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Copepod-chondrichthyan coevolution : a cladistic consideration Deets, Gregory B. 1995-12-31

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COPEPOD-CHONDRICHTHYANCOEVOLUTION:A CLADISTIC CONSIDERATIONGREGORY B. DEETSB.Sc., California State UniversityLong Beach,M.Sc., California State University Long Beach,19801985A THESIS SUBMITTED IN PARTIALFULFILLMENT OFTHE REQUIREMENTS FOR THE DEGREEOFDOCTOR OF PHILOSOPHYWe accept this thesis as conformingto the required standardbyinTHE FACULTY OF GRADUATE STUDIESDepartment of ZoologyTHE UNIVERSITY OF BRITISH COLUMBIAAUGUST 1994©Gregory B. Deets, 1994In presenting this thesisin partial fulfilment of the requirements foran advanceddegree at the University of British Columbia,I agree that the Library shall make itfreely available for reference andstudy. I further agree that permissionfor extensivecopying of this thesis forscholarly purposes may be grantedby the head of mydepartment or by his or her representatives.It is understood that copyingorpublication of this thesis for financialgain shall not be allowed withoutmy writtenpermission.(Signature)_____________________________Department of___________________The University of British ColumbiaVancouver, CanadaDate_______________________DE.6 (2/88)ABSTPCTA revision of the species ofEudactylina (Eudactylinidae :Siphonostomatoida)and Kroyeria (Kroyeriidae :Siphonostomatoida) was conducted,based on type andother specimens of parasiticcopepods from museums and personalcollections. Adescription of the external morphologyof each genus is included. Taxonomic,phylogenetic, and functional significanceof the morphology of thegeneral habitus, first andsecond antennae, oral andthoracic appendages arediscussed.The taxonomic account ofthe above genera recognized all nominalspecies inthe literature. Illustrationsand phylogenetic analyses,however, were necessarilyrestricted to only the material examinedin an attempt to standardize theabstractionsand interpretations associatedwith character observation. Detailedredescriptions aregiven of E. acuta, E. aspera,E. chilensis, E. corrugata, E. indivisa,E. insolens, E.Iongispina, E. myliobatidos, E.oliveri, E papillosa, E. peruensis, E. pollex, E.pusilla,E. similis, E. spinifera, E. squamosa,E tuberifera, E. turgipes, and new descriptions(all in press) are givenof, E. aphiloxenous, E. dactylocerca, E. diabolophila,E. epaktoIampte’ E. hornbosteli, E. nykterimyzon,E. pristiophori, E. urolophi, and E. vaquetillaefollowed by the detailed reclescriptionsof K. carchariaeglauci, K. caseyi, K. dispat K.elongata, K. gemursa, K. lineata,K. longicauda, K. papillipes, K. spatulata, K. sphyrnae, K. triakos and new descriptions (allin press) of K. branchioecetes, Kcresseyi, K.decepta, K. procerobscena, and K. rhophemophaga.In an attempt to unravel evolutionaryrelationships of their elasmobranch hostsand themselves a phylogeneticanalysis of each genus is presented. In the heuristicanalysis of Eudactyilna, 75 morphologicalcharacters resulted in a single tree with aconsisitency index of 0.77 anda retention index of 0.88, indicating a high degree ofcharacter congruence. An exact searchof nine species of Eudactylina with 55 characters resulted in a single tree witha consistency index of 0.88 and a retention index of0.88. The Eudactylina-derivedhost cladograms positmonophyly of the shark-likesqualoids , squatinids, pristiophorids,and batoids. This suggeststhat shark-likesqualoids, angelsharks, andsawsharks are more closelyrelated to rays than to othergaleomorph sharks, whereasthe pristiophorids represent the sistertaxon to batoids.The eudactylinid dadefound on the rhinopterids andmobulids appears to representacolonization event followedby tight cospeciation. Eudactylina-derivedcarcharhinidrelationships approximateconventional or currently acceptedhypotheses. Eudactylinaderived phyogenetic relationshipsof a subset of species fromSquatina and Myliobatisindicate speciation patterns consistentwith major vicariant events associatedwith thebreakup of Pangaea during theJurassic period approximately160 MY.The phylogenetic analysis of Kroyeria,using 44 morphological charactersresulted in a single tree with a consistencyindex of 0.75 and a retentionindex of 0.75. TheKroyeria-derived and Kroyeria-Kroeyerina-derivedhost cladograms posit an unconventional placement for Galeocerdo.Galeocerdo diverges at the bottom of thetree beforethe Triakidae. A sphyrnid dadefollows, functioning as the sister taxonto remainingmembers of the Carcharhinidae.The genus Carcharhinus appears paraphyleticwithNegaprion and Prionaceimbedded within this dade, corroboratingsimilarly held viewsby other systematists.Congruent host and parasite cladogramtopologies from both holocephalan andelasmobranch hosts suggestthe existence of well-established and specifichost-parasite associations as earlyas the late Devonian, approximately 400 MY.IIITABLE OF CONTENTSAbstract.List of TablesixList of FiguresxAcknowledgementsxiiiINTRODUCTION1MATERIALS AND METHODS12HISTORICAL REVIEW15EUDACTYLINA18EXTERNAL MORPHOLOGY18GENERAL HABITUS18CAUDALRAMUS19FIRSTANTENNA19SECOND ANTENNA20ORAL CONE AND MANDIBLE20FIRST MAXILLA20SECOND MAXILLA21MAXILLIPED21LEG ONE22LEG TWO22LEGSTHREEANDFOUR23LEG FIVE23LIFE HISTORY24GENERAL DESCRIPTION24REPRODUCTION25ivHOST-PARASITE RELATIONSHIPS26DELETERIOUS EFFECTS!FEEDING26SPECIFICITY26SYSTEMATIC ACCOUNT27GENUS EUDACTYLINAvan Beneden, 185327Eudactylina acanthil A. Scott,190128Eudactylina acuta van Beneden,185330Eudactylina aphiloxenossp. nov33Eudactylina aspera HelIer,186535Eudactylina chilensis Ho andMcKinney, 198138Eudactylina corrugata Bere,193040Eudactylina dactylocercasp. nov 42Eudactylina diabolophila sp. nov45Eudactylina dolifusi, Brian, 192447Eudactylina epaktolamptersp. nov 50Eudactylina hornbosteli sp.nov 54Eudactylina indivisa Castro and Baeza,1991 56Eudactylina insolens Scott andScott, 1913 58Eudactylina longispina Bere,193660Eudactylina myliobatidos Luque andFarfan, 1991 62Eudactylina nykterimyzon sp. nov65Eudactylina oliveri Laubier, 196868Eudactylinapapillosa Kabata, 197972Eudactylinaperuensis Luqueand Farfan,1991 74Eudactylina pollex Cressey, 196776Eudactylinapristiophori sp. nov79Eudactylina pus//Ia Cressey, 196781Eudactylina s/mills Scott, 190283vEudactylina squamosaBere, 1936• 85Eudactylina tuberifera Castroand Baeza, 198788EudactylinaturgipesBere, 193690Eudactylina urolophi sp. nov92Eudactylina vaquetillaesp. nov95REMAINING UNOBTAINABLENOMINAL SPECIES 97PHYLOGENETIC ANALYSIS101CLADOGRAM CONSTRUCTION101PARASITE-DERIVED HOSTCLADOGRAM102HISTORICAL BIOGEOGRAPHY105KROYERIA109EXTERNAL MORPHOLOGY109GENERAL HABITUS109CAUDAL RAMUS110DORSAL AND INTERPODALSTYLETS110FIRST ANTENNA112SECOND ANTENNA112MANDIBLE AND ORAL CONE113FIRST MAXILLA113SECOND MAXILLA113MAXILLIPED114LEG ONE114LEGTWO 115LEG THREE 115LEG FOUR116LEGSFIVEANDSIX116LIFE HISTORY117GENERAL DESCRIPTION117viREPRODUCTION.117HOST-PARASITE RELATIONSHIPS118DELETERIOUS EFFECTS!FEEDING118SPECIFICITY119SYSTEMATIC ACCOUNT119GENUS KROYERIA vanBeneden, 1853119Kroyeria branchioecetessp. nov 121KroyeriacarchariaeglauciHesse, 1879 123Kroyeria caseyl Benz andDeets, 1986 127Kroyeria cresseylsp. nov129Kroyeria decepta sp. nov132Kroyeriadispar Wilson, 1932135Kroyeriaelongata Pillai, 1967138Kroyeria gemursa Cressey,1967140Kroyeria lineata van Beneden,1853143Kroyeria longicauda Cressey,1970 146Kroyeriapapillipes Wilson, 1932149Kroyeriaprocerobscenasp. nov151Kroyeria rhophemophaga sp. nov154Kroyeriaspatulata Pearse, 1948157Kroyeria sphyrnae Rang nekar, 1957160KroyeriatriakosFukui, 1965163REMAINING UNOBTAINABLE NOMINALSPECIES 166PHYLOGENETIC ANALYSIS168CLADOGRAM CONSTRUCTION168PARASITE-DERIVED HOSTCLADOG RAM 169COMBINING PARASITE CLADOGRAMS170COMPETING HOST CLADOGRAMS172viiSUMMARY AND CONCLUSIONS.176REFERENCES178FIGURES196APPENDICESAPPENDIX A -435DATA MATRIX AND DEFINITIONOF CHARACTERSFOR EUDACTYLINA436DATA MATRIX FOR EUDACTYLINASUBSET 441DATA MATRIX AND DEFINITION OF CHARACTERSFOR KROYERIA445APPENDIX B -APPENDIX C -v”List of TablesI. RECODED EUDACTYLINAPHYLOGENY BY HOST MATRIX430II. RECODED EUDACTYLINASUBSET PHYLOGENY BY HOST MATRIX.. .431III. RECODEDKROYERIA PHYLOGENYBY HOST MATRIX 432IV. RECODEDKRQEYERINA BY HOST MATRIX433V. COMBINEDRECODED KROYERIA-KROEYERINABY HOST MATRIX. . . . 434ixList of Figures1. Eudactylina attached to gillfilament in situ1972-3 EudactylinaacanthiiA.Scott, 19011994-5. Eudactylina acuta vanBeneden, 18532036-7. Eudactylinaaphiloxenos sp. nov2078-9. Eudactylina aspera HelTer,186521110-11. Eudactylina chilensisHo and McKinney, 198121512-13. Eudactylinacorrugata Bere, 193021914-15. Eudactylina dactylocercasp. nov 22316-17. Eudactylinadiaboiophiiasp. nov22718-19. Eudactylina dolifusi, Brian, 192423120-23. Eudactylina epaktoiamptersp. nov 23524-25. Eudactylina hornbostelisp. nov24326-27. Eudactylina mdivisa Castro and Baeza,1991 24728-29. Eudactylina insolensScott and Scott, 1913 25130-31. Eudactylina longispina Bere, 193625532-33. Eudactylina myliobatidos Luque and Farfan,1991 25934-35. Eudactylina nykterimyzon sp. nov26336-39. Eudactylina oilyen Laubier, 196826740-41. Eudactylina papillosa Kabata, 197927542-43. Eudactylinaperuensis Luqueand Farfan, 199127944-45. EudactylinapoliexCressey, 196728346-47. Eudactylina pristiophorisp. nov 28748-49. Eudactylina pusilla Cressey, 1967 29150-51. Eudactylina simiiisScott, 1902 29552-53. Eudactylina squamosa Bere, 1936 29954-55. Eudactylina tuberifera Castro and Baeza, 1987 303x56-57. Eudactylina turgipes Bere,193630758-59. Eudactylina urolophisp.nov31160-61. Eudactylina vaquetillae sp. nov31562. Eudactylinidae Cladogram31963. Eudactylinidae Host-SummaryCladogram32164. Eudactylina Cladogram(Heuristic)32365. Eudactylina Cladogram(Exact)32566. Eudactylina-derived Host Cladogram(Heuristic) 32767. Eudactylina-derived HostCladogram (Exact) 32968. Area-Summary Cladogram for Squat/na33169. Area-Summary Cladogram for Myliobatis33370. Kroyeria attached to gill filament (in situ)33571-73. Kroyeria branchioecetes sp. nov33774-76. Kroyeria carchariaeglauci Hesse, 187934377-78. Kroyeria caseyi Benz and Deets, 198634979-80. Kroyeria cressey/sp. nov35381-84. Kroyeria decepta sp. nov35785-86. KroyeriadisparWilson, 193236587-88. Kroyeria elongata Pillai, 196736989-90. Kroyeria gemursa Cressey, 196737391-92. Kroyeria lineata van Beneden, 185337793-95. Kroyeria longicauda Cressey, 1970 38196-97. Kroyeria papillipes Wilson, 193238798-99. Kroyeria procerobscenasp. nov 391100-102. Kroyeriarhophemaphagasp. nov 395103-104. Kroyeria spatulata Pearse, 1948401105-106. Kroyeria sphyrnae Rangnekar, 1957 405107-108. KroyeriatriakosFukui, 1965 409xi109. Kroyeriidae Cladogram413110. KroyeriaCladogram415111. Kroyeria-derived host cladogram417112. Kroeyerina attached to olfactory lamellae (insitu) 419113. Kroeyerina-derived host cladogram421114. Combined Kroyeria-Kroeyerina-derivedhost cladogram 423115. Competing cladogram of Squalea and Hypnosqualea425116. Competing morphologicaland molecular cladograms of carcharhinids 427xiiAcknowledgementsThe completion of this dissertationwas far from a single-handed effort. I thankmy graduate advisor and friendDr. Martin Adamson for his guidance,friendship, criticalreview of the manuscript, humor,patience and especially his cheap home-made winefermented under his house. Thank youDr. Kabata (committee member) for your friendship, specimens, and world renownprofessionalism and ability, standards I’ll nevermeet. And thank you to committeemembers Dr. Al Lewis and Dr. Geoff Scudder forkindly serving on my committee,offering ideas and advice and editing thismonstrousthing.Additional thanks go to Dr. Ju-SheyHo and Dr. Masahiro Dojiri for use of theirlibraries and insightful discussions.Dr. Roger Cressey deserves morethan a species named after him! His help forthe arrangement of loans for theseparasitic copepods from the National Museum ofNatural History, Smithsonian lnstitutuion,coupled with the donation of his vast personalcollection for my inspection resulted in the discoveryof many new species and is largely responsible for the comprehensivenessof this effort. His sponsorship and kindnessduring my short-term appointment atthe Smithsonian will never be forgotten.I appreciate the assistance of Dr. Geoffrey Boxshall,The Natural HistoryMuseum, London, England, and Dr. Frank Ferrari,National Museum of Natural History,Smithsonian Institution, for shipping additional parasiticcopepods to me for this revision.I must thank George Benz forso many hilarious memories that again will neverbe forgotten, and who kept me going during someof the most testing and strenuoustimes in my life. His assistance onour co-authorships, his critical eye, insight andincredible potential for being such a S.O.B.is deeply appreciated.I thank Bernard Horn-Bostel (father-in-law equivalent) for financiallyassisting thisproject with a generous donation ofa Mac II, allowing the phylogenetic analyses andword processing to be done.The wild uncontrollable spendingby Patricia Horn-Bostel (mother-in-law-equivalent) on new software andhardware allowed many of the final figures to be completedas our hard drive was busting at the sectors.1 will be forever grateful to the Friends ofthe Provincial Museum, BritishColumbia for a generous awardin Zoology and Museum Studies, literally keeping meafloat when everything else was sinking.XIIEMostly, I thank Deborah Allison Horn-Bostel(spousal-unit equivalent) for helpingwith the trivial, such as the computer figures,some of the in situ illustrations,and tolerating some absolutely hideous field conditions. Butmore importantly, I thank her foralways being there, objective andsupportive in every episode of life while thiswasbeing attempted.Finally, this is dedicatedto the memory of my parents, Hubert and FlorenceDeets, both passing awayduring this long process.xiv(hen I was a boy, world wasbetter spot.what was so was so,what was not was not.Now I am a manworld have changed a lotSome things nearly so, others nearly not.There are times I almost thinkI am not sure of what I absolutely know.Very often find confusion in conclusionI concluded long ago.In my head are many facts that as a studentI have studied to procure.In my head are many facts of whichI wish I was more certain I was sure.Oscar Hammerstein ItThe King & IA Puzzlement1956xvINTRODUCTIONCoevolution has become an increasingly popular sub-disciplinewithin the field ofevolutionary biology. Ehrlich and Raven(1964) defined coevolution as an ecologicalphenomenon, a matter of “stepwise reciprocal response”between any two species with“close and evident” ecological relationships.Parasitologists citing von lhering (1891)have recognized a more restricted sense ofcoevolution; the historical relationshipsbetween hosts and parasites. Coevolution embodiestwo components, togetherness(co-) and history (evolution) (Mitter and Brooks, 1983).The study of togetherness (thefunctional fit of organisms to their environment) is usually restrictedto the field of ecology. The study of history (phylogeny) is enveloped within the realm of systematics(Brooks, 1 985a). Although this historical component ismissing from most of the earlierassessments of putatively coevolved systems (Brooks, 1979a; Brooks and Mitter, 1984;Mitter and Brooks, 1983), recent efforts have attemptedto incorporate it (Brooks andMcLennan, 1991,1993; Paterson, Gray & Wallis, 1993).Whether one studies free-living or parasitic organisms, the species associationsa given taxon exhibits are usually well defined, quite specific and broughtabout by acombination of descent and proximal or contemporaneous causes (colonization).Hence, it is of interest to determine whether most species associations are maintainedas equilibrium systems fueled by constant dispersal or maintained as historically constrained associations.Recent advances in systematics, among them the development of cladistics,allow one to approach the question froma phylogenetic standpoint. Specific questionsregarding coevolution, the evolution of ecological life-history traits, historical biogeography and classification canbe addressed if an explicit phylogenetic hypothesis is available.Cladistics, or phylogenetic systematics, attempts to reconstruct genealogical1relationships among taxaby determining the sequence of the originof their distinguishing features (Hennig, 1966; Wiley, 1981). Themajor interest in cladistics iscentered onconstruction of branching sequencesand defining monophyleticgroupings or naturaltaxa. In practicing cladistics,phylogeneticists subscribe strictlyto a structural, asopposed to a functional, approachto systematics. Observations of characterstructureare used to construct a hierarchicalpattern of taxa, then from these patterns thevalidityof hypotheses concerningevolutionary mechanisms is scrutinized (Brooksand Wiley,1986). Therefore, cladisticanalyses attempt to dissociate inferences of organismicrelationships and evolutionarypattern from assumptions concerning process(Ho andSaunders, 1984). Indeed, one of themost important contributions of cladistics has beento focus attention on the vitalimportance of pattern analysis, for it is only by havingsome aspect of pattern that science hassomething to explain (Cracraft, 1983).Formal statements concerning modern cladisticmethods were made by Hennig(1950, 1966). He proposed a general referencesystem for comparative biology basedon two major points. First he distinguishedbetween special reference systems and general reference systems in biologicalclassifications. Special reference systems emphasize a particular kind of relationshipamong different species. For example, a classification that placed all parasitic copepods inhabitingshark gills in one category, and allthose on batoid gills in anotherwould be a special reference system useful for categorizing parasiticcopepods in given host assemblages. This classificatory procedurewould place distantly related organisms in thesame taxonomic category. A general reference system should provide the most efficient summaryof the maximum amount ofinformation about the taxa beingclassified (Brooks, 1985b). Hennig suggested that thegeneral reference system in biology should bebased on the genealogical or phylogenetic relationships of the species involved. The choiceof genealogy was based on twoobservations: (1) the one attribute of any organismor species that would always beconstant was its history, so phylogenetic historyshould be the most stable criterion forclassifying and (2)genealogical relationships, like classifications, are inherentlyhierar2chical.Secondly, Hennig argued fora formal method of deducing phylogeneticrelationships. He objected to phylogenetic schemes thatwere based on hypothetical idealizedarchetype ancestors. Since species are compositesof ancestral and derived traits, it isunlikely such things as archetypesexist. Thus, homologous traits shared by two ormore species will be indicators of phylogeneticrelationship. Shared primitive traits indicate general phylogenetic relationships whileshared derived traits indicate more particular phylogenetic relationships.The terms plesiomorphy (plesio - near the source)andapomorphy (apo - away from the source) referto these relatively primitive and relativelyderived traits. Two taxa that share derivedhomologous characters (synapornorphies)are each other’s closest relatives and are calledsister taxa (Hennig, 1966).There are two comparative methods forevaluating the degree of primitiveness ofcharacters. These are the “ontogenetic criterion”and the “outgroup criterion”. Bothapproaches seek to establish the direction or pathof transformation from the primitiveto a more derived character state. A primitive orplesiomorphic character is more general because it defines a group thatis more inclusive than one defined by a less general,more derived or apomorphic condition of thatcharacter. Hence, the diagnostic featuresof each grouping in the genealogical hierarchy wouldbe those traits viewed as apomorphic at that level of that particular grouping (Wiley, 1981).The outgroup criterion states that any trait found inat least one member of thegroup being studied that also occurs in taxa outside thestudy group is plesiomorphic.Since outgroups are not archetypes and,therefore, may possess derived characterstates, it is often necessary to use more than one outgroup (composite outgroup)toestablish enough apomorphic traits