan, 2007 Small sized spiders mainly found in leaf litter. Male abdomen sometimes with a relatively sclerotized area on the dorsum. Chelicera with two promarginal teeth and one retromarginal tooth. Embolus of male palp usually not coiled; distal hematodocha sometimes reduced. Epigynal window usually absent; epigynum often with groove or transverse roof. Nine species have been reported from the South America (Makhan 2007; Ruiz 2011). An additional two species from Ecuador and Brazil are described here. 7.4.7.1 Soesilarishius micaceus sp. nov. Fig 7.19 Type material. Holotype: male, ECUADOR: Sucumbios: Reserva Faunistica Cuyabeno, Laguna Grande, Neotropic cabins; camp area & trail to Sendero La Hormiga, 22-27 April 1994, coll. W. Maddison, WPM#94-025 (QCAZ). Paratypes: 1 female, same data as holotype; 7 males, same data as holotype; 3 females, ECUADOR: Napo: Reserva Faunistica de Cuyabeno, Laguna Grande, PUCE field station, Sendero Campamento, trail through forest, 25-29 June 1988, coll. W. Maddison, WPM#88-007. Etymology. The specific epithet is derived from the Latin mica (crumb), referring to the small size of this species. Diagnosis. Differs from Soesilarishius amrishi Makhan, 2007 in the long embolus. Similar in the color pattern and markings to S. ruizi, but can be distinguished by the shape of the embolus and epigynum. Description. Male (holotype, UBC-SEM AR00161). Carapace length 1.0 (variation 0.9-1.1, n=8); abdomen length 0.7. Chelicera (Fig. 7.19E): yellow brown; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.19C-D): yellow brown. Embolus long and coiled, coming from prolateral side of bulb; retrolateral tibial apophysis short and finger-like. First tibia with three pairs of ventral macrosetae; first metatarsus with two pairs of ventral macrosetae. ! 393! Measurements of legs: I 1.7, II 1.4, III 2.0, IV 1.7. Color in alcohol (Fig. 7.19A): carapace dark brown to dark yellow brown; abdomen gray brown with light yellow spots and markings, anterior and median part with a relatively sclerotized area; legs brown to light yellow. Female (paratype, UBC-SEM AR00162). Carapace length 0.9 (variation: 0.9-1.1, n=4); abdomen length 1.0. Chelicera: promargin with two teeth and retromargin with one tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 1.7, II 1.4, III 1.8, IV 1.8. Epigynum (Fig. 7.19F): with a median transverse “roof”; opening to copulatory duct anteriorly. Vulva (Fig. 7.19G): copulatory duct long and convoluted; spermatheca small, not strongly swollen. Color in alcohol (Fig. 7.19B): similar to that of male. 7.4.7.2 Soesilarishius ruizi sp. nov. Fig 7.20 Type material. Holotype: male, BRAZIL: Amazonas: Manaus, Campus UFAM, synantropic, 3.098° S, 59.972° W, 22-23 May 2009, coll. G. Ruiz, T. Gasnier & B. Machado (IBSP, #INPA5033). Paratypes: 1 female, same data as in holotype (IBSP, #INPA5036). Etymology. The specific epithet is a patronym in honor of Dr. Gustavo Ruiz, who collected the specimens of this species and has provided much help in getting materials from Brazil for this study. Diagnosis. Differs from Soesilarishius micaceus by the shorter and wider embolus, the large and round depressions at the anterior part of the epigynum, the absence of a “roof” but the presence of a longitudinal groove on the epigynum, and the kidney-shaped spermatheca of the vulva. This species is also similar in the male palp shape to S. aurifrons (see Ruiz 2011), and in the color and marking patterns to S. minimus Ruiz, 2011. It can be distinguished from S. aurifrons by the wider embolus of the male palp; and from S. minimus by the longer and wider embolus of the male palp, the larger anterior depression and the presence of the groove on the epigynum, and the smaller spermatheca of the vulva. Description. Male (holotype, UBC-SEM AR00163). Carapace length 1.0; abdomen length 0.8. ! 394! Chelicera: dark brown; back surface with a depression; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.20C-D): gray brown. Embolus relatively short and wide; retrolateral tibial apophysis short and finger-like. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.2, II 1.7, III 2.3, IV 2.1. Color in alcohol (Fig. 7.20A): carapace dark, scattered with white scales; abdomen with a relatively sclerotized area, dorsum of abdomen dark with two pairs of white spots, anterior pair smaller and round, posterior pair larger; legs gray brown proximally and brownish to cream distally. Female (paratype, UBC-SEM AR00164). Carapace length 1.1; abdomen length 0.8. Chelicera: with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae; first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.1, II 1.6, III 2.2, IV 2.1. Epigynum (Fig. 7.20E): with two round depressions anteriorly and one elongate depression in the middle. Vulva (Fig. 7.20F): copulatory duct slightly coiled; spermatheca swollen and kidney-shaped. Color in alcohol (Fig. 7.20B): similar to that of male except paler in color with legs all cream and dorsal abdomen without sclerotized area. Natural history. Specimens were found on humid leaf litter in a recovering rainforest. 7.4.8 Genus Tylogonus Simon, 1902 Medium sized spiders usually found on foliage. Body form and color pattern more like dendryphantine. Most species greenish with iridescent scales. Male abdomen usually with white anterior margin and paired large white spots. Male chelicera modified, with front spur and concaved internal margin. Chelicera with two promarginal teeth and one retromarginal tooth. Embolus of male palp fixed and uncoiled. Epigynum without distinct window structure, but usually with small posterior depressions. Vulva anterior to the depressions. Nine species have been reported from the Central and South America (Platnick 2011). Two new species from Ecuador are described here. ! 395! 7.4.8.1 Tylogonus parvus sp. nov. Fig 7.21 Type material. Holotype: female, ECUADOR: Morona Santiago: km 36 from Limón towards Gualaceo, 3.0151° S, 78.5982° W, elev. 2500 m, cloud forest, 11 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, WPM#04-027 (QCAZ). Etymology. Latin parvus (small) and refers to the small depressions of the epigynum. Diagnosis. Differs from the female of Tylogonus viridimicans (see Galiano 1963) and T. yanayacu by the much smaller depressions of the epigynum. It also differs from T. yanayacu in the shape of the spermatheca. Description. Female (holotype, UBC-SEM AR00165). Carapace length 1.4; abdomen length 1.7. Chelicera: yellow brown; with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.8, II 2.5, III 2.8, IV 3.1. Epigynum (Fig. 7.21B): with two very small depressions posteriorly. Vulva (Fig. 7.21C): copulatory duct thin and short; spermatheca almost oval. Color in alcohol (Fig. 7.21A): carapace brownish, with iridescent green scales; abdomen brown with many iridescent green scales and an indistinct dark brown marking; legs pale yellow. Male. Unknown. 7.4.8.2 Tylogonus yanayacu sp. nov. Fig 7.22 Type material. Holotype: male, ECUADOR: Napo: Caucheras, Estación Yanayacu, 0.6049° S, 77.8886° W, elev. 2150 m, 20 July 2004, coll. Maddison, Agnarsson, Iturralde, Salazar, Avilés, WPM#04-052 (QCAZ). Paratypes: 1 female, same data as holotype; 1 male, same data as holotype. Etymology. The specific name refers to the type locality. ! 396! Diagnosis. Similar to Tylogonus auricapillus (see Galiano 1960), but differs in the shorter embolus, the shape of the embolic division and the retrolateral tibial apophysis of the male palp. This species differs from T. viridimicans (see Galiano 1963) by the shorter embolus, and the shape of the retrolateral tibial apophysis,of the male palp and the epigynal form. Description. Male (holotype, UBC-SEM AR00166). Carapace length 1.6 (variation 1.6-1.8, n=2); abdomen length 1.7. Chelicera (Figs 7.22E-F): pale yellow; front surface with a long spur; promargin with two teeth and retromargin with one tooth. Palp (Figs 7.22C-D): pale yellow. Proximal tegular lobe absent; embolic haemotodocha reduced; embolus short and dagger-like, with a wide ridged apophysis near its base; tetrolateral tibial apophysis branched with the ventral branch thick and further forked, and the dorsal one thin and finger-like. First tibia with nine ventral macrosetae, 4-5 aligned, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.4, II 3.2, III 3.5, IV 3.7. Color in alcohol (Fig. 7.22A): carapace brown, with a white stripe at each side of eye area composed of white setae extending obliquely to almost posterior end of carapace; abdomen brown with white anterior margin and two pairs of lateral white patches; legs pale yellow. The male paratype specimen paler in color than the male holotype. Female (paratype, UBC-SEM AR00167). Carapace length 1.5; abdomen length 2.2. Chelicera (Fig. 7.22H): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.2, II 2.8, III 3.2, IV 3.4. Epigynum (Fig. 7.22I): with two large depressions posteriorly. Vulva (Fig. 7.22J): copulatory duct thin and short; spermatheca almost kidney-shaped. Color in alcohol (Fig. 7.22B): paler than that of male, carapace yellowish brown, white markings absent; abdomen brownish, without white anterior margin or white patches. ! 397! Figure 7.1. Amphidraus complexus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 398! Figure 7.2. Belliena ecuadorica sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicerae, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. ! 399! Figure 7.3. Chapoda angusta sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female carapace, lateral view; F. female left chelicera, back view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-H, 0.2 mm. ! 400! Figure 7.4. Chapoda fortuna sp. nov. A. male paratype, dorsal view; B. male left palp, ventral view; C. male left palp (with patella and femur), retrolateral view. Scale bars: A, 0.5 mm; B-C, 0.2 mm. ! 401! Figure 7.5. Chapoda gitae sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-F, 0.1 mm. ! 402! Figure 7.6. Ecuadattus elongatus sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; D. male left palp, retrolateral view. Scale bars: A, 2.0 mm; B-C, 0.2 mm. ! 403! Figure 7.7. Ecuadattus napoensis sp. nov. A. male holotype, dorsal view; B. male left chelicera, back view; C. male left palp, ventral view; D. male left palp, retrolateral view. Scale bars: A, 1.0 mm; B-D, 0.2 mm. ! 404! Figure 7.8. Ecuadattus pichincha sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-E, 0.2 mm; F-G, 0.1 mm. ! 405! Figure 7.9. Ecuadattus typicus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. female left chelicera, back view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-F, 0.2 mm; G-H, 0.1 mm. ! 406! Figure 7.10. Ilargus foliosus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.2 mm. ! 407! Figure 7.11. Ilargus galianoae sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.2 mm. ! 408! Figure 7.12. Ilargus macrocornis sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view; D. male left chelicera, internal view. Scale bars: A, 1.0 mm; B-D, 0.2 mm. ! 409! Figure 7.13. Ilargus moronatigus sp. nov. A. female holotype, dorsal view; B. female left chelicera, back view; C. epigynum, ventral view; D. cleared epigynum, dorsal view. Scale bars: A, 0.5 mm; B, 0.2 mm; C-D, 0.1 mm. ! 410! Figure 7.14. Ilargus pilleolus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female carapace, dorsal view; F. female carapace, lateral view; G. female left chelicera, back view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-I, 0.2 mm. ! 411! Figure 7.15. Ilargus serratus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female carapace, dorsal view; F. female left chelicera, back view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A, 0.5 mm; B, 1.0 mm; C-H, 0.2 mm. ! 412! Figure 7.16. Maeota dorsalis sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view; D, male left palp, prolateral view. Scale bars: A, 0.5 mm; B-D, 0.2 mm. ! 413! Figure 7.17. Maeota flava sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view. Scale bars: A, 0.5 mm; B-C, 0.2 mm. ! 414! Figure 7.18. Maeota simoni sp. nov. A. male paratype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view. Scale bars: A, 0.5 mm; B-C, 0.2 mm. ! 415! Figure 7.19. Soesilarishius micaceus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicerae, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.1 mm. ! 416! Figure 7.20. Soesilarishius ruizi sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-F, 0.1 mm. ! 417! Figure 7.21. Tylogonus parvus sp. nov. A. female holotype, dorsal view; B. epigynum, ventral view; C. cleared epigynum, dorsal view. Scale bars: A, 0.5 mm; B-C, 0.1 mm. ! 418! Figure 7.22. Tylogonus yanayacu sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male right chelicera, front view; F. male right chelicera, back view; G. female carapace; dorsal view; H. female right chelicera, back view; I. epigynum, ventral view; J. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-J, 0.2 mm. ! 419! Figure 7.23. Living spider photos of Chapoda gitae sp. nov. A-B. male holotype; C-D. female (subadult). ! 420! Figure 7.24. Living spider photos of Ecuadattus pichincha sp. nov. A-B. male; C-D. female. ! 421! Figure 7.25. Living spider photos of Ecuadattus typicus sp. nov. A-D. males; E-F. females. ! 422! Figure 7.26. Living spider photos. A-B. male of Ilargus pilleolus sp. nov.; C. male of Ilargus macrocornis sp. nov.; D. male of Maeota simoni sp. nov.. ! ! 423! 8 New euophryine jumping spiders from Southeast Asia and Africa (Araneae: Salticidae: Euophryinae) 1 8.1 Synopsis Sixteen new species and four new genera of euophryine jumping spiders from the Old World (China, Malaysia and South Africa) are described. The new genera are Chinophrys (type species C. pengi sp. nov.), Foliabitus (type species F. longzhou sp. nov.), Parabathippus (type species Bathippus shelfordi Peckham & Peckham; P. cuspidatus sp. nov., P. kiabau sp. nov., P. magnus sp. nov.) and Parvattus (type species P. zhui sp. nov.). The other new species belong to the genera Colyttus (C. robustus sp. nov.), Emathis (E. gombak sp. nov.), Lagnus (L. edwardsi sp. nov.), Laufeia (L. concava sp. nov. and L. eximia sp. nov.), Thiania (T. latibola sp. nov. and T. tenuis sp. nov.) and Thyenula (T. laxa sp. nov., T. nelshoogte sp. nov. and T. wesolowskae sp. nov.). Following species from the Southeast Asia once described as Bathippus are transferred to Parabathippus: Bathippus birmanicus, B. digitalis, B. macilentus, B. petrae, B. rectus, B. sedatus and B. shelfordi. Laufeia liujiapingensis Yang & Tang is transferred to Chinophrys. Laufeia scutigerus Zabka is transferred to Foliabitus. Diagnostic illustrations are provided for all of the new species. Photographs of living spiders are also provided for some new species. 8.2 Introduction The subfamily Euophryinae is one of the most diverse groups in the Salticidae, with about 900 described species and the majority found in the tropics of both the Old and New World (Prószy!ski 1976; Maddison & Hedin 2003; Platnick 2011). In the Old World, euophryine jumping spiders are most diverse in the Australasian region and the Eurasian region, but less diverse in Africa (Platnick 2011; Prószy!ski & Deeleman-Reinhold 2012). In this Chapter, I report four new genera: Chinophrys (one new species), Foliabitus (one new species), Parabathippus (three new species) and Parvattus (one new species). An additional ten new species are described and included in the genera Colyttus (one species), Emathis (one species), Lagnus (one species), Laufeia (two species), Thiania (two species) and Thyenula (three !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! \"!A version of this chapter has been submitted for publication: Zhang, J. and Maddison, W. P., New euophryine jumping spiders from Southeast Asia and Africa (Araneae: Salticidae: Euophryinae). Names of new taxa are not finalized for nomenclature. ! ! 424! species). I transfer seven species previously described as Bathippus from the Southeast Asia (Bathippus birmanicus, B. digitalis, B. macilentus, B. petrae, B. rectus, B. sedatus and B. shelfordi) to the newly erected genus Parabathippus. I also transfer Laufeia liujiapingensis Yang & Tang to Chinophrys, and Laufeia scutigerus Zabka to Foliabitus based on their genitalic structures. Species described here were chosen in order to give names for taxa included in the molecular phylogenetic study on the subfamily Euophryinae (see Chapter 2). 8.3 Material and methods Photographs of living specimens were taken with a Pentax Optio 33WR digital camera. For macro capability, a small lens was glued to it. Photographs of preserved specimens were taken under a Leica MZ16 dissecting microscope with Leica Application Suite version 3.1.0. Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Terminology is standard for Araneae. All measurements are given in millimeters. Descriptions of color pattern are based on the alcohol-preserved specimens. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; AME, anterior median eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the \"small eyes\"). Depository of type specimens are as following: Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM); Florida State Collection of Arthropods, USA (FSCA); California Academy of Sciences, USA (CAS). Specimens are deposited in UBC-SEM unless it is indicated otherwise. 8.4 Taxonomy 8.4.1 Genus Chinophrys new genus Type species: Chinophrys pengi Zhang & Maddison, sp. nov. Etymology. “Chin” in the generic name is derived of “China” where the type species has been ! 425! reported; “ophrys” is taken from the jumping spider genus “Euophrys”, which has similar color pattern and body form as this genus; feminine in gender. Diagnosis. Medium sized spiders. Chelicera with multiple (four to six) promarginal teeth and one fissident retromarginal tooth of five to six cusps. Embolus of male palp long and coiled; tegulum with a proximal lobe over tibia. Epigynal window present with a median septum. Copulatory duct not highly convoluted; spermatheca swollen and divided into head and body. Chinophrys is similar in the body form and color pattern to Euophrys C. L. Koch, but differs in the genitalic structure and the teeth pattern of the chelicera. Remarks. Based on the original description, Chinophrys liujiapingensis (Yang & Tang, 1997) is transferred to this genus from Laufeia Simon because of its similarities in the cheliceral teeth pattern and the palpal structure with the type species (NEW COMBINATION). 8.4.1.1 Chinophrys pengi sp. nov. Fig 8.1 Type material. Holotype: male, CHINA: Guangxi: Fangchenggang City, around Banba Village, 21.683° N, 107.649° E, elev. 159 m, 20 May 2006, coll. J. Zhang, M. S. Zhu, W. G. Lian & H. Q. Ma, JXZ06#011. Paratypes: 1 female, CHINA: Guangxi: Pingxiang City, Longzhou County, Daqingshan Forestry Center, 22.299° N, 106.700° E, elev. 806-989 m, 15 May 2006, coll. J. Zhang, M. S. Zhu, W. G. Lian & H. Q. Ma, JXZ06#005; 1 female, same data as previous. Etymology. The specific epithet is a patronym in honor of Dr. Xianjin Peng, who has made great contributions in studying the jumping spider fauna of China. Diagnosis. Resembles Chinophrys liujiapingensis (Yang & Tang, 1997) in the body form and palpal shape, but differs in the presence of six promarginal teeth on the chelicera (four in C. liujiapingensis), the shape of the proximal tegular lobe and the longer retrolateral tibial apophysis of the male palp. Description. Male (holotype, UBC-SEM AR00168). Carapace length 2.5; abdomen length 2.4. Chelicera (Fig. 8.1E): yellow brown; promargin with six small teeth, retromargin with a six- ! 426! cusped fissident tooth. Palp (Figs 8.1C-D): gray brown. Embolus long and coiled for about a circle; palpal tibia with a long finger-like retrolateral apophysis and a round ventral bump. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 5.1, II 4.5, III 5.1, IV 5.2. Color in alcohol (Fig. 8.1A): carapace dark brown, with some white scales; abdomen dark gray brown, with light yellow dots and chevron-like markings, anterior and median part with a relatively sclerotized area; legs dark brown. Female (paratype, UBC-SEM AR00169). Carapace length 2.4 (variation: 2.4-2.5, n=2); abdomen length 2.9. Chelicera: with four small teeth on promargin and one four-cusped tooth on retromargin. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.7, II 3.5, III 4.5, IV 4.7. Epigynum (Fig. 8.1H): median septum narrow with posterior end wider; with coiled grooves within window. Vulva (Fig. 8.1I): copulatory duct long and thin; spermatheca with a spherical head region and a kidney-shaped body region. Color in alcohol (Fig. 8.1B): similar to that of male, but legs paler in color with brown annuli. Natural history. Specimens were found by beating foliage in forest and around the edge of forest. 8.4.2 Genus Colyttus Thorell, 1891 Medium sized, bright and colorful spiders. Chelicera with two promarginal teeth; retromargin with one unident or bicuspid tooth. Embolus of male palp usually wide and only slightly coiled; embolic disc large; proximal tegular lobe present. Epigynal window present with a median septum. Copulatory duct short; vulva with a pair of secondary spermathecae in addition to the primary spermathecae. Two species have been recorded from Southeast Asia (Prószy!ski 1984, 1987; Zabka 1985). A new species from Malaysia is described here. ! 427! 