to classify a taxon(Maddison,et al., 1984).The ontogenetic criterion states that, where twoorganisms possess differentadult states, if one organism exhibits the other’sadult trait during development, its adulttrait is apomorphic and that of the other adult traitis plesiomorphic. This approach ismore limited than outgroup analysis,since it works only for cases in which evolution3has proceeded by adding characteristics tothe ancestral developmental program(Brooks and Wiley, 1986).After determining which traits are apomorphicand which are plesiomorphic, oneis sometimes faced with apomorphic traits thatsuggest conflicting groupings. The reason for this is parallel and/orconvergent evolution, given the general name homoplasy,where similarity causes one to misattribute homology.Truly homologous traits of various taxa will yield congruent groupings. As longas homoplasious traits do not co-varyin larger numbers than the homologous traits,parsimony analyses will pinpoint theproper phylogenetic relationships. The possibleoccurrence of great amounts of parallelevolution requires only that many traitsbe used in the analysis, since the pattern ofrelationships indicated by a plurality of traits is thebest estimate of phylogenetic relationships (Brooks, 1985b). Since this requiresthat large numbers of traits be analyzedsimultaneously, phylogenetic computer packages such asPAUP (PhylogeneticAnalysis Using Parsimony) and MacClade have been developedto generate and analyze phylogenetic trees.As previously stated, concordance between phylogenetic relationships ofparasites and their hosts has been recognized since the nineteenthcentury (von Ihering,1891). Hennig (1966) discussed the concordance briefly and theresultant possibility ofinferring host phylogenies from parasitedata. The continued discovery of co-varyingassociations between parasites and their hosts has ledto the formulation of various“rules” of coevolution. The rules form a small group of interrelated concepts:1. Eichler’s Rule (for review see, Inglis, 1971): The more genera of parasites ahost harbors, the larger the systematic group to which the host belongs.2. Manter’s Rules (for review see Inglis, 1971): (1) Parasites evolve more slowlythan their hosts; (2) the longer the association witha host group, the more pronouncedthe specificity exhibitedby the parasite group; (3) a host species harbors the largestnumber of parasite species inthe area where it has resided longest, so if the same ortwo closely related speciesof host exhibit a disjunct distribution and possess similar4parasite faunas, the areas in whichthe hosts occur must have been contiguousat apast time.3. Szidat’s Rule (see Szidat, 1956; 1960): The moreprimitive the host, the moreprimitive the parasite it harbors.Finally, and probably thebest known:4. Farenholz’s Rule (for reviewssee Brooks, 1979a, 1981, 1985a; Inglis, 1971):Parasite phylogeny mirrors host phylogeny.Brooks (1981) developed this latter concept of coevolution interms of phylogenetic systematics (cladistics)and suggested that host-parasite coevolution canarisethrough processes comparableto those that produce homologous and homoplasiouscharacters. This analogous relationship betweencharacter state transformation seriesand parasite phylogenies allows one to view cospeciated orhistorically associated parasites as homologies or autapomorphies of theirhosts and colonizing parasite speciesas homoplastic characters of their hosts. Thus,evolutionary relationships can be takeninto consideration by first doinga phylogenetic analysis of the parasites and then treating that cladogramas a multistate character tree of the hosts that are inhabited by theparasite species.Such a tree is constructed with a source of information thatis not available instandard multistate analyses, namely the cladisticanalysis of the parasites themselves.The characters have characters,so to speak, that are used to infer their relationship.Various methods exist for converting the topologyof a phylogenetic into a matrixof numerical characters. This permitsmultiple parasite taxa to be analyzed simultaneously (a common data matrix), in order to generate host phylogenies (O’GradyandDeets, 1987).Additionally, historical approaches to ecology rely on an a priori phylogeneticanalyses. Replacing the names of the terminal taxa from the parasite cladogram withtheir respective ecological life historytraits produces an ecological summary cladogram. The result is a spatiotemporalinterpretation of the evolution of the parasites5infection-site associations (Brooks, 1985a; Deets,1987). In the same vein, historicalapproaches to biogeography are analyzed by replacingthe names of the terminal taxawith their distributions.Phylogenetic systematics has been applied mostlyto free-living taxa. Onlyrecently has this method entered the field ofparasitology. The first study to demonstratethe feasibility and applicability ofcladistics with parasitic taxa was made by Brooks(1977). The phylogenetic relationships of plagiorchioid trematodes in thisanalysis wereshown to be congruent with their anuranhosts as well as exhibiting a definite vicariantdistribution consistent with the Pangaean breakup.This was followed by a series ofpapers utilizing cladistic methods on parasitic taxa including crocodiliansand their digenean parasites (Brooks, 1979a), vicariancebiogeography, potamotrygonid stingraysand their helminth parasites (Brooks, et. al, 1981),pinworms and primates (Brooks andGlen, 1982), nematodes and primates (Glen andBrooks, 1985; 1986), and otherpapers formulating other applications with cladistics and evolutionarytheory (Brooks,1979b; 1980). Recently, Adamson and van Waerebeke(1985) employed cladistics toanalyze parasitic nematode classification and evolution, and Boeger and Kritsky (1989)tested Compagno’s (1977) various hypotheses of elasmobranch evolution with theircladistic analysis of the genera within the Hexabothriidae (Monogenea).In the midst of this blitzkrieg of phylogenetic analyses on worms, workers on theparasitic copepods (Crustacea), staging more of a sitzkrieg, gradually entered thecladistic arena. The first authors to investigate the phylogenetic relationships and historical zoogeography of a host group of fishes (Merluccius)as inferred from the phylogenetic relationships of their parasitic copepods were Kabata and Ho (1981). Though theirwork is free of any formal cladistic analysis, and devoidof associated jargon, it is phylogenetic in its approach. Both authors were keenly aware that when drawing conclusionson host zoogeographical or phylogenetic problems, one must be conscious of the phylogenetic relationships of the parasites considered. Hence, the relatively plesiomorphicor apomorphic conditions ofthe characters of their “indicator” species were taken into6consideration.Cressey, Collette and Russo (1983),specialists on parasiticcopepods andscombrid teleosts, were the first authorsto investigate phylogenetic relationshipsof parasitic copepods using formalquantitative cladistic methodology. Ho’s (1984) discoveryof the Spiophanicolidae,a family of highly modified copepods on polychaetes promptedhim to analyze phylogeneticallya certain suite of poecilostomatoid families [informallytermed the Nereicoliform Groupby Gooding (1963), IlIg, (1970) and Gotto (1979)] thatare parasitic on variousinvertebrate phyla. This resulted in the clarification of someclassificatory problems embedded in theliterature for nearly 20 years. A year later, Hoand Do (1985) analyzed the genera in theLernanthropidae, highly derived parasitesofteleost gills. That same year, Collette and Russo (1985)looked into the phylogeny ofthe Spanish Mackerels and theircopepod parasites in order to determine which evolutionary events of the parasitescould be explained by the evolutionary events of theirhosts. Following this, Deets (1987) conducteda phylogenetic analysis and systematicrevision of the genus Kroeyerina anda higher level analysis of the genera within thefamily Kroyeriidae. The apparent static nature of the parasites’life-history traits (specificinfection sites) coupled with the congruent phylogeneticpattern with their elasmobranchhosts suggested both hosts and parasites experiencedthe same vicariant events andsubsequent allopatric speciation. After this worka phylogenetic analysis of theEudactylinidae (Deets and Ho, 1988), resulted in the resurrection ofthe previously synonomized genus Protodactylina Laubier, 1966. A Tethyan distributionwas speculatedfor a single monophyletic subset(dade) of 3 monotypic genera parasitic on batoids.Next, Benz and Deets (1988), with therediscovery of a rare parasite specific to mobulidbranchial filters, carried out a phylogenetic analysis of the generawithin Cecropidae.Finally, Dojiri and Deets (1988) withthe finding of a new genus Norkus (Sphyriidae)phylogenetically analyzed the sphyriid genera.Again tight phylogenetic congruencewith the hosts and parasites werethe result. Additionally, parasite life history traits mirrored the cladogram’s topology.7In order to apply these methodsand aforementioned concepts itis imprative topossess detailed knowledge of the studied taxon’s morphology.At present the morphology of parasitic copepods is only poorly known. It appearsmorphological details ofminute animals are often ignored just becauseof their dimunitiveness (Kabata, 1979).Kabata (1979) redefined the familyEudactylinidae and removed from it Kroyeriavan Beneden, 1853 and KroeyerinaWilson, 1932 to form a new family Kroyeriidae.Recently, Deets (1987) established a new genusProkroyeria to accommodateKroeyerina meridionalis Ramirez,1 975. Therevised Eudactylinidae then consisted ofseven genera, but since then three havebeen discovered (Deets and Benz, 1987;Deets and Ho, 1988) bringing the totalto ten, namely Bariaka Cressey, 1966;Carnifossorius Deets and Ho, 1988, Eudactylinavan Beneden, 1853, EudactylinellaWilson, 1932, Eudactylinodes Wilson, 1932, EudactylinopsisPillai, 1966, HeterocladiusDeets and Ho, 1988, Jusheyus Deetsand Benz, 1987, Nemesis Risso, 1826, andProtâdactylina Laubier, 1966.This elasmobranch-copepod system is a good model to work on for manyreasons. The host group has been shown tobe monophyletic (Compagno,1977;Maisey,1984) , and this coupled with their antiquityas shown by paleontological data(Maisey, 1984) and molecular data (Davies, et al., 1986) should resultin a host-parasite system with a strong historical core. The phylogenetic relationshipsof living sharksand rays remain unsettled partly becausetoo few of the taxa have received investigation beyond the superficial statementsneeded for taxonomic identification (not unlikethe parasitic copepods), and partlydue to the depauperate fossil record of extant andextinct elasmobranchs (Compagno, 1977). Other problems obscuring elasmobranchinterrelationships are their morphological conservatism andthe fact that characterpolarity in elasmobranchs is difficultto define due to a lack of understanding of the characters in plesiomorphic outgroup taxa (Fechhelmand McEachran, 1984). Though thereare some phylogeneticanalyses in the literature (Compagno,1977, 1984a,b, 1988;Maisey, 1984; Heemstra and Smith,1980; Nishida, 1990 and Shirai 1992a, 1992b),8there is little agreement on any one phylogenyand it appears that more morphologicalwork and other modes of investigation (such asphylogenetic analyses of their parasites) are needed to to aid in resolutionand/or corroboration of these issues.Additionally, the parasite speciesassociated with this host group have not had sufficientattention paid to those morphologicaldetails that should be used for the specific discriminants in phylogenetic analyses, and thereforeare in need of this revision.Specifically, this effort involves the previouslydiscussed coevolutionary conceptsand cladistic methods, and empiricallyrevolves around this fascinating complex(Eudactylinidae and Kroyeriidae) of parasiticcrustaceans that inhabit olfactory andbranchial lamellae of an equally intriguing hostgroup, the elasmobranchs. I intendtoprovide a comprehensive taxonomic revisionand phylogenetic analysis for the twogenera Eudactyilna and Kroyeria. Specific rationaleregarding the choice of these twogenera follow.Eudactylina, world-wide in distribution, is found amongstthe branchial lamellaeof a systematically broad rangeof elasmobranchs. It is the most species-rich genus ofany gill-dwelling group of parasitic copepodson elasmobranchs. The principal attachment organ is the large chetate maxilliped,reminiscent of the second antenna inKroyeria. Eudactylina exhibitsa definite host preference for squaloid, squatinoid, pristiophoroid and batoid elasmobranchs (only 6 speciesfound on carcharhiniform hosts).This host association for Eudactylina is very interestingas some elasmobranch systematists (Maisey, 1984; Shirai, 1992a, 1992b)have radically hypothesized that thissqualoid-squatinoid-pristiophoroid “shark” lineageto be more closely related to batoidsthan to the other “sharks”. The conventionalnotion of shark monophyly would therefore, degrade into paraphyly. Recently, this complex hasbeen elevated to a monophyletic superorder the Squalea Shirai, 1992 when batoids and hexanchoids areincluded (Shirai, 1 992a). The Hypnosqualea of Shirai(1 992a) is a monophyletic subunit within the Squalea composedof the squatinoids plus pristiophoroids plus thebatoids, all hosts of Eudactylina.Hence, a taxonomic revision and systematic analysis9of Eudactyilna could aid in corroboration ofthis recently formulated, novelhypothesisthat not all sharks are sharks.Kroyeria occurs worldwide and is the secondmost species-rich genusof gill-dwelling copepods parasitic onelasmobranchs. All but one of the species are foundattached to the gill lamellae of theirhosts (Benz and Dupre, 1987; Deets, 1987).Kroyeria caseyl Benz andDeets, 1986 is atypical of the entire family in that the femaleis mesoparasitic (fossorial, anteriorportion of the animal usually modifiedinto a hold-fast and rooted into the host tissue,posterior portion exposed andfreely trailing),deeply embedded within the host’sinterbranchial septa. The male of K. caseyl, likethose of other members of thegenus, primarily attach themselves to the secondarylamellae and secondarily to the underlyingexcurrent water channels of their host’s gills(Benz and Dupre, 1987) by means of theirmodified chelate second antennae. The longvermiform body trails behind, nestled between the gillfilaments of its host. The carcharhiniform families Carcharhinidae (requiem sharks),Sphyrnidae (hammerheadsharks) and the Triakidae (hound sharks or whiskerysharks) are the primary hostsreported for this genus. Although these familiesare closely related (Compagno, 1977,1988) phylogenetic relationships within this carcharhiniformcomplex are considered tobe in a state of disarray (Maisey, 1984). In fact,both morphological and molecular evidence continues to mount suggesting thepossible paraphyly of the Carcharhinidae,Carcharhinus, and Triakidae (Compagno, 1988;Lavery, 1992; Naylor, 1992). A taxonomic revision and systematic analysis of Kroyeria, specificto the Carcharhiniformesmay assist in answering these aforementionedquestions.This research therefore, is an effort to apply phylogenetic systematics or cladistics to this parasitic crustacean-elasmobranch hostsystem in order to reveal and hopefully resolve the uncertainphylogenetic relationships of these hosts and parasites. Thiseffort proceeds by initial historical andbiological reviews and the subsequent taxonomIc revision and redescription of two parasiticcopepod genera, Eudactylina and Kroyeria.Additionally, both genera are in need ofrevision. Each revision is followed by a phylo10genetic analysis of that genus, from which parasite-derivedhost cladograms are constructed, and in two instances, area summary-cladogramsare produced. Competingindependent host phylogenies are then compared withthe parasite-derived host cladograms. Finally, the Kroyeria cladogram willbe numerically recoded and combined withyet another recoded tree from a previous revision and phylogeneticanalysis of the parasitic copepod genus Kroeyerina(cf Deets, 1987), into a common matrix to generate asingle host phylogeny based on all the parasite data from the Kroyeriidae.11MATERIAL AND METHODSChondrichthyan hosts were captured inthree general localities. Specimens fromthe southern Californiabight (San Diego to Point Conception,California) were.obtainedfrom the San Pedro, Californiabased commercial fishermen using set coastalgill netsand pelagic drift nets.Material from the Sea of Cortez or Gulf ofCalifornia was caughtwith gill nets, long lines andharpoon by the fishermen at Bahia de Los Angeles andPunta Arena de Ia Ventana.Sampling proceeded intermittently from October1980,through January 1994.Additionally, type and non-type materialwas obtained from the U.S. NationalMuseum of Natural History, Washington, D.C.,U.S.A., The Natural History Museum,London, England, and the MuseumNational D’Histoire Naturelle, Paris, France.Specimens were also received from the Institutode Biologia Marina, Mar del Plata,Argentina and Instituto de InvestigacionesOceanologicas, Universidad de Antofagasta,Antofagasta, Chile. The CaliforniaAcademy of Sciences in San Franciscq and theNational Museum of Natural History,Smithsonian Institution’s support center allowedme to inspect preserved elasmobranchs forparasitic copepods during my short-termvisitor appointment in June of 1 988.Additional specimens were donatedto me from the personal collections ofGeorge Benz, (Tennessee Aquarium);Dr. Roger Cressey, (National Museum of NaturalHistory, Smithsonian Institution),Dr. Ju-Shey Ho, (California State University, LongBeach), Dr. Z. Kabata, (Pacific BiologicalStation, Nanaimo, British Columbia), and RaulCastro Romero, (Universidad deAntofagasta, Antofagasta, Chile).Parasites recovered from thehost’s branchial and olfactory lamellae were immediately preserved and subsequentlystored in 70% ethanol. Later, copepods werecleared in 85% lactic acid, lightlystained with lignin pink, and transferred to wooden12slides according to procedures of Humes andGooding (1964). The parasitic copepodswere then dissected to permit a detailed morphologicalexamination of the appendages.All drawings were made with the aid of a cameralucida. Illustrations were drawn onCanson Vidalon Tracing Vellum, no. 110 (extra heavy),and inked with RotringRapidograph technical pens.Phylogenetic analyses were carriedout using the following protocol:Eudactylinodes Wilson, 1932, recently revealedto be the sister taxon of Eudactylina,and at times Eudactylineila,Carnifossorius, and Eudactylinopsis, sister group to theEudactylina-Eudactylinodes dade (Deets and Ho,1988), functioned as outgroups forEudactylina. Kroeyerina, the sister taxonto Kroyeria, and at times, Prokroyeria mendlonalis (Ramirez), the most basally placed member of theKroyeriidae (Deets, 1987)were chosen as outgroups for Kroyeria.These outgroups were selected in order todetermine character polarity for their respectiveingroups.Character data were defined, produced and coded (see appendices). All character data were treated and analyzed as unorderedto avoid risk of predetermining thetopology of the resultant cladogram (O’Grady and Deets, 1987;Dojiri and Deets, 1988).Non-linear, multistate transformation series (the recoded cladograms)were standardized by coding techniques reviewed and outlinedby O’Grady and Deets (1988). Uponcompletion of the character data matrix the computer program PAUP (PhylogeneticAnalysis Using Parsimony; D. Swofford, U.S. National Museumof Natural History,Smithsonian Institution, Washington, D.C., 20560) version 3.OS was used to analyzethe data. Specifically, the exact search algorithm BRANCH AND BOUND for smalldatasets and the heuristic search algorithm TREE-BISECTION AND RECONNECTION(TBR) for large data sets, were utilizedto generate the most parsimonious hierarchy ofparasite and parasite-derived host relationships. The computer program MacGlade (W.Maddison and D. Maddison, University of Arizona) version 3.0 was used interactivelyas a tool to increase insight on character evolution.Parasite cladograms were perceived as characterstate trees of their hosts, con-13verted into numericalcodes, combined and placedinto a data matrix andphylogeneticafly analyzed in orderto generate the final host phylogenies(Brooks, 1981; Brooksand McClennan 1991, 1993).Elasmobranch figureswere lifted and modified fromLast and Stevens (1994),and from Stevens (1987).14HISTORICAL REVIEWSince their inception in 1853 neitherEudactyilna van Beneden or Kroyeria vanBeneden have suffered anynomenclatural restructuring. However,they have experienced some shuffling at thefamilial level. Originally, both genera were assignedto theconvenient catch-all taxon Dichelesthiidae,the “tribu des Dichelestiens” of Edwards,1840. (For a comprehensive review of thehistory of Dichelesthiidae,see Kabata,1979). In 1853 the familywas composed of the genera Anthosoma Leach,1816,Dichelesthium Hermann, 1804and Nemesis Risso, 1826.Streenstrup and Lutken (1861) accepted“Dichelestiner” as a valid higher taxon.They included in it more generathan any previous authors, namely: Anthosoma,Congericola van Beneden, 1854, Dichelesthium,Eudactylina, Kroyeria, Lamproglenavan Nordmann, 1832, Lernanthropus de Blainville,1822, Nemesis and Pagodina(=Nemesis). The concept of Dichelesthiidaewas similar in the work of Krøyer (1863,1864), and von Nordmann (1864)used the structure of the egg sacs as the basic division between the genera. Hisgroup “Dichelestini” with filiform uniseriate egg sacs (asdisfinct from saccular multiseriate) consistedof: Anthosoma, Congericola,Dichelesthium, Donusa Nordmann, 1 864, Ergaslilna vanBeneden, 1851(=Nemesis),Eudactyilna, Kroyeria, Lamproglena(a cyclopoid), Lernanthropus, Nemesis, Pagodinaand Stalagmus Nordmann, 1864 (Donusa, Ergasillna,Pagodina, and Stalagmus are nolonger valid taxa).It wasn’t until Heller (1865) provideda key to the “Familia Dichelestina” thatsomeone offered an idea of intrafamilial groupingsby way of a key to the genera.Pagodina was concomitantly synonomized withNemesis.A similar approach to dichelesthiid systematicswas followed by Gerstaeker(1866-1879). His key to the family “Dichelesthiina”included the following genera:Aethon Krøyer, 1857, Anthosoma, Bacculus Lubbock,1860 (larval stage of PennellaOken, 1816), ClavelIa(=Hatschekia Poche, 1902), Congericola, Dichelesthium,Donusa (a polychaete parasite, probablya cyclopoid), Epachthes Nordmann, 1832(syn. of Lernanthropus), Ergasilina (syn.of Nemesis), Eudactylina, Kroyeria,Lamproglena, Lernanthropus, Nemesis, Norion Nordmann, 1864, PhilichthysSteenstrup, 1862 (type genus of Philichthyidae),Pseudocycnus Heller, 1865,Stalagmus (syn. of Lernanthropus) and Tucca Kreyer, 1837(a poecilostome).Brian (1 906) along withScott and Scott (1 91 3) included subsets of above in theirtreatment of Dichelesthiidae with representatives from their restrictedareas of study,15Italian and British waters respectively.Therefore, Dichelesthiidae withits vague and over inclusivediagnostic boundaries functioned as a depository for manynew genera and speciesthat could not beaccommodated in other existingwell-defined families.Wilson (1922), offered thefirst review of intrafamilial organizationof theDichelesthiidae by generating a keyto subfamilies. In his Copepoda of the Woods HoleRegion, Wilson (1932) upgraded thestatus of his four subfamilies and established thefollowing families with their respectivegenera:Anthosomidae (Anthosoma, Lernanthropus);E udacty Ii n idae (Kroyeria,Kroeyerina, Nemesis, Eudactyilna, Eudactylinodes,Eudactylinella);Pseudocycnidae (Pseudocycnus);Dich e lesth i idae (Hatschekia,Pseudohatschekia, Pseudocongericola, Lamproglena)Dichelesthium was excluded dueto its abscence from the Woods Hole Region.Yamaguti (1939) accepted the familial rankof these four families, whileMarkevich (1956) kept them as subfamiliesand transferred Lamproglena (thecyclopoid) to Eudactylinidae from Dichelesthiidae.Later Yamaguti (1963) reunitedWilson’s four families by establishing thesuperfamily Dichelesthioidea.Kabata (1979), at wits end, realized the lack ofrelationship between the variousgenera within these families, and the lackof affinity amongst the families embedded inYamaguti’s Dichelesthioidea and systematicallyrestructured this taxonomic complex.Using body segmentation for the first timeas the primary criterion for classification Kabata, (1 979) distinguishedsix groups of genera within this miscellaneous assemblage previously recognized asDichelesthiidae. Additionally, the cephalothoracicappendages that each of these groups posess wereshown to be unique. Witness thechelate second antennae of Kroyeriidae or thechelate maxiliipeds of Eudactylinidae.Coupled to this, one find similar patterns withthe swimming legs. Compare the fourpairs of well developed, non-modified biramous,trimerite swimming legs of Kroyeriidaewith the four pairs of Hatschekiidae,which have very reduced third and fourth legs andoften only bimerite rami of legs one and two asexamples. These pieces of evidenceadd layers of justification or corroboration to thegroups distinguished by Kabata’s tagmatic criterion. On the basis ofthese and many other arguments outlined in Kabata(1979), the following family unitswere proposed in accordance with this key:161. Four distinct segments between cephalothorax andgenital segment EudactylinidaeThree segments (exceptionally four)between cephalothorax and genital trunk2Segmentation indistinct, neck present between cephalothoraxand genital trunk3No free segments or neck between cephalothorax andgenital trunk, dorsal plate usually presentontrunkLernanthropidae2. Free segments distinct, cephalothoraxwith caligiform dorsal shield, four pairs of biramous legsKroyeriidaeFree segments rather indistinct,dorsal shield of cephalothorax not caligiform, three pairs of variouslymodified legs, elytra absentDichelesthiidae3. Second maxilla with bifid claw, maxilliped absentHatschekiidaeSecond maxilla with simple, denticulated claw, maxilliped present,subchelate. . PseudocycnidaeOne genus, Pseudohatchekia Yamaguti, 1939,cannot be accommodated in anyof these previous six families. If Yamaguti’sillustrations accurately represent the morphological attributes of this genus, it would seema new family (Pseudohatchekiidae)should formally be established upon inspectionand revision of the species therein.So, after 140 years of a tightly coupled and shared systematic history, in spiteoftheir obvious morphological disparities (perhapsdue to both being parasites of elasmobranchs) the paths of Eudactylinaand Kroyeria diverge, finally finding their way intotheir own families, Eudactylinidae and Kroyeriidae,respectively.17EUDACTYLINAEXTERNAL MORPHOLOGYGENERAL HABITUSThe body of Eudactylina is typicallysub-cylindrical with seven distinct tagmata(Figure 2a): the cephalothorax (whichconsists of the somites bearing thefirst antenna,second antenna, mandible, first maxilla,second maxilla, maxilliped, and thefirst pedigerous somite), the well-developedpedigerous somites two, three, and four(bearingbiramous and usually trimerous legs),a reduced pediger five (bearing a reduced fifthleg), a genital segment (bearingthe genital orifices), and a multi-segmentedabdomen(posteriorly bearing the caudal rami).The cuticle is equipped with posteriorlydirected cuticular expansions. Thesecuticular flaps or outgrowths vary insize and shape from species to species, therebypossessing some taxonomic importance.Eudactyilna orients itself upstream relative tothe flow of water over the gills(Figure 1), and these posterorly directed flapsmay function as tiny brakes keeping theparasite relatively secured at that location on the gill.The cephalothorax houses the main “organs”(appendages) of attachment forEudactyilna. These are the firstantenna with a few setae modified into large claws onthe second and third segments,the second antenna with its uncinate apical segment,and primarily the huge robustchelate maxilliped.The major articulation of the bodyis between the fifth pedigerous somite and thegenital somite.The genital somite of the female appearsto be located on the sixth thoracicsomite. Close examination of the ventrolateralarea adjacent to the oviducal openingsreveal three well developed albeit smallspines (Fig. 3D detail) which most likely represent the vestigial sixth legs. Additionally,the position of this putative sixth leg roughlycorresponds to the position that thereduced sixth leg occupies on the male. If thishomology in structure and position istrue, then it is likely that the abbreviated numberof segments in the abdomenof the female (two) relative to the supposedly more pIesiomorphic condition of the male(three or four segments), is due to the suppresion notincorporation (into a genital complex)of these somites during ontogeny.18CALJDALRAMUSThe caudal rami originateposteriorly from the last abdominal somite. Thecaudal ramus is undoubtably anorgan of slippage prevention. The parasite plungesthepaired rami down intothe secondary lamellae of the host’s gills(Figure 1), helping towedge the entire animalin place amongst the secondary lamellae.It seems the posteriorly-directed cuticular flaps onthe ventral surface of the rami coupled with thewelldeveloped, often digitiform terminalspines (modified setae) suggesta stopping function. Within the genus caudal ramivary markedly in form, from the relatively unmodified state possessing four relativelylong pinnate apical setae plus two relativelylongnaked setae as seen in themale Eudactylina epaktolampter (Figure 22B),to theextremely derived condition of possessing fourstout apical spines barely larger thanthe adjacent cuticular flaps asin female Eudactylina oliveri (Figure 36B), and thebizarrely modified digitiform conditionfound in the female of Eudactylina dactylocerca(Figure 14B). The striking differencesexhibited by the caudal rami between the different species of Eudactyilna make thisa useful taxonomic character.FIRST ANTENNAThe first antenna of female Eudactylina(Figure 2C) is indistinctly four-, five- orsix-segmented, exhibiting geniculate flexion betweenthe second and third segments.Although no physiological work has been doneon this appendage it is safe to presumesimilar (homologous) innervation exists hereas seen in other siphonostome copepodsindicating an organ capable of chemosensoryand tactile functions (see Kabata, 1979).Additionally, the large dorsally-directed claw-likespine (often toothed) on the secondsegment would seem to be an auxiliary attachment structure especially in lightof theway the parasite lodges its cephalothoraxinto the secondary lamellae of the gills(Figures 1A, 1B). The first antenna of the male hasa greater number of more clearlydelimited segments. The males of E. epaktolampterand E. oliveri (Figures.22C and38C, respectively) have eight or nine segments, andmore setae are found on the segments. In general (with regards to the female) the first or basalsegment always bearsone small seta on the outer margin. The apical segment appearsto possess a maximum of 14 setae plus one aesthete. Although somedescriptions in this revision and byother authors show fewer setae Ibelieve some of this may be an artifact of specimendamage coupled with the fact we are pressingthe limits of light microscopy with theminutiae involved.19The remaining segments between the basalsegment and apical segment exhibit different degrees of fusion or segment incorporation inthe different species.SECOND ANTENNAThe second antenna (Figure 2D) isfive-segmented, not sexually dimorphic.The basal segment is small, unarmedand pedunculate. The second segment is relatively elongate, sometimes armed witha styliform process (appears to havea prehensile function), and sometimes possessing cuticularflaps. The third segment usuallybears a large claw-like extension and alwayshas two slender setae at its base andcuticular flaps. The fourth segment sometimesexhibits cuticular flaps. The fifth segment forms a large uncinate terminal claw bearinga large accesory spine on the lateralsurface. Two small slender setae are always foundat the base of the fifth segment.The many different combinations andstates of these characters listed abovemake the second antenna an important taxonomicdiscriminant.ORAL CONE and MANDIBLEThe mandible is typically siphonostome beinga uniramous subcylindrical structure with a dentiferous distal end(Figure 2E). The mandible also appears to be dividedinto two parts, their boundaries demarcated bya suture. The dentiferous margin bearsfrom five to eight teeth.The oral cone (Figure 36D), consisting of labium and labrum housesthemandible. The possession of the mouthtube is the distinguishing characteristic of thesuborder Siphonostomatoida. The structure is ratheruniform throughout the genuswith some differences seen in the nature of the cuticular flapsfound on both the labiumand labrum. The use of this structure for systematic purposes was not pursued.FIRST MAXILLAThe first maxilla found adjacentto the mandible is the first segmental appendagefound in the post-nauplial stages of copepods (Kabata, 1979). It is biramous consistingof an endopod and an exopod(Figure 2F). The exopod is armed with two setae whichmay bear setules or denticulations.The endopod terminates with one long seta and twoshort setae the former bearing setules or small denticulations in certain species.The20body of the first maxilla also bears small spinulationsor cuticular flaps in some species.SECOND MAXILLAThe second maxilla isa large brachiform appendage (Figure 2G). Theorifice ofthe maxillary gland is presentat the base as is the basal process,a small fleshy extension of cuticle. The most proximalsegment is the lacertus. The lacertus is typically “fortified” by the presence of welldeveloped sclerites, and isarmed with cuticular flaps.This segment articulates with thebrachium by means of the cubital joint. Thebrachiumis also armed with the cuticular flapsand distally bears what appears tobe a tuft orpaired tuft of setae, and apatch of prickles or denticles termed the crista at the base ofwhat may represent a thirdsegment the calamus or claw. The claw usually bearsaseries of comb-like serrated membranesor, as in a few species, well developed teeth.Although the comb-like membranessuggest a grooming function, the closely relatedspecies Nemesis robusta (van Beneden,1851) also a member of Eudactylinidae hasbeen shown to use the second maxillato assist in feeding on the secondary lamellae ofthe thresher shark (A/op/as vulpinus(Bonnaterre, 1758)). Feeding by these gill parasites involves the mechanical raspingof host tissue and the second maxillae wereshown in histological sections to be theappendage responsible for host tissue excavation (Benz and Adamson,1990). The second maxilla has also been implicated inmanipulation of the frontal filament during thedevelopmental stages of most siphonostomes in which they are known (Kabata,1979). Unfortunately, the specific function ofthe second maxilla in Eudacty/ina isunknown.MAXILLIPEDThe maxilliped is the posteriormost oral appendage borneupon the first thoracicsegment incorporated into the cephalon. Sexually dimorphic,the male maxillipedoccurs as a sub-chelate structure (Figure22D), while the female form exists as aremarkable, fully chelate structure (Figure 2H). The femalemaxilliped appears to consist of four segments. The unarmed pedunculatebasal segment supports the mainbody or corpus maxillipedis. The corpus maxillipedisis always armed with cuticularflaps, a small spine-likeseta, at times a lateral flange, and a greatly produced posterolateral region, the myxa, forming alarge scoop-like receptacle. By means of the cubitaljoint the corpus articulates with thesubchela. The subche!a is composed of the proximal shaft bearing oneseta approximately midway along the outer margin and another21located distally near the base of the clawalong the inner margin. A membraneis alsofound adjacent to the latter along theinner margin of the most distalreaches of theshaft. The shaft may bear cuticular flaps. Themost terminal segment of thesubchela isthe claw. This is an uncinate structure accompaniedalong the lateral surfaceby aquadrangular expansion of varying sizes. Whenclasped, the claw and quadrangularexpansion are perfectly accommodatedby the receptacle-like myxa, forming a pincer.This structure is the primaryattachment appendage that the copepoduses to grip thesecondary lamellae of the host’s gills(Figure 1).LEG ONEThe first thoracic leg is the only leg incorporatedinto the cephalothorax (Figures2A and 3A). The pair of legs is connectedby an interpodal bar allowing their synchronous movement. The sympod is welldelimited by a media’ suture separating the proximal coxa from the adjacent basis. Bothcoxa and basis are armed ventrally with cuticular flaps or scales. The basis bearsone lateral and one somewhat medialseta along itsdistal margin. The leg is alwaysbiramous, composed of a lateral exopod and a medialendopod. Both rami are usually three-segmented(trimerous) and rarely bimerous. Theproximal (first) and middle (second) segmentsof the exopod bear a distolateral spine-like seta (pinnate setae are found in legs onethrough four in the male). The terminal(third) segment typically bears threeor four variously modified setae. The endopod isalso a trimerous or rarely bimerousramus with only the terminal segment armed withtwo setae.LEG TWOThe second thoracic legs (Figure7B) are united by an interpodal bar. In a fewspecies the interpodal bar of leg twohas a non-articulated medial extension creating asingle ventroposteriorly directed interpodal stylet (Figure 15C).The coxa and basis aredistinct, with the basis armed withone seta along the distolateral edge. The secondthoracic appendage is also biramous, with bothrami trimerous except in a few cases inwhich the endopod is two-segmented.The terminal segment of the endopod carries twosetae. The exopod with few exceptions(e.g., E. acanthi,) is a bizarrely modified ramus.It has few if any cuticularflaps, in contrast to the other three leg pairs. The proximalsegment is greatly enlarged, oftentremendously elongate and bears a single curved22spine distolaterally. The middle segment is armedwith a single spine-likeseta of various sizes. The terminal segment is reduced in sizebearing a small lateral seta, an oddquadrangular or thick seta is found terminally and a stronglyhooked seta located onthe medial margin of the segment. The sometimes strongly producedspine on the middle segment and the strongly hooked seta on the medial edge of theterminal segmentwhen viewed in context with the in situ illustration of Figure 1, suggeststhe ramus functions in slippage prevention if notas an auxiliary attachment appendage.LEGS THREE AND FOURThe third and fourth pairs of legs (Figure3C) are nearly identical in all species ofEudactylina with the exception of E. doilfusi where the endopodof the fourth leg ismodified into a robust uncinate process (Fgure 19D). Each leg pair isjoined by aninterpodal bar. In a few species the interpodal bar of legthree has a non-articulatedmedial extension creating a single ventroposteriorlydirected interpodal stylet (Figure33E). Both pairs of legs are adorned with cuticular flaps.The sympod is well delimitedinto a proximal coxa and more distal basis. Thebasis bears a single spiniform seta onthe lateral margin. Both legs are biramous and the ramitrimerous with the appearanceof a few bimerous endopods on leg four being theexception. The terminal segmentbearing a single seta is the only setal armature found on theendopod. The exopodbears a single spine-like seta on both segment one and two, and three spine-like setaeon the terminal segment. The third and fourth pair of legs with theirposteriorly directedcuticular flaps probably aid in slippage prevention. Additionally, Figure 1 showsEudactylina using their endopods and exopods independently, in fact,at right angles toeach other to what appears to be separating the secondarylamellae in order to moresecurely wedge itself amongst the lamellae. This would seem to cause the secondarylamellae to attempt to return to their normal positions possibly pressuring or pushingback down and around the parasite aiding in its attempt to remain securely affixed tothe host tissue.LEG FIVEThe fifth leg is a uniramous, one-segmented appendage found on the last andfourth free thoracic somite (Figure 2A and 3D). The sympod consists ofa single segment and a small seta is found distodorsally. The ramus bears three naked or pinnate23slender setae distally. Dependingon the species, bothsympod and ramus maybeunarmed or covered with cuticu tar flaps.The sixth leg of the female is rarelyseen (or looked for)as it appears to be composed of three minute stout spine-likesetae adjacent to the opening of theoviducal orifice (Figure 3D detail) onthe genital segment. The sixth leg ofthe male is found on thedistolateral edge of the genitalsegment. It is represented by twoor three setae arisingfrom a small flap, presumablythe sympod (Figure 22A and 23G).LIFE HISTORYGENERAL DESCRIPTIONThe first mention ofa larval eudactylinid was given by Wilson (1922) forEudactylinodes nigra. Devoid ofillustrations, the narrative description tells us littlemore than that the organism wasa nauplius. The next time any mention ofaeudactylinid larva appears is by Kabata (1976)for Eudactylina similis. Unfortunately,Kabata was unable to culture the parasitebeyond the first naupliar stage. Hence, nothing is known about the infectivestage of Eudactylina.In general, most fish-parasitizingsiphonostomatoid copepods with known onto-genies possess a three part post-embryonic developmentconsisting of the nauplius,copepodid, and adult (Raibaut, 1985) . The numberof naupliar stages varies from oneto six. Next is a single copepodid stage,followed by four modified copepodite stages,chalimus I - IV, that possesses, in manybut not all species, the remarkable anchoringdevice, the frontal filament. Accordingto Raibaut (1985), the frontal filament is formedduring the copepodid stage and is extruded forattachment to the host immediately priorto the molt into the first chalimusstage. In contrast, Wilson (1911, P1. 