8.4.2.1 Colyttus robustus sp. nov. Figs 8.2, 8.17E Type material. Holotype: male, MALAYSIA: Pahang: Genting Highlands, 3.400° N, 101.777° E, elev. 1000 m, 15-16 May 2005, coll. W. Maddison, D. Li, I. Agnarsson, J. Zhang, WPM#05- 023. Etymology. Latin robustus (robust), referring to the vigorous body of the species. Diagnosis. Differs from Colyttus bilineatus Thorell (Prószy!ski 1987) in the absence of dark stripes on the abdomen, the wider male palpal bulb, the larger embolic disc, the shape of the embolus and the wider proximal tegular lobe of the male palp. Description. Male (holotype, UBC-SEM AR00170). Carapace length 3.2; abdomen length 3.8. Chelicera: promargin with two teeth, retromargin with one tooth. Palp (Figs 8.2B-C): cream to light yellow brown. Proximal tegular lobe obvious; embolus short and relatively thick; retrolateral tibial apophysis finger-like. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of macrosetae. Ventral patella, tibia and metatarsus of leg I with fringes. Measurements of legs: I 7.3, II 6.3, III 7.1, IV 6.9. Color in alcohol (Fig. 8.2A): eye area orange, other part light orange, with a wide cream stripe behind fovea; abdomen light brown, with a large sandy yellow foliage-like marking medially and a brown spot on each side of the foliage-like marking near the center; legs cream to dark brown with first pair darker in color. Female. Unknown. 8.4.3 Genus Emathis Simon, 1899 Medium sized spiders. Chelicera with a fissident retromarginal tooth of multiple cusps. Embolus of male palp coiled for at least half a circle; tegulum lacking proximal lobe. Window of epigynum without obvious median septum. Copulatory duct sometimes convoluted (Prószy!ski 1984; Barrion & Litsinger 1995). Five species have been reported from Southeast Asia (Prószy!ski & Deeleman-Reinhold 2012). ! 428! The species previously described as Emathis from the Caribbean Islands have been transferred to the genus Petemathis (Prószy!ski & Deeleman-Reinhold 2012). A new species from Malaysia is described here. 8.4.3.1 Emathis gombak sp. nov. Figs 8.3, 8.17A-B Type material. Holotype: male, MALAYSIA: Selangor: Ulu Gombak Field Station, 3.325° N , 101.753° E, elev. 250 m, 16-19 May 2005, coll. W. Maddison, D. Li, I. Agnarsson, J. Zhang, WPM#05-026. Paratype: female, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Similar in the body form and cheliceral teeth pattern to other Emathis species, but differs in the short and not strongly convoluted copulatory duct. This species resembles Emathis coprea (see Prószy!ski 1984) in the short embolus of the male palp, but can be distinguished by the narrower palpal bulb and the much narrower embolus. Description. Male (holotype, UBC-SEM AR00171). Carapace length 2.7; abdomen length 2.7. Chelicera (Fig. 8.3D): dark yellow brown; with two promarginal teeth and one retromarginal tooth of five cusps. Palp (Fig. 8.3C): yellow brown. Embolus relatively short and coiled for about half a circle; retrolateral tibial apophysis long and finger-like. First tibia with three pairs of ventral macrosetae, first metatarsus with four pairs of ventral macrosetae. Measurements of legs: I 8.2, II 5.6, III 6.6, IV 5.3. Color in alcohol (Fig. 8.3A): carapace orange in eye area and yellowish brown behind fovea, posterior part dark yellow brown; abdomen sandy yellow, with indistinct grayish brown markings; first two pairs of legs yellow brown, last two pairs of legs cream to yellow brown. Female (paratype, UBC-SEM AR00172). Carapace length 2.3; abdomen length 2.3. Chelicera: with three promarginal teeth and one retromarginal tooth of five cusps. First tibia with three pairs of ventral macrosetae, first metatarsus with four pairs of ventral macrosetae. Measurements of legs: I 4.9, II 4.3, III 5.3, IV 4.5. Epigynum (Fig. 8.3E): window at anterior part of epigynal plate; median septum absent; opening to copulatory duct at posterior end of ! 429! window. Vulva (Fig. 8.3F): copulatory duct short and not convoluted; spermatheca almost spherical. Color in alcohol (Fig. 8.3B): similar to that of male except legs paler in color. 8.4.4 Genus Foliabitus new genus Type species: Foliabitus longzhou Zhang & Maddison, sp. nov. Etymology. The generic name is from the combination of the Latin folia (leaf) and habito (dweller), referring to its habitat; masculine in gender. Diagnosis. Medium sized spiders. Chelicera with two promarginal teeth and one retromarginal tooth. Abdomen with light colored stripes on dorsum. Embolus of male palp long and coiled for less a circle; proximal tegular lobe relatively small; retrolateral tibial apophysis finger-like. Epigynal window present, with a median septum. Vulva with only one pair of spermathecae. Similar in the body form to Donoessus Simon and Colyttus Thorell, but differs from them in the presence of light colored stripes on the abdomen; the longer and more coiled embolus, the smaller embolic disc, the indistinct proximal tegular lobe of the male palp; the presence of grooves within the epigynal window. Remarks. Foliabitus scutigerus (Zabka, 1985) is transferred to this genus from Laufeia Simon because of its similar morphology with the type species (NEW COMBINATION). 8.4.4.1 Foliabitus longzhou sp. nov. Fig 8.4 Type material. Holotype: male, CHINA: Guangxi: Pingxiang City, Longzhou County, Daqingshan Forestry Center, 22.299° N, 106.700° E, elev. 806-989 m, 15 May 2006, coll. J. Zhang, M. S. Zhu, W. G. Lian & H. Q. Ma, JXZ06#005. Paratypes: 1 female, same data as holotype; 2 females, same data as holotype. Etymology. A noun in apposition taken from the type locality. Diagnosis. Differs from Foliabitus scutigerus (Zabka, 1985) in the wider palpal bulb, the ! 430! thicker embolus and the narrower retrolateral sperm duct loop of the male palp. Description. Male (holotype, UBC-SEM AR00173). Carapace length 2.3; abdomen length 2.8. Chelicera (Fig. 8.4E): red brown; promargin with two teeth and retromargin with one tooth. Palp (Figs 8.4C-D): grayish brown. Proximal tegular lobe not distinct; embolus long and coiled for almost a circle; retrolateral tibial apophysis curved with a small cusp near distal end. First tibia with four pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 4.4, II 4.3, III 5.0, IV 5.0. Color in alcohol (Fig. 8.4A): carapace dark brown, relatively smooth; abdomen also dark brown, with an oval slightly sclerotized area, lateral margins with light yellow stripes; first pair of legs reddish brown, other legs yellowish with grayish brown markings. Female (paratype, UBC-SEM AR00174). Carapace length 2.0 (variation: 2.0-2.2, n=3); abdomen length 3.8. First tibia with four pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 4.9, II 4.3, III 4.6, IV 4.7. Epigynum (Fig. 8.4H): with curved grooves within window; median septum narrow and almost reaching anterior edge of window. Vulva (Fig. 8.4I): copulatory duct well sclerotized; spermatheca long and oval. Color in alcohol (Fig. 8.4B): similar to that of male, but abdomen with a medial longitudinal light stripe in addition to the two lateral stripes; first pair of legs yellowish to light orange and other legs without grayish brown markings. Natural history. Specimens were collected by beating foliage in forest. 8.4.5 Genus Lagnus L. Koch, 1879 Medium sized spiders. Chelicera with two promarginal teeth and one fissident retromarginal tooth of multiple cusps. Legs, especially in males, extremely elongate and spiny, with first tibia having more than seven pairs of ventral macrosetae and first metatarsus having more than five pairs of ventral macrosetae. Male palp elongate; embolus slender and coiled for about half a circle; proximal tegular lobe and ventral tibial bump absent; retrolateral sperm duct loop wide. Epigynal window present with a median septum. Vulva with relatively short copulatory duct and only one pair of spermathecae. Lagnus can be easily distinguished from other euophryine genera by the elongate, slender and spiny legs especially in male and the elongate male plap. ! 431! Two species have been recorded from Fiji (Wanless 1988; Patoleta 2008). Another new species from Philippines is described here. 8.4.5.1 Lagnus edwardsi sp. nov. Fig 8.5 Type material. Holotype: male, PHILIPPINES: Luzon, Laguna Prov., Mt. Makiling, December 1993 (FSCA). Paratypes: 1 female, PHILIPPINES: Luzon, Laguna Prov., Mt. Makiling, December 1993; 1 male, PHILIPPINES: Luzon, Laguna Prov., Mt. Makiling, March 1993; 1 male, PHILIPPINES: Luzon, Laguna Prov., Mt. Makiling For. Res., 17 December 1993, coll. G. B. Edwards et al. (FSCA). Etymology. The specific epithet is a patronym in honor of Dr. G. B. Edwards, who collected and provided specimens of the species for this study. Diagnosis. Male of this species can be distinguished from Lagnus longimanus (see Wanless 1988) by the much shorter palpal femur and patella, the longer retrolateral tibial apophysis, and the absence of a crescent-like light colored marking at the posterior end of the carapace. Female of this species differs from L. monteithorum Patoleta, 2008 by the shape of the epigynal window and the oval spermatheca. Description. Male (holotype, UBC-SEM AR00175). Carapace length 3.2; abdomen length 3.9. Chelicera (Fig. 8.5D): red brown; with two promarginal teeth and one retromarginal tooth of 5-6 cusps; back surface with a big depression; fang with a few denticles at upper surface. Palp (Fig. 8.5C): femur and patella yellow brown, tibia and cymbium sandy yellow. Embolus long and coiled for about half a circle; retrolateral sperm duct loop wide occupying more than three quarters of bulb width; cymbium with one prolateral and one relateral macrosetae; retrolateral tibial apophysis long finger-like, reaching the proximal end of embolic disc. First pair of legs slender, with nine pairs of ventral macrosetae on tibia, with six ventral macrosetae on the prolateral side and eight ventral macrosetae on the retrolateral side of metatarsus. Measurements of legs: I 13.9, II 10.3, III 10.8, IV 11.6. Color in alcohol (Fig. 8.5A): carapace dark yellow brown with sandy yellow lateral margins and a middle stripe behind fovea; abdomen gray brown with a light yellow medial marking; venter gray brown; legs sandy yellow to dark yellow brown. ! 432! Female (paratype, UBC-SEM AR00176). Carapace length 2.7 (variation 2.9-3.2, n=3); abdomen length 3.9. Teeth pattern on chelicera similar to that of male. Tibia of first leg with eight pairs of ventral macrosetae, metatarsus of first leg with six pairs of ventral macrosetae. Measurements of legs: I 6.7, II 5.8, III 6.8, IV 7.4. Epigynum (Fig. 8.5E): window wide with opening to copulatory duct close to its posterior end; median septum relatively wide. Vulva (Fig. 8.5F): copulatory duct short with accessory gland; spermatheca oval. Color in alcohol (Fig. 8.5B): similar to that of male. Natural history. Specimens were found from large buttressed trees on the sides of the large buttresses, where they had lots of flat surface to move on rapidly. 8.4.6 Genus Laufeia Simon, 1889 Small spiders. Body brown or dark brown, without distinctive large markings or stripes; carapace and abdomen covered with light colored scales. Male usually with a slightly sclerotized area on the dorsal abdomen. Chelicera usually with a bicuspid tooth on retromargin. Male palp with rrelatively wide retrolateral sperm duct loop; embolus long or short; embolic disc large or relatively small; proximal tegular lobe present. Epigynal window sometimes indistinct. Spermatheca of various shapes. Laufeia, Orcevia and Junxattus share similar somatic characters such as body form, color and cheliceral teeth pattern, but show considerate difference in genital structure (Prószy!ski & Deeleman-Reinhold 2012). Unpublished molecular data suggest they fall into one clade. Two species previously described as Laufeia are transferred to other genera in this paper: Laufeia liujiapingensis Yang & Tang, 1997 to Chinophrys; Laufeia scutigera Zabka, 1985 to Foliabitus. Two new species from China and Malaysia are described here. 8.4.6.1 Laufeia concava sp. nov. Figs 8.6, 8.17C-D Type material. Holotype: male, MALAYSIA: Pahang: Gunung Brinchang, Trail 1 to Brinchang, 4.515° N , 101.383° E, elev.1950-2030 m, 20 May 2005, coll. W. Maddison, D. Li, ! 433! I. Agnarsson & J. Zhang, WPM#05-029. Paratypes: 1 female, same data as holotype; 4 females and 3 males, same data as holotype; 3 females, MALAYSIA: Pahang: Gunung Brinchang, Trail 1 to Brinchang, 4.511° N, 101.385° E, elev. 1800-1950 m, 20 May 2005, coll. W. Maddison, D. Li, I. Agnarsson & J. Zhang, WPM#05-030; 1 female, MALAYSIA: Pahang: Gunung Brinchang, near peak, 4.525° N, 101.383° E, elev. 1980 m, 23 May 2005, coll. W. Maddison, WPM#05-036; 2 males, MALAYSIA: Pahang: Cameron Highlands, Arcadia, Trail 3, 4.482° N, 101.388° E, elev. 1550 m, 21-23 May 2005, coll. W. Maddison, D. Li, I. Agnarsson & J. Zhang, WPM#05-033. Etymology. Latin concava (concave), referring to the notch at the tip of the retrolateral tibial apophysis of the male palp. Diagnosis. Similar in the appearance to Laufeia aenea Simon, but differs in the thicker retrolateral tibial apophysis, the longer and thinner embolus of the male palp; the wider epigynal median septum and the presence of rim leading to the opening of copulatory duct of the epigynum. Description. Male (holotype, UBC-SEM AR00177). Carapace length 1.3 (variation 1.1-1.3, n=6); abdomen length 1.2. Chelicera (Fig. 8.6E): dark brown; front surface with a longitudinal ridge; promargin and retromargin each with one bicuspid tooth. Palp (Figs 8.6C-D): yellowish brown. Proximal tegular lobe present. Embolus relatively short and coiled for less a circle; embolic disc small, with a process along the embolus; retrolateral sperm duct loop occupying almost half of bulb width; retrolateral tibial apophysis short and wide, not reaching the distal end of tegulum, with a notch at distal end. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.6, II 2.0, III 2.1, IV 2.3. Color in alcohol (Fig. 8.6A): carapace dark red brown, with many white sclaes; dorsal abdomen brown or gray brown with sandy yellow markings, with a relatively sclerotized area; ventral abdomen brownish; first two pairs of legs dark yellow brown, only coxa, trochanter and tarsus sandy yellow, last two pairs of legs sandy yellow, with brown annuli on tibia and metatarsus. Female (paratype, UBC-SEM AR00178). Carapace length 1.2 (variation: 1.1-1.2, n=9); abdomen length 1.3. Chelicera (Fig. 8.6G): promargin and retromargin each with one bicuspid ! 434! tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 1.9, II 1.8, III 1.8, IV 2.3. Epigynum (Fig. 8.6H): window small with relatively wide median septum; opening to copulatory duct anterior lateral to window with rim. Vulva (Fig. 8.6I): copulatory duct wide, spermatheca small and oval. Color in alcohol (Fig. 8.6B): similar to that of male except legs paler in color. 8.4.6.2 Laufeia eximia sp. nov. Fig 8.7 Type material. Holotype: male, CHINA: Guangxi: Tianlin County, Langping Village, from Aokou to Matishi, 24.467° N, 106.367° E, elev. 1260-1454 m, 26 May 2006, coll. J. Zhang, M. S. Zhu, W. G. Lian, H. Q. Ma, JXZ06#014. Paratype: 1 female, same data as holotype. Etymology. Latin eximia (special), referring to the extraordinary male palpal structure of the species. Diagnosis. Differs from other Laufeia species by the unique genital structures: the long and thick embolus, the large process on the embolic disc, and the large retrolateral extension of the cymbium of the male palp; the spermatheca anterior to the opening of copulatory duct. Description. Male (holotype, UBC-SEM AR00179). Carapace length 1.3; abdomen length 1.2. Chelicera (Fig. 8.7H): dark brown; promargin with two teeth, retromargin with one bicuspid tooth. Palp (Figs 8.7C-E): reddish brown. Cymbium with a large retrolateral extension; proximal tegular lobe present; embolus relatively wide and coiled for less a circle; embolic disc with a large branched process; retrolateral tibial apophysis long and pointed at the tip. First tibia with one ventral macroseta on prolateral side, two ventral macrosetae on retrolateral side; first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.2, II 1.6, III 1.7, IV 2.1. Color in alcohol (Fig. 8.7A): body dark, with some white scales on carapace; abdomen with a pair of sandy yellow markings laterally; legs brown to sandy yellow. Female (paratype, UBC-SEM AR00180). Carapace length 1.1; abdomen length 1.1. Chelicera: with two promarginal teeth and one bicuspid retromarginal tooth. First tibia with one ventral macroseta on prolateral side, two ventral macrosetae on retrolateral side; first metatarsus with ! 435! two pairs of ventral macrosetae. Measurements of legs: I 1.5, II 1.4, III 1.7, IV 1.7. Epigynum (Fig. 8.7I): window relatively small, with a wide median septum. Vulva (Fig. 8.7J): copulatory duct thin, with accessory gland near spermatheca; spermatheca oval. Color in alcohol (Fig. 8.7B): similar to that of male. Natural history. Specimens were collected by beating dry leaves at the edge of a forest. 8.4.7 Genus Parabathippus new genus Type species. Bathippus shelfordi Peckham & Peckham, 1907. Etymology. The generic name is reminiscent of the name Bathippus; masculine in gender. Diagnosis. Medium sized spiders. Female chelicera with two promarginal teeth and one unident retromarginal tooth. Male chelicera usually elongate. Tegulum of male palp without proximal lobe; retrolateral sperm duct loop usually wide occupying more than half of bulb width; embolus slender and usually coiled for more than a circle; ventral tibial bump absent. Epigynal window large with a median septum. Spermatheca swollen and distinctive from copulatory duct. Parabathippus resembles Bathippus Thorell in the body form, color patten and the elongate male chelicera, but differs in the long and coiled embolus with the plane of spiral parallel to the longitudinal axis of the palpal bulb (the plane of the embolic spiral is usually perpendicular to the longitudinal axis of the palpal bulb in Bathippus), the oval embolic disc of the male palp; and the longer and usually less sclerotized copulatory duct. Remarks. The Southeast Asian Bathippus species have different genitalic structures from the species from Papua New Guinea, Pacific Islands and Australia including the type of Bathippus, B. macrognathus (Thorell). The molecular data also indicate that they fall into distinct clades (see Chapter 2). Thus the new genus namely Parabathippus is described here and the following species are transferred to Parabathippus from Bathippus: Parabathippus birmanicus (Thorell, 1895) (New Combination) Parabathippus digitalis (Zhang, Song & Li, 2003) (New Combination) Parabathippus macilentus (Thorell, 1890) (New Combination) Parabathippus petrae (Prószy!ski & Deeleman-Reinhold, 2012) (New Combination) Parabathippus rectus (Zhang, Song & Li, 2003) (New Combination) ! 436! Parabathippus sedatus (Peckham & Peckham, 1907) (New Combination) Parabathippus shelfordi (Peckham & Peckham, 1907) (New Combination) 8.4.7.1 Parabathippus cuspidatus sp. nov. Figs 8.8, 8.18A-C Type material. Holotype: male, MALAYSIA: Pahang: Tanah Rata, Jungle Trail 9 from Robinson Falls, 4.46° N, 101.40° E, elev.1200-1500 m, 21-22 May 2005, coll. W. Maddison, D. Li, I. Agnarsson, J. Zhang, WPM#05-035. Paratypes: 2 males, same data as holotype; 1 male, same data as holotype; 1 female, MALAYSIA: Pahang: Genting Highlands, 3.400° N, 101.777° E, elev. 1000 m, 15-16 May 2005, coll. W. Maddison, D. Li, I. Agnarsson, J. Zhang, WPM#05- 023; 4 females, same data as previous; 1 male, same data as previous. Etymology. Latin cuspidatus (having a cusp), referring to the small lateral cusp on the front surface of the male chelicera. Diagnosis. Male can be distinguished from other Parabathippus species by the cheliceral shape and dentition, and the palpal shape. Female is similar in the epigynal shape to Parabathippus shelfordi, but differs in the absence of dark patches on the dorsal abdomen; the wider window of the epigynum, the shorter copulatory duct and the more round spermatheca. Description. Male (holotype, UBC-SEM AR00181). Carapace length 3.2 (variation: 2.8-3.2, n=5); abdomen length 3.3. Chelicera (Figs 8.8E-F): dark red brown; elongate; promargin with three teeth, retromargin with one tooth and one mound (some specimens without this mound); front surface with a cusp laterally. Palp (Figs 8.8C-D): yellow brown to brown. Embolic disc almost round; embolus coiled for about a circle; retrolateral tibial apophysis finger-like. First tibia with three pairs of ventral macrosetae, first metatarsus with five ventral macrosetae (3-2 aligned). First and second legs with ventral fringes. Measurements of legs: I 7.4, II 6.3, III 8.2, IV 6.6. Color in alcohol (Fig. 8.8A): carapace dark red brown, posterior eyes with dark surroundings, with a short orange stripe behind eye area; abdomen gray brown without distinct marking; first two pairs of legs red brown, last two pairs yellow brown. Some male specimens lighter in color than holotype. ! 437! Female (paratype, UBC-SEM AR00182). Carapace length 2.6 (variation: 2.6-2.9, n=5); abdomen length 3.3. First tibia with three pairs of ventral macrosetae, first metatarsus with five ventral macrosetae (3-2 aligned). Measurements of legs: I 5.4, II 4.9, III 6.9, IV 5.7. Epigynum (Fig. 8.8G): median septum continuous with anterior margin of window. Vulva (Fig. 8.8H): copulatory duct partly membranous, anterior margin of copulatory duct loop far in front of the anterior margin of window, with accessory gland on the sclerotized part of the copulatory duct; spermatheca oval. Color in alcohol (Fig. 8.8B): carapace light yellow, eye area orange, with two brownish stripes behind eye area; abdomen grayish brown, with a distinct yellowish leaf-like marking medially; legs light orange, with brownish markings. 8.4.7.2 Parabathippus kiabau sp. nov. Fig 8.9 Type material. Holotype: male, MALAYSIA: Sabah: Village of Kiabau, Central Sabah, 5.832° N, 117.225° E, coll. K. Ober, 23 November 2000, #00.437. Etymology. A noun in apposition taken from the type locality. Diagnosis. Similar in the male palpal shape to Parabathippus magnus and P. macilentus Thorell (Prószy!ski 1984) in the male palpal shape. Differs from P. magnus in the markings on the dorsal abdomen, the male cheliceral dentition, and the narrower embolus of the male palp; and from P. macilentus in the wider embolic spiral and the larger embolic disc. Description. Male (holotype, UBC-SEM AR00183). Carapace length 2.2; abdomen length 2.1. Chelicera (Figs 8.9D-E): slightly elongate; promargin and retromargin each with three teeth. Palp (Figs 8.9B-C): yellow brown. Embolic disc large and round; embolus coiled for one and a half circle; retrolateral tibial apophysis long and finger-like. First tibia with seven ventral macrosetae (3-4 aligned), first metatarsus with five ventral macrosetae (3-2 aligned). Measurements of legs: I 5.3, II 4.3, III 6.0, IV 4.7. Color in alcohol (Fig. 8.9A): carapace dark red brown, with a tri-forked orange marking behind posterior lateral eyes; abdomen gray brown with yellowish irregular markings; first two pairs of legs red brown, last two pairs light orange with brownish annuli. ! 438! Female. Unknown. 8.4.7.3 Parabathippus magnus sp. nov. Figs 8.10, 8.18D-F Type material. Holotype: male, MALAYSIA: Pahang: Tanah Rata, Jungle Trail 9 from Robinson Falls, 4.46° N, 101.40° E, elev.1200-1500 m, 21-22 May 2005, coll. W. Maddison, D. Li, I. Agnarsson, J. Zhang, WPM#05-035. Paratypes: 1 female, same data as holotype; 2 males and 3 females, same data as holotype; 2 males, same data as holotype; 1 female, same data as holotype. Etymology. Latin magnus (large), referring to the large second tooth-like protrusion on the promargin of the male chelicera. Diagnosis. Similar in the body form and markings to Parabathippus macilentus (see Prószy!ski 1984), but differs in the thicker embolus of the male palp. This species differs from P. cuspidatus in the presence of large tooth-like protrusions near the fang base of the promargin, the absence of a lateral cusp on the front surface of the male chelicera; the narrower epigynal window, the presence of transverse ridges within window, the wider median septum; and the irregular-shaped spermatheca. Description. Male (holotype, UBC-SEM AR00184). Carapace length 3.2 (variation: 2.7-3.5, n=5); abdomen length 3.6. Chelicera (Figs 8.10E-F): yellow brown; elongate; promargin with four teeth with the second one from the fang base large, retromargin with two teeth. Palp (Figs 8.10C-D): yellow brown. Embolus thick and coiled for about one and a half circle; retrolateral sperm duct loop occupying about two thirds of bulb width; retrolateral tibial apophysis finger- like with a few cusps from ventral view. First tibia with three pairs of ventral macrosetae, first metatarsus with five ventral macrosetae (3-2 aligned). First leg with ventral fringes on tibia and patella. Measurements of legs: I 9.4, II 7.3, III 9.9, IV 8.1. Color in alcohol (Fig. 8.10A): carapace red brown, posterior eyes with dark surroundings; abdomen brown with indistinct yellowish speckles and a sandy yellow leaf-like marking medially; legs red brown to orange. Female (paratype, UBC-SEM AR00185). Carapace length 2.7 (variation 2.7-3.3, n=5); ! 439! abdomen length 3.2. Chelicera (Fig. 8.10G): with two promarginal teeth and one retromarginal tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with five ventral macrosetae (3-2 aligned). Measurements of legs: I 6.4, II 5.7, III 9.1, IV 6.6. Epigynum (Fig. 8.10H): median septum of window wide and short, not reaching anterior margin of the window; with transverse ridges within window. Vulva (Fig. 8.10I): most of copulatory duct membranous, only a short region before spermatheca well sclerotized; spermatheca irregular in shape. Color in alcohol (Fig. 8.10B): carapace light orange, eye area brownish, with two brownish stripes behind eye area; abdomen gray brown, with some yellowish streaks and speckles, and a big yellowish leaf-like marking medially; legs light orange. 8.4.8 Genus Parvattus new genus Type species: Parvattus zhui Zhang & Maddison, sp. nov. Etymology. The generic name is from the combination of parvus (small) and attus (a common ending for salticid genera), referring to the extremely small embolic division of the male palp; masculine in gender. Diagnosis. Small leaf litter dwelling jumping spiders. Male chelicera with two promarginal teeth and one retromarginal tooth. Male palpal bulb large; embolic division very small with embolus short and wide; retrolateral sperm duct loop wide; proximal tegular lobe and ventral tibial bump present; retrolateral tibial apophysis finger-like. Differs from other leaf litter dwelling euophryine genera by the peculiar shape of the embolic division of the male palp. 8.4.8.1 Parvattus zhui sp. nov. Fig 8.11 Type material. Holotype: male, CHINA: Guangxi: Ningming County, about 3km north west of Aidian Village, 21.833° N, 107.017° E, elev. 406-538 m, 19 May 2006, coll. J. Zhang, W. G. Lian & H. Q. Ma, JXZ06#010. Etymology. The specific epithet is a patronym in honor of Prof. Mingsheng Zhu, who made great contributions in studies of Chinese spider fauna, and tragically died of lung cancer in 2010. Prof. Zhu organized and executed the expedition to Guangxi Province, China in 2006, ! 440! during which this species was first found. Diagnosis. See diagnosis of the genus. Description. Male (holotype, UBC-SEM AR00186). Carapace length 1.3; abdomen length 1.0. Chelicera (Fig. 8.11D): dark brown; promargin with two teeth and retromargin with one tooth. Palp (Figs 8.11E-F): proximal tegular lobe and ventral tibial bump present; embolus short and wide and slightly coiled; retrolateral sperm duct loop occupying about three quarters of bulb width; retrolateral tibial apophysis finger-like. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 2.3, II 1.9, III 2.4, IV 2.4. Color in alcohol (Fig. 8.11A): carapace red brown, scattered with light colored scales, eye area dark brown; abdomen also red brown, with yellowish brown symmetrical markings; legs light yellow with gray brown wide annuli. Female. Unknown. Natural history. Specimens were found in leaf litter at the edge of forest. 8.4.9 Genus Thiania C. L. Koch, 1846 Medium sized spiders. Body usually flattened. Carapace almost rectangular. Male palpal bulb usually large; embolus usually wide and short and slightly curved (with the exception of T. tenuis); proximal tegular lobe present or absent; retrolateral sperm duct loop present. Epigynal window present with a median septum. Thiania is similar in the flattened body form and the male palpal structures to Nicylla Thorell (Prószy!ski 1968) and Thianitara Simon (Prószy!ski 1984). The molecular data suggest they fall in the same clade, and Nicylla and Thianitara probably need be synonymized with Thiania (see Chapters 2 and 3). Twenty species of Thiania have been reported (Platnick 2011). Two new species from Malaysia are described here. Placement of T. tenuis within Thiania is partly based on unpublished molecular data. ! 441! 8.4.9.1 Thiania latibola sp. nov. Figs 8.12, 8.17F Type material. Holotype: male, MALAYSIA: Pahang: Cameron Highlands, Arcadia, Trail 3, 4.482° N, 101.388° E, elev. 1550 m, 21-23 May 2005, coll. W. Maddison, D. Li, I. Agnarsson & J. Zhang, WPM#05-033. Etymology. The specific epithet is from the combination of lati (wide) and bola (derived from embolus), referring to the wide embolus of the male palp. Diagnosis. Similar to Thiania bhamoensis (see Prószy!ski 1984; Zabka 1985), but differs in the absence of iridescent scales on the dorsal abdomen and the axe-like embolus. The species can also be distinguished from Nicylla sundevalli (see Prószy!ski 1968) by the pointed retromarginal tooth on the male chelicera and the shape of the embolus of the male palp. Description. Male (holotype, UBC-SEM AR00187). Carapace length 2.8; abdomen length 3.2. Chelicera (Fig. 8.12D): red brown; promargin with one bicuspid tooth, retromargin with one tooth. Palp (Figs 8.12B-C): brownish. Palpal bulb wide with an obvious proximal tegular lobe; embolic disc large; embolus short and wide; retrolateral tibial apophysis long fand inger-like. First tibia with four pairs of ventral macrosetae, first metatarsus with three pairs of ventral macrosetae. Measurements of legs: I 7.2, II 5.3, III 5.3, IV 5.2. Color in alcohol (Fig. 8.12A): carapace brown, covered with white setae, eye area dark; abdomen brown, with three light yellow bands at posterior part; legs light yellow to brown. Female. Unknown. 8.4.9.2 Thiania tenuis sp. nov. Fig 8.13 Type material. Holotype: male, MALAYSIA: Sabah, coll. K. Ober, 2000, 00.418. Etymology. Latin tenuis (slender), referring to the thin and long embolus of the male palp. ! 442! Diagnosis. Differs from other Thiania species by the long and whip-like embolus of the male palp. Description. Male (holotype, UBC-SEM AR00188). Carapace length 3.9; abdomen length 4.6. Chelicera: dark brown; promargin with one bicuspid tooth, retromargin with one tooth. Palp (Figs 8.13B-C): yellow brown to gray brown. Palpal bulb narrow; proximal tegular lobe absent; embolic disc small; embolus long and whip-like; retrolateral tibial apophysis long and finger- like. First tibia with four pairs of ventral macrosetae. Color in alcohol (Fig. 8.13A): carapace dark brown, covered with white setae; abdomen dark brown, only with a few yellowish speckles; legs light yellow to dark brown. Female. Unknown. 8.4.10 Genus Thyenula Simon, 1902 Medium sized spiders. Body usually brown or dark brown with light colored markings on dorsum of abdomen. Some species with a relatively sclerotized area on dorsal abdomen in male. Embolus of male palp narrow or wide, long and coiled; proximal tegular lobe present; retrolateral sperm duct loop occupying more than half of bulb width. Epigynal window present with a median septum. Spermatheca oval or kidney-shaped. Of the ten described species (Platnick 2011), six are reported from South Africa (Prószy!ski 1987; Wesolowska 1993; Wesolowska 2001; Wesolowska & Haddad 2009), three are from Zimbabwe (Wesolowska 2000; Wesolowska & Cumming 2008), and one is from Egypt (Platnick 2011). Three new species from South Africa are described here. 8.4.10.1 Thyenula laxa sp. nov. Fig 8.14 Type material. Holotype: male, SOUTH AFRICA: Northern Province: NE of Ebenezer Dam, George's Valley, ca. 100 km E of Pietersburg, 23 March 2001, coll. G. S. Bonder, GSB#23.111.01c. ! 443! Etymology. Latin laxa (wide), referring to the wide embolus of the male palp. Diagnosis. Differs from other Thyenula species by the wide embolus of the male palp. Description. Male (holotype, UBC-SEM AR00189). Carapace length 2.3; abdomen length 1.9. Chelicera (Fig. 8.14D): dark brown; with two promarginal teeth and one retromarginal tooth. Palp (Figs 8.14B-C): yellow brown. Embolus wide and coiled almost one circle, branched at the tip; retrolateral sperm duct loop wide occupying about three quarters of bulb width; proximal tegular lobe present; retrolateral tibial apophysis long and finger-like. First leg with three pairs of ventral macrosetae on tibia and two pairs of ventral macrosetae on metatarsus. Measurements of legs: I 6.3, II 5.0, III 5.7, IV 5.5. Color in alcohol (Fig. 8.14A): carapace dark brown to yellow brown, with a ‘U’-shaped marking behind eye area composed of light yellow scales; abdomen dark brown, with sandy yellow speckles and marking; legs brown. Female. Unknown. 8.4.10.2 Thyenula nelshoogte sp. nov. Fig 8.15 Type material. Holotype: female, SOUTH AFRICA: Mpumalanga, Nelshoogte forest, 25.803° S, 30.830° E, elev. 1435m, 12 December 2003, coll. W. Moore, WM03.055. Etymology. A noun in apposition taken from the type locality. Diagnosis. Differs from other Thyenula species in the relatively small epigynal window, which is far anterior to the epigynal groove; and the spermathecae, which are posterior to the epigynal window. Description. Female (holotype, UBC-SEM AR00190). Carapace length 2.7; abdomen length 2.9. Chelicera with two promarginal teeth and one retromarginal tooth. Epigynum (Fig. 8.15B): window relatively small with median septum wide. Vulva (Fig. 8.15C): copulatory duct narrow; spermatheca almost kidney-shaped. First leg with three pairs of ventral macrosetae on tibia and two pairs of ventral macrosetae on metatarsus. Measurements of legs: I 4.3, II 3.8, III 4.5, IV ! 444! 5.1. Color in alcohol (Fig. 8.15A): carapace with eye area dark brown, posterior part yellow brown to dark yellow brown; dorsal abdomen sandy yellow, with brown spots and dark brown markings; legs light yellow to light yellow brown. Male. Unknown. 8.4.10.3 Thyenula wesolowskae sp. nov. Fig 8.16 Type material. Holotype: male, SOUTH AFRICA: Mpumalanga: Mariepskop State Forest, Drakensberg, 1.8km N Mariepskop Chalets (Ranger Station), moist montane indigenous forest, 14 Oct. 1999, coll. D. Ubick & S. Prinsloo (CAS). Paratypes: 1 male and 4 females, same data as holotype (CAS). Etymology. The specific epithet is a patronym in honor of Dr. W. Wesolowska, who has made great contributions in studies of jumping spiders from Africa. Diagnosis. Similar to Thyenula oranjensis Wesolowska, 2001, but differs in the smaller embolic disc, the thicker embolus, and the wider proximal tegular lobe of the male palp; the wide copulatory duct and the round spermatheca of the vulva. Description. Male (holotype, UBC-SEM AR00199). Carapace length 1.5; abdomen length 1.6. Chelicera: with two promarginal teeth and one retromarginal tooth. Palp (Figs 8.16C-D): yellow brown. Embolus long and coiled for more than half a circle; retrolateral sperm duct loop very wide almost occupying full width of bulb; proximal tegular lobe relatively large; retrolateral tibial apophysis long and finger-like. First leg with three pairs of ventral macrosetae on tibia and two pairs of ventral macrosetae on metatarsus. Measurements of legs: I 3.5, II 2.5, III 2.6, IV 3.2. Color in alcohol (Fig. 8.16A): carapace with eye area dark brown, posterior part dark yellow brown; dorsal abdomen gray brown to dark brown, with sandy yellow speckles and markings; first pair of legs dark brown, other legs light yellow brown with brown annuli. Female (paratype, UBC-SEM AR00200). Carapace length 1.8 (variation 1.5-1.8, n=4); abdomen length 2.3. Chelicera (Fig. 8.16E): with two promarginal teeth and one retromarginal ! 445! tooth. First tibia with three pairs of ventral macrosetae, first metatarsus with two pairs of ventral macrosetae. Measurements of legs: I 3.1, II 2.6, III 3.0, IV 3.5. Epigynum (Fig. 8.16F): window large with a narrow median septum; opening to copulatory duct close to anterior margin of window. Vulva (Fig. 8.16G): copulatory duct relatively wide; spermatheca almost spherical. Color in alcohol (Fig. 8.16B): similar to that of male. ! 446! Figure 8.1. Chinophrys pengi sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. male endites and labium, ventral view; G. female endites and labium, ventral view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-I, 0.2 mm. ! 447! Figure 8.2. Colyttus robustus sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view. Scale bars: A, 2.0 mm; B-C, 0.2 mm. ! 448! Figure 8.3. Emathis gombak sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, back view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A, 2.0 mm; B, 1.0 mm; C-F, 0.2 mm. ! 449! Figure 8.4. Foliabitus longzhou sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, back view; F. female carapace, dorsal view; G. female left chelicera, back view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A-B, 1.0 mm; C-I, 0.2 mm. ! 450! Figure 8.5. Lagnus edwardsi sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left chelicera, back view; E. epigynum, ventral view; F. cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-F, 0.2 mm. ! 451! Figure 8.6. Laufeia concava sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male right chelicera, back view; F. male right chelicera, front view; G. female left chelicera, back view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; E-G, 0.2 mm; C-D, H-I, 0.1 mm. ! 452! Figure 8.7. Laufeia eximia sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left palp, dorsal view; F. female carapace; dorsal view; G. male endites and labium, ventral view; H. male left chelicera, internal view; I. epigynum, ventral view; J. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; F-H, 0.2 mm; C-E, I-J, 0.1 mm. ! 453! Figure 8.8. Parabathippus cuspidatus sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, retrolateral view; F. male left chelicera, prolateral view; G. epigynum, ventral view; H. cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-H, 0.2 mm. ! 454! Figure 8.9. Parabathippus kiabau sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view; D. male left chelicera, prolateral view; E. male left chelicera, retrolateral view. Scale bars: A, 1.0 mm; B-E, 0.2 mm. ! 455! Figure 8.10. Parabathippus magnus sp. nov. A. male paratype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. male left chelicera, prolateral view; F. male left chelicera, retrolateral view; G. female left chelicera, back view; H. epigynum, ventral view; I. cleared epigynum, dorsal view. Scale bars: A-B, 2.0 mm; C-J, 0.2 mm. ! 456! Figure 8.11. Parvattus zhui sp. nov. A. male holotype, dorsal view; B. male carapace, dorsal view; C. male carapace, lateral view; D. male endites, labium and chelicerae, ventral view; E. male left palp, ventral view; F. male left palp, retrolateral view. Scale bars: A, 0.5 mm; B-D, 0.2 mm; E-F, 0.1 mm. ! 457! Figure 8.12. Thiania latibola sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view; D. male left chelicera, back view. Scale bars: A, 2.0 mm; B-D, 0.2 mm. ! 458! Figure 8.13. Thiania tenuis sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view. Scale bars: A, 2.0 mm; B-C, 0.2 mm. ! 459! Figure 8.14. Thyenula laxa sp. nov. A. male holotype, dorsal view; B. male left palp, ventral view; C. male left palp, retrolateral view; D. male right chelicera, back view. Scale bars: A, 1.0 mm; B-D, 0.2 mm. ! 460! Figure 8.15. Thyenula nelshoogte sp. nov. A. female holotype, dorsal view; B. epigynum, ventral view; C. cleared epigynum, dorsal view. Scale bars: A, 1.0 mm; B-C, 0.2 mm. ! 461! Figure 8.16. Thyenula wesolowskae sp. nov. A. male holotype, dorsal view; B. female paratype, dorsal view; C. male left palp, ventral view; D. male left palp, retrolateral view; E. female left chelicera, back view; F. epigynum, ventral view; G. cleared epigynum, dorsal view. Scale bars: A-B, 0.5 mm; C-G, 0.2 mm. ! 462! Figure 8.17. Living spider photos. A-B. Emathis gombak sp. nov.: A. male; B. female. C-D. Laufeia concava sp. nov.: C. male; D. female. E. male of Colyttus robustus sp. nov.. F. male of Thiania latibola sp. nov.. ! 463! Figure 8.18. Living spider photos. A-C. Parabathippus cuspidatus sp. nov.: A-B. male; C. female. D-F. Parabathippus magnus sp. nov.: D-E. male; F. female. ! ! ! ! 464! 9 Conclusions and future directions 9.1 Major conclusions One of the major goals of this thesis is to provide a phylogenetic framework for euophryine jumping spiders. In Chapter 2, the phylogeny of Euophryinae is reconstructed using molecular data. The monophyly of Euophryinae is strongly supported by the molecular phylogeny with an extensive sample of euophryine taxa. An interesting finding from the molecular phylogeny is that Diolenius and its relatives, which were once placed in the subfamily Dioleninae (Simon 1901a, 1903; Gardzinska & Zabka 2005), fall into a clade within Euophryinae. Similar to Bodner (2009), the molecular phylogeny suggests Euophryinae falls in a large clade with various salticoid groups including Hasarieae, Heliophaninae, Leptorchesteae, Aelurilloida, the Philaeus group, Plexippoida and some unclassified genera (Nannenus, Bristowia, Cheliceroides, Salticus). However, the accurate position of Euophryinae within this clade is still uncertain. The molecular phylogeny also shows that “Bathippus” pahang Zhang, Song & Li is not closely related to other Southeast Asian “Bathippus” spp. (transferred to Parabathippus in the thesis) and falls outside of the euophryine clade. Close relationships of some euophryine genera, such as Anasaitis and Corythalia, Cytaea and Euryattus, etc., are suggested. The other major goal of this thesis is to clarify the taxonomy of the Euophryinae. The molecular phylogeny in Chapter 2 provides a framework for a more natural classification of Euophryinae. I then extensively studied morphological characteristics of a broad range of euophryine genera and species in order to place on the phylogeny genera for which molecular data were unavailable (Chapter 3). Even though morphology alone struggles to resolve the phylogeny of euophryines, it helps to extend the molecular phylogeny, allowing 36 genera without molecular data to be placed within Euophryinae based on similar morphology. In addition, optimization of the morphological characters on the phylogeny provides a better understanding of the euophryine generic groups and the delimitations of many euophryine genera. Certain characters, especially on the male palp, such as the presence of a lamella on the tegular shoulder, presence of a sperm duct loop on the proximal or prolateral proximal side of the palpal bulb, show great value in delimiting various euophryine groups. Although numerous studies have been done on various euophryine taxa, little systematic work on the subfamily Euophryinae as whole has been conducted. In the thesis, I provide a full list of ! 