30, Fig. 10)shows a nauplius of the Iernaeopodid,Achtheres ambloplitis with an already formed,coiled frontal filament. Although this chalimuslarva had been considered to be presentin all of the siphonostomecopepods of fish, Cabral, Coste, and Raibaut (1984•) demonstrated this was not the case. Experimentalinfestations of Lernanthropus kroyeri vanBeneden, 1852 on Dicentarchus labrax (Linne,1758)revealed that this species possessed two free swimming naupliarstages but lacked the chalimus stages. It did haveone free living copepodidstage followed by the infectious second fixed copepdid. Fixedcopepodids Ill, IV and V preceededthe two sub-adult stages before finally transforminginto the adult. Furthermore,Kabata and Khodorevski (1977) described a copepodid,not a chatimus, from another gillinhabiting siphonostome Dichelesthium oblongum24(Abildgaard, 1794). The fact that I could notfind any evidence of any frontal filamentremnants or frontal gland scars on the manyeudactylinids I have examined suggeststhat a similar life cycle prevails in this group ofcopepods. It can be presumbed thatthey become infective as copepodids, attachthemselves presumably with the secondantennae and maxillipeds, suppress or skip the chalimusstage and molt finally into thepre-adult or adult stage. Indeed, Kabata (1981)suggests with regard to the absence ofthe frontal filament in therelatively primitive (minimal cephalization) siphonostomeDissonus nudiventris that the frontal filament is probablya relatively derived character.Additionally, the majority of fish parasitizingsiphonostome copepods exhibit aholoxene or direct life-cycle, requiring only asingle host to live out their life. Again theevidence suggests this is probably the case with theeudactylinids, if not all of the gilldwelling families of possible dichelesthoidaffinities (Eudactylinidae, Kroyeriidae,Hatchekiidae, Lernanthropidae and Dichelesthiidae).REPRODUCTIONCopulation apparently occurs between the adultstages of the parasites. I haveobserved preserved specimens in the presumedcopulatory embrace similar to whathas been described by Benz and Adamson (1990)for Nemesis robusta (van Beneden,1851), also a member of the Eudactylinidaeparasitizing the common thresher shark(Alopias vulpinus (Bonnaterre, 1758)). The male attachesto the female using the prehensile second antennae and subchelate maxillipeds. The male thensomehow transfers two spermatophores to the lateral surfaces ofthe female’s genital segment whichattach at the female’s oviducal openings (Figure 3D and 16A) (see Benz and Adamson,1990 for a detailed description of the morphology and attachment of the brownbodyand seminal vesicle in N. robusta). When the eggs exit the oviducal opening,they arefertilized by the males gametes from the seminal receptacle. Ovigerousfemales produce uniseriate egg sacs, presumably secreted by cement glands located in the genitalsegment.25HOST-PARASITE RELATIONSHIPSDELETERIOUS EFFECTS / FEEDINGEudactyilna always attach themselvesto the secondary Iamellae of their elasmobranch host’s gill, principally by their clasping chelatemaxilliped (Figure 1). Whenextracting these parasites from the gills Ihave never observed gill tissue pathologiesinduced by Eudactyilna. Certainly, some damage is beingdone since most of the specimens still had gill tissue gripped in their maxillipedswhich had to be carefully removedbefore the microscopic examination andillustrations could proceed.Additionally, over the last 14 years of collectingEudactylina from elasmobranchgills I have observed a general trend forthis genus to exhibit a rather low parasite loadrelative to other elasmobranch gill infecting siphonostomegenera such as Nemesis andKroyeria. For example, the maximum numberof Eudactylina acanthil A. Scott, 1901 Icollected from a spiny dogfish (Squalus acanthiasLinnaeus, 1758), was 60 (ayerage offive specimens was 35). Similarly, a single Pacificelectric ray (Torpedo cailfornica,Ayres) yielded a maximum of 77 Eudactylina similisT. Scott, 1902. These maximumnumbers are reported here because it is very commonto recover less than five individual Eudactylina from a single batoid orsqualoid host.Because individual Eudactylina seem to be attachedto the secondary lamellae,presumably they feed directly on these respiratory surfaces. Unfortunately, no studieshave been done on gut contents in Eudactylina. Benz and Adamson(1990) studiedhistological sections in the closely related eudactylinid Nemesis robusta, andfounddark staining granules (partially digested blood?)reminiscent of those commonly foundin the diverticula of haematophagus monogenetictrematodes.SPECIFICITYEudactylina exhibits a high degree of both ecological and host specificity.General observations reveal Eudactylina to distribute itself in no apparent pattern orpreferred areas across the host’s hemibranchs, and Eudactyilna will onlybe found onelasmobranch gills. Regarding host specificity, the majority of the species ofEudactylina are specific to a given host species or genus, making them good biologicaltags. A few species are apparently more flexible with their choice of hosts but still showspecifity at a more general level in thehierarchy, restricting themselves to hosts within a26given family. Specifics are detailed in both thetaxonomic account andphylogeneticanalysis below.SYSTEMATIC ACCOUNTGenus Eudactylina van Beneden, 1853Eudactylinidae: Female. Cephalothorax coveredby well demarcated dorsal shield; foursucceeding thoracic segments bearing cuticular flaps onterga. Genital segment small,quadrate and bearing oviducal openings and reduceduniramous leg five. Abdomentwo-segmented. Caudal ramus bearing from fourto six sometimes modified setae.Posteriorly directed cuticular flaps present on ventralsurfaces of genital segment,abdomen, and caudal ramus.First antenna indistinctly four to six segmented with geniculateflexion betweensecond and third segments. Second segment bearinglarge curved prehensile claw.Terminal segment generally bearing one medialseta, one lateral aesthete, plus an additional 13 to 14 slender setae. Second antenna five-segmentedwith prehensile terminalclaw. Mouth tube siphonostome. Mandibletwo-segmented; distal end dentiferous,bearing five to eight teeth. First maxilla biramous withendopod and exopod armed withthree and two apical setae respectively. Second maxillabrachiform, two- possiblythree-segmented. Maxilliped chelate, myxa produced intoa large expanded receptacle.Legs one through four biramous, rami bimerousto trimerous. Exopod of leg two usually,though not always, modified (cf. E. acanthi,). Legfive one-segmented and uniramousbearing three distal setae. Leg six represented by threeminute spines at oviducal orifice.Male: Similar to female. Abdomen three- to four-segmented. Caudal ramus lessmodified. Maxilliped subchelate, myxal area bearinga strong spinous process. Firstfour pairs of less modified legs, with more plesiomorphicarmature (long pinnate setae).Leg six on posterolateral edge of genital segment representedby two or three setae.TYPE-SPECIES: Eudactylina acuta van Beneden, 1853.COMMENTS: In spite of the general uniformity of habitus, the species in thisgenus differ from another in a multitude of structural details suchas morphology of thecuticular flaps, armature of the second antenna,armature and segmentation of thoracic27legs and in specific characterattributes of the caudal ramus.Currently, Eudactylina consists ofthe 26 species illustratedand phylogeneticallyanalyzed herein plus 12 nominal but unfortunatelyfor this investigationunobtainablespecies, bringing the total to 38. The unobtainablenominal species willbe reviewed atthe end of the following section. An additionalfour species (E. carchariaeglauciHesse,1884, E. mustelilaevis Hesse,1884, E. puriensis Tripathi, 1956, and E squatinaeangeliHesse, 1884) have been inadequatelydescribed to be recognized and are consideredspecies inquirendae. The majority ofthe species described are parasites of squaloids(dogfish, lantern sharks), squatinids(angel sharks), pristiophorids (sawsharks),andbatoids (skates, guitarfish andrays). The remaining species are found ontheCarcharhinidae (requiemsharks) and Sphyrnidae (hammerheads). There are approximately 350 species of “sharks”(Compagno, 1984a), and 425-450 species ofrays(Eschmeyer et al, 1983), and only38 species of the highly host specific Eudactylina areknown. This information suggeststhis genus is potentially enormous with many morespecies waiting to be discovered.Eudactylina acanthil A. Scott, 1901(Figures 2-3)Material examined. Severalco-type females, BMN H 1911 .11 .8.48318-322,1913.9.18.272-281, 1963.4.29.15, 1975.379-392from British waters. Many femalesfrom the Vancouver Island region onloan from Dr. Z. Kabata, Pacific Biological Station,Nanaimo, British Columbia. Numerous femalesfrom the southern California Bight. Allspecimens were found attached to the branchiallamellae of the spiny dogfish, Squalusacanthias Linnaeus (1758).DescriptionFemale (Figure 2A)Overall length in lateral view approximately 2.25 mm.Cephalothorax longer thanwide, lateral margin notched accomodatingthe second maxillae. Ventrolateral marginof cephalothorax bearing small cuticularflaps. Tergum of first free thoracic somite withsmall cuticular flaps on ventrolateral margin.Second and third free thoracic somiteswith naked and indistinct terga.Fourth free thoracic somite bearing fifth le smallerthan previous two. Genital segmentsmaller than preceeding somite. Abdomen two-segmented, second segment ventrally bearing cuticular flaps.Caudal ramus (Figure25) longer than wide, bearingfour terminal naked setae, one slender, medial seta, andone lateral nakedseta; ventral surface armed with posteriorly directed cuticular flaps.28First antenna (Figure 2C) indistinctlyfive- or six-segmented,armature (proximalto distal) as follows: one stout seta, one smallspinule, eight stout setae plusone largecurving (prehensile) claw, ninestout setae, one short seta, 14 slendersetae plus oneaesthete. Second antenna (Figure 2D)five-segmented, uncinate, prehensile.Basalsegment stout, second and thirdsegments with sparse cuticular flaps, thirdsegmentbearing two slender setaearising from atypically reduced spinous process.Fourth segment elongate and unarmed;fifth segment an unciformterminal claw bearing two slender setae and onestout auxiliary spine approximately 1/4length of claw. Mouth tubesiphonostome and similarto that of other species. Mandible(Figure 2E) of two parts,dentiferous margin with six teeth.First maxilla (Figure 2F) biramous; endopod bearingtwo apical setae; exopodlonger surmounted by two stout setae andone longer bilaterally denticulated seta. Secondmaxilta (Figure 2G) brachiform, lacertuslarger thanbrachium armed with triangular cuticular flaps,brachium with triangular cuticular flapsand a tuft of coarse, sparse setaeat base of terminal claw (calamus). Claw bearingtwo rows of denticlesand proximal serrated membrane. Maxilliped (Figure 2H)chelate, indistinctly segmented, proximal segmentpedunculate; corpus maxillipedisrobust, bearing small stout spine ondistal margin; myxal area expanded into largereceptacle to accommodate claw of opposablechela. Shaft with single spine on lateralconvex margin and strip of membranealong concave distal margin. Claw unciform withquadrangular cuticular expansion producing lateralshield.First four pairs of legs biramous with three-segmentedrami and two-segmentedsympods. All basipods with lateralslender seta; first basipod bears additional distomedial seta. Ventral surfaces of all fourlegs bearing triangular cuticular flaps. Armature oframi as follows: (non-pinnatesetae in Roman numerals, pinnate setae (bearingsetules) in Arabic numerals).Leg one Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg two Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0 Ill Endopod0-0 0-0 ILeg one (Figure 3A) terminal segment of exopod compressedbearing two smallspiniform seta, lateral seta with singledenticle; terminal segment of endopod with twomost lateral setae denticulated, most medial setapapilliform. Leg two (Figure 3B) withatypically unmodified exopod, all setae on exopodand endopod spiniform and smooth.Legs three and four (Figure 3C)similar;all setae spine-like and denticulated3terminalsegment of endopod compressed.Leg five (Figure 3D) suboval; distally bearing three29long slender setae plus one similarseta arising dorsally from the base. Leg six (Figure3D detail) represented by three small, stout setaeon a posterodorsal ridge adjacenttothe oviducal opening.Male: UnknownComments: Kabata (1979)mentions the possibility of misinterpreting cuticularflaps for the very tinyspine-like setae and vice-versa on the swimming legs of E.acanthiL In spite of this word of caution,I have found the number of apical elements onendopods one, two, three, and four tobe two, two, one, and one respectively, in contrast to Kabata’s three, three, three, and three, theformer being generally more consistent with the endopodal formula exhibitedby the genus.Eudactylina typically exhibits a bizarrely modified exopodon leg two. Thisspecies however stands out as the anomaly withinthe genus in possessing a normalendopod (resembling endopods of legs one, three, andfour).All previous host records of Eudactylina acanthii comespecifically from thebranchial lamellae of the spiny dogfish Squalusacathias. Like most parasitic cope-pods, the geographic range of this parasite is coincident with that of itshost. This cope-pod has been reported from the Irish Sea, eastern and western North Atlantic,Sea ofJapan, Vancouver Island region, coastal Angola inthe southern Atlantic (see Kabata,1979), Quehui, Chiloe’, Chile (Castro andBaeza, 1991), and now from southernCalifornia waters.Eudactylinaacuta van Beneden, 1853(Figures 4-5)Syn: Eudactylina complexa Brian, 1924 (see Kabata, 1979)Material examined. Several females, MNHN, CP 156 and CP 173 from thebranchial lamellae of the angelshark, Squatina squatina (Linnaeus, 1758)restricted tothe western North Atlantic and the Meditteranean (no specific site collectiondata).DescriptionFemale (Figure 4A)Overall length in lateral view approximately 3.3 mm. Cephalothorax longer thanwide, lateral margin notched accomodating lacertus of second maxillae. Ventrolateral30and dorsal surfaces of cephalothorax bearing smallcuticular flaps. Tergum of first freethoracic somite with small cuticular flaps on dorsalsurface. Second and third free thoracic somites with cuticular flaps on the dorsal surfaceand distinct terga. Fourth freethoracic somite smaller than previous two,bearing leg five. Genital segment smallerthan preceeding somite, with cuticular flaps onventral surface. Abdomen two-segmented, ventral surface bearing cuticular flaps. Caudalramus (Figure 4B) longer than wide,bearing two terminal stout setae, one slendermedial seta, and one lateral naked seta;ventral surface armed with patches of posteriorly directed cuticularflaps. Oviducalopening dorsal, egg strings uniseriate (Figure 4C).First antenna (Figure 4D) indistinctly five-segmented, armature(proximal to distal) as follows: one stout seta, four stoutsetae plus one large serrated, curving (prehensile) claw, three slender setae, two short setae plustwo well developed denticulatedclaws, two large stout setae and 12 slendersetae plus one aesthete. Second antenna(Figure 4E) five-segmented, prehensile. Basal segment stout(not shown), secondsegment with stout spiniform process, third segment bearing rectangular cutici.jlar flaps,two slender setae arising from large, curving spinousprocess. Fourth segment elongate and unarmed; fifth segment an unciform terminal claw bearing two slender setaeand one stout auxiliary spine nearly reaching theend of the claw. Mouth tubesiphonostome and similar to that of other species. Mandible (Figure 4F) of two parts,dentiferous margin with seven teeth. First maxilla (Figure 4G) biramous; endopodbearing two apical setae; exopod longer surmountedby two stout setae and one longerseta. Second maxilla (Figure 4H) brachiform, lacertus larger than brachium armed withcuticular flaps, brachium with crescent-shaped cuticular flaps and two tufts of setae(one coarse and one fine) at base of terminal claw (calamus). Claw bearing three distal rows of serrated membranes and one proximal serrated membranous flap.Maxilliped (Figure 41) chelate, indistinctly segmented, proximal segment pedunculate(omitted in illustration); corpus maxillipedis robust bearing small stout spine on distalmargin, rectangular cuticular flaps and a large transverse cuticular flange; myxal areaexpanded into large receptacle to accommodate claw of opposable segment. Shaftwith two setae: one near midpoint on lateral convex margin and the other adjacent to astrip of membrane along concave distal margin. Claw unciform with quadrangular cuticular expansion producing lateral shield.First four pairs of legs biramous with three-segmented rami and two-segmentedsympods. All basipods with lateral slender seta; first basipod bears additional distomedial slender seta. Ventral surfaces of all four legs bearing crescent shaped cuticularflaps. Armature of rami as follows: (non-pinnate setae in Roman numerals, pinnatesetae (bearing setules) in Arabic numerals).31Leg one Exopod 1-0 1-0 IVEndopod 0-0 0-0 IILeg two (modified) Exopod 1-01-0 III Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-01-0 III Endopod 0-0 0-0 ILeg one (Figure 5A) all setae denticulated.Exopod two (Figure 5B) typicallymodified without cuticular flaps, firstsegment greatly enlarged bearing one smoothseta, second segment smaller with hugestout spine, third segment compressed apically armed with one stout seta, onetruncate seta and one recurving slender seta.Legtwo endopod (Figure5C) with naked proximal segment, segments two andthree withcuticular flaps and two finely spinulatedsetae on apical segment. Legs three and four(Figure 5D) similar; endopodalseta denticulated, exopodal setae curving with branching tips (Figure 5E). Leg five (Figure5F) oval; lateral surface covered with rectangularcuticular flaps, distally bearing three long slendersetae plus one similar seta arisingdorsally from the base.Male: Not obtained (see Kabata, 1979)Comments: E. acuta appears tobe specific to the angeishark Squat/na squat/na. Kabata (1979) mentionsthis parasite has been recorded (Valle, 1880; Brian, 1906;Oorde-de-Lint and Schuurmans Stekhoven,1936) from the spiny dogfish Squalusacanthias, but neither he nor I have ever seenthis copepod on that host (hundreds ofwhich I have examined) suggesting asuspect association.The geographical distribution of Squat/nasquat/na ranges from southernNorway, Sweden and Shetland Islandto Morocco off the western Sahara, CanaryIslands, and the Mediterranean (Compagno, 1984a). Predictably, thecopepod hasbeen reported from most of this range.Additionally, Kabata (1979) has tentatively synonymizedE. complexa (Brian,1924) with E. acuta. Besides similaritiesbetween descriptions of the two species, bothare found on the same host in thesame area. Hence, the records of E. complexa (=E.acuta) from the host genera Torpedo, Pteromylaeus, Raja,and Myliobatis reported byEssafi and Raibaut (1977) from theMediterannean will be treated as uncertain hostassociations.This species is easily distinguishedby the nature of the caudal ramus, the huge,stout spine on the second segment of the modifiedexopod of leg two, and the branching, digitiform claw-like setae on theexopods of legs three and four.32Eudactylina aphiloxenossp. nov.