465! euophryine genera and review the generic groups and the delimitations for some genera. The generic review of euophryines will form a foundation for future taxonomic studies on this subfamily. Some problems in the traditional taxonomy of euophryines are also corrected. Twenty-two new synonyms of euophryine genera and 191 new combinations of euophryine species are proposed. In addition, 14 new genera and 96 new species of euophryines are described (Chapters 3, 5-8). The Euophryinae is peculiar among the subfamilies of jumping spiders because they are well represented in both the Old and New World. The molecular phylogeny in Chapter 2 shows that euophryines from different continents usually form their own clades on the phylogeny with few cases of mixture. This finding is concordant with results from previous studies (Maddison & Hedin 2003a; Maddison et al. 2008; Bodner 2009), which suggest that much of salticid diversification occurred after the separation of continents of the Old and New World (Maddison & Hedin 2003a; Maddison et al. 2008). The results from temporal divergence analyses (Chapter 2) further show that most divergences within Euophryinae are after the Eocene, when most continents had already separated. Several intercontinental dispersal events are required to explain the distribution of euophryines, given the age. Early dispersals between the Old and New World may have been facilitated by the Antarctic land bridge. Ancestors of extant euophryines may have been able to survive better under low temperature than other salticid groups of similar age, which may have been crucial to their dispersal and diversification in both the Old World and New World. Euophryines have independently developed two “hot spots” of diversity: New Guinea (Old World) and the Caribbean Islands (New World). Most euophryines from New Guinea fall into one clade and represent a single radiation, with the only exceptions being the Cytaea-Euryattus clade and the Thorelliola clade. In contrast, the Caribbean euophryines are embedded within two big clades, and seem to represent as many as five independent radiations (Chapter 2). Most euophryine genera have typical palpal and epigynal structures (Prószy!ski 1976; Maddison & Hedin 2003a): the palp with a coiled embolus at the distal end of the tegulum, the plane of the embolic spiral more or less parallel to the longitudinal axis of the palp, and a loop of the sperm duct projecting towards the centre of the tegulum; the female epigynum commonly shows two spiral grooves that frame two circular areas of relatively transparent and flat ! 466! integument (referred to as “epigynal window” in this thesis). However, some genera that fall in the clade Euophryinae clearly do not fit into these delimitations. For instance, the embolus of Diolenius comes from the proximal end of the bulb instead of from the distal end; the sperm duct of Chalcolecta does not form the retrolateral loop; the plane of embolus spiral of some Coccorchestes species is perpendicular to the longitudinal axis of the tegulum but not parallel; Tylogonus spp. have fixed embolus; Anasaitis has no obvious window in the epigynum. The molecular phylogeny indicates that these abnormal genitalic forms are derived from typical euophryine-like genitalic structures (Chapter 2). Myrmecophagy (ant-eating) is relatively rare in jumping spiders but has been found in some euophryines (Clark et al. 2000; Jackson & Li 2001): Chalcotropis (6 spp.), Xenocytaea (2 spp.), Zenodorus (=Omoedus, 3 spp.), Anasaitis (1 sp.), Naphrys (1 sp.). The evolution of myrmecophagy in jumping spiders had never been studied before. My molecular phylogeny seems to suggest that the myrmecophagic behavior in Euophryinae may have evolved not only independently in the Old World and the New World, but also separately in different euophryine lineages in each continent. However, more ecological data on whether or not other euophryines are ant-feeding specialists are needed to determine the evolution of myrmecophagy in Euophryinae (Chapter 2). The evolution of genitalia in various animal groups has attracted attention of many researchers (e.g. Eberhard et al. 1998; Arnqvist & Rowe 2002b; Kuntner et al. 2009). However, no quantitative study has been conducted on genital evolution of jumping spiders. The embolus of the male palp and the copulatory duct of the female vulva are key components of genitalia that directly interact during copulation. In Chapter 4, I investigate the correlated evolutionary pattern of the lengths of male embolus and female copulatory duct for the first time in jumping spiders under the phylogenetic context. The results show that they are positively correlated among euophryine species. This correlation could be explained either by sexual selection mechanisms, such as cryptic female choice that affect male paternity success (Eberhard 1996) and sexual conflict that propels sexually antagonistic coevolution (Arnqvist & Rowe 2002a), or by species recognition mechanisms (“lock-and-key”). Further investigations on the intra-specific variation in two euophryine species (Chapoda recondita and Antillattus cambridgei) that show considerable ! 467! difference in sexual dimorphism, are of little help in narrowing down possible mechanisms. Although both male embolus and female copulatory duct show negative intra-specific allometries when compared to other somatic traits, which may appear to favor the “lock-and- key” hypothesis, it could also occur via post-copulatory sexual selection. The size-corrected intra-specific variation is high for genitalia, which has been argued to indicate sexual selection. Nevertheless, the high size-corrected phenotypic variation for most genitalic traits is due to high dispersion of points around the regression line instead of steep allometric slope. The high dispersion may represent high intra-specific variability subject to sexual selection, or may result purely from decoupled developmental mechanisms of genitalic traits and somatic traits. In principle, sexual selection and non-sexual selection mechanisms may coexist, and the divergent genitalia across euophryines species may be the result of a balance between the sexual and non- sexual selection forces (House & Simmon 2003). Unlike the genitalic traits directly associated with copulation, the pre-copulatory sexually- selected traits (e.g. male chelicerae) tend to show positive intra-specific allometry, and thus may have evolved under strong directional selection. The fact that the species with stronger somatic sexual dimorphism (Antillattus cambridgei) has lower intra-specific variation in genitalia may imply a trade-off between pre- and post-copulatory sexual selection. 9.2 Future directions Although progress has been made in understanding phylogeny and evolution of the subfamily Euophryinae in this thesis, our understanding of this group is far from complete. The molecular phylogeny in Chapter 2 provides the basic phylogenetic framework of Euophryinae, but some relationships, especially the deeper ones, have yet to be resolved, and the accurate position of the Euophryinae in salticid phylogeny is still uncertain. Future work on providing a more robust and resolved phylogeny of Euophryinae is particularly important for further investigations in evolutionary processes such as historical biogeography and character evolution. In the molecular phylogenetic study, the gene regions amplified have been widely used in the phylogenetic analyses of jumping spiders (e.g. Maddison & Hedin 2003a, b; Maddison et al. 2008; Bodner 2009). In order to test the monophyly of Euophryinae and have a general idea of relationships of a broad range of euophryine taxa, I mainly focused on collecting data of those known DNA markers from an extensive sampling of potential euophryine taxa, but put less effort on finding ! 468! new DNA markers for phylogenetic reconstruction. Therefore, more work is needed on development of primers for retrieving different genes, especially nuclear protein coding genes, to find suitable markers for reconstructing a more robust and well-resolved phylogeny of Euophryinae. The content of Euophryinae is dramatically extended in this thesis with 96 valid genera included. However, some genera that may actually belong to the Euophryinae were not considered (e.g. Baviola) because of the difficulty in making a decision only by morphology. Besides, some euophryine genera are hard to place within known generic groups based on morphology (e.g. Caribattus and Magyarus). Thus, collecting molecular data from those genera will also help to complete the picture of euophryine phylogeny. The temporal divergence of euophryines sheds lights on the historical biogeography of this group. However, questions such as the continental origin and the events (dispersals and/or vicariance) resulting in the current distributional pattern of euophryine jumping spiders remain uncertain. A more robust and well-resolved phylogeny of euophryines and salticids is crucial to answer those questions. In addition, new discoveries of salticid fossils may dramatically push back the timeline of major groups in Salticidae and change our present understanding of their historical biogeography. The classification of Euophryinae is messy. Although Chapter 3 reviews euophryine genera in detail, it is only a beginning in clarifying the systematics of Euophryinae. A more comprehensive phylogeny will aid in forming a more natural classification of this group. Also, thorough comparative studies of certain morphological traits, such as cleared male palps and SEMs of female vulva, may reveal more reliable features to delimit euophryine generic groups or genera. The Euophryinae is also one of the largest groups within jumping spiders with much of its diversity still undiscovered. Most new euophryine taxa in the thesis are chosen to provide names for the terminal taxa in the molecular phylogeny. A vast number of species have yet to be described. Genital evolution is an intriguing area and Chapter 4 is only the first step of exploration. Although the study reveals a positively coevolved pattern of female copulatory duct and male embolus, the underling mechanisms are still a mystery, and much more data, especially from experimental and behavioral studies, are needed to untangle the puzzle. Comparative studies on genitalia and somatic morphology of closely related sympatric species are essential to uncover if ! 469! the genitalia is species-specific and has the potential to function as the “lock-and-key”. Evidence from experiments on testing the relationship between the paternity and the size of a male genitalic structure would be particularly useful to test sexual selection hypothesis for genitalic evolution in euophryines. 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Agobardus bahoruco JXZ324 DOMINICAN REPUBLIC: Pedernales: P.N.Sierra de Bahoruco (N18.128 W71.558) ! ! ! Agobardus cf. brevitarsus JXZ311 DOMINICAN REPUBLIC: Pedernales: P.N.Sierra de Bahoruco (N18.128 W71.558) ! ! ! ! Agobardus cordiformis JXZ358 DOMINICAN REPUBLIC: Pedernales: east of Pedernales (N17.965 W71.635) ! ! ! ! Agobardus gramineus JXZ314 DOMINICAN REPUBLIC: Pedernales: east of Pedernales (N17.965 W71.635) ! ! ! ! Agobardus oviedo JXZ312 DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo (N17.802 W71.349) ! ! ! ! Agobardus phylladiphilus JXZ313 DOMINICAN REPUBLIC: Pedernales: P.N.Sierra de Bahoruco (N18.128 W71.558) ! ! ! ! Amphidraus complexus JXZ035 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! ! Anasaitis adorabilis JXZ359 DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo (N17.802 W71.349) ! ! ! ! Anasaitis brunnea JXZ316 DOMINICAN REPUBLIC: Barahona: Highway 44 south of Barahona (N18.138 W71.070) ! ! ! Anasaitis canosa [USA] JXZ193 USA: Florida: Alachua Co., Gainesville ! ! ! ! Anasaitis canosa [Panama] JXZ248 PANAMA: Colón: Punta Galeta (N9.40376 W79.86354) ! ! ! Anasaitis hebetata JXZ318 DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García (N18.424 W71.112) ! ! ! Anasaitis laxa JXZ319 DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García (N18.424 W71.112) ! ! ! Anasaitis sp. [Peblique] JXZ317 DOMINICAN REPUBLIC: Pedernales: Peblique (N18.059 W71.638) ! ! ! Antillattus cf. applanatus JXZ336 DOMINICAN REPUBLIC: Barahona: Cachote (N18.101 W71.194) ! ! ! ! Antillattus gracilis JXZ320 DOMINICAN REPUBLIC: La Vega: P.N.Armando Bermúdez (N19.06 W70.86) ! ! ! Asaphobelis physonychus JXZ179 BRAZIL: São Paulo: São Paulo ! ! ! ! Athamas nitidus JXZ142 PAPUA NEW GUINEA: Madang, Adalbert Mts., Sewan- Keki (S4.704 E145.419) ! ! ! ! ! \"##! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Athamas cf. whitmeei JXZ345 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 3, Tompoi (S5.344 E151.315) ! ! ! Athamas sp. JXZ181 MICRONESIA: Pohnbe, Pahn Takai (S6.9457 E158.2735) ! ! Bathippus directus [Tualapa] JXZ205 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Bathippus cf. directus [Suyang] JXZ259 PAPUA NEW GUINEA: Enga Province: Suyan Camp, Porgera (S5.4833 E143.1337) ! ! ! ! Bathippus gahavisuka JXZ228 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.015 E145.412) ! ! ! ! Bathippus korei JXZ284 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! ! Bathippus macrognathus JXZ372 PAPUA NEW GUINEA: Southern Highlands Province: Putuwé, junction of Lagaip & Uruwabwa Rivers (S5.231 E142.532) ! ! ! Bathippus madang JXZ143 PAPUA NEW GUINEA: Madang, Adalbert Mts., Sewan- Keki (S4.704 E145.419) ! ! ! ! Belliena ecuadorica (F) JXZ041 ECUADOR: Morona Santiago: km 15 from Limón towards Gualaceo (S3.0090 W78.5024) ! ! ! Belliena ecuadorica (M) JXZ060 ECUADOR: Morona Santiago: km 20 from Limón towards Gualaceo (S3.0044 W78.5142) ! ! ! Belliena sp. [JatunSacha] JXZ061 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! ! Belliena sp. [Yanayacu] JXZ042 ECUADOR: Napo: Caucheras, Estación Yanayacu (S0.6049 W77.8886) ! ! ! ! Bulolia excentrica JXZ283 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! ! Bythocrotus crypticus JXZ322 DOMINICAN REPUBLIC: Barahona: Parque Nacional Sierra Martín García (N18.424 W71.112) ! ! ! Bythocrotus cf. crypticus JXZ323 DOMINICAN REPUBLIC: El Seibo: Pedro Sanchez (N18.86 W69.11) ! ! ! ! Canama extranea JXZ258 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Canama fimoi JXZ265 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Canama cf. forceps JXZ210 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! ! Canama hinnulea JXZ089 AUSTRALIA: Palm Island (S18.733 E146.583) ! Canama hinnulea JXZ161 AUSTRALIA: Palm Island (S18.733 E146.583) ! ! Canama triramosa JXZ277 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.016-6.017 E145.417- 145.416) ! ! ! Chalcolecta prensitans JXZ296 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Chalcolemia nakanai JXZ346 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 1, Lamas (S5.614 E151.408) ! ! ! Chalcoscirtus alpicola d294 CANADA: Alberta: Rock Lake (N53.47157 W118.23852) ! ! ! ! Chalcoscirtus diminutus JXZ365 USA: Arizona: Pima Co., Santa Catalina Mountains, San Pedro Vista (N32.399 W110.690) ! ! ! Chalcoscirtus infimus JXZ293 SPAIN: Andalucia: Frigiliana (N36.78 W3.90) ! ! ! Chalcotropis cf. caeruleus JXZ099 MALAYSIA: Sabah: Mt. Kinabalu, Liwagu Trail (N6.008 E116.543) ! ! ! Chalcotropis luceroi JXZ368 PHILIPPINES: Mt. Makiling ! Chalcotropis luceroi s153 PHILIPPINES: Luzon (755)AY297257 (393)AY297320; (594)AY296677 (969)AY297418 Chapoda angusta JXZ059 ECUADOR: Manabí: Puerto Lopez (S1.5497 W80.8104) ! ! ! ! Chapoda fortuna [Panama] JXZ253 PANAMA: Chiriqui: Fortuna, Quebrada Samudio (N8.73464 W082.24839) ! ! ! ! Chapoda cf. fortuna [CostaRica] JXZ114 COSTA RICA: Prov. San José: Masizo de la Muerte (N9.667 W83.85) ! ! ! ! Chapoda gitae JXZ251 PANAMA: Panamá: Panama City, Parque Metropolitano (N8.99442 W79.54301) ! ! ! Chapoda cf. inermis [Panama] JXZ241 PANAMA: Panamá: Gamboa, Pipeline Road (N9.15840 W79.74252) ! ! ! ! Chapoda cf. inermis [CostaRica] JXZ115 COSTA RICA: Prov. San José: Aprox. 10km from Santa Maria de Dota towards Naranjillo Village (N9.65 W83.97) ! ! ! ! Chapoda peckhami JXZ242 PANAMA: Chiriqui: Fortuna, Casa Verde Smithsonian Field Station (N8.72205 W082.23746) ! ! ! ! Chapoda sp. [Fortuna] JXZ254 PANAMA: Chiriqui: Fortuna, near Casa Verde Smithsonian Field Station (N8.72205 W82.23854) ! ! ! cf. Chapoda sp. [Arizona] JXZ366 USA: Arizona: Santa Cruz Co., Santa Rita Mountains (N31.6705 W110.9156) ! ! ! ! Chinophrys pengi JXZ145 CHINA: Guangxi: Tianlin County, Langping Village (N24.467 E106.367) ! ! ! Cobanus cf. cambridgei JXZ122 COSTA RICA: Prov. San José: Aprox. 10km from Santa Maria de Dota towards Naranjillo Village (N9.65 W83.97) ! ! ! Cobanus cf. electus JXZ054 ECUADOR: Pichincha: near El Cisne (N0.1493 W79.0317) ! ! ! ! Cobanus extensus JXZ051 ECUADOR: Pichincha: near El Cisne (N0.1493 W79.0317) ! ! ! ! Cobanus mandibularis JXZ245 PANAMA: Panamá: Gamboa, Pipeline Road (N9.15840 W79.74252) ! ! ! Cobanus unicolor JXZ244 PANAMA: Chiriqui: Fortuna, Quebrada Samudio (N8.73464 W82.24839) ! ! ! Cobanus sp. [CostaRica] JXZ113 COSTA RICA: Prov. San José: Masizo de la Muerte (N9.667 W83.85) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Cobanus sp. [Ecuador] JXZ116 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! Cobanus sp. [Panama] JXZ243 PANAMA: Chiriqui: Fortuna, Quebrada Samudio (N8.73464 W82.24839) ! ! ! ‘Cobanus' cambridgei JXZ321 DOMINICAN REPUBLIC: La Vega: Reserva Científica Ébano Verde (N19.033 W70.543) ! ! ! ! Coccorchestes cf. aiyura JXZ289 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Coccorchestes cf. ildikoae JXZ208 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.016-6.017 E145.417- 145.416) ! ! ! Coccorchestes cf. inermis JXZ139 PAPUA NEW GUINEA: Madang, Adalbert Mts., Sewan- Keki (S4.704 E145.419) ! ! ! ! Coccorchestes clavifemur JXZ288 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! Colyttus bilineatus JXZ070 MALAYSIA: Johor: Teluk Mahkota (N1.9 E104.104) ! ! ! ! Colyttus robustus JXZ071 MALAYSIA: Pahang: Genting Highlands (N3.400 E101.777) ! ! ! ! Compsodecta haytiensis JXZ325 DOMINICAN REPUBLIC: Barahona: Highway 44 south of Barahona (N18.138° W71.070) ! ! ! ! Compsodecta peckhami JXZ327 DOMINICAN REPUBLIC: Pedernales: Rio Mulito (N18.155 W71.758) ! ! ! Corticattus guajataca JXZ305 PUERTO RICO: Isabela: Bosque de Guajataca (N18.421 W66.966) ! ! ! ! Corticattus latus JXZ337 DOMINICAN REPUBLIC: Pedernales: Laguna de Oviedo (N17.802 W 71.349) ! ! ! ! Coryphasia fasciiventris JXZ201 BRAZIL: São Paulo: São Paulo ! ! ! cf. Coryphasia sp. [Brazil] JXZ200 BRAZIL: São Paulo: São Paulo ! ! ! Corythalia cf. albicincta JXZ239 PANAMA: Bocas del Toro: Isla Colón, Bocas Smithsonian Field Station (N9.35302 W82.25746) ! ! ! Corythalia bicincta JXZ237 PANAMA: Panamá: Gamboa, Pipeline Road (N9.15840 W79.74252) ! ! ! Corythalia broccai JXZ331 DOMINICAN REPUBLIC: El Seibo: Pedro Sanchez - Miches Road (N18.958 W69.069) ! ! ! Corythalia bromelicola JXZ330 DOMINICAN REPUBLIC: Barahona: Cachote-La Cienega Road (N18.0562 W71.1416) ! ! ! ! Corythalia coronai JXZ343 DOMINICAN REPUBLIC: Pedernales: Rio Mulito (N18.155 W71.758) ! ! ! ! Corythalia electa JXZ031 ECUADOR: Morona Santiago: km 7 from Limón towards Gualaceo (S2.9962 W78.4558) ! ! ! ! Corythalia cf. latipes JXZ196 BRAZIL: São Paulo: São Paulo ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Corythalia peblique JXZ344 DOMINICAN REPUBLIC: Pedernales: Peblique (N18.059 W71.638) ! ! ! Corythalia porphyra JXZ112 COSTA RICA: Prov. Heredia: 6km ENE Vara Blanca (N10.183 W84.117) ! ! ! ! Corythalia sulfurea JXZ236 PANAMA: Panamá: Gamboa, Pipeline Road (N9.15840 W79.74252) ! ! ! Corythalia cf. valida JXZ131 ECUADOR: Morona Santiago: km 15 from Limón towards Gualaceo (S3.0090 W78.5024) ! ! ! Corythalia sp. [Baeza] JXZ049 ECUADOR: Napo: NE of Baeza (S0.2712 W77.7587) ! ! ! ! Corythalia sp. [JatunSacha] JXZ032 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! ! Corythalia sp. [PuertoLopez] JXZ132 ECUADOR: Manabí: Puerto Lopez (S1.5497 W80.8104) ! ! ! ‘Corythalia' banksi JXZ299 PUERTO RICO: Guánica: Bosque Seco de Guánica (N17.975 W66.872) ! ! ! ! ‘Corythalia' cf. canalis JXZ247 PANAMA: Bocas del Toro: Isla Solarte (N9.33335 W82.22012) ! ! ! ! ‘Corythalia' elegantissima JXZ328 DOMINICAN REPUBLIC: La Altagracia: Punta Cana (N18.516 W68.379) ! ! ‘Corythalia' gloriae JXZ301 PUERTO RICO: Maricao: Bosque de Maricao (N18.150 W66.994) ! ! ! ‘Corythalia' locuples JXZ315 DOMINICAN REPUBLIC: La Vega: Reserva Científica Ébano Verde (N19.033 W70.543) ! ! ! ! Cytaea mitellata JXZ280 PAPUA NEW GUINEA: National Capital District: Port Moresby (S9.443 E147.179) ! ! ! Cytaea nimbata JXZ229 PAPUA NEW GUINEA: Eastern Highlands Province: Goroka (S6.07 E145.40) ! ! ! ! Cytaea oreophila JXZ045 SINGAPORE: Labrador Park (N1.27 E103.80) ! ! ! ! Cytaea cf. rai JXZ182 MICRONESIA: Pohnbe, Pahn Takai (S6.9457 E158.2735) ! ! ! Cytaea cf. sinuata JXZ269 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Cytaea sp. [Wewak] JXZ144 PAPUA NEW GUINEA: East Sepik Province: Wewak ! ! ! ! Dinattus minor JXZ329 DOMINICAN REPUBLIC: Pedernales: Rio Mulito (N18.155 W71.758) ! ! ! Diolenius cf. decorus d237 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! ! Diolenius varicus JXZ349 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 2, Vouvou (S5.446 E151.464) ! ! ! ! Donoessus striatus JXZ148 MALAYSIA: Selangor: canyon near Ulu Gombak (N3.325 E101.765) ! Donoessus striatus JXZ066 MALAYSIA: Selangor: canyon near Ulu Gombak (N3.325 E101.765) ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Donoessus striatus JXZ044 MALAYSIA: Selangor: Ulu Gombak Field Station (N3.325 E101.753) ! Ecuadattus napoensis JXZ083 ECUADOR: Napo: Cocodrilo (S0.6490 W77.7927) ! ! ! ! Ecuadattus pichincha JXZ184 ECUADOR: Pichincha: Mindo, Mindo Garden forest (S0.0732 W78.7542) ! ! ! Efate albobicinctus JXZ183 MICRONESIA: Pohnbe, Pahn Takai (S6.9457 E158.2735) ! ! ! ! Emathis gombak JXZ072 MALAYSIA: Selangor: Ulu Gombak Field Station (N3.325 E101.753) ! ! ! Euophrys frontalis JXZ137 GERMANY: Saxony: Authausen (N51.607 E12.711) ! ! ! Euophrys monodnock JXZ162 CANADA: Nova Scotia: Barneys River at Route 104 (N45.5862 W62.2271) ! ! ! Euophrys cf. proszynskii JXZ133 SPAIN: Sitges (N41.2383 E1.8244) ! ! ! ‘Euophrys' a-notata JXZ190 CHILE: Reg. X, Chiloé: P.N. Chiloé (S42.57627 W74.07820) ! ! ! ! ‘Euophrys' cf. patagonica JXZ191 CHILE: Reg. X, Chiloé: P.N. Chiloé (S42.57627 W74.07820) ! ! ! ! Euryattus bleekeri JXZ086 AUSTRALIA: Queensland: Cow Bay, Crocodylus Village ! ! ! ! Euryattus cf. porcellus JXZ270 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Euryattus cf. venustus JXZ212 PAPUA NEW GUINEA: Eastern Highlands Province: Goroka (S6.07 E145.40) ! ! ! Euryattus sp. [Queensland] JXZ094 AUSTRALIA: Queensland: Stratbroke Is. ! ! ! ! Euryattus sp.1 [Gahavisuka] JXZ273 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Euryattus sp.2 [Gahavisuka] JXZ215 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.015 E145.412) ! ! ! Foliabitus longzhou JXZ076 CHINA: Guangxi: Pingxiang City, Daqingshan Forestry Center (N22.299 E106.700) ! ! ! ! Foliabitus sp. [Malaysia] JXZ079 MALAYSIA: Pahang: Genting Highlands (N3.400 E101.777) ! ! ! Hypoblemum cf. albovittatum JXZ164 AUSTRALIA: South Australia: Glenelg (S34.973 E138.511) ! ! ! ! Hypoblemum sp. [NewSouthWales] JXZ085 AUSTRALIA: New South Wales: Gerringong ! ! ! ! Ilargus coccineus JXZ195 BRAZIL: São Paulo: São Paulo ! ! ! Ilargus foliosus JXZ123 ECUADOR: Morona Santiago: km 15 from Limón towards Gualaceo (S3.0090 W78.5024) ! ! ! Ilargus galianoae JXZ036 ECUADOR: Morona Santiago: km 16 from Limón towards Gualaceo (S3.0060 W78.4997) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Ilargus macrocornis JXZ057 ECUADOR: Morona Santiago: Cordillera de Cutucú (S2.8058 W78.2462) ! ! ! ! Ilargus moronatigus JXZ043 ECUADOR: Morona Santiago: km 38 from Limón towards Gualaceo (S3.0108 W78.6150) ! ! ! ! Ilargus pilleolus JXZ037 ECUADOR: Azuay: SE of Gualaceo (S2.968 W78.700) ! ! ! ! Ilargus serratus JXZ055 ECUADOR: Morona Santiago: km 13 from Limón towards Gualaceo (S3.0093 W78.4939) ! ! ! ! Jotus auripes JXZ090 AUSTRALIA: Queensland: Oakview State Forest (S26.167 E152.333) ! ! ! ! cf. Jotus sp. [Queensland] JXZ155 AUSTRALIA: Queensland: Cow Bay, Crocodylus Village ! ! ! Junxattus daiqini JXZ024 SINGAPORE: Bukit Timah Nature Reserve (N1.355 E103.78) ! ! ! ! Lagnus edwardsi s152 PHILIPPINES: Luzon (769)AY297283 (393)AY297346; (580)AY296701 (1047)AY297411 Lagnus edwardsi JXZ369 PHILIPPINES: Makiling ! Laufeia concava JXZ046 MALAYSIA: Pahang: Gunung Brinchang (N4.515 E101.383) ! ! ! ! Laufeia eximia JXZ077 CHINA: Guangxi: Tianlin County, Langping Village (N24.467 E106.367) ! ! ! Lepidemathis haemorrhoidalis JXZ367 PHILIPPINES: IRRI ! Lepidemathis haemorrhoidalis s262 PHILIPPINES: Luzon (693)AY297260 (393)AY297323; (580)AY296680 (1041)AY297389 Leptathamas paradoxus JXZ207 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Lycidas cf. griseus JXZ092 AUSTRALIA: New South Wales: Buderoon ! ! ! ! Lycidas cf. karschi JXZ157 AUSTRALIA: New South Wales: Gerringong ! ! ! Lycidas cf. vittatus JXZ146 AUSTRALIA: Queensland: Cow Bay (S16.233 E145.437) ! ! ! Maeota dichrura JXZ180 BRAZIL: São Paulo: São Paulo ! ! ! ! Maeota dorsalis JXZ198 BRAZIL: São Paulo: São Paulo ! ! ! Maeota flava JXZ199 BRAZIL: São Paulo: São Paulo ! ! ! ! Maeota simoni JXZ252 PANAMA: Panamá: Panama City, Parque Metropolitano (N8.99442 W79.54301) ! ! ! ! Maeota sp. [Cutucú] JXZ117 ECUADOR: Morona Santiago: Cordillera de Cutucú (S2.7708 W78.2570) ! ! ! ! Maeota sp. [JatunSacha] JXZ130 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! ! ! \"#\"! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Maeota sp. [Manabi] JXZ058 ECUADOR: Manabí: Puerto Lopez (S1.5497 W80.8104) ! ! ! ! Maeota sp. [MoronaSantiago] JXZ127 ECUADOR: Morona Santiago: road between Limón and Patuca (S2.8310 W78.3607) ! ! ! ! Maeota sp. [Napo] JXZ048 ECUADOR: Napo: NE of El Chaco (S0.2025 W77.7015) ! ! ! ! cf. Maeota sp. [Panama] JXZ256 PANAMA: Chiriqui: Fortuna, Quebrada Honda (N08.75176 W82.23953) ! ! ! ! Maileus cf. fuscus JXZ098 MALAYSIA: Pahang: Taman Negara ! ! ! ! Maratus cf. amabilis JXZ160 AUSTRALIA: Western Australia: Mt. Cooke ! ! ! ! Maratus sp. [SouthAustralia] JXZ159 AUSTRALIA: South Australia: Glenelg (S34.973 E138.511) ! ! ! ! ‘Margaromma' cf. semirasum JXZ226 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.015 E145.412) ! ! ! ! ‘Margaromma' cf. torquatum JXZ279 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Marma nigritarsis JXZ353 BRAZIL: Amazonas: Manaus (S3.098 W59.972) ! ! ! ! Mexigonus arizonensis JXZ163 USA: Arizona: Yavapai Co., Iron Springs (N34.58476 W112.57071) ! ! ! ! Mexigonus cf. minutus [Ecuador] d117 ECUADOR: Pichincha: Quito ! ! ! ! Mexigonus morosus JXZ362 USA: California: San Mate Co. (N37.434 W122.311) ! ! ! Naphrys pulex JXZ081 CANADA: Ontario: St Williams ! ! ! ! Naphrys xerophila JXZ257 USA: Florida: Marion Co., Ocala National Forest ! ! ! Nebridia cf. semicana JXZ052 ECUADOR: Pichincha: Mindo, Mindo Garden forest (S0.0732 W78.7542) ! ! ! ! Neonella vinnula JXZ178 USA: Florida: Duval Co., Ft. George Island ! ! Ohilimia scutellata JXZ295 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! Omoedus brevis JXZ276 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! ! Omoedus cf. danae JXZ274 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! Omoedus darleyorum JXZ262 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Omoedus cf. durvillei JXZ294 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Omoedus ephippigera JXZ022 SINGAPORE: Nee Soon Swamp Forest (N1.39 E103.81) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Omoedus cf. metallescens JXZ154 AUSTRALIA: Palm Island (S18.733 E146.583) ! ! ! ! Omoedus meyeri JXZ290 PAPUA NEW GUINEA: Enga Province: Kai-ingri (S5.574 E143.048) ! ! ! ! Omoedus omundseni JXZ221 PAPUA NEW GUINEA: Enga Province: near Suyan Village (S5.495 E143.144) ! ! ! Omoedus orbiculatus JXZ136 AUSTRALIA: Queensland: Stratbroke Is. ! ! ! Omoedus orbiculatus JXZ088 AUSTRALIA: Queensland: near Atherton ! Omoedus papuanus JXZ286 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.016-6.017 E145.416- 145.417) ! ! ! ! Omoedus cf. piceus JXZ206 PAPUA NEW GUINEA: Southern Highlands Province: Wanakipa Station (S5.2571 E142.5216) ! ! ! ! Omoedus cf. ponapensis JXZ138 PAPUA NEW GUINEA: Sandaun (S2.740 E141.251) ! ! ! ! Omoedus swiftorum JXZ266 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Omoedus tortuosus JXZ282 PAPUA NEW GUINEA: Enga Province: near Suyan Village (S5.495 E143.144) ! ! Omoedus sp. [Gahavisuka] JXZ223 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.015 E145.412) ! ! ! Omoedus sp. [Adalbert] JXZ140 PAPUA NEW GUINEA: Madang, Adalbert Mts., Sewan- Keki (S4.704 E145.419) ! ! ! Orcevia keyserlingi JXZ025 MALAYSIA: Pahang: Genting Highlands (N3.400 E101.777) ! ! ! ! Palpelius beccarii JXZ209 PAPUA NEW GUINEA: Southern Highlands Province: Putuwé (S5.231 E142.532) ! ! ! Palpelius cf. discedens JXZ352 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 2, Vouvou (S5.446 E151.464) ! ! ! ! Parabathippus cuspidatus JXZ150 MALAYSIA: Pahang: Genting Highlands (N3.400 E101.777) ! ! ! Parabathippus kiabau JXZ097 MALAYSIA: Sabah: Village of Kiabau (N5.832 E117.225) ! ! ! ! Parabathippus cf. macilentus JXZ151 MALAYSIA: Selangor: Ulu Gombak Field Station (N3.325 E101.753) ! ! ! Parabathippus magnus JXZ064 MALAYSIA: Pahang: Tanah Rata (N4.46 E101.40) ! ! ! Parabathippus magnus JXZ026 MALAYSIA: Pahang: Genting Highlands (N3.400 E101.777) ! Parabathippus shelfordi JXZ065 SINGAPORE: Nee Soon Swamp Forest (N1.39 E103.81) ! ! ! ! Paraharmochirus tualapaensis JXZ297 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Parvattus zhui JXZ078 CHINA: Guangxi: Ningming County (N21.833 E107.017) ! ! ! ! Pensacola signata JXZ371 GUATEMALA: Depto. Petén: Reserva Natural Ixpanpajul ! ! ! ‘Pensacola' darlingtoni JXZ341 DOMINICAN REPUBLIC: La Vega: Reserva Científica Ébano Verde (N19.033 W70.543) ! ! ! ! ‘Pensacola' maxillosa JXZ335 DOMINICAN REPUBLIC: La Vega: road Constanza to Ocoa, Valle Nuevo (N18.700 W70.606) ! ! ! ! ‘Pensacola' tuberculotibiata JXZ056 ECUADOR: Napo: NE of El Chaco (S0.2025 W77.7015 ! ! ! ! Petemathis minuta JXZ302 PUERTO RICO: Maricao: Bosque de Maricao (N18.150 W66.994) ! ! ! Petemathis portoricensis [Maricao] JXZ303 PUERTO RICO: Maricao: Bosque de Maricao (N18.150 W66.994) ! ! ! ! Petemathis portoricensis [Adjuntas] JXZ306 PUERTO RICO: Adjuntas: HWY143 to Cerro Punta (N18.167 W66.576) ! ! ! ! Petemathis tetuani JXZ308 PUERTO RICO: Río Grande: El Yunque National Forest (N18.294 W65.792) ! ! ! ! Phasmolia elegans JXZ225 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Popcornella furcata JXZ334 DOMINICAN REPUBLIC: La Vega: Reserva Científica Ébano Verde (N19.04 W70.518) ! ! ! Popcornella nigromaculata JXZ304 PUERTO RICO: Maricao: Bosque de Maricao (N18.150 W66.994) ! ! ! Popcornella spiniformis JXZ339 DOMINICAN REPUBLIC: Barahona: Cachote (N18.098 W71.187) ! ! ! Popcornella yunque JXZ309 PUERTO RICO: Río Grande: El Yunque Nat. Forest (N18.3174 W65.8314) ! ! ! Pristobaeus cf. jocosus JXZ347 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 2, Vouvou (S5.446 E151.464) ! ! ! ! Prostheclina sp. JXZ095 AUSTRALIA: New South Wales: Jarvis Bay ! ! ! ! Pseudeuophrys erratica JXZ194 USA: Massachusetts: Cambridge ! ! ! Saitis barbipes JXZ147 SPAIN: Sitges (N41.2383 E1.8244) ! ! ! ! cf. Saitis sp. [WestAustralia] JXZ091 AUSTRALIA: Western Australia: Lake King/Peak, Charles Road ! ! ! ! ‘Saitis' cf. mundus JXZ106 SOUTH AFRICA: Mpumalanga Province: Songimvelo Nature Reserve (S26.042 E31.013) ! ! ! ! ‘Saitis' leighi JXZ105 SOUTH AFRICA: KwaZulu-Natal Province: Ngome State Forest (S27.820 E31.417) ! ! ! Servaea vestita JXZ093 AUSTRALIA: New South Wales: Jarvis Bay ! ! ! ! Sidusa sp.1 [FrenchGuiana] JXZ128 FRENCH GUIANA: Commune Règina, les Nourages Field Station (N4.069 W52.669) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Sidusa sp.2 [FrenchGuiana] JXZ100 FRENCH GUIANA: Commune Règina, les Nourages Field Station (N4.069 W52.669) ! ! ! ! ‘Sidusa' recondita JXZ111 COSTA RICA: Heredia: Est. Biol. La Selva (N10.433 W84.017) ! ! ! ! Siloca cf. campestrata JXZ029 ECUADOR: Morona Santiago: km 7 from Limón towards Gualaceo (S2.9962 W78.4558) ! ! ! ! Siloca cf. sanguiniceps JXZ053 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! ! ‘Siloca' electa JXZ326 DOMINICAN REPUBLIC: La Vega: road Constanza to Ocoa, Valle Nuevo (N18.848 W70.720) ! ! Sobasina wanlessi JXZ298 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Soesilarishius cf. amrishi JXZ354 BRAZIL: Amazonas: Manaus (S3.098 W59.972) ! ! ! Soesilarishius micaceus JXZ120 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! ! ! Soesilarishius micaceus JXZ062 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) ! Soesilarishius ruizi JXZ356 BRAZIL: Amazonas: Manaus (S3.098 W59.972) ! ! ! ! ‘Stoidis' placida JXZ300 PUERTO RICO: Maricao: Bosque de Maricao (N18.150 W66.994) ! ! ! Talavera minuta JXZ202 CANADA: British Colombia: Vancouver, Iona Beach ! ! ! ! Tariona cf. bruneti JXZ197 BRAZIL: São Paulo: Campos do Jordño ! ! ! Thiania bhamoensis JXZ135 SINGAPORE: Labrador Park (N1.27 E103.80) ! ! Thiania bhamoensis JXZ020 SINGAPORE: Labrador Park (N1.27 E103.80) ! Thiania latibola JXZ067 MALAYSIA: Pahang: Gunung Brinchang (N4.515 E101.383) ! ! ! ! Thiania cf. suboppressa JXZ021 MALAYSIA: Selangor: Ulu Gombak Field Station (N3.325 E101.753) ! ! ! ! Thiania tenuis JXZ084 MALAYSIA: Sabah ! ! ! Thiania cf. viscaensis JXZ080 MALAYSIA: Selangor: Ulu Gombak Field Station (N3.325 E101.753) ! ! ! ! Thianitara spectrum JXZ019 MALAYSIA: Selangor: Ulu Gombak Field Station (N3.325 E101.753) ! ! ! ! Thorelliola aliena JXZ285 PAPUA NEW GUINEA: Southern Highlands Province: Umgé (S5.304 E142.512) ! ! ! Thorelliola crebra JXZ217 PAPUA NEW GUINEA: Enga Province: Suyan Camp (S5.4833 E143.1337) ! ! ! Thorelliola ensifera JXZ063 SINGAPORE: Nee Soon Swamp Forest (N1.39 E103.81) ! ! ! ! ! \"#$! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Thorelliola Joannae JXZ264 PAPUA NEW GUINEA: Southern Highlands Province: Umgé (S5.304 to 5.305 E142.510 to 142.512) ! ! ! ! Thorelliola mahunkai JXZ156 PAPUA NEW GUINEA: Enga Province: foot of Mt. Hagen (S5.47.5488 E143.58.761) ! ! ! Thorelliola cf. mahunkai JXZ291 PAPUA NEW GUINEA: Eastern Highlands Province: Goroka (S6.07 E145.40) ! ! ! ! Thorelliola tamasi JXZ281 PAPUA NEW GUINEA: Eastern Highlands Province: Mt. Gahavisuka Provincial Park (S6.016-6.017 E145.417- 145.416) ! ! ! Thorelliola tualapa JXZ287 PAPUA NEW GUINEA: Southern Highlands Province: Tualapa, near Wanakipa (S5.283 E142.498) ! ! ! Thyenula cf. aurantiaca JXZ107 SOUTH AFRICA: KwaZulu-Natal Province: Ngome State Forest (S27.820 E31.417) ! ! ! ! Thyenula laxa JXZ103 SOUTH AFRICA: Mpumalanga, Uitsoek forest (S25.256 E30.551) ! ! ! ! Thyenula nelshoogte JXZ108 SOUTH AFRICA: Mpumalanga, Nelshoogte forest (S25.803 E30.830) ! ! ! ! Thyenula wesolowskae JXZ192 SOUTH AFRICA: Limpopo, Entabeni Nature Reserve (S22.996 E30.264) CAS ! ! ! ! Thyenula sp. [SouthAfrica] JXZ149 SOUTH AFRICA: Northern Province: Strydpoort Mountains (S24.17 E29.93) ! ! Thyenula sp. [SouthAfrica] JXZ104 SOUTH AFRICA: Northern Province: Strydpoort Mountains (S24.17 E29.93) ! ! Truncattus cachotensis JXZ338 DOMINICAN REPUBLIC: Barahona: Cachote (N18.101 W71.194) ! ! ! ! Truncattus dominicanus JXZ340 DOMINICAN REPUBLIC: La Vega: P.N.Armando Bermúdez (N19.06 W70.86) ! ! ! ! Truncattus flavus JXZ332 DOMINICAN REPUBLIC: La Vega: P.N.Armando Bermúdez (N19.06 W70.86) ! ! ! ! Tylogonus cf. auricapillus JXZ050 ECUADOR: Pichincha: Mindo, Mindo Garden forest (S0.0732 W78.7542) ! ! ! ! Tylogonus parvus JXZ034 ECUADOR: Morona Santiago: km 36 from Limón towards Gualaceo (S3.0151 W78.5982) ! ! ! ! Tylogonus cf. viridimicans JXZ118 ECUADOR: Morona Santiago: km 38 from Limón towards Gualaceo (S3.0108 W78.6150) ! ! ! ! Tylogonus yanayacu JXZ033 ECUADOR: Napo: Caucheras, Estación Yanayacu (S0.6049 W77.8886) ! ! ! ! Variratina minuta JXZ222 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! Viribestus suyanensis JXZ261 PAPUA NEW GUINEA: Enga Province: Suyan Camp (S5.4833 E143.1337) ! ! ! ! ‘Wallaba' decora JXZ342 DOMINICAN REPUBLIC: Pedernales: P.N. Sierra de Bahoruco (N18.148 W 71.620) ! ! ! ! Xenocytaea agnarssoni JXZ350 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 2, Vouvou (S5.446 E151.464) ! ! ! ! \"##! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Xenocytaea albomaculata JXZ351 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 1, Lamas (S 5.614 E 151.408) ! ! ! Xenocytaea proszynskii JXZ348 PAPUA NEW GUINEA: New Britain: Nakanai Mts, Camp 3, Tompoi (S5.344 E151.315) ! ! ! Zabkattus brevis JXZ260 PAPUA NEW GUINEA: Southern Highlands Province: Umgé (S5.304-5.305 E142.510-142.512) ! ! ! Zabkattus furcatus JXZ218 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! Zabkattus richardsi JXZ216 PAPUA NEW GUINEA: Eastern Highlands Province: Goroka (S6.07 E145.40 ! ! ! Zabkattus trapeziformis JXZ275 PAPUA NEW GUINEA: Central Province: Varirata National Park (S9.436 E147.364) ! ! ! Euophryine sp. [GentingHighlands] JXZ074 MALAYSIA: PAHANG: Genting Highlands (N3.400° E101.777) ! ! ! ! Other salticid groups cf. Acragus MRB137 ECUADOR: Napo (S1.0466 W77.7430) Bodner 2009 Bodner 2009 Bodner 2009 Aelurillus cf. ater d140 KAZAKHSTAN: Almaty Region (N43.643 E75.805) (753) EU815504 Bodner 2009 (757) EU815564 (968) EU815615 Arasia mollicoma d046 AUSTRALIA: New South Wale: Munmorah Area (659) EU815483 Bodner 2009 (597) EU815550 (976) EU815598 Attidops youngi s97 USA: Missouri (762)AF327933 (393)AF328020; (579)AF327961 (1047)AF327990 ‘Bathippus’ pahang JXZ028 MALAYSIA: PAHANG: Tanah Rata (N4.46 E101.40) ! ! ! ! Bristowia sp. [Gabon] JXZ363 GABON: Woleu-Ntem: Monts de Cristal, Tchimbélé (N0.62 to 0.63 E10.4) ! ! Cheliceroides sp. [China] d222 CHINA: Guangxi: Tianlin County, Langping Village (638) EU815524 Bodner 2009 (782) EU815579 Chinattus parvulus d009 USA: North Carolina (N35.341 W83.878) (755) EU815464 Bodner 2009 (965) EU815581 Cotinusa sp. MRB029 ECUADOR: Pichincha (N0.1493 W79.0317) Bodner 2009 Bodner 2009 Freya decorata d211 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) (757) EU815521 Bodner 2009 Bodner 2009 Galianora bryicola d124 ECUADOR: Napo: Estación Biológica Jatun Sacha (S1.067 W77.617) (756) DQ665771 Bodner 2009 (692) DQ665727 (972) DQ665758 Ghelna canadensis d005 USA: North Carolina (N35.70446 W82.37339) (744) DQ665767 Bodner 2009 Habronattus decorus d004 CANADA: Nova Scotia Bodner 2009 Bodner 2009 Habronattus decorus HA051 USA: Massachusetts: Dedham AF477279 Havaika sp. s127 USA: Hawaii (748)AY297252 (581)AF477249 (1047)AY297382 ! \"##! Species Voucher Locality 28S Actin 5C 16S-ND1 COI Heliophanus cupreus d044 POLAND: Mielik (N52.331 E23.042) (534) DQ665769 Bodner 2009 (975) DQ665756 Holcolaetis sp. d036 SOUTH AFRICA: Kwazulu-Natal (S28.2369 E32.4100) (659) DQ665770 Bodner 2009 (971) DQ665757 Hurius vulpinus s213 ECUADOR: Pichincha (743)AY297239 (393)AY297306; (578)AY296662 (1047)AY297368 Jollas sp. s162 ECUADOR: Sucumbios (734)AY297241 (393)AY297308; (580)AY296664 (1047)AY297370 Lyssomanes viridis d129 USA: Mississippi Bodner 2009 Lyssomanes viridis s160 USA: Florida (754)AY297231 (390)AY297297; (577)AY296652 (891)AY297360 Mantisatta longicauda s209 PHILIPPINES: Luzon (767)AY297270 (390)AY297333; (583)AY296689 (1047)AY297399 Nannenus lyriger d105 MALAYSIA: Pahang: Taman Negara (N4.381 E102.399) (777) EU815493 Bodner 2009 (898) EU815558 Neon nelli s310 USA: Massachusetts (748)AF327931 (390)AF328018; (578)AF327959 (1047)AF327988 Pachyballus sp. [S.Afr.] d141 AOUTH AFRICA: Kwazulu-Natal Province (S28.237 E32.410) (752) EU815505 Bodner 2009 Philaeus chrysops d025 ITALY: Calabria: Gozza (N38.413 E16.335) (720) EU815475 Bodner 2009 (925) EU815545 (958) EU815590 Plexippus paykulli MRB016 SINGAPORE: Nee Soon Swamp Forest (N1.39 E103.81) Bodner 2009 Bodner 2009 Bodner 2009 Bodner 2009 Portia labiata s206 PHILIPPINES: Luzon (752)AY297232 (387)AY297298; (594)AY296653 (960)AY297361 Salticus scenicus d003 CANADA: British Columbia: Mission (699) DQ665777 Bodner 2009 Bodner 2009 Salticus scenicus s107 USA: Washington, Missouri (393)AY297352; (579)AY296707 Thrandina parocula d123 ECUADOR: Morona Santiago (S2.9227 W78.4079) (773) DQ665779 Bodner 2009 (793) DQ665726 (970) DQ665761 Tomocyrba sp. MRB243 GABON: Tchimbélé, Woleu-Ntem, Monts de Cristal (N0.629 E10.404) Bodner 2009 Bodner 2009 Bodner 2009 Yllenus arenarius JXZ173 POLAND: Kozki Bodner 2009 Bodner 2009 Bodner 2009 Thomisidae Xysticus sp. s316 USA: Colorado (744)AY297296 Bodner 2009 (387)AY297359; (582)AY296714 (1047)AY297296 ! 501! Appendix 2. List of morphological characters scored for phylogenetic analyses. Character 1. Body shape: 0) normal; 1) beetle-like; 2) ant-like; 3) debris-like. Most euophryines are normal in the body form, whereas some develop mimicry to beetles (Coccorchestes), ants (Efate, Paraharmochirus and Sobasina) and debris (Leptathamas). Character 2. Raptorial first leg: 0) no; 1) yes (e.g. Figs 3.34A-B, H-I). The raptorial first leg refers to the feature typical in Diolenius and Ohilimia, in which metatarsi of the first legs are much slender and equipped with paired very short macrosetae, while tarsi are robust with many pairs of long and thick macrosetae, and trochanters are usually elongate. Althhough in “Thianitara” spectrum Simon, the trochanters of the first legs are not elongate, other features are very similar to those of Diolenius and Ohilimia, and thus scored as 1. Character 3. Carapace length to height ratio: 0) less than 2.5; 1) equal or more than 2.5. Character 4. Carapace hump at posterior part: 0) absent; 1) present. A small bump at the posterior part of carapace is exhibited in Paraharmochirus and some species of Sobasina (see Wanless 1978). Character 5. Carapace posterior end concave: 0) no; 1) yes (e.g. Figs 3.32B, I-J). More or less concaved posterior part of carapace is seen in Bulolia, Coccorchestes, Leptathamas, and Omoedus. Character 6. Male cheek expanded laterally: 0) no; 1) yes (e.g. Fig. 3.7L). The male carapace of some species of Agobardus and Corythalia expands laterally right behind ALEs. This feature is also found in males of Ascyltus, a euophryine genus from Australasia (Berry et al. 1997) but was not included in this study because of lacking the materials. Character 7. Carapace punctures: 0) absent; 1) present (e.g. Figs 3.36A-B, H-I). Small and pit- like punctures on the carapace are seen in Paraharmochirus and Sobasina. Character 8. Guanine deposit in eye area: 0) absent; 1) present (e.g. Figs 3.18A-B, I). The white appearance in the eye area of some euophryines, such as Chapoda spp. and Thorelliola spp., is 502! probably due to the guanine deposit in the prosomal section of the diverticula (Oxford & Gillespie 1998). Character 9. Male endite anterior-lateral small cusp: 0) absent; 1) present (e.g. Fig. 3.19D). Character 10. Male clypeus elongate setae: 0) relatively fine; 1) robust horns (Fig. 3.54A). Most euophryines only have similar fine setae as in females on the clypeus in males. However, males of some Thorelliola species have a pair of robust horn-like projections probably derived from the fine setae directly on the clypeus or on a basal truncus (Gardzinska & Patoleta 1997; Szüts & De Bakker 2004). Character 11. Eye rows: 0) 3 rows (less than half of ALE overlap with AME); 1) 4 rows (more than half of ALE overlap with AME, e.g. Figs 3.32I-J). Most euophryines have eyes arranged in three rows: the first eye row with AMEs and ALEs, the second with PMEs, and the third with PLEs. However, in some genera such as Bulolia and Leptathamas, their ALEs strongly move backward and inward which makes the eyes appear in four rows. The four-row arrangement of eyes also appears in Athamas, which is not included in this study. Character 12. Female ocular area length to PLEs width ratio: 0) < 1; 1) >= 1. Character 13. Female AMEs width and PMEs width: 0) AMEs narrower than PMEs; 1) AMEs as wide as or wider than PMEs. Character 14. Female PME-PLE distance to PLE diameter ratio: 0) <= 0.8; 1) > 0.8 and < 1.5; 2) >= 1.5. Character 15. Longitudinal fovea: 0) present; 1) absent. Character 16. Longitudinal fovea position: 0) around PLEs; 1) far behind PLEs. Character 17. Male chelicerae: 0) projecting ventrally; 1) projecting oblique forward (e.g. Fig. 3.8A); 2) projecting straight forward (e.g. Fig. 3.8H). 