(Figures 6-7)Material examined. Numerous femalesfrom the branchial lamellae of the Pacificangelshark Squatina californicaAyres, 1859 from the southern California bight. Femaleholotype (USNM 266519) and 7 female paratypes(USNM 266520) deposited at theUnited States National Museum of NaturalHistory.Etymology : The specific name aphiloxenos is derived fromthe greek aphilo, forunwanted or hateful and from the greek xenos forguest. Thus, the unwanted guest.DescriptionFemale (Figure 6A)Overall length in lateral view approximately 2.2 mm. Cephalothoraxlonger thanwide, lateral margin notched near base of maxillipedand lacertus of second maxillae.Ventrolateral and dorsal surfaces of cephalothorax bearing smalltriangular cuticularflaps. Tergum of first free thoracic somite with small triangular cuticularflaps on dorsalsurface. Second and third free thoracic somites with triangularcuticular flaps on thedorsal surface and distinct terga. Fourth free thoracicsomite smaller than previous two,with distinct tergum and triangular cuticularflaps, bearing leg five. Genital segmentsmaller than preceeding somite, with cuticular flaps on ventral surface.Abdomen two-segmented, ventral surface bearing cuticular flaps. Caudal ramus(Figure 6B) slightlylonger than wide, bearing two stout terminal setae, one naked dorsalseta, and one lateral naked seta; ventral surface armed with posteriorly directedtriangular cuticularflaps.First antenna (Figure 6C) indistinctly five-segmented, armature (proximal to distal) as follows: one stout seta; six smooth slendersetae, two denticulated setae (oneshort (ghosted in near lateral margin), one long) plus one large serrated,curving (prehensile) claw; four slender setae plus one stout bilaterally denticulatedseta; two shortsetae plus two well developed denticulated spines; 14 slender setae, one unilaterallydenticulated seta plus one aesthete. Second antenna (Figure 6D) five-segmented, prehensile. Basal segment stout, second segment with stout spiniform process, third segment bearing triangular cuticular flaps, two slendersetae arising near base of largestyliform process. Fourth segment elongate with small cuticular flaps along convexmargin; fifth segment an unciform terminal clawbearing two slender setae and onestout auxiliary spine. Mouth tube siphonostome and similar to that of other species.33Mandible (Figure 6E) of two parts,dentiferous margin withseven teeth. Firèt maxilla(Figure 6F) biramous; endopod bearingtwo apical denticulated setae;exopod longersurmounted by two stout setae and onelonger denticulated seta.Second maxilla(Figure 6G) brachiform, lacertus slightly largerthan brachium armed withtriangularcuticular flaps, brachium with triangularcuticular flaps and two tufts of setaeand possibly a small reduced spineat base of terminal claw (calamus). Medialsurface of clawbearing three distal rowsof serrated membranes andone proximal serrated membranous flap, lateral surface onlyarmed with two serrated membranous flaps. Maxilliped(Figure 6H) chelate, indistinctlysegmented, proximal segment pedunculate(omitted inillustration); corpus maxillipedisrobust bearing small stout spine on distal margin,triangular cuticular flaps; myxalarea expanded into large receptacleto accommodate clawof opposable segment. Shaft withtwo setae: one near midpointon lateral convex margin and the otheradjacent to a strip of membrane along concavedistal margin. Clawunciform with transverse cuticular flangebearing quadrangular cuticular expansion producing lateral shield.First four pairs of legs biramouswith three-segmented rami and two-segmentedsympods. All basipods with lateral slender seta;first basipod bears additional distomedial slender seta. Ventralsurfaces of all four legs bearing triangular cuticular flaps.Armature of rami as follows:Leg one Exopod I-CI-C IV Endopod 0-0 0-0 IILeg two (modified) Exopod I-C I-C IllEndopod 0-0 0-0 IILeg three Exopod 1-C1-0 III Endopod 0-0 0-0 ILeg four Exopod 1-01-0 III Endopod 0-0 0-0 1Leg one (Figure7A) all setae (except mostmedial seta on terminal segment ofexopod) denticulated. Exopod two(Figure 7B) typically modified without cuticularflaps, first segment greatly enlargedbearing one smooth seta, second segment smallerwith slender seta, third segmentcompressed apically armed with one short seta, onetruncate seta and one recurving slenderseta. Leg two endopod (Figure 7B) with twofinely denticulated slendersetae (one long, one short) on apical segment. Legs threeand four (Figure 7C) similar; endopodalseta denticulated, exopodal setae curving withbranching tips (Figure 7C detail). Legfive (Figure 7D) elongate; lateral surface coveredwith triangular cuticular flaps, distallybearing three long slender setae plus one setaarising dorsally from thebase.Male: Not found34Comments: E. aphiloxenos appears tobe specific to the Pacific angeisharkSquat/na californica.Squatina californica ranges from southeasternAlaska to the Gulf of California(Compagno, 1984a). Interestingly, Kato, Springer andWagner (1967) synonomizedthe southern angelote Squat/na armata (Philippi, 1887) from offthe eastern SouthPacific shores of South America with this hostspecies. Compagno (1984a) mentionsthere is evidence against this taxonomicinterpretation and adds that the Gulf ofCalifornica angelshark may in fact be differentfrom the Pacific angelshark. Evidence insupport of Compagno’s position is added here withparasite data. The southernangelote originally Squat/na armata is parasitized by Eudactyl/natuberifera Castro andBaeza (1987) from off the West coast of Chile,a species unmistakeably distinct (andredescribed later herein) from the new species infectingSquat/na californ/ca.E. aph/loxenos is similar to E. acuta and E. tuber/fera, both parasites ofangelsharks. All three species possess relatively large (almostdigitiform) branchingtips on the setae of exopods three and four. E.aphiloxenous can be distinguished fromits other two allies by its relatively elongate fifth leg,the relatively small and slenderseta on the second segment of the modified second exopod, its very small and numerous triangular-shaped cuticular flaps, and its lack of the transverse cuticular flange onthe corpus of the maxilliped.Eudactylinaaspera Heller, 1865(Figures 8-9)Material exam/ned. One female, BMNH 1968.1.5.3 from the branchial lamellaeof the brownbanded bamboo shark, Chiloscyllium punctatum Muller and Henle, 1838collected from Moreton Bay, Queensland. Several females USNM 153636 from thebranchial lamellae of the milk shark, Rhizopr/onodon acutus (Ruppel, 1837), severalfemales USNM 153634 from the spinner shark, Carcharhinus brev/pinna (Muller andHenle, 1839), (=Carcharh/nus maculipinn/s (Poey, 1865)), and numerous femalesUSNM 153639 from the scalloped hammerhead, Sphyrna lewini (Griffith and Smith,1834), all collected from the Indian Ocean near Nosy Be, Madagascar.Descr/pt/onFemale (Figure 8A)Overall length in lateral view approximately 1.5 mm. Cephalothorax longer than35wide, lateral margin notched accomodatinglacertus of second maxillae.Ventrolateraland dorsal surfaces of cephalothorax coveredwith cuticular flaps. Tergumof first, second, third, and fourth free thoracic somites coveredwith cuticular flaps.Fourth free thoracic somite smaller than previous three, bearingleg five. Genital segmentsmaller thanpreceeding somite, with cuticular flapson ventral surface. Abdomentwo-segmented,ventral surface bearing cuticularflaps. Caudal ramus (Figure 8B) longer thanwide,bearing two stout setae, one dorsalnaked seta, and one lateral naked seta; ventralsurface armed with posteriorly directedcuticular flaps.First antenna (Figure 8C) five-segmented,armature (proximal to distal) as follows: one slender seta; sevensmooth slender setae, one small denticulatedseta plusone large denticulated, curving(prehensile) claw; seven smooth setae, one largespineplus one well developed denticulated spine;one large denticulated spine; terminalsegment with 12 smooth slender setae, two denticulatedsetae plus one aesthete. Secondantenna (Figure 8D) five-segmented,prehensile. Basal segment short and unarmed,second segment with well developed spiniformprocess and triangular cuticular flaps,third segment bearing wavy quadrangularcuticular flaps, two slender setae arising fromlarge, curving spinous process. Fourthsegment elongate and unarmed; fifth.segmentan unciform terminal clawbearing two slender setae and one stout auxiliary spine.Mouth tube siphonostome andsimilar to that of other species. Mandible (Figure BE) oftwo parts, dentiferous margin with six teeth. Firstmaxilla (Figure BE) biramous; endopod bearing triangular cuticular flaps and two apicaldenticulated setae; exopod longersurmounted by two small setae and one longer denticulatedseta. Second maxilla(Figure 8G) brachiform, lacertus larger than brachiumarmed with cuticular flaps (omitted in illustration), brachium with crescent-shapedcuticular flaps and two tufts of setae(one coarse or rope-like (possiblyfused setae) and one fine or hair-like) at base of terminal claw (calamus). Claw bearingone serrated membrane parallelling the distal halfof the claw and two pendulous stripsof membrane, one hanging from each side.Maxilliped (Figure 8H) chelate, indistinctly segmented, proximalsegment pedunculate;corpus maxillipedis robust bearing small stout spine on distal margin,narrow rectangular cuticular flaps and a large transversecuticular flange; myxal area expanded intolarge receptacle to accommodate clawof opposable segment. Shaft bearing many finetriangular cuticular flaps with two setae: one near midpoint on lateralconvex marginand the other adjacentto a strip of membrane along concave distal margin. Claw unciform with quadrangularcuticular expansion producing lateral shield.First four pairs of legs biramous with three-segmented ramiand two-segmentedsympods. All basipods withlateral slender seta; first basipod bears additional distomedial seta. Ventral surfaces ofall four legs bearing crescent to sub-triangular shaped36cuticular flaps. Armature of ramias follows:Leg one Exopod0-0 0-0 IV Endopod0-0 0-0 IILeg two (modified) Exopod 1-01-0 III Endopod 0-00-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-01-0 III Endopod 0-00-0 ILeg one (Figure 9A) allsetae slender and denticulated. Exopod two (Figure9C)typically modified, first segmentgreatly enlarged with proximal crescentic cuticularflappatch, one distal stout seta,second segment smaller with stout seta, thirdsegmentarmed with one small sub-apicalseta, one apical slightly curving seta plus one largeserrated curving claw-like seta. Leg twoendopod (Figure 9B) completely covered withcrescentic and sub-triangular cuticularflaps with two finely denticulated slender setaeon terminal segment. Legs threeand four (Figures 9D and 9E) similar; endopodalseta(Figure 9D) claw-like and unilaterallydenticulated with a single setule or flagelliformprocess arising midpoint on the seta, exopodalsetae (Figure 9E) stout with largestmost apical seta curving with bifid tip. Leg five(Figure 9F) longer than wide; lateral surface covered with crescent shaped cuticularflaps, distally bearing three long bifurcateslender setae plus one seta arisingdorsally from the base.Male: UnknownComments: E. aspera was originally reported fromCarcharias pleurotaenia,Bleeker, 1852 (=Carcharhinus limbatus(Valenciennes, 1839)), the blacktip shark fromJava (Heller, 1865). Since thenthis parasite has been reported from the branchiallamellae of the sharpnose shark, Rhizoprionodonterraenovae, (Richardson, 1836), thesmooth tooth or fine tooth shark Aprionodonisodon (=Carcharhinus isodon(Valenciennes, 1839)), collected from Lemon Bay, Florida (Gulfof Mexico) (Bere, 1936).Cressey (1967) reported E. aspera from the spinner shark, Carcharhinusbrevipinna(Muller and Henle, 1839), the scalloped hammerhead, Sphyrnalewini (Griffith andSmith, 1834), and from Rhizoprionodonacutus (ROppel, 1837) all collected from theIndian Ocean near Nosy Be,Madagascar. Kabata (1970) added to the host list anunidentified requiem shark, Carcharhinussp., and a member of the Hemiscyliidae, thebrownbanded bamboo shark, Chiloscyllium punctatumMuller and Henle, 1838 bothfrom Moreton Bay, Queensland. Finally,Essafi and Raibaut (1977) collected the parasite from the spinner shark from Tunisianwaters. The parasite seems to have an affinity for hosts of the family Carcharhinidae, with onlythe one record from the Sphyrnidae37and one record from theHemiscyliidae, an orectolobid(carpet sharks)as the hostgroup exceptions.Heller (1865) shows pointedprocesses arising fromthe lateral margins of thecephalothorax. This appearsto be an erroneous interpretationof the notches in the lateral margins of the dorsalshield.This species is easilydistinguished by the largespatulate process on the secondsegment of the secondantenna, the pendulousmembranous flaps on the claw of thesecond maxilla, the elongateproximal segment and largedenticulated claw-like setaon the distal segmentof the second exopod, and thebranching setae of leg five.Eudactylina chilensisHo and McKinney, 1981(Figures 10-11)Material examined. Severalfemales, from the personalcollections of Dr. JuShey Ho, CaliforniaState University, Long Beachand Raul Castro Romero,Universidad de Antofagasta,Antofagasta, Chile, all specimenscollected from thebranchial lamellae ofthe black shark or hooktooth dogfish,Aculeola nigra De Buen,1959 collected fromCoquimbo, Chile (eastern South Pacific).DescriptionFemale (Figure 1 OA)Overall length in lateral view approximately2.3 mm. Cephalothorax longer thanwide, lateral margin notchedaccomodating area of lacertus of second maxillae.Ventrolateral and dorsal surfacesof cephalothorax bearing small crescentic cuticularflaps. Tergum of first, second,and third free thoracic somites coveredwith small cuticular flaps on dorsal surface.Fourth free thoracic somite notas densely covered withflaps and smaller than previoussomites and bearing leg five. Genital segmentsmallerthan preceeding somite, withcuticular flaps on ventral surface. Abdomen two-segmented, ventral surface bearingcuticular flaps. Caudal ramus (Figure lOB) longer thanwide, distal margin bearingthree large clenticulated setae and one smallsmooth stoutseta, dorsal surface with onesmooth slender seta, ventral surface armed with triangularshaped posteriorly directed cuticularflaps and one tiny seta.First antenna (Figure 1OC) five-segmented,armature (proximal to distal) as follows: first segment with oneunilaterally denticulated seta, second segment with sixsmooth slender setae, onethickspine, one unilaterally denticulated seta plus one largedenticulated curving (prehensile)claw, third segment with nine smooth setae plus one38well developed unilaterallydenticulated seta, fourthsegment bearing oneunilaterallydenticulated seta, fifth segmentwith one unilaterallydenticulated seta, 13slender setaeplus one aesthete. Second antenna(Figure 1OD) five-segmented,prehensile. Basalsegment short, second segment withsub-triangular cuticularflaps, third segment bearing rectangular cuticularflaps, two slender setae arising nearbase of spinous process.Fourth segment elongatewith small triangular cuticular flaps;fifth segment an unciformterminal claw bearing twoslender setae and onestout auxiliary spine. Mouth tubesiphonostome andsimilar to that of other species.Mandible (Figure 1 OE) of two parts,dentiferous margin with eight teeth.First maxilla (Figure 1 OF)biramous; endopod withtriangular cuticular flaps andbearing two apical spinulatedsetae; exopod longer surmounted by two stout setaeand one longer slender seta.Second maxilla (Figure 1 OG)brachiform, lacertus larger than brachiumarmed with crescent shaped cuticular flaps,basal process locatedat base of lacertus, brachium withcrescent-shaped cuticularflaps and two tufts of setae,one tuft composed of tine hair-likesetules the other consisting of coarse rope-like,possibly fused setules at base of terminalclaw (calamus).Claw bilaterally bearing twostrips of serrated membranes. Maxilliped(Figure 1CH)chelate, indistinctly segmented,proximal segment pedunculate (omittedin illustration);corpus maxillipedis robustbearing small stout spine on distal margin, rectangularandsemicircular cuticular flaps; myxalarea expanded into large receptacleto accommodate claw of opposablesegment. Shaft bearing triangular cuticularflapswith twosetae: one near midpoint on lateralconvex margin and the other along concavedistalmargin. Claw unciform withquadrangular cuticular expansion producing lateralshield.First four pairs of legs biramouswith three-segmented rami and two-segmentedsympods. All basipods with lateralslender seta; first basipod bears additional distomedial slender seta. Ventralsurfaces of all four legs bearing sub-triangular shaped cuticular flaps. Armature of ramias follows:Leg one Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IllEndopod 0-0 0-0 ILeg four Exopod 1-01-0 III Endopod 0-0 0-0 ILeg one (Figure hA) all setae onrami slender and denticulated. Exopod two(Figure 11 B) typically modified generallydevoid of cuticular flaps, first segment greatlyenlarged proximally bearing smallpatch of cuticular flaps and distally bearing onesmooth, slender seta, secondsegment smaller with slender seta, third segment armedwith two small curvingseta and one large bilaterally denticulated curvingseta. Leg two39endopod (Figurel 1 B) with two finelydenticulated slender setae on apicalsegment.Legs three and four similar; exopod three andfour (Figure 11C) with stoutsetae andone large bilaterally denticulatedseta on terminal segment, endopod threeand four(Figure liD) bearing a single bilaterallydenticulated seta on terminal segment.Legfive (Figure 11 E) oval; lateral surfacecovered with fine triangular cuticular flaps, distallybearing three slender setaeplus one seta arising dorsally from thebase.Male: Not obtained(see Ho and McKinney, 1981)Comments: This parasite has notbeen reported since its initial discovery onAculeola nigra (Dalatiiformes:Etmopteridae) from Chilean waters by Ho andMcKinney(1981).This species can be distinguishedby the prescence of cuticular flaps on thesecond, third, and fourth segmentsof the second antenna with the reduced spiniformprocess on the third segment, the three apical slender,bilaterally, denticulated setaeplus the unique, tiny lateralseta on the caudal ramusEudactylina corrugata Bere, 1930(Figures 12-13)Material examined. One female, USNM 60469 fromthe branchial lamellae of thelittle skate, Raja erinacea Mitchell collectedfrom St. Andrews, New Brunswick, onefemale, USNM 79619 from the branchiallamellae of Raja erinacea collected fromWoods Hole, Massachusetts July 17, 1914.DescriptionFemale (Figure 12A)Overall length in lateral view approximately 1.7 mm. Specimen unnaturallybloated due to lactic acid absorption during microscopicexamination. Cephalothoraxlonger than wide, lateral margin notched accomodating lacertus ofsecond maxillae.Ventrolateral and dorsal surfaces of cephalothorax covered with cuticular flaps. Terga offirst, second, third, and fourth free thoracic somitessparsely covered with cuticularflaps; terga of third and fourth free thoracic somitesindistinct. Second and third freethoracic somites bearing posteriorly directed cuticular flaps on ventralsurface. Fourthfree thoracic somite smaller thanprevious three, bearing leg five. Genital segmentsmaller than preceeding somite,with cuticularflaps on ventral surface. Abdomen two40segmented, ventral surface bearingcuticular flaps. Egg string (Figure 12B)uniseriate.Caudal ramus (Figure 12C) suboval,bearing two terminal slightly curvedstout setae,one medial naked seta, and one lateralnaked seta; ventral surface armedwith posteriorly directed cuticular flaps.First antenna (Figure 1 2D) indistinctlyfive-segmented, armature (proximalto distal) as