503! Character 18. Cheliceral size sexual dimorphism: 0) no; 1) yes. Character 19. Female cheliceral teeth size: 0) normal; 1) very small (e.g. Fig. 3.7D). Character 20. Female cheliceral promarginal tooth count: 0) 1 tooth; 1) 2 teeth; 2) 1 bicuspid tooth; 3) 3 teeth; 4) 4 teeth; 5) 5 teeth; 6) 6 teeth; 7) 7 teeth. Character 21. Female cheliceral retromarginal tooth count: 0) no tooth; 1) 1 tooth; 2) 1 bicuspid tooth; 3) multiple cusps; 4) 3 teeth. Character 22. Male cheliceral promarginal tooth count: 0) no tooth; 1) 1 tooth; 2) 2 teeth; 3) 1 bicuspid; 4) multiple cusps; 5) 3 teeth; 6) 4 teeth; 7) 5 teeth; 8) 6 teeth; 9) 8 teeth. Character 23. Male cheliceral retromarginal tooth count: 0) no tooth; 1) 1 tooth; 2) 2 teeth; 3) 1 bicuspid tooth; 4) multiple cusps; 5) 3 teeth; 6) 4 teeth. Character 24. Male cheliceral fang furrow: 0) normal; 1) widened. Character 25. Male cheliceral anterior surface longitudinal ridge: 0) absent; 1) present (e.g. Fig. 3.13E). Character 26. Male cheliceral posterior surface depression: 0) absent; 1) present (e.g. Figs 3.10K, Q). Character 27. Male cheliceral anterior surface projection: 0) absent; 1) present (e.g. Figs 3.8E, L). Character 28. Male cheliceral ectal surface projection: 0) absent; 1) present (e.g. Figs 3.9D-E). Character 29. Male cheliceral mesal margin concave: 0) no; 1) yes (e.g. Figs 3.9D-E). Character 30. Male fang spur: 0) absent; 1) present (e.g. Fig. 3.39E). 504! Character 31. Female longest leg: 0) 3rd leg longest; 1) 4th leg longest; 2) 1st leg longest. Character 32. Male longest leg: 0) 3rd leg longest; 1) 1st leg longest; 2) 4th leg longest. Character 33. Female longest leg length to cephalothorax length ratio: 0) <2; 1) >=2 and <=3; 2) >3. Character 34. Male leg I fringe: 0) absent; 1) present. Character 35. Male leg II fringe: 0) absent; 1) present. Character 36. Male leg III fringe: 0) absent; 1) present. Character 37. Male leg IV fringe: 0) absent; 1) present. Character 38. Female tibia I ventral macroseta number: 0) 0 pair; 1) 2 pairs or less; 2) 2.5 pairs; 3) 3 pairs (including 3.5p with one lateral moving ventrally); 4) 4 pairs; 5) 5 pairs; 6) more than 5 pairs. Character 39. Male tibia I ventral macroseta number: 0) 0 pair; 1) 2 pairs or less; 2) 2.5 pairs; 3) 3 pairs (including 3.5p with one lateral moving ventrally); 4) 4 pairs; 5) 5 pairs; 6) more than 5 pairs. Character 40. Female metartarsus I ventral macroseta number: 0) 0 pair; 1) 2 pairs (including 2.5p with one lateral moving ventrally); 2) 3 pairs (including 3.5p with one lateral moving ventrally); 3) 4 pairs; 4) more than 4 pairs. Character 41. Male metatarsus I ventral macroseta number: 0) 0 pair; 1) less than 2 pairs; 2) 2 or 2.5 pairs; 3) 3 pairs (including 3.5p with one lateral moving ventrally); 4) 4 pairs; 5) more than 4 pairs. 505! Character 42. Male abdominal dorsal scutum: 0) absent; 1) present (e.g. Figs 3.25A, E). In most cases that I scored as male abdominal dorsal scutum present, the scutum appears to be a relatively hard oval area rather than a fully sclerotized armor. Character 43. Male epiandrous spigots: 0) absent; 1) present. Character 44. Female prespiracular bump: 0) absent; 1) present. Character 45. Male prespiracular bump: 0) absent; 1) present. Some euophryine species have a bump in front of the male and/or female spiracle. The prespiracular bump appears to be either bare and sclerotized like in Antillattus gracilis Bryant, or less sclerotized and with many stiff setae like in Parabathippus shelfordi (Peckham & Peckham). Character 46. Embolus coil: 0) no; 1) equal or less than half a circle; 2) more than half a circle but no more than one and a half circle; 3) more than one and a half circle. Most euophryines have curved or coiled embolus, which has been suggested to be an important character to define the subfamily (Prószy!ski 1976; Maddison & Hedin 2003a). However, some species (e.g. Anasaitis spp., Sobasina spp., Soesilarishius spp., Tylogonus spp., etc.) have short and not curved embolus, which is probably due to the reversal evolution. Character 47. Tegular lobe over tibia: 0) absent (e.g. Figs 3.12C, I); 1) present (e.g. Figs 3.7C, J, Q). Tegular lobe refers to the proximal projection of the tegulum usually over palpal tibia, and appears to be quite common within euophryines. In the cases that the lobe is present but not very obvious, this character was scores as 1 (e.g. Antillattus darlingtoni Bryant, Fig. 3.9C). Character 48. Palpal tibia ventral bump: 0) absent (e.g. Figs 3.8C, J); 1) present (e.g. Figs 3.6C, J, Q). This feature refers to a small ventral bump-like process at the distal end of the palpal tibia. Character 49. Lamella on tegular shoulder: 0) absent; 1) present (e.g. Figs 3.43C-D, H-I). This feature refers to the small distal process at the retrolateral shoulder of the tegulum common in the genus Saitis. Different names, such as “anterior lamella” (Zabka & Pollard 2003), “proximal tegular lobe” (Richardson & Zabka 2007) and “anterior tegular bulge” (Waldock 1995), have 506! been used for this character in the literatures. Similar structure is also seen in the genus Sondra of Astioida (Davies & Zabka 1989), which may represent a convergent evolution. Character 50. Palpal femur shape: 0) straight; 1) curved (e.g. Fig. 3.30N). In the lineages with long and developed male chelicerae, the palpal femora are usually also elongate and bent (e.g. Bathippus, Canama, etc.). Character 51. Palpal femur dorsal macrosetae: 0) no macrosetae; 1) 1 macroseta; 2) 2 macrosetae; 3) 3 macrosetae; 4) 4 macrosetae; 5) 5 macrosetae; 6) 6 macrosetae; 7) 8 macroseate or more. Character 52. Palpal femur dorsal long hairs: 0) absent; 1) present. Character 53. Palpal patella and tibia length dorsally: 0) patella longer than tibia; 1) patella shorter or equal to tibia. Character 54. Palp tibia and cymbium length dorsally: 0) tibia shorter than cymbium; 1) tibia longer or equal to cymbium. Character 55. RTA shape: 0) simple, long and finger-like (e.g. Fig. 3.7J); 1) complex (Fig. 3.20D); 2) simple and small (hook, e.g. Fig. 3.28C). Most euophryines have RTAs that are finger-like. However, some species have RTAs rather wide (e.g. Ohilimia scutellata), or even with multiple apophyses (e.g. Amphidraus complexus); and some species have very small and hook-like RTAs (e.g. Popcornella nigrimaculata). The RTA is completely missing in Talavera Peckham & Peckham (Logunov and Kronestedt 2003), and was scored here as “-” (inapplicable). Character 56. Sperm duct loop on retrolateral side of bulb: 0) present (e.g. Fig. 3.6C); 1) absent (e.g. Fig. 3.15H). Most euophryines have a sperm duct loop on the retrolateral side of the palpal bulb. And it has been considered as an important character to distinguish euophryines from other salticoids (Maddison & Hedin 2003a). However the retrolateral sperm duct loop is missing in some genera such as Bulolia, Coccorchestes, Sobasina, Marma, Neonella etc., which may be secondary lost in these lineages. Besides, similar sperm duct loop is also present in the 507! subfamily Dendryphantinae (e.g. Macaroeris, see Prószy!ski 2010 online database) and Ballinae (e.g. Leikung, see Benjamin 2004), which probably results from convergent evolution. Character 57. Sperm duct loop on proximal or prolateral proximal side of bulb: 0) absent; 1) present (e.g. Fig. 3.15H). The sperm duct loop on the proximal or prolateral proximal side of the tegulum is quite common in Ballinae (Benjamin 2004), and it is also present in a few lineages of Euophryinae (e.g. Coccorchestes, Bulolia, Leptathamas, etc.). Character 58. Retrolateral sperm duct loop width: 0) less than half of bulb width; 1) about half of bulb width; 2) more than half of bulb width. Character 59. Apophysis on palpal femur: 0) absent; 1) present (e.g. Figs 3.18D, K). Character 60. Apophysis on palpal patella: 0) absent; 1) present (e.g. Fig. 3.14B). Character 61. Palpal tibia prolateral apophysis: 0) absent; 1) present (e.g. Fig. 3.54C). It refers to a protrusion on the prolateral side of the palpal tibia, sometimes armed with a large macrosetae like in some species of Thorelliola. Character 62. Palpal tibia macrosetae: 0) absent; 1) present (e.g. Figs 3.12I-J). Character 63. Embolus position: 0) distal (e.g. Fig. 3.7J); 1) prolateral (e.g. Fig. 3.34J); 2) proximal (e.g. Fig. 3.34C). Most euophryines have emboli located at the distal end of the bulb. However, in some lineages the emboli either come from the prolateral side (e.g. Ohilimia) or from the proximal end of the bulb (e.g. Diolenius). Character 64. Plane of spiral of embolus: 0) parallel to longitudinal axis of bulb (e.g. Fig. 3.8C); 1) perpendicular to longitudinal axis of bulb (e.g. Fig. 3.30L). The spiral of the embolus with its plane parallel to the longitudinal axis of the bulb has been proposed to be an important character to define the subfamily Euophryinae (Maddison & Hedin 2003). However, some lineages of Euophryinae, e.g. Canama and Coccorchestes, have the plane of the spiral of embolus more perpendicular to the longitudinal axis of the bulb. 508! Character 65. Lamella along embolus: 0) absent; 1) present. The feature refers to an independent process along the embolus found in several lineages in Euophryinae, e.g. Sidusa (Fig. 3.16C), Neonella (Fig. 3.23C), Pristobaeus (Fig. 3.37C), Colyttus (Fig. 3.45Q). The term “conductor” has been applied to this structure in the literature (Davies & Zabka 1989; Bodner 2002). Although “conductor” has been widely used in the other spider families for the male palpal structure, it usually refers to an apophysis associated with the tegulum (Coddington 1990), and thus is not homologous with the feature I refer here. Hence, I prefer the term “lamella along embolus” to avoid any confusion. Character 66. Embolic disc: 0) present (e.g. Fig. 3.7J); 1) very reduced or absent (e.g. Fig. 3.6C). It refers to the expanded sclerite at the end of distal hematodocha where the embolus usually comes from (e.g. Fig. 3.58C). The embolic disc is very reduced or even completed lost in a few lineages of Euopheyinae, e.g. Anasaitis, Tylogonus, Talavera (see figs. 23-25 in Logunov & Kronestedt 2003), etc. Character 67. Process on embolic disc: 0) absent; 1) present. It refers to a large apophysis from the embolic disc independent from the embolus found in Amphidraus (Fig. 3.20C), Marma (Fig. 3.20I) and Laufeia eximia. Character 68. Plane of embolic disc: 0) dorsal of plane of front of tegulum (e.g. Fig. 3.7J); 1) ventral of plane of front of tegulum. All euophryines have the plane of embolic disc at the dorsal of the plane of the front surface of the tegulum except in Thiania (Figs 3.47C, I) and Colyttus (Figs 3.45C, J, Q). Character 69. Retrolateral cymbial extension: 0) absent; 1) present. Character 70. Distal hematodocha: 0) developed; 1) reduced; 2) no trace. Character 71. Salticid radix: 0) present; 1) absent (not distinguishable from distal hematodocha); 2) fused with tegulum. Salticid radix is a feature that recently found widely present in Hispaninae and Salticoida. It is common in euophryines and usually appears as a separate sclerite on the distal hematodocha between the embolic disc and the tegulum. Sometimes it rotates to the back of the bulb and thus is not visible unless the palp is expended. 509! In a few lineages, no distinct sclerite is noticed on the distal hematodocha, e.g. Chapoda peckhami, and scored as “1”; in Tylogonus, the radix seems to become fused to the tegulum and distinguishable from the latter by the much narrowed sperm duct in the radix, and then scored as “2”. Character 72. Position of copulatory opening in relation to vulva: 0) anterior (e.g. Fig. 3.13P); 1) median (e.g. Fig. 3.8G); 2) posterior (e.g. Fig. 3.7N). Character 73. Fertilization duct position: 0) at base of copulatory duct (e.g. Fig. 3.40K); 1) far away from copulatory duct (e.g. Fig. 3.20L). Character 74. Epigynal median septum of window: 0) present (e.g. Fig. 3.7M); 1) absent. Most euophryines have a “window”-like structure (see Character 80 below) separated by a septum in the middle. However, the septum is absent in some lineages of euophryines, e.g. Thorelliola (Fig. 3.54G), Emathis (Fig. 3.48E), etc.. In the cases where the “window” is absent, this character is scored as “-”. Character 75. Spermatheca size: 0) obviously large (e.g. Fig. 3.7N); 1) small as copulatory duct (e.g. Fig. 3.36N). Character 76. Number of spermathecae: 0) 1 pair (e.g. Fig. 3.12F); 1) 2 pairs (e.g. Fig. 3.21G). In addition to the primary spermathecae, some euophryines have a pair of secondary spermathecae, of which the exact function is still unclear. In cases when the copulatory ducts have apparent swollens, e.g. Cobanus, or have a large and separate sacs, e.g. Coryphasia physonycha (Simon), this character was scored as “1”. Character 77. Opening with spiral guide: 0) no (e.g. Fig. 3.16F); 1) yes (e.g. Fig. 3.16S). Character 78. Barrier ridge at opening: 0) absent; 1) present. Character 79. Window length: 0) occupying much less than 1/2 of epigynal plate; 1) occupying about 1/2 of epigynal plate; 2) occupying much more than 1/2 of epigynal plate. Here I refer the 510! epigynal plate as the area from the anterior end of the window or the vulva whichever is anterior most, to the epigynal furrow. Character 80. Window of epigynum: 0) present (e.g. Fig. 3.7M); 1) absent (e.g. Fig. 3.6F). The “window” refers to the circular area(s) of relatively transparent and flat integument, which is common in euophryines. In a few lineages of euophryines, e.g. Anasaitis (e.g. Figs 3.6F, M), Talavera (Fig. 3.41I; see figures in Logunov & Kronestedt 2003), no obvious window is present. Character 81. Primary spermatheca shape: 0) spherical or ovoid (e.g. Fig. 3.7N); 1) kidney- shaped (e.g. Fig. 3.41E); 2) irregular shaped; 3) narrow coiled (e.g. Fig. 3.31Q). Character 82. Primary spermatheca position in relation to translucent window: 0) anterior to window (e.g. Fig. 3.33L); 1) within window (e.g. Fig. 3.7N); 2) posterior to window (e.g. Fig. 3.13O).! ! \"##! Appendix 3. Scored morphological matrix. “-” indicates the character is not applicable in the species; “?” indicates the character state is ambiguous in the species. 1 . B o d y s h a p e 2 . R a p to r ia l fi r st l e g 3 . C a r a p a c e l e n g th t o h e ig h t r a ti o 4 . C a r a p a c e h u m p a t p o st e r io r p a r t 5 . C a r a p a c e p o st e r io r e n d c o n c a v e 6 . M a le c h e e k e x p a n d e d l a te r a ll y 7 . C a r a p a c e p u n c tu r e s 8 . C a r a p a c e g u a n in e d e p o si t in e y e a r e a 9 . M a le e n d it e a n te r io r -l a te r a l sm a ll c u sp 1 0 . M a le c ly p e u s e lo n g a te s e ta e 1 1 . E y e r o w s 1 2 . F e m a le e y e a r e a l e n g th t o P L E s w id th r a ti o 1 3 . F e m a le A M E s w id th a n d P M E s w id th 1 4 . F e m a le P M E -P L E d is ta n c e t o P L E d ia m e te r r a ti o 1 5 . L o n g it u d in a l fo v e a 1 6 . L o n g it u d in a l fo v e a p o si ti o n 1 7 . M a le c h e li c e r a 1 8 . C h e li c e r a l si z e s e x u a l d im o r p h is m 1 9 . F e m a le c h e li c e r a l te e th s iz e 2 0 . F e m a le c h e li c e r a l p r o m a r g in a l to o th c o u n t 2 1 . F e m a le c h e li c e r a l r e tr o m a r g in a l to o th c o u n t 2 2 . M a le c h e li c e r a l p r o m a r g in a l to o th c o u n t 2 3 . M a le c h e li c e r a l r e tr o m a r g in a l to o th c o u n t 2 4 . M a le c h e li c e r a l fa n g f u r r o w 2 5 . M a le c h e li c e r a l a n te r io r s u r fa c e l o n g it u d in a l r id g e 2 6 . M a le c h e li c e r a l p o st e r io r s u r fa c e d e p r e ss io n 2 7 . M a le c h e li c e r a l a n te r io r s u r fa c e p r o je c ti o n 2 8 . M a le c h e li c e r a l e c ta l su r fa c e p r o je c ti o n 2 9 . M a le c h e li c e r a l m e sa l m a r g in c o n c a v e 3 0 . M a le f a n g s p u r 3 1 . F e m a le l o n g e st l e g 3 2 . M a le l o n g e st l e g 3 3 . F e m a le l o n g e st l e g l e n g th t o c e p h a lo th o r a x l e n g th r a ti o 3 4 . M a le l e g I f r in g e 3 5 . M a le l e g I I fr in g e 3 6 . M a le l e g I II f r in g e 3 7 . M a le l e g I V f r in g e 3 8 . F e m a le t ib ia I v e n tr a l m a c r o se ta n u m b e r 3 9 . M a le t ib ia I v e n tr a l m a c r o se ta n u m b e r 4 0 . F e m a le m e ta r ta r su s I v e n tr a l m a c r o se ta n u m b e r 4 1 . M a le m e ta ta r su s I v e n tr a l m a c r o se ta n u m b e r Ghelna canadensis 0 0 0 0 0 0 0 0 1 0 0 0 0 2 0 0 0 0 0 1 1 2 1 0 1 1 0 0 0 0 1 ? 1 0 0 0 0 3 3 1 2 Heliophanus cupreus 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 2 1 0 0 1 0 0 0 0 1 2 1 0 0 0 0 1 1 1 2 \"Bathippus\" pahang 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Chinattus parvulus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 ? 1 0 0 ? 0 3 3 1 2 Freya decorata 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Plexippus paykulli 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Salticus scenicus 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 1 1 2 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 Agobardus bahoruco 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 2 2 4 1 0 0 0 0 0 1 1 1 1 1 0 0 0 4 3 2 3 Agobardus cf. anormalis montanus 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1 1 0 1 2 2 3 1 0 0 0 0 0 1 1 1 1 0 0 0 0 3 3 2 3 Agobardus cf. brevitarsus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 2 3 Agobardus cordiformis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 2 3 Agobardus gramineus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 2 3 Agobardus oviedo 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 1 0 0 0 0 0 1 1 2 1 0 0 0 0 3 3 2 3 Agobardus phylladiphilus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 2 3 Amphidraus complexus 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 1 0 0 0 1 3 2 4 0 0 0 0 0 0 0 1 2 0 0 0 0 0 3 3 1 2 ! \"#$! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Anasaitis adorabilis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 0 1 0 1 1 0 0 3 2 1 2 Anasaitis banksi 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 0 0 1 1 1 1 0 3 3 1 2 Anasaitis brunnea 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 3 3 1 2 Anasaitis canosa [USA] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 1 1 0 1 1 0 0 2 2 1 2 Anasaitis cf. canalis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 1 0 1 1 1 0 0 3 3 1 2 Anasaitis elegantissima 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 1 0 1 1 1 1 0 1 1 1 2 Anasaitis gloriae 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 1 0 0 0 1 0 0 1 1 0 0 0 0 3 3 1 2 Anasaitis hebetata 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 3 1 0 1 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 2 ? 2 Anasaitis laxa 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 ? ? 1 0 0 0 0 1 1 1 2 Anasaitis locuples 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 1 0 1 1 0 1 1 1 2 Anasaitis placida 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 1 0 0 0 1 0 0 1 0 1 1 0 0 3 3 1 2 Anasaitis sp. [Peblique] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 1 2 Antillattus cambridgei 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 1 3 2 6 0 0 0 1 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Antillattus cf. applanatus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 2 1 0 1 0 0 0 1 0 1 1 1 1 0 0 0 3 3 1 2 Antillattus darlingtoni 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 2 1 0 0 0 1 1 1 0 1 1 1 1 0 0 0 3 3 1 2 Antillattus gracilis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 3 3 5 4 0 1 0 1 0 0 0 1 2 1 1 0 0 0 3 3 2 3 Antillattus maxillosus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 3 1 0 1 0 0 1 0 0 1 1 1 1 0 0 0 3 3 1 2 Antillattus scutiformis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 1 0 0 0 1 0 1 2 1 1 1 1 1 3 3 1 2 Bathippus directus [Tualapa] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 1 1 6 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 3 3 3 4 Bathippus gahavisuka 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 1 5 1 0 0 0 1 0 0 1 0 1 1 0 0 0 0 3 3 3 4 Bathippus korei 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 7 1 9 1 1 0 0 0 0 0 1 0 1 1 0 0 0 0 3 3 3 4 Bathippus macrognathus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 1 5 1 0 0 0 0 0 0 1 0 1 1 1 0 0 0 3 3 3 4 Bathippus madang 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 1 1 ? ? ? 1 1 1 0 0 1 0 0 1 ? 1 ? 0 0 0 0 ? 3 ? 4 Belliena ecuadorica 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Bulolia excentrica 0 0 0 0 1 0 0 0 0 0 1 1 1 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 2 1 1 0 0 0 0 3 3 1 2 Bythocrotus cf. crypticus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 1 1 0 0 0 0 2 1 1 0 0 0 0 3 3 1 2 Bythocrotus crypticus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 1 1 0 0 0 0 2 1 1 0 0 0 0 3 3 1 2 Canama cf. forceps 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 1 1 ? ? ? 