follows: first segmentbearing one slender seta; second segment with sevensmooth setae, one denticulatedseta plus one large denticulated, curving (prehensile)claw; third segment withnine smooth setae plus one large uncinate spine; fourthsegment with one slender seta; terminalsegment with 1 4 smooth setae plus one aesthete.Second antenna (Figure 12E)five-segmented, prehensile. Basal segment short,second segment with stout spiniformprocess, third segment bearing quadrangular cuticularflaps, two slender setae arising from nearbase of large curving spinous process.Fourth segment elongate and unarmed; fifth segmentan unciform terminal claw bearingtwo slender setae and one stout auxiliary spine.Mouth tube siphonostome and similarto that of other species. Mandible (not illustrated)of two parts, dentiferous margin witheight teeth. First maxilla (Figure 1 2F) biramous;endopod bearing two apical spinulated setae; exopod longer surmounted by two small slendersetae and one longer spinulated seta. Second maxilla (Figure 12G) brachiform,lacertus larger than brachium,brachium with sub-triangular and crescent-shapedcuticular flaps and two tufts of setae(one coarse or rope-like (possibly fusedsetae) and one fine or hair-like) at base of terminal claw (calamus). Claw bearing two pairsof serrated membranes parallelling theclaw plus one distal strip of membrane alongthe convex margin. Maxilliped (Figure12H) chelate, indistinctly segmented, proximalsegment pedunculate; corpus maxillipedis robust bearing small stout spine on distal margin, narrow rectangularcuticularflaps and region of small triangular flaps; myxalarea expanded into large receptacle toaccommodate claw of opposable segment. Shaft bearing twosetae: one near midpointon lateral convex margin and the other adjacentto a strip of membrane along concavedistal margin. Claw unciform with quadrangular cuticular expansion producing lateralshield.First four pairs of legs biramous with two-segmented endopods, three-segmented exopods and two-segmented sympods. Allbasipods with lateral slender seta; firstbasipod bears additional distomedial slenderseta. Ventral surfaces of all four legsbearing rectangular to sub-triangular shaped cuticular flaps. Armature of ramias follows:Leg one Exopod 1-0 1-0 IV Endopod0-0 - IILeg two (modified) Exopod 1-0 1-0 Ill Endopod0-0 - II41Leg three Exopod 1-0 1-0III Endopod0-0 -Leg four Exopod 1-0 1-0 IIIEndopod 0-0 - ILeg one (Figurel3A) exopodindistinctly three-segmented; exopodalsetae slender unilaterally denticulated withor without spinules, endopodal setae bilaterallyspinulated. Exopod two (Figure 13B)typically modified, first segment greatly enlargedwithproximal triangular cuticular flaps alongthe medial margin, one distal stout seta, secondsegment smaller withstout spine, third segment armed with onesmall sub-apical seta,another sub-apical recurving slenderseta plus one large, blunt apicaltruncate seta. Legtwo endopod (Figure 1 3C) with twofinely spinulated slender setae on terminalsegment. Legs three and four (Figure13D) similar; setae slender and denticulated.Legfive (Figurel3E) longer than wide;lateral surface covered with rectangular cuticularflaps, distally bearing three slendersetae plus one seta arising dorsally from thebase.Male: UnknownComments: E. corrugatato has only been reportedby Bere (1930) occurring onthe skates Raja erinacea Mitchell, 1825 collectedfrom St. Andrews, New Brunswickand Woods Hole, Massachusettsand from Raja scabrata Garman, 1913 (=Raja radiataDonovan, 1807) collected fromWoods Hole, Massachusetts.This species is easily distinguished bythe large truncate seta on the terminalsegment of the second exopod andby the two-segmented endopods of legs onethrough four.Eudactylina dactylocerca sp. nov.(Figures 14-15)Material examined. Several females from thebranchial lamellae of the shovel-nose guitarfish Rhinobatus productus (Ayres)collected from inshore waters from thesouthern California Bight. Female holotype (USNM266521) and 4 female paratypes(USNM 266522) deposited at the United StatesNational Museum of Natural History.Etymology: The specific name dactylocerca isderived from the Greek dactylos meaning finger or digit andcerco from kerkos meaning tail, referring to the digitiform. processes on the caudal rami.42DescriptionFemale (Figure 1 4A)Overall length in lateral view approximately1.9 mm. Cephalothoraxlonger thanwide, lateral margin notched accomodating lacertusof second maxillae.Anterolateraland dorsal surfaces of cephalothorax covered withcuticular flaps. Dorsal andventrolateral surfaces of tergum of first, second, andthird free thoracic somites coveredwithcuticular flaps; terga of first andsecond free thoracic somites aliform. Fourth freethoracic somite smaller than previousthree, bearing leg five. Genital segment smallerthanpreceeding somite. Abdomen(Figure 14C) two-segmented, ventral surface bearingapair of blunt tubercles on eachsegment, and a pair of semicircular cuticular flaps ontheposterior segment. Egg string (Figure 14D)uniseriate. Caudal ramus (Figure 14B)beautifully modified into a tridentate digitiformstructure composed of three stronglysclerotized tuberculous processes(fused modified setae?), one tiny proximal seta (sensilla?), one dorsomedial slenderseta, one small stout ventral seta plus two ventrodistalslender setae; ventral surface armed witha single posteriorly directed semicircularcuticular flap.First antenna (Figure 14E) five-segmented,armature (proximal to distal) as follows: first segment bearing one slender seta; secondsegment with four smooth slender setae, one large smoothseta, three denticulated slender setae plus one large denticulated, curving (prehensile) claw; third segmentwith eight smooth setae, one denticulated slender seta plus one large uncinate denticulatedspine; fourth segment withone slender seta and an atypical conicalprocess; terminal segment with 13 smoothspiniform plus one aesthete. Second antenna(Figure 14F) five-segmented, prehensile. Basal segment short, secondsegment with unciform process and truncated quadrangular cuticular flaps, third segment bearing quadrangularcuticular flaps and twoslender setae arising from near base of well produced spinousprocess. Fourth segment elongate andunarmed; fifth segment an unciform terminal claw bearing two slender setae and one stout uncinate auxiliary spine. Mouth tube siphonostome andsimilarto that of other species. Mandible (Figure 14G) of two parts,dentiferous margin withfive teeth. First maxilla (Figure 1 4G) biramous; endopod bearingtiny triangular cuticular flaps and two apical denticulatedsetae; exopod longer surmounted by two smallsetae and one longer slender seta. Second maxilla (Figure 14H)brachiform, lacertuslarger than brachium, brachium with crescent-shapedcuticular flaps and two tufts ofsetae (one coarse or rope-like (possibly fused setae) and one fine or hair-like)at baseof terminal claw (calamus). Claw bearing tworows of serrated membranes parallellingthe claw plus one distal pendulousstrip of membrane along the convex margin of otherside. Maxilliped (Figure 141) chelate,indistinctly segmented, proximal segment pedun43culate; corpus maxillipedis robustbearing small seta on distal marginand narrow rectangular cuticular flaps; myxal area expandedinto large receptacleto accommodateclaw of opposable segment. Shaft bearing twosetae: one withenlarged proximalregion near midpoint on lateral convexmargin and the other adjacentto a strip of membrane along concave distal margin.Claw (Figure 14J) complex,unciform with tiny proximal seta, cuticular expansionsproducing a fused complex of claw andcuticle.First four pairs of legs biramous,except leg four with two-segmented endopod,three-segmented exopods and endopodson remaining rami, sympodstwo-segmented.All basipods with lateralslender seta; first basipodbears additional distomedial slenderseta. Ventral surfaces of all fourlegs bearing rectangular to sub-triangularshapedcuticular flaps. Armature of ramias follows:Leg one Exopod1-0 1-0 Ill Endopod0-0 0-0 IILeg two (modilied) Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IllEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0Ill Endopod 0-0Leg one (Figurel5A) rami three-segmented;exopodal setae unilaterally bearingserrated flange with medial denticulations onlargest apical seta on terminal segment,largest endopodal seta unilaterally bearing serratedflange, smaller bilaterally denticulated. Exopod two (Figure 1 5B)typically modified, first segment greatly enlarged withquadrangular cuticular flaps along the lateral margin,one distal seta, second segmentsmaller with seta, third segment armed withtwo smooth curving setae plus one largecurving bilaterally denticulatedseta. Leg two endopod (Figure 1 5B) with one slenderseta and one large bilaterally serrated seta onterminal segment. Sclerite bar betweenleg two and leg three (Figure 150)with single, large medial stylet. Legs three and four(Figure 15D) similar; except legfour has a two-segmented endopod; exopodal setaeclaw-like and unilaterally denticulated, endopodalseta unilaterally bearing denticles andserrated membranous flange. Leg five (Figurel5E)slightly longer than wide; lateralsurface smooth, distally bearing three pinnatesetae plus one seta arising dorsally fromthe base.Male: UnknownComments: E. dactylocerca isspecific to the shovelnose guitarfish Rhinobatusproductus. It is the thirdeudactylinid to be reported from this genus of host.Eudactylina rhinobati Raibaut and Essafi, 1979 hasbeen found from Rhinobatus rhino44batus (Linne’, 1758), and from Rhinobatuscern/cu/us (Geoffrey Saint-Hilaire, 1817) collected from Tunisia, and recently Luque and Farfan(1991) acquired Eudactylina peruensis from Rhinobatus planiceps Garman, 1880 fromeastern South Pacific waters offthe west coast of Peru.Some of the illustrations of E.rhinobati (Raibaut and Essafi, 1979) lack detail butimply something atypical, yet vaguelysimilar (digitiform elements) among the caudalrami of these three rhinobatid-infestingspecies. All three species share the dorsolateral aliform expansions of free thoracicsomites one and two, suggesting a very closerelationship between these species despite thegreat geographical distances that separate them.This species is easily distinguished fromall other species in the genus by themodified caudal rami. The interesting finedenticulations along the lateral margin of thesetae on exopods three and four are shared by E. peruensis.Eudactylina diabolophi!a sp. nov.(Figures 16-17)Material examined. Two females from the branchial lamellae of the Manta orDevil Ray Manta birostris (Donndorff, 1798), (California Academyof Sciences FishCollection) collected August 20, 1951 during the George Vanderbilt Equatorial PacificExpedition from station 49 located at551.9’ N X 162 7.6’ N, near Sand and LineIslands. Female holotype (USNM 266523) depositedat the United States NationalMuseum of Natural History.Etymology: The specific name diabolophi/a is derived from the Greek diabolos meaning devil and phil/as meaning loving, referring to this species predilection for feedingupon the devil ray.DescriptionFemale (Figure 16A)Overall length in lateral view approximately 3.8 mm. Cephalothorax longer thanwide, lateral margin notched. Anterolateral and dorsal surfaces of cephalothorax covered with cuticular flaps. Dorsolateral surfaces of first, and second free thoracic somitescovered with cuticular flaps. Fourth free thoracic somite smaller than previous three,bearing leg five. Genital segment smaller than preceeding somite. Abdomen two-segmented; lateral surface of posterior segment with cuticular flaps. Caudal ramus (Figure451 6B) atypically elongate with six distal,relatively unmodified naked setae,a distal areaof anteriorly directed rectangular flaps and the lateral marginarmed with subtriangularcuticular flaps.First antenna (Figure 16C) six-segmented, armature (proximalto distal) as follows: first segment atypically bearing a lateral patch ofsemicircular cuticular flaps andbearing one reduced stout seta; secondsegment with five naked and one large denticulated, curving (prehensile) claw;third segment with three naked setae; fourth segment with three small nakedseta plus one large styliform seta, fifth segment with a single naked seta; terminal segment with13 setae plus one aesthete. Second antenna(Figure 16D) five-segmented, prehensile. Basal segmentshort, second segment elongate and unarmed, third segment bearing thin rectangularcuticular flaps and two slender setae arising from nearbase of greatly reduced spinous process (this may only bethe distomedial corner of this segment). Fourth segment elongatewith fine triangularcuticular flaps along convex margin; fifth segment an elongate, unciform terminal clawbearing two slender setae and one small auxiliaryspine. Mouth tube siphonostomeand similar to that of other species. Mandible (Figure 16E) of two parts, dentiferousmargin with five teeth. First maxilla (Figure 1 6F) biramous; sympodwith three rows offine triangular cuticular flaps, endopod bearing tiny triangular cuticular flaps and twostout apical setae; exopod longer with tiny triangular cuticular flaps and surmountedbytwo small stout setae and one longerseta. Second maxilla (Figure 14G) brachiform,lacertus larger than brachium bearing small triangular cuticular flaps and basal process,brachium with triangular cuticular flaps and two tufts of setae (one coarse or rope-like(possibly fused setae)) and one fine or hair-like) at base of terminal claw (calamus).Claw bearing two rows of denticles proximally plus a row of denticles along both theconcave and convex margins. Maxilliped (Figure 16H) chelate, indistinctly segmented,proximal segment pedunculate; corpus maxillipedis robust bearing small naked seta ondistal margin and tiny triangular cuticular flaps; myxal area proximally bearing triangularcuticular flaps and expanded into a receptacle to accommodate claw of opposable segment. Shaft with proximal patch of small triangular flaps and bearing two setae: onevery small near midpoint (not illustrated) on lateral convex margin and the other alongconcave distal margin. Claw (Figure 16H) complex, unciform with a very small lateralshield.First four pairs of legs biramous and trimerite, sympods two-segmented. Allbasipods with lateral slender seta; first basipod bears additional distomedial slenderseta. Ventral surfaces of all four legs bearing triangular shaped cuticular flaps.Armature of rami as follows:46Leg one Exopod 1-0 1-0III Endopod 0-00-0 IILeg two Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg one (Figurel7A) ramithree-segmented; exopodal setae unilaterally to bilaterally denticulated, largest endopodalseta bilaterally denticulated, smaller smooth. Legtwo (Figure 17C) atypicallyunmodified, setae stout with and without denticulations.Legs three and four (Figure 17B) similar;distolateral areas on first and secondsegments of exopods greatly extendingpast segmental boundaries, with ventral facedevoid of cuticular flaps, largest of threestout setae on terminal segment bilaterallydenticulated. Leg five (Figurel7D) subquadrate, slightly wider thanlong, distally bearing three slender setae plus one similarseta arising dorsally from the base; lateral surface with small triangular cuticular flapsMale: UnknownComments: E. diabolophila is the first record of Eudactylina from the genusManta. Although Pacific and Atlantic mantasare presently considered conspecific, itwould be of interest to examine this host fromthe Atlantic for additional corroborativeevidence.This species is easily distinguished from all other species in the genusby thegreatly extended distolateral regions ofthe first and second segments on the third andfourth exopods, the unusually elongate caudal rami, the very large stronglycurved clawof the second antenna, and the greatly reduced“lateral shield” of the claw of the maxilliped. The unmodified condition of the exopodof leg two found in this species is similarly found in E. acanthii and E. squamosa.Eudactylina doilfusi Brian, 1924(Figures 18-19)Syn: Eudactylina spinifera Wilson, 1932, syn nov.Eudactylina spinifera Wilson, 1932; of Bere (1936)Eudactylina spinifera Wilson, 1932; of Yamaguti (1963)Eudactylina spinifera Wilson, 1932; of Cressey (1970)47Material examined. Several females(types?) MNHN CP 174 fromthe branchialIamellae of the host squale (Marao);Several female specimens USNM63915 from thebranchial lamellae of Carchariascommersoni(=?) collected from Wood’sHole, July25, 1927; female “holotype” USNM 56621from gills of Carcharhinus milberti;fromWood’s Hole; numerous femalesUSNM 153650, 153651, 153652,63915, 79087 fromthe gills of Carcharhinus milberti (Valenciennes, j..Muller and Henle, 1839),(=Carcharhinus plumbeus (Nardo, 1827)).DescriptionFemale (Figure 18A)Overall length in lateral viewapproximately 1.8 mm. Cephalothorax longer thanwide, lateral margin notchedaccomodating lacertus of second maxillae.Anterolateraland dorsal surfaces of cephalothorax sparselycovered with spiniform cuticular flaps.Dorsal and ventrolateral surfaces ofterga of first, second, third, and fourth free thoracicsomites sparsely covered with spiniform cuticularflaps. Fourth free thoracic somitesmaller than previous three, bearing legfive. Genital segment smaller than pr.eceedingsomite and sparsely covered withspiniform cuticular flaps. Abdomen (Figures 18A, B)two-segmented, ventral surface bearinga pair of slender setae on anterior segment,and posteriorly directedtriangular cuticular flaps on both segments. Caudal ramus(Figure 1 8B) longer than wide with posteriorly directedtriangular cuticular flaps on ventral surface, rami bearing one lateraland one medial seta, distally one stout unilaterallydenticulated seta, two terminal finely denticulated orspinulated seta and possibly onetiny unilaterally denticulated or spinulated seta.First antenna (Figure 18C) indistinctly five-segmented, armature(proximal todistal) as follows: first segmentbearing one short seta; second segment with eightsmooth stout setae of various sizes, plus onelarge denticulated, curving (prehensile)claw; third segment with nine smoothstout setae; fourth segment with one stout seta;terminal segment with 14 smooth slendersetae plus one aesthete. Second antenna(Figure 1BD) five-segmented, prehensile.Basal segment short, second segmentnaked, third segment bearing six small semicircularcuticular flaps and two slendersetae arising from near base of well producedspinous process. Fourth segment elongate and unarmed; fifth segment an unciformterminal claw bearing two slender setae,one very small stout seta, and onestout uncinate auxiliary spine. Mouth tube siphonostome and similar to that of otherspecies. Mandible (Figure 1 8E) of two parts, dentiferous margin with five teeth. First maxilla (Figure 18F) biramous; endopod bearingtinytriangular cuticular flaps, one apicaldenticulated seta, and one smooth with papilliformtip; exopod longer surmountedby two small setae and one longer slender seta all48tipped with tiny setule. Second maxilla(Figure 18G) brachiform, lacertuslarger thanbrachium with triangular cuticular flaps,brachium with triangular-shaped cuticularflapsand two tufts of setae (one coarse orrope-like (possibly fused setae) and onefine orhair-like) at base of terminal claw(calamus). Claw bearing two rows of serratedmembranes parallelling the claw plus six claw-likedenticles along the concave distal surface. Maxilliped (Figure 18H)chelate, indistinctly segmented, proximalsegmentpedunculate (not illustrated); corpus maxillipedisrobust apparently devoid of typicalsmall spiniform seta on distal margin,two patches of triangular cuticular flaps;myxalarea expanded into largereceptacle to accommodate claw of opposablesegment.Shaft bearing two setae: one smallslender seta near midpoint on lateral convex marginand the other adjacentto a strip of membrane along concave distal margin. Clawcomplex, unciform with tiny membraneor membranous flange along concave margin.First four pairs of legs biramous and trimerite except fortwo-segmented endopodof leg one and the endopodof leg four fused into a large claw, sympods two-segmented. All basipods with lateral slender seta;first basipod bears additional distomedialslender seta. Ventral surfaces of allfour legs bearing small triangular shaped cuticularflaps. Armature of rami as follows:Leg one Exopod 1-0 1-0 IllEndopod 0-0 - IILeg two (modified) Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0 III Endopod0-0Leg one (Figurel 9A) rami three-segmented; exopodalsetae bearing bilaterallyor unilaterally serratedmembranous flange, denticulations on medial edge of largestapical seta on terminal segment, largest endopodalseta bilaterally denticulated, smallerseta smooth. Leg two (Figure 1 9B) with exopod typically modified, first segmentgreatlyenlarged with proximal patch of triangular cuticular flaps, distally bearing onenaked,seta, second segment smaller with naked seta, third segmentarmed with two stoutsetae each bearing a single denticle, plus one smaller nakedseta. Leg two endopod(Figure 19B) with one slender seta and one largebilaterally spinulated seta on terminalsegment. Leg three (Figure 19C) exopodal setae stout and unilaterally denticulated,largest seta on terminal segment bilaterally denticulated; endopodalseta stout andbilaterally denticulated. Leg four (Figure 19D) exopod similarto leg three but withsmooth setae except for large bilaterally denticulated on terminalsegment, endopodmodified into a heavily sclerotized, fused unciform claw witha proximal patch of cuticular flaps. Leg five (Figure 1 9E)slightly longer than wide; lateral surface with few cuticu49lar flaps, distally bearing three spiniformsetae plus one setaarising dorsally from thebase.Male: UnknownComments: E. dolifusi Brian, 1924was originally reported from the gills of a shark(squale (Marao)) collected fromMauritius. It was later discovered on the branchiallamellae of the brown or sandbar shark,Carcharhinus milberti (Valenciennes, in Mullerand Henle, 1839), (=Carcharhinusplumbeus (Nardo, 1827)) from the Wood’sHoleregion but was describedas a new species, E. spiniferaby Wilson (1932). Bere (1936)reported E. spinifera from the duskyshark C. obscurus (LeSueur, 1818). Yamaguti(1963) transferred the errors intohis compilation, and finally Cressey (1970) reportedWilson’s E. spinifera from the sandbarshark, C. plumbeus and the blacknose sharkC.acronotus (LeSueur, 1818).Examination of several specimensconfirms E. spinifera Wilson, 1932 is a juniorsynonym of E. dolifusi Brian, 1924.Eudactylina dolifusi seems to bespecific to the few aforementioned species ofsharks of the Carcharhinidae, witha preference for the sandbar shark Carcharhinusplumbeus.This species is readily identifiedby the huge, modified claw-like fused endopodof leg four.Eudactylina epaktolampter sp. nov.(Figures 20-23)Material examined. Two females and one malefrom the branchial lamel!ae of thesmooth lanternshark Etmopterus pusillus (Lowe,1839), (California Academy ofSciences, CAS 1967 Vll:6) collected from off theMississippi Delta, August 24, 1962 onthe R/V Oregon, station 3724, located29204’N X 8831; and two females from the gillsof E. pusillus (USNM 220344) collectedfrom Atlantic Liberia at a depth of 400 m, station location 06208’N X 10257’W .Female holotype (USNM 266524) deposited at theUnited States National Museum ofNatural History.Etymology: The specific name epaktolampterisderived from the Greek epakter meaning hunter and lampter meaninglantern, referring to the predilection of this parasite for50lantern sharks.DescriptionFemale (Figure 20A)Overall length in lateral view approximately 3.1 mm.Cephalothorax longer thanwide, lateral margin notched. Surface of cephalothorax, first, second,and third freethoracic somites covered with cuticular flaps.Fourth free thoracic somite smaller thanprevious three, bearing leg five.Genital segment slightly smaller than preceedingsomite. Abdomen two-segmented; ventrodistalsurface of posterior segment with cuticular flaps. Caudal ramus (Figure 20B) with threestout setae, one very tiny setule, plustwo elongate slender setae (one medial, one lateral).First antenna (Figure 20C) five-segmented, armature (proximalto distal) as follows: first segment bearing onestout seta; second segment with seven slender setae,one large stout seta, and one large smooth, curving (prehensile)claw; thir&segmentwith ten slender setae; fourth segment with one long slenderseta; terminal segmentwith 14 slender setae plus one aesthete. Second antenna (Figure20D) five-segmented, prehensile. Basal segment short, second segment elongate and unarmed, thirdsegment bearing proximally directed triangular cuticular flaps and two slender setae,segment apparently devoid of typical spinous process. Fourth segment elongate andunarmed; fifth segment an elongate, unciform terminal claw bearing two slender setaeand one small auxiliary spine. Mouth tube siphonostome and similarto that of otherspecies. Mandible (Figure 20E) of two parts, dentiferous margin with seven teeth. Firstmaxilla (Figure 20F) biramous; endopod bearing tiny triangular cuticular flaps and twostout apical setae, one unilaterally denticulated and one with proximal patch of spinules;exopod longer with tiny triangular cuticular flaps and surmountedby two small stoutsetae (one omitted in illustration) and one longer seta. Second maxilla (Figure2OG)brachiform, lacertus larger than brachium bearing semicircular cuticular flaps, brachiumwith semicircircular cuticular flaps and two tufts of setae (one coarse or rope-like (possibly fused setae) and one fine or hair-like) at base of terminal claw (calamUs). Clawatypically bearing a patch or bibbed patch of fine spinulations along concave surface.Maxilliped (Figure 20H) chelate, indistinctly segmented, proximal segment pedunculate(not shown); corpus maxillipedis robust bearing small spiniform seta on distal marginand small rectangular cuticular flaps; myxal area expanded into a receptacle to accommodate claw of opposable segment. Shaft bearing two setae: one near midpoint andthe other along concave distal margin near membranous strip(see detail). Claw(Figure 20H detail) complex, unciform with a large orbicular lateral shield51First leg biramous and bimerite,legs two, three and four biramousand trimerite,sympods two-segmented. All basipods withlateral slender seta; first basipodbearsadditional distomedial slenderseta. Ventral surfaces of all four legs bearingsemicircular cuticular flaps. Armature of rami as follows:Leg one Exopod I-C - IVEndopod 0-0 - IILeg two (modified) Exopod1-0 1-0 III Endopod 0-0 0-0 IILeg three Exopod 1-01-0 III Endopod 0-0 0-0 ILeg four Exopod 1-01-0 lii Endopod 0-0 0-0 ILeg one (Figure 21A) rami two-segmented(possibly indistinctly three); exopodaland endopodal setae bilaterallydenticulated. Leg two exopod (Figure 21B)typicallymodified, first segment greatly elongatedwith proximal patch of flaps, all setae smoothand stout, endopod (Figure 21C) tipped withtwo slender setae. Legs three and four(Figure 21D) similar; all setae smooth andstout except bilaterally denticulated middleseta on terminal segment of exopod. Leg five(Figure2l E) subquadrate, longer thanwide distally bearing three slendersetae plus one similar seta arising dorsally from thebase; lateral surface with small triangular cuticularflaps.Male: (Figure 22A)Overall length in lateral view approximately 1 .8 mm. Similarto female, exceptabdomen four-segmented, leg five andleg six arising from genital complex. Caudalramus (Figure 22B) bearing fourpinnate setae and two smooth slender setae.First antenna (Figure 22C) indistinctly of nine segments; armature (proximaltodistal) as follows: first segment bearingone stout seta, second segment with nine slender setae plus one large curving (prehensile)claw, third segment armed with four slender setae, fourth segment with twosetae, fifth segment with one seta, sixth segmentwith four slender setae, seventh segment armedwith two setae, eighth segment bearing a single small stout seta andlong aesthete, ninth segment bearing 12 slendersetae. Maxilliped (Figure 22D) subchelate; corpus bearingmany small rectangularcuticular flaps, myxal area bearing a well produced styliform process,subchela bearingtwo stout setae, claw produced intoa large curving process proximally producing a bifidspine which in concert with the claw creates a concavity for themyxal process to actupon.First pair of legs (Figures 23A and 23B) biramous and bimerite, legs two, threeand, four biramous and trimerite.Sympods two-segmented. Semicircular cuticular flapson ventral surfaces of legs I - IV.Armature of rami as follows:52Leg oneExopod 1-0- V Endopod0-0 - IILeg twoExopod I-II-I VII Endopod0-0 0-I IIILeg threeExopod I-II-I VII Endopod0-I 0-I IVLeg fourExopod I-II-I VII Endopod0-0 0-0IIIExopod one(Figure 23A) two-segmented;first segment withsingle smoothseta,second segmentbearing twolong pinnate setaeplus three smallernaked setae.Endopod one(Figure 23B) tippedwith two nakedsetae. Leg two(Figure 23C)exopodalsetae predominatelypinnate mediallyand smoothand along lateralmargin, proximalsegment ofendopod withlarge scierotizedlateral process,remaining setaepinnate.Leg three(Figure 23D) exopodaland endopodalsetae smoothalong lateralmargin,pinnate alongmedial margin.Leg four similarto leg threeexcept forterminal segmentof endopod (Figure23E) bearing onlythree insteadof four setae.Leg five (Figure23F)armed with threeslender setaeon distal edgeand one pinnateseta at base.Leg six(Figure 23G)consists of threeslender nakedsetae on distaledge of finelyspinulatedlobe.Comments:E. epaktolampteris the first recordof Eudactyilnafrom the genusEtmopterus. Bothof my finds comefrom the smoothlanternshark,Etmopteruspusillus(Lowe, 1839),collectedin the Gulf of Mexicooff the MississippiDelta andoff theLiberian Atlantic.The maleof this speciesexhibits a veryunusual bluntspinous processon thelateral marginof the basal segmentof the secondendopod.This characterhas onlybeen reportedonce before fromthe entire orderSiphonostomatoida,predictablyfromanother male eudactylinid,Eudactylinachi/ensis Hoand McKinney,1981 from Theblackshark Aculeolanigra De Buen,1959.This sharedderived characterindicates a closephylogeneticrelationship betweenthese speciesof parasites.Interestingly,both hostshave been removedfrom the familySqualidae andrecentlyplaced in a commonfamilyEtmopteridae.The parasitessupport thissystematic restructuring.This species iseasily distinguishedfrom all othercongeners bythe spinulationscovering the concavesurface of the secondmaxilla, theminute apicomedialseta on thecaudal rami,the large orbicularlateral shieldon the clawof the maxilliped.Theabsence ofthe spiniform processon the thirdsegment ofthe secondantennais sharedby E. acanthii,E. diabolophi/a,E. insolens,E. Iongispina,E. oliveri, E.pollex, andE.vaquetillae, allstrikingly differentfrom this speciesin a multitudeof characteristics.53Eudactylina hornbostell sp. nov.(Figures 24-25)Material examined. Several females from the branchial lamellae of the bat rayMyliobatis sp., captured from waters off Nosy Be’, Madagascar. Donated from the personal collection of Dr. Roger Cressey, National Museum of Natural History, SmithsonianInstitution, Washington, D.C. Female holotype (USNM 266525) and 5 female paratypes(USNM 266526) deposited at the United States National Museum of Natural History.Etymology: The specific name hornbosteli is in honor of Mr. Bernard HornBostel for computer hardware support for this effort.DescriptionFemale (Figure 24A)Overall length in lateral view approximately 1.6 mm. Cephalothorax longerthanwide, lateral margin slightly concave. Surface of cephalothorax, first, second,thirdand, fourth free thoracic somites covered with wavy cuticular flaps.Fourth free thoracicsomite, genital segment and, two-segmented abdomen all bearing wavy cuticularflapson ventral surface. Caudal ramus (Figure 24B) with two stout curving setae,plus twoelongate slender setae (one dorsal, one lateral).First antenna (Figure 24C) apparently four-segmented,armature (pràximal todistal) as follows: first segment bearing one tiny seta; second segmentwith four smallsetae, three elongate blunt setae and one large denticulated,curving (prehensile) claw;third segment with seven slender setae plus two largedenticulated spines; fourth segment an elongate process with 14 setae of various shapes andsizes plus one aesthete.Second antenna (Figure 24D) five-segmented, prehensile.Basal segment short, second segment elongate with a very long, proximallydirected spiniform process, thirdsegment elongate bearing proximally directed wavyrectangular cuticular flaps and twoslender setae arising from near base of long curving spiniformprocess. Fourth segment elongate and unarmed; fifth segment an elongate,unciform terminal claw bearingtwo slender setae and one extremely elongate auxiliary spine.Mouth tube siphonostome and similar to that of other species. Mandible(Figure 24E) of two parts, dentiferous margin with five teeth. First maxilla(Figure 24F) biramous; endopod more robustbearing two slender apical setae, one unilaterally denticulatedand one smooth; exopodlonger and surmounted by two small stout setae andone longer slender seta. Secondmaxilla (Figure 24G) brachiform, lacertus larger than brachiumbearing wavy cuticularflaps, brachium with wavy cuticular flaps (some triangular),distal patch of triangular54prickles two tuftsof setae (one coarseor rope-like (possiblyfused setae), onefine orhair-like (omittedin illustration)) at baseof terminal claw (calamus).Claw bearingfourstrips of serratedmembrane plusone serrated stripon other side. Maxilliped(Figure24H) chelate,indistinctly segmented,proximal segment pedunculate;corpus maxillipedis robustbearing small setaon distal marginand large wavy cuticularflaps; myxalarea expandedinto a receptacleto accommodateclaw of opposable segment.Shaftwith long membranousflange, bearing twosetae: one elongatenear midpoint andtheother along concavedistal margin near membranousstrip (see detail).Claw (Figure20H detail) complex,unciform with alarge orbicular lateralshield.First four pairs oflegs biramous andtrimerite, sympodstwo-segmented.Allbasipodswith lateral pinnateslender seta; firstbasipod bearsadditional distomedialslender seta.Ventral surfaces oflegs two, three,and four bearingwavy cuticularflaps.Armature oframi as follows:Leg oneExopod 1-0 1-0IV Endopod 0-00-0 IILeg two (modified)Exopod 1-01-0 III Endopod0-0 0-0IILeg threeExopod 1-01-0 III Endopod0-0 0-0 ILeg fourExopod 1-0 1-0III Endopod0-0 0-0 ILeg one (Figure25A) with singlerow of fine triangularcuticular flapsfringing distal margins ofall segments exceptterminal segmentof exopod; lateralexopodal setaeunilaterally bearingserrated membranousflange, apicalsetae smoothand slender,endopoda! setae nakedand slender. Legtwo exopod(Figure 25B) typicallymodified,first segment greatlyenlarged withstout seta,second segmentswollen with stoutseta,terminal segmentwith two small slendersetae plus onetruncate seta,endopod (Figure25B) tipped withtwo slender setae.Legs threeand four (Figure25C) similar;lateralmargins of endopodalsegments oneand two with fringesof fine setae,all exopoda)setae stout bearinglarge subapicaltines, endopodtipped by singlefinely spinulatedseta. Leg five(Figure25D) longerthan wide;lateral surface withwavy cuticülarflapsand distally bearingthree semipinnateslender setae plusone pinnate setaarising dorsally from the base.Male: UnknownComments: E. hornbosteliis easilydistinguished fromits congeners bytheextremely elongatesetae on the second,third and fourthsegments ofthe first antenna,the extremely elongateand spiniformapical segmentof the first antenna,the elongated55segments of the second antenna, the very long, curvingclaw and auxiliary spine on thesecond antenna, the elongate spiniform process found on thesecond segment of thesecond antenna, the large digitiform subapical tines found onthe setae of exopodsthree and four, and the fringe of spinules or cuticularflaps along the distomedial margins of the exopodal and endopodal segments of legone.Eudactylina indivisa Castro and Baeza, 1991(Figures 26-27)Material examined. Several females from the branchial lamellae of theeasternSouth Pacific bat ray, Myliobatis peruvianus (Garman, 1913) coflectedfromAntofagasta, Chile. Donated from the personal collection of Mr. Raul CastroRomero,Universidad de Antofagasta, Chile.DescriptionFemale (Figure 26A)Overall length in lateral view approximately 1.8 mm. Cephalothorax longer thanwide, lateral margin notched accomodating lacertus of second maxillae. Surface ofcephalothorax covered with cuticular flaps. Terga of first, second, third, and fourth freethoracic somites covered with cuticular flaps. Fourth free thoracic somite, genitalsegment and abdomen bearing posteriorly directed cuticular flaps on ventral surface.Fourth free thoracic somite smaller than previous three, bearing leg five. Genital segment smaller than preceeding somite. Abdomen two-segmented. Caudal ramus(Figure 26B) longer than wide, bearing two distal apically curved stout setae, one dorsalslender seta, one lateral slender seta, and one proximal setule; ventral surface armedwith posteriorly directed cuticular flaps.First antenna (Figure 260) apparently four-segmented, armature (proximal todistal) as follows: first segment bearing one slender seta; second segment with foursmall setae, one stout seta, one slender denticulated seta, one long slender seta, plusone large denticulated, curving (prehensile) claw; third segment bearing eight slendernaked setae plus one large spine armed with rectangular flaps; terminal segment with14 smooth spiniform setae plus one aesthete. Second antenna (Figure 26D) five-segmented, prehensile. Basal segment short, second segment with spiniform process,third segment bearing six rectangular cuticular flaps, two slender setae arising fromnear base of elongate styliform process. Fourth segment elongate and unarmed; fifth56segment an unciform terminal claw bearing two slendersetae and one elongate auxiliary spine. Mouth tube siphonostome and similar to thatof other species. Mandible(Figure 26E) of two parts, dentiferous margin with seventeeth. First maxilla (Figure26F) biramous; endopod with tiny triangular cuticular flapsand bearing two apical bilaterally denticulated setae; exopodlonger surmounted by two small setae and onelonger seta. Second maxilla (Figure 26G)brachiform, lacertus larger than brachiumbearing large round rectangular cuticularflaps, brachium with wavy, rectangular, andsemicircular cuticular flaps, a distal patch of triangularprickles and two tufts of setae(one coarse or rope-like (possibly fused setae) and one fine or hair-like))at base of terminal claw (calamus). Claw bearing four strips of serratedmembranes parallelling thedistal portion of the claw plus one proximal stripof membrane along the convex margin,two strips present on other side. Maxilliped (Figure26H) chelate, indistinctly segmented, proximal segment pedunculate; corpus maxillipedisrobust bearing small stout spineon distal margin, narrow wavy, rectangular cuticular flaps;myxal area expanded intolarge receptacle to accommodate claw of opposablesegment. Shaft bearing two setae:one near midpoint on lateral convex margin and the other adjacentto a stripof membrane along concave distal margin. Claw unciform with quadrangularcuticular expansion with thinner cuticular expansion on distal edgeproducing lateral shield.First four pairs of legs biramous and trimerite, two-segmented sympods. Allbasipods with lateral slender seta; first basipod bears additionaldistomedial slenderseta. Ventral surfaces of all four legs bearing various shaped cuticular flaps. Armatureof rami as follows:Leg one Exopod 1-0 1-0 IV Endopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 III Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 Ill Endopod 0-0 0-0 ILeg four Exopod 1-0 1-0 III Endopod 0-0 0-0 ILeg one (Figure 27A) lateral exopodal setae unilaterally bearing serrated membranous flange, apical setae slender with small setules (spinules), endopodal setaebilaterally spinulated. Exopod two (Figures 27B and 27C) modified, first segmentgreatly enlarged with a few tiny triangular cuticular flaps along the medial margin, onedistal denticulated seta, second segment smaller with large, stout spine, third segmentarmed with one small subapical seta, another subapical recurvingseta plus one bluntapical truncate seta. Leg two endopod (Figure 27B) with subtriangular expansion in distolateral corner of first segment, one small spinulatedseta and one large bilaterallydenticulated seta on terminal segment. Legs three and four (Figures 27Dand 27E)57similar;exopodsthreeandfour(Figure27E)with