2 3 0 1 0 1 0 0 1 ? ? ? 0 ? 0 0 ? 3 ? 3 Canama extranea 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 3 3 2 3 Canama fimoi 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 1 0 0 0 1 0 0 0 0 1 1 0 0 0 0 3 3 2 3 Canama hinnulea 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 1 1 ? ? ? 2 3 0 1 0 1 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 3 Canama triramosa 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 2 2 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 3 3 1 2 ! \"#$! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Petemathis minuta 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 3 2 4 1 0 1 1 0 0 0 1 2 1 1 0 0 0 3 3 1 2 Petemathis portoricensis [Adjuntas] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 3 2 4 0 0 1 0 0 0 0 0 1 1 0 0 0 0 3 3 1 2 Petemathis portoricensis [Maricao] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 3 2 4 0 0 1 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Petemathis tetuani 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 4 0 0 1 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 cf. Coryphasia sp. [Brazil] 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 3 0 1 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 cf. Maeota sp. [Panama] 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Chalcolecta prensitans 0 0 1 0 0 ? 0 1 ? ? 0 0 0 2 0 0 ? ? 0 1 2 ? ? ? ? ? ? ? ? ? 2 ? 2 ? ? ? ? 6 ? 2 ? Chalcolemia nakanai 0 0 1 0 0 ? 0 0 ? ? 0 1 0 1 1 - ? ? 0 1 3 ? ? ? ? ? ? ? ? ? 2 ? 2 ? ? ? ? 6 ? 3 ? Chalcoscirtus diminutus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 3 2 1 2 Chalcoscirtus infimus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 - 0 0 0 1 0 2 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 3 1 1 2 Chalcotropis luceroi 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 1 1 ? ? ? 2 3 0 1 0 1 0 0 0 ? 0 ? 0 0 0 0 ? 4 ? 4 Chalcotropis cf. caeruleus 0 0 0 0 0 0 0 0 0 ? 0 ? ? ? 0 0 0 0 ? ? ? 3 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 3 Chapoda angusta 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 0 2 1 0 0 0 0 3 3 1 2 Chapoda cf. inermis [CostaRica] 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 1 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Chapoda cf. inermis [Panama] 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 1 0 0 0 0 0 1 2 1 1 0 0 0 3 3 1 2 Chapoda fortuna [Panama] 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Chapoda gitae 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 1 3 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Chapoda montana 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Chapoda peckhami 0 0 0 0 0 0 0 1 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 3 0 1 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Chapoda recondita 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 1 0 0 0 1 0 1 2 0 0 0 0 0 3 3 1 2 Chinophrys pengi 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 4 3 8 4 0 0 0 0 0 0 0 1 2 1 1 0 0 0 3 3 1 2 Coccorchestes cf. aiyura 1 0 0 0 1 0 1 0 1 0 0 0 0 1 1 - 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 1 1 2 Coccorchestes cf. ildikoae 1 0 0 0 1 0 1 1 1 0 0 ? ? ? 1 - 0 0 ? ? ? 2 3 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 1 ? 2 Coccorchestes cf. inermis 1 0 0 0 1 0 1 0 1 0 0 0 0 1 1 - 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 0 1 Coccorchestes clavifemur 1 0 0 0 1 0 1 0 1 0 0 ? ? ? 1 - 0 0 ? ? ? 6 3 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 1 ? 2 Colyttus bilineatus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 3 3 1 2 Colyttus striatus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 1 0 0 0 0 0 1 1 0 0 0 0 3 3 1 2 Compsodecta haytiensis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Compsodecta peckhami 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 1 0 1 0 0 0 0 1 1 0 0 0 0 3 3 1 2 Corticattus latus 0 0 1 0 0 0 0 0 1 0 0 1 0 1 0 1 0 0 0 2 1 3 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 0 1 0 Coryphasia cf. campestrata 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 1 0 0 0 0 0 1 2 1 1 0 0 0 4 4 3 4 Coryphasia fasciiventris 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 1 0 0 0 0 0 1 1 1 0 0 0 0 3 3 2 3 ! \"#$! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Coryphasia physonycha 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 1 1 1 0 0 0 4 4 3 4 Corythalia bicincta 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 1 2 1 1 1 1 0 1 1 1 2 Corythalia broccai 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 1 0 0 0 0 0 0 1 1 1 1 0 0 3 3 2 3 Corythalia bromelicola 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 2 3 Corythalia coronai 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 1 0 0 0 0 0 1 2 1 1 1 0 0 3 3 2 3 Corythalia decora 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 1 0 0 0 0 0 1 2 1 1 1 0 0 3 3 2 3 Corythalia electa 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 1 2 Corythalia minor 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 1 0 0 0 1 0 0 1 1 1 0 0 0 3 3 2 3 Corythalia peblique 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 1 0 0 0 0 0 1 1 1 1 1 0 0 3 3 2 3 Corythalia porphyra 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 ? 0 0 1 1 1 1 3 3 1 2 Corythalia sulfurea 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 2 1 3 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Cytaea mitellata 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 6 3 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Cytaea nimbata 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 3 2 5 3 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Cytaea oreophila 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 4 2 6 3 0 0 0 0 0 0 0 1 0 1 0 0 0 0 3 3 1 2 Diolenius varicus 0 1 0 0 0 0 0 0 0 0 0 1 0 2 1 - 0 0 0 1 3 2 4 0 0 0 0 0 0 0 2 1 2 1 0 0 0 6 6 4 5 Ecuadattus napoensis 0 0 0 0 0 0 0 1 0 0 0 ? ? ? 0 0 0 0 ? ? ? 3 1 0 1 1 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Ecuadattus pichincha 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 1 3 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Efate albobicinctus 2 0 1 0 0 0 0 0 0 0 0 ? ? ? 1 - 0 0 ? ? ? 1 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 5 ? 3 Emathis gombak 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 3 3 2 4 0 0 0 0 0 0 0 0 0 1 0 0 0 0 3 3 3 4 Euophryine sp. [GentingHighlands] 0 0 0 0 0 ? 0 0 ? ? 0 0 0 0 0 0 ? ? 0 1 2 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 2 ? Euophrys frontalis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 1 0 0 0 3 3 1 2 Euophrys monodnock 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 1 1 0 0 3 3 1 2 Euryattus bleekeri 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 5 2 5 3 0 1 0 0 0 0 0 2 1 1 0 0 0 0 3 3 1 2 Euryattus sp.1 [Gahavisuka] 0 0 0 0 0 0 0 0 ? ? 0 0 0 1 0 0 ? ? 0 6 2 ? ? ? ? ? ? ? ? ? 2 ? 1 ? ? ? ? 3 ? 1 ? Euryattus sp.2 [Gahavisuka] 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 7 3 0 1 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Foliabitus longzhou 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 1 0 0 0 0 0 2 0 1 1 1 0 0 4 4 1 2 Foliabitus sp. [Malaysia] 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? 1 ? 1 1 ? ? ? 3 ? 2 Ilargus coccineus 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Ilargus foliosus 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Ilargus galianoae 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Ilargus macrocornis 0 0 0 0 0 0 0 1 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 1 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Ilargus moronatigus 0 0 0 0 0 0 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 1 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 1 ? ! \"#\"! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Ilargus pilleolus 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Ilargus serratus 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Lagnus edwardsi 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 3 2 4 0 1 1 0 0 0 0 1 1 1 0 0 0 0 6 6 4 5 Laufeia daiqini 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 2 3 3 0 0 0 0 0 0 0 1 1 0 1 1 0 0 3 3 1 2 Laufeia eximia 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 1 2 2 3 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 1 2 Laufeia keyserlingi 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 2 3 3 0 1 0 0 0 0 0 1 1 0 0 0 0 0 1 1 1 2 Lepidemathis haemorrhoidalis 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 3 2 4 0 1 0 0 0 0 0 0 1 1 0 0 0 0 3 3 3 4 Leptathamas paradoxus 3 0 0 0 1 0 0 0 1 0 1 1 1 2 1 - 0 0 0 1 2 2 3 0 0 0 1 0 0 0 1 1 1 1 0 0 0 3 3 1 2 Maeota dichrura 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Maeota dorsalis 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Maeota flava 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Maeota simoni 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota sp. [JatunSacha] 0 0 0 0 0 0 0 1 ? ? 0 0 0 1 0 0 ? ? 0 1 1 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 1 ? Maeota sp. [Manabi] 0 0 0 0 0 0 0 1 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? ? ? ? 0 0 0 ? ? ? ? Maeota sp. [MoronaSantiago] 0 0 0 0 0 0 0 1 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 3 ? 2 Maeota sp. [Napo] 0 0 0 0 0 ? 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 1 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 1 ? Maeota tuberculotibiata 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Marma nigritarsis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 3 2 4 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 2 1 2 Mexigonus arizonensis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 2 1 0 1 0 0 0 1 0 1 1 1 0 0 0 0 3 3 1 2 Mexigonus morosus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Mopiopia cf. bruneti 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Naphrys pulex 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 1 1 1 2 Neonella vinnula 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 ? 1 2 Ohilimia scutellata 0 1 0 0 0 0 0 0 0 0 0 0 0 2 1 - 0 0 0 1 2 2 3 0 0 0 0 0 0 0 2 1 2 1 0 0 0 6 6 2 3 Omoedus brevis 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 3 1 0 1 0 0 0 0 0 ? ? ? ? 0 ? 0 ? 2 ? 2 Omoedus cf. danae 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 3 3 0 1 0 0 0 1 0 0 0 1 1 0 0 0 3 3 1 2 Omoedus cf. piceus 0 0 0 0 1 0 0 0 0 0 0 ? ? ? 1 - 0 0 ? ? ? 3 1 0 0 0 0 0 0 0 ? 2 ? 0 0 0 0 ? 1 ? 1 Omoedus cf. semirasus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 1 0 1 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Omoedus cf. torquatus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 3 3 0 1 0 0 0 1 0 0 0 1 1 0 0 0 3 3 1 2 Omoedus darleyorum 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 1 0 0 0 1 0 1 1 1 0 0 0 0 3 ? 2 ? Omoedus ephippigera 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 - 0 0 1 2 1 3 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 1 1 2 Omoedus meyeri 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 1 3 1 0 1 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 ! \"#$! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Omoedus omundseni 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 1 0 1 0 1 0 1 1 1 0 0 0 0 3 3 1 2 Omoedus orbiculatus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 1 ? 0 0 0 0 1 2 1 0 0 0 0 1 2 1 2 Omoedus papuanus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 2 1 0 1 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Omoedus swiftorum 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 1 3 1 0 1 0 1 0 1 0 1 2 1 0 0 0 0 3 3 1 2 Parabathippus cf. macilentus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 1 5 2 0 0 0 0 0 0 0 0 ? 1 0 0 0 0 3 3 1 2 Parabathippus cuspidatus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 1 5 2 0 0 0 0 1 0 0 0 0 1 1 1 0 0 3 3 1 2 Parabathippus kiabau 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 1 1 ? ? ? 5 5 0 0 0 0 0 0 0 ? 0 ? 1 0 0 0 ? 3 ? 2 Parabathippus magnus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 5 2 0 0 0 0 0 0 0 0 0 2 1 1 0 0 3 3 1 2 Parabathippus shelfordi 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 1 5 1 0 0 0 0 0 0 1 0 0 2 1 0 0 0 3 3 2 3 Paraharmochirus tualapaensis 2 0 1 1 0 0 1 0 0 0 0 0 0 2 1 - 0 0 0 1 2 2 3 0 1 0 0 0 0 0 2 1 0 1 0 0 0 6 6 2 3 Parvattus zhui 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 1 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 0 ? 0 0 0 0 ? 3 ? 2 Pensacola signata 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 1 0 1 1 0 0 0 0 1 1 0 0 0 3 3 1 2 Phasmolia elegans 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 3 2 5 3 0 0 0 0 0 0 0 1 0 1 0 0 0 0 5 4 2 3 Popcornella furcata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 2 1 0 0 0 0 0 3 3 1 2 Popcornella nigromaculata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 ? 1 0 0 0 0 0 3 3 1 2 Popcornella spiniformis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Pristobaeus beccarii 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 3 3 1 2 Pristobaeus cf. jocosus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 4 2 5 0 1 0 0 0 0 0 0 1 1 0 0 0 0 3 3 1 2 Pseudeuophrys erratica 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 1 1 2 1 0 1 0 0 0 1 0 1 1 1 0 0 0 0 3 3 1 2 Saitis auripes 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 ? 0 0 0 0 1 2 1 1 0 0 0 3 3 1 2 Saitis barbipes 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 1 0 1 0 0 1 0 3 3 1 2 Saitis cf. fuscus 0 0 0 0 0 ? 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 2 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 2 ? Saitis cf. griseus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 3 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Saitis sp. [NewSouthWales] 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 3 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Saitis sp. [SouthAustralia] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 1 ? 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 3 2 1 2 Saphrys a-notata 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 2 3 1 2 Saphrys cf. patagonica 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Servaea vestita 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 4 3 0 1 0 1 0 1 0 ? 1 ? 0 0 0 0 ? 2 ? 2 Sidusa extensa 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 2 2 2 0 0 0 0 0 0 0 1 1 1 1 0 0 0 4 4 2 3 Sidusa sp. [CostaRica] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 1 1 1 1 1 0 4 4 2 3 Sidusa sp.1 [FrenchGuiana] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 ? ? 1 0 0 0 0 4 4 2 3 Sidusa sp.2 [FrenchGuiana] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 0 1 1 0 0 0 4 4 2 3 ! \"#$! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Sidusa unicolor 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 2 2 2 0 0 0 0 0 0 0 1 1 1 1 0 0 0 4 4 2 3 Sobasina wanlessi 2 0 1 1 0 0 1 0 0 0 0 1 0 2 1 - 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 1 1 1 0 0 0 5 5 2 3 Soesilarishius cf. amrishi 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Soesilarishius minutus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 0 1 0 0 0 0 3 3 1 2 Soesilarishius ruizi 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 2 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 3 3 1 2 Talavera minuta 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 - 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 2 1 1 2 Thiania bhamoensis 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 2 1 3 1 0 0 0 0 0 0 0 2 1 0 0 0 0 0 3 3 1 2 Thiania latibola 0 0 1 0 0 0 0 0 0 0 0 ? ? ? 0 1 0 0 ? ? ? 3 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 3 Thiania spectrum 0 1 1 0 0 0 0 0 0 0 0 ? ? ? 0 1 0 ? ? ? ? 2 3 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 5 ? 1 Thiania tenuis 0 ? 1 0 0 0 0 0 0 0 0 ? ? ? 0 1 0 0 ? ? ? 3 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? ? ? ? Thorelliola aliena 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 2 3 0 1 0 1 0 0 0 1 1 1 0 0 0 0 4 4 3 4 Thorelliola cf. mahunkai 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 3 2 4 0 0 0 1 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Thorelliola crebra 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 0 1 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Thorelliola ensifera 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 3 2 4 0 0 0 1 0 0 0 1 1 0 1 1 0 0 3 3 1 2 Thorelliola Joannae 0 0 0 0 0 ? 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 3 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 1 ? Thorelliola tamasi 0 0 0 0 0 0 0 0 0 1 0 ? ? ? 0 0 0 0 ? ? ? 2 4 0 0 0 1 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 3 Thorelliola tualapa 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 3 2 4 0 0 0 1 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Thyenula cf. aurantiaca 0 0 0 0 0 ? 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 1 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 1 ? Thyenula cf. mundus 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? ? ? 1 1 ? ? ? 3 ? 2 Thyenula laxa 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Thyenula leighi 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 1 0 0 0 0 0 ? ? ? 0 0 ? ? ? 3 ? 2 Thyenula nelshoogte 0 0 0 0 0 ? 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 1 ? ? ? ? ? ? ? ? ? 1 ? 0 ? ? ? ? 3 ? 1 ? Thyenula sp. [SouthAfrica] 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 ? 1 1 ? 0 0 3 3 1 2 Thyenula wesolowskae 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 3 3 1 2 Truncattus cachotensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 2 3 0 0 1 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Truncattus dominicanus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 2 3 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 3 1 2 Truncattus flavus 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 2 2 3 0 0 1 0 0 0 0 1 1 0 1 0 0 0 3 3 1 2 Tylogonus cf. auricapillus 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 1 0 0 ? ? ? 2 1 0 0 0 1 0 1 0 ? 1 ? 0 0 0 0 ? 4 ? 2 Tylogonus yanayacu 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 2 1 0 0 0 1 0 1 0 1 2 1 0 0 0 0 3 4 1 2 Variratina minuta 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 2 1 0 0 0 0 0 0 0 1 2 1 0 0 0 0 3 ? 1 2 Viribestus suyanensis 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 3 0 1 0 1 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Xenocytaea agnarssoni 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 1 2 1 0 0 0 0 0 0 0 0 2 1 0 0 0 0 2 3 1 2 ! \"#$! 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 2 7 2 8 2 9 3 0 3 1 3 2 3 3 3 4 3 5 3 6 3 7 3 8 3 9 4 0 4 1 Xenocytaea albomaculata 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? ? 2 1 0 0 0 0 0 0 0 ? 0 ? 0 0 0 0 ? 3 ? 2 Xenocytaea proszynskii 0 0 0 0 0 0 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 1 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 3 ? 1 ? Zabkattus brevis 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 2 2 3 0 0 0 1 0 0 0 1 1 1 0 0 0 0 3 3 1 2 Zabkattus furcatus 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 3 0 0 0 1 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 3 Zabkattus richardsi 0 0 0 0 0 0 0 0 1 0 0 ? ? ? 0 0 0 0 ? ? ? 2 3 0 0 0 1 0 0 0 ? 1 ? 0 0 0 0 ? 3 ? 2 Zabkattus trapeziformis 0 0 0 0 0 ? 0 0 ? ? 0 0 0 1 0 0 ? ? 0 1 2 ? ? ? ? ? ? ? ? ? 2 ? 1 ? ? ? ? 3 ? 1 ? ! \"#$! 4 2 . M a le a b d o m in a l d o r sa l sc u tu m 4 3 . M a le e p ia n d r o u s sp ig o ts 4 4 . F e m a le p r e sp ir a c u la r b u m p 4 5 . M a le p r e sp ir a c u la r b u m p 4 6 . E m b o lu s c o il 4 7 . T e g u la r l o b e o v e r t ib ia 4 8 . P a lp a l ti b ia b u m p 4 9 . L a m e ll a o n t e g u la r s h o u ld e r 5 0 . P a lp a l fe m u r s h a p e 5 1 . P a lp a l fe m u r d o r sa l m a c r o se ta e 5 2 . P a lp a l fe m u r l o n g h a ir s 5 3 . P a lp a l p a te ll a a n d t ib ia l e n g th d o r sa ll y 5 4 . P a lp t ib ia a n d c y m b iu m l e n g th d o r sa ll y 5 5 . R T A s h a p e 5 6 . S p e r m d u c t lo o p o n r e tr o la te r a l si d e o f te g u lu m 5 7 . S p e r m d u c t lo o p o n p o st e r io r o r p r o la te r a l p o st e r io r s id e o f te g u lu m 5 8 . R e tr o la te r a l sp e r m d u c t lo o p w id th 5 9 . A p o p h y si s o n p a lp f e m u r 6 0 . A p o p h y si s o n p a te ll a 6 1 . P a lp a l ti b ia p r o la te r a l a p o p h y si s 6 2 . P a lp a l ti b ia m a c r o se ta e 6 3 . E m b o lu s p o si ti o n 6 4 . P la n e o f sp ir a l o f e m b o lu s 6 5 . L a m e ll a a lo n g e m b o lu s 6 6 . E m b o li c d is c 6 7 . P r o c e ss o n e m b o li c d is c 6 8 . P la n e o f e m b o li c d is c 6 9 . R e tr o la te r a l c y m b ia l e x te n si o n 7 0 . D is ta l h e m a to d o c h a 7 1 . S a lt ic id r a d ix 7 2 . P o si ti o n o f c o p u la to r y o p e n in g i n r e la ti o n t o v u lv a 7 3 . F e r ti li z a ti o n d u c t p o si ti o n 7 4 . E p ig y n a l m e d ia n s e p tu m o f w in d o w 7 5 . S p e r m a th e c a s iz e 7 6 . N u m b e r o f sp e r m a th e c a e 7 7 . O p e n in g w it h s p ir a l g u id e 7 8 . B a r r ie r r id g e a t o p e n in g 7 9 . W in d o w l e n g th 8 0 . W in d o w s o f e p ig y n u m 8 1 . P r im a r y s p e r m a th e c a s h a p e 8 2 . P r im a r y s p e r m a th e c a p o si ti o n i n r e la ti o n t o t r a n sl u c e n t w in d o w Ghelna canadensis 0 0 0 0 0 1 0 0 0 2 0 0 0 0 1 0 - 0 0 0 0 0 - 0 0 0 0 0 0 0 0 1 - 1 0 0 0 - 1 3 - Heliophanus cupreus 0 1 0 0 0 1 0 0 0 1 0 0 0 1 1 0 - 1 0 0 0 0 - 0 1 - - 0 2 2 0 1 1 0 0 0 0 2 0 1 2 \"Bathippus\" pahang 0 1 0 0 1 0 0 0 0 1 0 0 0 1 1 0 - 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 2 0 0 2 Chinattus parvulus 0 1 0 0 0 1 0 0 0 0 1 0 0 0 1 0 - 0 0 0 0 0 - 0 1 - - 0 0 0 1 1 - 1 0 0 0 - 1 3 - Freya decorata 0 1 0 0 0 1 0 0 0 1 1 0 0 1 1 0 - 0 0 0 0 0 - 1 1 - - 0 2 0 0 1 - 1 0 0 0 - 1 3 - Plexippus paykulli 1 1 0 0 0 1 0 0 0 3 0 0 0 0 1 0 - 0 0 0 0 0 - 0 1 - - 0 2 0 0 1 - 0 0 0 0 - 1 0 - Salticus scenicus 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 - 0 0 0 0 0 - 0 1 - - 0 2 0 0 1 - 0 0 0 0 - 1 0 - Agobardus bahoruco 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 2 Agobardus cf. anormalis montanus 0 0 0 0 2 0 0 0 1 0 1 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 1 0 1 0 0 1 Agobardus cf. brevitarsus 0 0 0 0 2 0 0 0 1 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 1 0 2 0 0 1 Agobardus cordiformis 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 2 Agobardus gramineus 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Agobardus oviedo 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 2 Agobardus phylladiphilus 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Amphidraus complexus 1 ? 0 0 1 1 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 - 0 0 0 0 - 1 0 - ! \"#$! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Anasaitis adorabilis 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 - 0 1 - - 0 1 0 0 1 - 0 0 0 0 - 1 1 - Anasaitis banksi 0 1 0 0 1 1 1 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 - 0 0 0 0 - 1 0 - Anasaitis brunnea 0 ? 0 0 0 1 0 0 0 0 1 0 0 0 0 0 2 0 0 0 0 0 - 0 1 - - 0 1 0 0 1 - 0 0 0 0 - 1 0 - Anasaitis canosa [USA] 0 1 0 0 0 1 1 0 0 0 1 0 0 0 0 0 2 0 0 0 0 0 - 0 1 - - 0 0 0 0 1 - 0 0 0 0 - 1 0 - Anasaitis cf. canalis 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 2 0 0 0 0 0 - 0 1 - - 0 1 0 0 1 - 0 0 0 0 - 1 0 - Anasaitis elegantissima 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 1 Anasaitis gloriae 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 - 0 0 0 0 0 1 0 1 1 - 0 0 0 0 - 1 0 - Anasaitis hebetata 0 ? ? 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 - 0 1 - - 0 1 0 ? ? ? ? ? ? ? ? ? ? ? Anasaitis laxa 0 ? 0 0 0 1 1 0 0 0 1 0 0 0 0 0 2 0 0 0 0 0 - 0 1 - - 0 1 0 2 0 - 0 0 0 0 - 1 0 - Anasaitis locuples 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 - 0 1 - - 0 2 0 2 1 - 0 0 0 0 - 1 0 - Anasaitis placida 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 - 0 0 0 0 0 1 0 1 1 ? 0 0 0 0 - ? 0 - Anasaitis sp. [Peblique] 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 - 0 1 - - 0 1 0 2 0 - 0 0 0 0 - 1 0 - Antillattus cambridgei 0 0 0 1 1 0 0 0 1 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 1 Antillattus cf. applanatus 0 ? 0 0 1 1 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Antillattus darlingtoni 0 0 0 0 1 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Antillattus gracilis 0 0 1 1 1 0 0 0 1 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 2 0 0 1 Antillattus maxillosus 0 0 0 0 1 1 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 1 0 0 1 0 0 1 Antillattus scutiformis 0 0 0 1 1 1 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Bathippus directus [Tualapa] 0 0 0 0 1 0 0 0 1 2 0 1 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 2 0 0 1 Bathippus gahavisuka 0 0 0 0 1 0 0 0 1 2 0 1 1 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 ? 0 1 0 0 0 0 0 1 0 0 2 Bathippus korei 0 0 0 0 1 0 0 0 1 2 0 1 1 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 2 0 1 1 Bathippus macrognathus 0 0 0 0 1 0 0 0 1 2 0 1 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 2 0 0 1 Bathippus madang 0 0 ? 0 1 0 0 0 1 2 0 1 1 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 1 ? ? ? ? ? ? ? ? ? ? ? Belliena ecuadorica 1 ? 0 0 ? 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Bulolia excentrica 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 1 - 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Bythocrotus cf. crypticus 0 1 0 0 1 0 1 0 0 2 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Bythocrotus crypticus 0 1 0 0 1 0 1 0 0 2 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Canama cf. forceps 0 0 ? 0 3 0 0 0 1 2 0 1 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Canama extranea 0 0 0 0 3 1 0 0 1 2 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 1 0 0 2 1 0 1 0 0 0 2 0 3 0 Canama fimoi 0 0 0 0 2 0 0 0 1 2 0 1 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 1 0 0 2 1 0 1 0 0 0 2 0 3 1 Canama hinnulea 0 0 ? 0 2 0 0 0 1 3 0 1 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Canama triramosa 0 0 0 0 3 0 0 0 1 2 0 1 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 2 1 0 1 0 0 0 2 0 3 1 ! \"#$! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Petemathis minuta 0 ? 0 0 1 1 0 0 1 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 1 Petemathis portoricensis [Adjuntas] 0 0 0 1 1 1 1 0 1 3 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 2 Petemathis portoricensis [Maricao] 0 1 0 1 1 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 2 0 0 1 Petemathis tetuani 0 1 ? 0 1 1 1 0 0 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? cf. Coryphasia sp. [Brazil] 0 ? ? 0 1 1 0 0 0 1 1 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? cf. Maeota sp. [Panama] 0 ? ? 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Chalcolecta prensitans ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 1 0 1 0 0 2 0 2 1 Chalcolemia nakanai ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 0 0 1 0 0 2 0 0 1 Chalcoscirtus diminutus 0 ? 0 0 0 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 - 0 0 0 0 0 0 0 2 1 - 1 0 0 0 - 1 ? - Chalcoscirtus infimus 1 ? 0 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Chalcotropis luceroi 0 1 ? 0 1 1 0 0 1 2 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Chalcotropis cf. caeruleus 0 0 ? 0 1 1 0 0 0 0 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Chapoda angusta 0 0 0 0 1 1 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Chapoda cf. inermis [CostaRica] 0 0 0 0 1 0 1 0 1 1 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 1 0 0 2 Chapoda cf. inermis [Panama] 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 1 0 0 2 Chapoda fortuna [Panama] 0 ? ? 0 1 1 1 0 0 1 1 0 0 0 0 0 2 1 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Chapoda gitae 0 1 0 0 1 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 2 Chapoda montana 0 ? ? 0 1 1 1 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Chapoda peckhami 0 0 ? 0 1 0 1 0 1 1 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 ? ? ? ? ? ? ? ? ? ? ? Chapoda recondita 0 0 0 0 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Chinophrys pengi 1 0 0 0 2 1 1 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 1 2 Coccorchestes cf. aiyura 1 0 0 0 ? 1 0 0 0 0 0 0 0 0 1 1 - 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 2 Coccorchestes cf. ildikoae 1 0 ? 0 ? 1 0 0 0 0 0 0 0 0 1 1 - 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Coccorchestes cf. inermis 1 0 0 0 ? 1 0 0 0 0 0 0 0 0 1 1 - 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 2 Coccorchestes clavifemur 1 0 ? 0 3 1 0 0 0 1 0 0 0 0 1 1 - 1 0 0 0 0 1 0 0 0 0 0 0 1 ? ? ? ? ? ? ? ? ? ? ? Colyttus bilineatus 0 1 0 0 1 1 0 0 1 3 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 0 0 1 0 0 2 Colyttus striatus 0 1 0 0 1 1 0 0 1 3 0 1 0 0 0 0 2 0 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 1 0 0 2 0 0 1 Compsodecta haytiensis 0 0 0 0 1 0 0 0 1 1 1 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 Compsodecta peckhami 0 0 0 0 1 0 0 0 0 2 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 1 Corticattus latus 0 1 0 0 2 1 1 0 0 0 0 0 0 0 1 1 - 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 1 0 0 2 Coryphasia cf. campestrata 0 1 0 0 2 1 0 0 0 3 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 Coryphasia fasciiventris 0 1 0 0 2 1 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 2 0 0 1 ! \"##! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Coryphasia physonycha 0 1 0 0 1 1 0 0 0 3 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0 0 1 0 0 2 0 0 2 Corythalia bicincta 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 2 0 0 1 Corythalia broccai 0 ? 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 0 2 0 0 1 Corythalia bromelicola 0 0 0 1 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 0 2 0 0 0 Corythalia coronai 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 ? 0 2 0 0 1 Corythalia decora 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 ? 0 1 0 0 2 Corythalia electa 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 1 Corythalia minor 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 0 2 0 0 1 Corythalia peblique 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 2 0 0 1 Corythalia porphyra 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 1 2 Corythalia sulfurea 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Cytaea mitellata 0 1 ? 0 2 0 0 0 0 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Cytaea nimbata 0 1 0 0 3 0 0 0 0 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Cytaea oreophila 0 1 0 0 2 0 0 0 0 4 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 0 0 1 Diolenius varicus 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 - - 0 0 0 0 1 0 0 0 0 0 2 0 0 1 Ecuadattus napoensis 0 1 ? 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ? ? ? ? Ecuadattus pichincha 0 1 0 0 1 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 2 Efate albobicinctus 1 ? ? 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 - 0 1 - - 0 0 ? ? ? ? ? ? ? ? ? ? ? ? Emathis gombak 0 0 0 0 1 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 2 Euophryine sp. [GentingHighlands] ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 1 0 2 0 1 1 Euophrys frontalis 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 2 Euophrys monodnock 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 2 Euryattus bleekeri 0 1 0 0 2 0 0 0 0 5 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 2 0 0 1 Euryattus sp.1 [Gahavisuka] ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 1 0 0 2 0 1 1 Euryattus sp.2 [Gahavisuka] 0 1 ? 0 2 0 0 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Foliabitus longzhou 1 0 0 0 2 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 2 0 1 1 Foliabitus sp. [Malaysia] 0 ? ? 0 1 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Ilargus coccineus 0 1 0 0 2 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 1 Ilargus foliosus 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 1 1 Ilargus galianoae 0 1 0 0 2 1 1 0 0 0 ? 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 2 0 1 2 Ilargus macrocornis 0 1 ? 0 1 1 0 0 0 0 ? 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Ilargus moronatigus ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 0 0 1 0 1 2 ! \"#$! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Ilargus pilleolus 0 1 0 1 1 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 2 Ilargus serratus 0 1 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 1 Lagnus edwardsi 0 0 0 0 2 0 0 0 0 4 0 1 0 0 0 0 2 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 Laufeia daiqini 1 1 0 0 0 1 0 0 0 3 0 0 0 0 0 0 1 0 0 0 0 0 - 0 1 - - 0 1 0 0 1 - 0 0 0 0 - 1 0 - Laufeia eximia 1 ? 0 0 2 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 1 0 1 1 0 0 0 0 0 - 1 0 - Laufeia keyserlingi 1 1 0 0 1 1 0 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 - 0 0 0 0 - 1 2 - Lepidemathis haemorrhoidalis 0 1 0 0 2 0 0 0 0 7 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Leptathamas paradoxus 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Maeota dichrura 0 0 ? 0 1 1 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota dorsalis 0 ? ? 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota flava 0 ? ? 0 2 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota simoni 0 ? ? 0 3 1 0 0 0 0 1 1 0 0 0 0 ? 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota sp. [JatunSacha] ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 0 0 2 0 0 2 Maeota sp. [Manabi] 0 ? ? 0 1 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota sp. [MoronaSantiago] 0 ? ? 0 2 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Maeota sp. [Napo] ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 0 0 1 0 0 2 Maeota tuberculotibiata 0 0 ? 0 3 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Marma nigritarsis 0 1 0 0 1 1 0 0 0 0 1 0 0 0 1 0 - 0 0 0 0 1 0 0 0 1 0 0 1 0 0 1 - 0 0 0 0 - 1 1 - Mexigonus arizonensis 0 1 0 0 1 1 1 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 1 Mexigonus morosus 0 0 0 0 1 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 2 0 3 1 Mopiopia cf. bruneti 0 1 ? 0 2 1 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Naphrys pulex 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 2 0 0 1 Neonella vinnula 1 ? 0 0 1 1 0 0 0 0 0 0 0 0 1 1 - 0 0 0 0 0 0 1 1 - - 0 0 ? 2 1 0 0 0 0 0 ? 0 0 1 Ohilimia scutellata 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 1 - - 0 0 0 0 1 - 0 0 0 0 - 1 2 - Omoedus brevis 0 ? ? 0 1 0 1 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Omoedus cf. danae 0 1 0 0 0 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 - 0 0 0 0 0 0 0 0 1 0 1 0 0 0 ? ? 3 2 Omoedus cf. piceus 0 1 ? 0 3 0 0 0 0 4 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Omoedus cf. semirasus 0 1 0 0 3 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus cf. torquatus 0 1 0 0 2 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus darleyorum 0 ? 0 1 3 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus ephippigera 0 1 0 0 3 0 0 0 0 4 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus meyeri 0 0 0 0 3 0 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 ! \"#$! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Omoedus omundseni 0 1 0 0 3 0 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus orbiculatus 0 1 0 0 3 0 0 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus papuanus 0 1 0 0 3 0 0 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Omoedus swiftorum 0 0 0 0 3 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0 0 1 0 3 2 Parabathippus cf. macilentus 0 0 1 1 2 0 0 0 1 3 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 2 0 0 1 Parabathippus cuspidatus 0 0 1 1 2 0 0 0 1 3 0 1 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 1 0 0 1 Parabathippus kiabau 0 0 ? 1 2 0 0 0 1 2 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Parabathippus magnus 0 0 1 1 2 0 0 0 1 3 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 1 1 0 2 1 Parabathippus shelfordi 0 0 1 1 2 0 0 0 1 1 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 1 0 0 1 Paraharmochirus tualapaensis 1 ? 0 0 1 0 0 0 0 1 0 0 0 0 0 0 2 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 2 0 0 2 Parvattus zhui 0 ? ? 0 1 1 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 ? 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Pensacola signata 0 0 0 0 1 0 0 0 1 0 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 2 0 0 1 Phasmolia elegans 0 0 1 1 1 0 0 0 0 2 0 1 0 0 1 0 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 1 Popcornella furcata 0 0 0 0 0 1 0 0 0 0 1 0 0 2 0 0 1 0 0 1 0 0 - 0 1 - - 0 1 0 2 1 - 0 0 0 0 - 1 1 - Popcornella nigromaculata 0 0 0 0 0 1 0 0 0 0 1 0 0 2 1 0 - 0 0 1 0 0 - 0 1 - - 0 1 0 0 0 - 0 0 0 0 - 1 0 - Popcornella spiniformis 0 0 0 0 0 1 0 0 0 0 0 0 0 2 0 0 1 0 0 0 0 0 - 0 1 - - 0 1 0 1 1 - 0 0 0 0 - 1 0 - Pristobaeus beccarii 0 1 0 0 1 1 0 0 1 3 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 2 Pristobaeus cf. jocosus 0 0 0 0 1 1 0 0 1 5 0 1 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Pseudeuophrys erratica 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 - - 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Saitis auripes 0 1 0 0 1 1 1 1 0 0 1 0 0 0 0 0 ? 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 2 Saitis barbipes 0 1 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 2 0 1 2 Saitis cf. fuscus ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 0 0 0 0 0 1 0 1 2 Saitis cf. griseus 1 1 0 0 1 1 1 1 0 0 1 0 0 0 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 2 Saitis sp. [NewSouthWales] 0 1 ? 0 1 1 1 1 0 0 1 0 0 0 0 0 ? 0 0 0 0 0 0 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Saitis sp. [SouthAustralia] 0 1 0 0 1 1 0 1 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 2 Saphrys a-notata 1 1 0 0 1 1 1 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 Saphrys cf. patagonica 1 1 0 0 1 1 1 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 2 Servaea vestita 0 1 ? 0 2 1 0 0 0 2 0 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Sidusa extensa 0 1 1 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 1 0 0 1 0 0 2 0 0 1 Sidusa sp. [CostaRica] 0 1 0 0 1 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 1 0 0 1 0 0 2 0 0 1 Sidusa sp.1 [FrenchGuiana] 0 1 1 1 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 2 0 0 1 Sidusa sp.2 [FrenchGuiana] 0 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 1 0 0 0 0 0 2 0 0 1 ! \"#\"! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Sidusa unicolor 0 1 0 1 1 0 0 0 1 2 0 1 1 0 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 0 2 1 0 0 1 0 0 2 0 0 1 Sobasina wanlessi 1 ? 0 0 0 0 0 0 0 0 0 0 0 0 1 0 - 0 0 0 0 0 - 0 1 - - 0 1 0 2 1 - 1 0 0 0 - 1 3 - Soesilarishius cf. amrishi 1 0 ? 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 - 0 1 - - 0 1 0 ? ? ? ? ? ? ? ? ? ? ? Soesilarishius minutus 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 1 - 0 0 0 0 0 1 0 0 1 - 1 0 0 0 - 1 3 - Soesilarishius ruizi 1 ? 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 - 0 1 - - 0 1 0 0 1 - 0 0 0 0 - 1 1 - Talavera minuta 0 0 0 0 0 1 0 0 0 0 0 0 0 - 0 0 2 0 0 0 0 0 - 0 1 - - 0 0 0 0 1 - 0 0 0 0 - 1 0 - Thiania bhamoensis 0 1 0 0 1 1 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 2 Thiania latibola 0 1 ? 0 1 1 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thiania spectrum 1 0 ? 0 1 0 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thiania tenuis 0 1 ? 0 2 0 0 0 0 5 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thorelliola aliena 0 1 0 0 2 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 1 2 Thorelliola cf. mahunkai 0 1 0 0 1 0 0 0 0 3 0 0 0 0 0 0 2 0 0 1 1 0 1 0 0 0 0 0 0 0 2 1 ? 0 0 0 0 ? ? 0 ? Thorelliola crebra 0 1 0 0 1 0 0 0 1 2 0 0 0 0 0 0 2 1 1 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 2 Thorelliola ensifera 0 1 0 0 1 0 0 0 0 3 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 0 0 0 0 0 2 1 1 0 0 0 0 2 0 0 1 Thorelliola Joannae ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 1 0 0 0 0 1 0 0 2 Thorelliola tamasi 0 0 ? 0 0 0 0 0 0 3 0 0 0 0 0 0 2 0 0 0 0 0 - 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thorelliola tualapa 0 1 0 0 0 0 0 0 0 3 0 0 0 0 0 0 2 0 0 1 1 0 - 0 0 0 0 0 0 0 2 1 ? 0 0 0 0 ? ? 0 ? Thyenula cf. aurantiaca ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 0 0 2 0 1 1 Thyenula cf. mundus 0 0 ? 0 2 1 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thyenula laxa 1 1 ? 0 2 1 1 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thyenula leighi 0 0 ? 0 2 0 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Thyenula nelshoogte ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 0 1 1 0 1 2 Thyenula sp. [SouthAfrica] 0 ? 0 0 1 1 1 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 1 Thyenula wesolowskae 1 1 0 0 2 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 1 Truncattus cachotensis 0 ? 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 2 Truncattus dominicanus 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 2 0 0 1 Truncattus flavus 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 2 Tylogonus cf. auricapillus 0 0 ? 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 0 0 - 0 1 - - 0 2 2 ? ? ? ? ? ? ? ? ? ? ? Tylogonus yanayacu 0 0 0 0 0 0 0 0 0 3 0 0 0 1 0 0 0 0 0 0 0 0 - 0 1 - - 0 2 2 2 1 0 0 0 0 0 ? 0 1 0 Variratina minuta 0 ? 0 0 3 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 1 0 0 0 2 0 3 0 Viribestus suyanensis 0 0 ? 0 1 0 0 0 1 2 0 1 0 1 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Xenocytaea agnarssoni 0 1 0 0 2 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 3 2 ! \"#$! 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1 5 2 5 3 5 4 5 5 5 6 5 7 5 8 5 9 6 0 6 1 6 2 6 3 6 4 6 5 6 6 6 7 6 8 6 9 7 0 7 1 7 2 7 3 7 4 7 5 7 6 7 7 7 8 7 9 8 0 8 1 8 2 Xenocytaea albomaculata 0 1 ? 0 1 0 0 0 0 0 1 0 0 0 1 1 - 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Xenocytaea proszynskii ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 1 0 0 0 0 1 0 0 2 Zabkattus brevis 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 3 2 Zabkattus furcatus 0 0 ? 0 2 1 0 0 0 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Zabkattus richardsi 0 0 ? 0 2 1 0 0 0 1 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? Zabkattus trapeziformis ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 1 0 0 0 1 0 3 2 "@en ; edm:hasType "Thesis/Dissertation"@en ; vivo:dateIssued "2012-11"@en ; edm:isShownAt "10.14288/1.0072804"@en ; dcterms:language "eng"@en ; ns0:degreeDiscipline "Zoology"@en ; edm:provider "Vancouver : University of British Columbia Library"@en ; dcterms:publisher "University of British Columbia"@en ; dcterms:rights "Attribution-NonCommercial-NoDerivatives 4.0 International"@en ; ns0:rightsURI "http://creativecommons.org/licenses/by-nc-nd/4.0/"@en ; ns0:scholarLevel "Graduate"@en ; dcterms:title "Phylogeny and systematics of the jumping spider subfamily Euophryinae (Araneae : Salticidae), with consideration of biogeography and genitalic evolution"@en ; dcterms:type "Text"@en ; ns0:identifierURI "http://hdl.handle.net/2429/42354"@en .