denticulatedsetae,endopodsthreeandfour(Figure27D)tippedwith asinglebilaterallydenticulatedseta.Legfive (Figure27F)subtriangular,longerthanwide,distallybearingthreeslendersetaeplusonepinnatesetaarisingdorsallyfromthe base;lateralsurfacecoveredwithtriangularcuticularflaps.Legsix (Figure27G)representedbythreeverytinyspineslocatedon therim oftheapertureof theoviducalopening.Male:UnknownComments:E. indivisaseemsto bea parasiteof temperateeasternSouthPacificbat rays.Sofar, ithasbeenreportedfromthegills ofMyliobatisperuvianus(Garman,1913)andMyliobatischilensisPhillipi,1892,fromnear Antofagasta,Chile.Theunusualpresenceof cuticularflapson thelargespineon thethird segmentof thefirstantenna,thedifficultto characterizeclawof themaxilliped,and thelongstraight,denticulatedspine-likesetaon thesecondsegmentof exopodtwo arereadydiscriminantsfor thisspecies.EudactylinainsolensScottand Scott,1913(Figures28-29)Materialexamined.Severalfemales(BMNH1913.9.18.292-293)fromthebranchiallamellaeof thetopeshark,Galeorhinusgaleus(Linnaeus,1758)collectedfrom theIrishSea.DescriptionFemale(Figure28A)Overalllengthin lateralviewapproximately1.8 mm.Cephalothoraxlongerthanwide,lateralmarginnotchedaccommodatinglacertusof secondmaxillae.Anterolateralsurfaceof cephalothoraxcoveredwithcuticularflaps.Tergaof first,second,third,andfourthfreethoracicsomitesatypicallydevoidof cuticularflaps.Genitalsegmentandabdomenbearingposteriorlydirectedcuticularflapson ventralsurface.Fourthfreethoracicsomitebearinglegfive. Genitalsegmentsmallerthanpreceedingsomite.Abdomentwo-segmented.Caudalramus(Figure28B)longerthanwide,bearingthreeterminaldenticulatedsetae,onedorsomedialseta,andonelateralnakedseta;ventralsurfacearmedwithposteriorlydirectedtriangularcuticularflaps.Firstantenna(Figure28C)five-segmented,armature(proximalto distal)as fol58lows:first segmentbearingone spiniformseta; secondsegmentwith six smoothsetae,two unilaterallydenticulatedspiniformsetae plusone largedenticulated,curving(prehensile)claw; thirdsegmentbearingnine smoothsetae; fourthsegmentbearing asingle seta.Terminalsegmentatypicallyexhibitingthe presenceof cuticularflaps, andarmedwith 13smoothslender setaeplus oneaesthete.Secondantenna(Figure28D)five-segmented,prehensile.Basal segmentshort,secondsegmentwith triangularcuticularflaps, thirdsegmentbearingthree rectangularcuticularflaps,two slendersetae arisingfrom nearbaseof reducedspiniformprocess.Fourthsegmentelongateand unarmed;fifth segmentan unciformterminalclaw bearingtwo slendersetaeandone tinyseta nearauxiliaryspine. Mouthtube siphonostomeand similarto thatofother species.Mandible(Figure28E) of twoparts,dentiferousmargin withsix teeth.First maxilla(Figure 28F)biramous;endopodwith tinytriangularcuticularflaps,bearing one apicalbilaterallydenticulatedseta, andone smoothseta;exopodlongersurmountedby twosmall setaeand onelonger seta.Secondmaxilla (Figure28G) brachiform, lacertussubequalto brachiumbearinglarge triangularcuticularflaps,brachiumwith triangularcuticularflaps, andone tuftof setae atbase ofterminalclaw(calamus).Claw bearingtwo rowsof denticlesparallellingthe concavemargin.Maxilliped(Figure28H) chelate,indistinctlysegmented,proximalsegmentpedunculate;corpusmaxillipedis robustbearingsmall stoutspine ondistal margin,and tinytriangularcuticularflaps; myxalarea expandedinto largereceptacleto accommodateclaw ofopposablesegment.Shaftbearing twosetae: onenear midpointon lateralconvex marginand theother adjacentto a stripof membranealong concavedistalmargin.Claw unciformwithovoid cuticularexpansionproducinglateral shield.First fourpairs oflegs biramousand trimeriteexcept two-segmentedexopodandthe fusedto partiallyfused endopodof legone, two-segmentedsympods.All basipodswith lateralslender seta;first basipodbears additionaldistomedialslenderseta.Ventralsurfacesof all fourlegs bearingtriangularshapedcuticularflapsexcept forthin rectangular flapsfound onthe basalsegmentof modifiedexopodtwo. Armatureof rami asfollows:Leg oneExopod1-0 1-0III Endopod0-0- IILeg two(modified)Exopod1-0 1-0III Endopod0-0 0-0IILeg threeExopod1-0 1-0III Endopod0-0 0-0ILeg fourExopod1-0 1-0III Endopod0-0 0-0ILeg one(Figure29A) exopodalsetaespiniform,bilaterallybearingserratedmembranousflange, smallerendopodalseta bilaterallyspinulated,largerunilaterally59bearingdenticles.Exopod two(Figure29B) modified,first segmentgreatly enlargedwith manythin rectangularcuticularflaps andone distalcurving seta,second segmentsmallerwith distallyextendinglaterallobe bearingsingle curvedseta, thirdsegmentarmedwith threecurved setae.Leg twoendopod(Figure 29C)with onesmall spinulated setaand one largebilaterallyspinulatedseta onterminalsegment.Legs threeandfour (Figures29D) similar;exopodsthree andfour withsmoothstyliformsetae,endopodsthree andfour tippedwith a singleunilaterallydenticulatedseta.Leg five(Figure29E) ovoid,barelylonger thanwide; lateralsurface coveredwith triangularcuticular flaps,distally bearingthree slendersetae plusone similarseta arisingdorsallyfrom thebase.Male: UnknownComments:E. insolensseems tobe a rarelyreportedparasiteof temperateeasternNorth Atlanticand Mediterraneansharks. Sofar, it hasbeen reportedfrom thegills of thetope shark,Galeorhinusgaleus (Linnaeus,1758) collectedfrom theIrish Sea(Scott andScott,1913), andoff Norfolkin the NorthSea (Hamond,1969).Essafi andRaibaut(1977)have foundthis parasiticcopepodon the gillsof thesmooth-houndMustelusmustelus(Lin naeus,1758),the blackspottedsmooth-houndMustelusmediterraneusQuignardand Capape,1972 (=MusteluspunctulatusRisso, 1826),andthe starrysmooth-houndMustelusasteriasCloquet,1821 allcollectedin theMediterraneanSea nearTunis.It shouldbe notedthat the generathis copepodparasitizes,GaleorhinusandMustelusare membersof the carcharhiniformfamily Triakidae.The unusualabsenceof cuticularflaps on thedorsalsurfacesof the cephalothorax and freethoracicsomitesone throughfour coupledwith thedistallyextendedlobeof the secondsegmentof the modifiedsecond endopodand denticulatedclaw ofthesecond maxillahelp to identifythis copepod.EudactylinalongispinaBere, 1936(Figures30-31)Materialexamined.One holotypefemale (USNM69839)from thebranchialIamellaeof the bonnetheadshark, Sphyrnatiburo(Linnaeus,1758)collectedfromLemon Bay,Florida inthe Gulf ofMexico,and onefemale (USNM153653)from the60same species of host from Tampa Bay, Florida.DescriptionFemale (Figure 30A)Overall length in lateral view approximately 1.1 mm. Cephalothorax longer thanwide, lateral margin notched accomodating lacertus of second maxillae. Lateral surfaceof cephalothorax covered with cuticular flaps. Cuticular flaps restrictedto lateral surfaces of first and second free thoracic somites. Third andfourth free thoracic somitesatypically devoid of cuticular flaps. Fourth free thoracic somitebearing leg five. Genitalsegment and abdomen bearing posteriorly directed cuticular flaps on ventral surface.Genital segment smaller than preceeding somite. Abdomen two-segmented. Caudalramus (Figure 30B) longer than wide, bearing three terminal slender setae, one dorsomedial slender seta, and one lateral slender seta; ventral surface armed with posteriorlydirected triangular cuticular flaps.First antenna (Figure 30C) five-segmented, armature (proximal to distal) as follows: first segment bearing one small stout seta; second segment with eight smoothstout setae, plus one curving (prehensile) claw; third segment bearing seven smoothsetae; fourth segment bearing a single seta. Terminal segment armed with 14 smoothslender setae plus one aesthete. Second antenna (Figure 30D) five-segmented, prehensile. Basal segment short, second segment naked, third segment devoid of cuticular flaps, two slender setae arising from distal margin. Fourth segment elongate andunarmed; fifth segment an unciform terminal claw bearing two slender setae and oneelongate auxiliary spine. Mouth tube siphonostome and similar to that of other species.Mandible (Figure 30E) of two parts, dentiferous margin with six teeth. First maxilla(Figure 30E) biramous; endopod bearing two slender setae, exopod longer surmounted by two small setae and one longer seta. Second maxilla (Figure 30F) brachiform,lacertus subequal to brachium bearing elongate, triangular cuticular flaps, brachiumwith long, triangular cuticular flaps, and one tuft of setae at base of terminal claw (calamus). Claw bearing two rows of denticles parallelling the concave margin. Maxilliped(Figure 30G) chelate, indistinctly segmented, proximal segment pedunculate; corpusmaxillipedis robust bearing small stout spine on distal margin, and large triangular cuticular flaps; myxal area expanded into large receptacle to accommodate claw of opposable segment. Shaft bearing two setae: one near midpoint on lateral convex marginand the other adjacent to a strip of membrane along concave distal margin. Claw unciform with ovoid cuticular expansion producing lateral shield.First four pairs of legs biramous and trimerite except two-segmented endopod ofleg two, two-segmented sympods. All basipods with lateral slender seta; first basipod61bearsadditionaldistomedialstoutseta.Ventralsurfacesof all fourlegs bearingtriangular shapedcuticularflaps.Armatureof ramias follows:Leg oneExopod1-01-0IllEndopod0-00-0IILeg two(modified)Exopod1-01-0IIIEndopod0-0-IILeg threeExopod1-01-0IllEndopod0-00-0 0-0Leg fourExopod1-0 1-0IIIEndopod0-00-00-0Leg one(Figure31A) exopodalsetaesmoothandblunt,endopodalsetaeslender, largestsetadenticulated,smallestseta smooth.Exopodtwo(Figure31 B)typicallymodified,first segmentgreatlyenlargedwithproximalpatchof largetriangularcuticularflaps andonedistalcurvedseta,secondsegmentsmallerbearingsinglecurvedseta,third segmentarmedwith threecurvedsetae,one armedwith asinglesetule.Leg twoendopod(Figure31C)with twosmoothslendersetae.Legs threeand foursimilar;sympodsand exopodsthreeand four(Figure31 D)with denticulatedstyliformsetae,terminalsegmentof endopodsthreeandfour (Figure31E)modifiedinto alongbluntscierotizedprocess,patchof largetriangularcuticularflapson basalsegment.Leg five(Figure31 F) longerthan wide,distallybearingthreeslendersetaeplus onesimilarseta(not illustrated)arisingdorsallyfrom thebase;lateralsurfacesparselycoveredwithtriangularcuticularflaps.Male:UnknownComments:E. longispinawasreportedoriginallybyBere(1936),andlater byPearse(1952)and Cressey(1970)from thegillsof thebonnetheadsharkSphyrnatiburo(Linnaeus,1758).All collectionswerefrom theWestcoastof Floridain theGulfof Mexico.The bizarrelymodifiedterminalsegmenton theendopodof legsthreeand fourforminga long,bluntprocessis uniqueto thisspecies.EudactylinamyliobatidosLuqueand Farfan,1991(Figures32-33)Materialexamined.One female(USNM251 291)fromthe branchiallarnellaeofthe batray MyliobatischilensisPhillippi,1892 collectedfrominshorewatersnear62Chorrillos,Peru.DescriptionFemale (Figures32A)Overall lengthin lateral view approximately1 .8 mm. Cephalothoraxlonger thanwide, lateralmargin slightlynotched accomodatinglacertus ofsecond maxiflae.Dorsolateralsurface of cephalothoraxcovered withwavy cuticularflaps.Wavy cuticularflaps presenton dorsolateralsurfaces of first,second, thirdand fourthfree thoracicsomites.Fourth freethoracic somitebearing legfive. Fourth freethoracic somite,genital segment,and abdomenbearing posteriorlydirected cuticularflaps onventral surface. Genitalsegmentsmaller thanpreceedingsomite. Abdomentwo-segmented.Caudal ramus(Figure 328)longer thanwide, bearingtwo relativelyelongate,apicallycurved terminalsetae, onedorsal slenderseta, andone lateralslender seta;ventralsurface armedwith posteriorlydirected triangularcuticular flaps.First antenna(Figure 32D)indistinctly four-segmented,armature(proximal todistal) as follows:first segmentbearing onesmall seta;second segmentwith fivesmooth setae(one elongate,two minute, twostout), onesmall, unilaterallydenticulated seta, plusone curving,denticulated(prehensile)claw; thirdsegmentbearing sixsmooth setae,plus two largedistal auxiliaryspines (oneunilaterally bearingteeth andthe other withfour semicircularcuticular flaps),Terminal segmentarmed with11 slendersetae, onelarge stout seta,plus one aesthete.Second antenna(Figure 32D)five-segmented,prehensile.Basal segmentshort, secondsegmentarmed withspinousprocess, thirdsegment bearingsix rectangularcuticular flapsand twoslender setaearising frombase of wellproducedspinous process.Fourth segmentelongate andunarmed; fifthsegment anunciform terminalclaw bearinga single slenderseta andoneelongate auxiliaryspine. Mouthtube siphonostomeand similar tothat of otherspecies.Mandible (Figure32E) of two(possibly three)parts, dentiferousmargin withsix teeth.First maxilla(Figure 32F)biramous;endopod bearingtwo setae, bothunilaterallydenticulated, exopodlonger surmountedby two smallnaked setaeand one longer,unilaterally denticulatedseta. Secondmaxilla(Figure 32G)brachiform,lacertus largerthanbrachiumbearing large,semicircularcuticularflaps, brachiumwith largeslender,rectangular cuticularflaps, a distalpatch of prickles,plus two tuftsof setae (onecoarseorrope-like (possiblyfused setae)and onefine or hair-like)at baseof terminalclaw (calamus). Clawbearing fourrows of serratedmembranesdistally, plusone longerserratedstrip proximally,two similarstrips parallellingthe concavemargin presenton otherside.Maxilliped (Figure32H) chelate,indistinctlysegmented,proximal segmentpedunculate; corpus maxillipedisrobust, typicalsmall stoutspine on distalmargin notobserved,63corpuswith rectangularcuticularflaps;myxalarea bearinglargetransversecuticularflangeand expandedinto largereceptacleto accommodateclawof opposablesegment.Shaftbearingtwo setae:one nearmidpointon lateralconvexmarginand theotheradjacentto a stripof membranealongconcavedistalmargin.Claw unciformbearingmembranousflangewith quadratecuticularexpansionproducinglateralshield(see detail).First fourpairsof legsbiramousand trimeriteexceptfor thetwo-segmentedendopodof thefirst leg,two-segmentedsympods.All basipodswith lateralslenderseta;first basipodbears additionaldistomedialslenderseta.Ventralsurfacesof all fourlegs bearingvariouslyshapedcuticularflaps.Armatureof ramias follows:Leg oneExopod1-0 1-0IVEndopod0-0- IILeg two(modified)Exopod1-0 1-0III Endopod0-00-0IILeg threeExopod1-0 1-0IllEndopod0-00-0ILeg fourExopod1-01-0IIIEndopod0-00-0ILeg one(Figure33A)exopodalsetaebearinglateralserratedmembranousflangeexceptfor thetwo mostdistomedialsetae(largestsetaappearsto bilaterallybear spinulesor tinydenticles,smallestsetanaked),endopodalsetaebilaterallydenticulated/ spinulated.Exopodtwo (Figure33B)modified,first segmentgreatlyenlargedwith twothin, rectangularcuticularflapsand onedistaldenticulatedseta,secondsegmentsmallerbearingsinglestoutseta,thirdsegmentarmedwith onesmallslenderseta,one slenderrecurvingseta,and onetruncateseta.Legtwo endopod(Figure33C) withtwo slendersetae,longestbilaterallybearingstoutsetulesor minutedenticlesand theothersmooth.Legs threeand foursimilar;exopodsthreeand four(Figure33D)with denticulatedstyliformsetae,terminalsegmentof endopodsthreeandfour(Figure33D) bearingsinglespinulatedseta.Medialstylet(Figure33E) stoutand blunt,locatedbetweenpedigersthreeand fourand notdirectlyarisingfromeithersomitesinterpodalbars.Leg five(Figure33F)longerthan wide;lateralsurfacesparselycovered withsmall triangularcuticularflaps,distallybearingthreeslendersetaeplusonepinnatesetaarisingdorsallyfrom thebase.Male:UnknownComments:This redescriptionof E. myliobatidosbringsattentionto detailmissing inthe originaldescriptionof thisspecies(LuqueandFarfan,1991).This speciessharesthe charactersof a four-segmentedfirstantenna,elongatedsegmentsof the64second antenna, and wavy cuticular flaps with E. hornbostell, E. mdivisa, and E. nykterimyzon all parasites of Myliobatis sp. from their respective waters.The large semicircular flaps and denticulations on the two large distal auxiliaryspines on the third segment of the first antenna, the denticulated seta on the proximalsegment of exopod two and the relatively elongate terminal setae on the caudalramiare found in E. mdivisa, also a parasite of Myliobatis chilensis. Both E. mdivisa andE.myliobatidos are so similar to one another that synonymization may be warranted.Although, the two large rectangular cuticular flaps on the proximal segment of the modified second exopod separate this species from E. indivisa, additional collecting of theseparasites may show this trait not to be as taxonomically unique as this present effortsuggests.Eudactylina nykterimyzon sp. nov.(Figures 34-35)Material examined. Several females from the branchiallamellae of the bat rayMyiobatis californica Gill, collected from inshore waters nearEl Segundo, Californica;and several females from the same host speciesfrom Punta Arena de Ia Ventana, inthe southern Sea of Cortez (Gulf of California).Female holotype (USNM 266527) and7 female paratypes (USNM 266528) deposited at theUnited States National Museumof Natural History.Etymology: The specific name nykterimyzon is derivedfrom the Greek nykteris meaning bat, and myzo meaning suck, referring to this species’predilection for feeding uponbat rays.DescriptionFemale (Figures 34A and 34B)Overall length in lateral view approximately 2.0 mm.Cephalothorax longer thanwide, lateral margin notched accomodating lacertusof second maxillae. Dorsolateralsurface of cephalothorax covered with cuticular flaps.Cuticular flaps present on dorsolateral surfaces of first, second, third and fourth freethoracic somites. Fourth free thoracic somite bearing leg five. Fourth free thoracicsomite, genital segment, andabdomen bearing posteriorly directed cuticular flapson ventral surface. Genital segment smaller than preceeding somite. Abdomen two-segmented.Caudal ramus65(Figure 34C) longer than wide, bearing two terminal, apically curved setae, one dorsomedial slender seta, and one lateral slender seta; ventral surfacearmed with posteriorlydirected triangular cuticular flaps.First antenna (Figure 34D) indistinctly four-segmented,armature (proximal todistal) as follows: first segment bearing one nakedseta; second segment with sevensetae (four small, three large), plus one curved,denticulated (prehensile) claw; thirdsegment bearing nine (six small, one elongate, andtwo large distally placed auxiliaryspines (setae)) setae, terminal segment armed with14 setae plus one aesthete.Second antenna (Figure 34E) five-segmented,prehensile. Basal segment short, second segment armed with spiniform process, third segment bearingthree cuticular flapsand two slender setae arising from base of well produced spiniformprocess. Fourthsegment elongate and unarmed; fifth segment an unciform terminal claw bearing twoslender setae and one elongate auxiliary spine. Mouthtube siphonostome and similarto that of other species. Mandible (Figure 34F) of two parts, dentiferous margin withsix teeth. First maxilla (Figure 34G) biramous; endopodbearing two slender setae,longer seta unilaterally denticulated, exopod longer surmounted by two small setae andone longer slender seta. Second maxilla (Figure 34H) brachiform,lacertus larger thanbrachium bearing large, wavy cuticular flaps, brachium with large wavy, and triangularcuticular flaps, a distal patch of prickles, possibly a small spine plus two tufts of setae(one coarse or rope-like (possibly fused setae) and one fine or hair-like)at base of terminal claw (calamus). Claw bearing three rows of serrated membranes distally, plusone longer serrated strip proximally, two similar strips parallelling the concave marginpresent on other side. Maxilliped (Figure 341) chelate, indistinctly segmented,. proximalsegment pedunculate; corpus maxillipedis robust bearing small stout spine on distalmargin, and large wavy cuticular flaps; myxal area expanded into large receptacletoaccommodate claw of opposable segment. Shaft bearing two setae: one near midpointon lateral convex margin and the other adjacent toa strip of membrane along concavedistal margin. Claw unciform bearing membranous flange with subquadrate cuticularexpansion producing lateral shield.First four pairs of legs biramous and trimerite, two-segmented sympods. Allbasipods with lateral slender seta; first basipod bears additional distomedial seta.Ventral surfaces of all four legs bearing variously shaped cuticular flaps. Armature oframi as follows:Leg one Exopod 1-0 1-0 IV Endopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 Ill Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 lii Endopod 0-0 0-0 I66Leg four Exopod 1-0 1-0III Endopod 0-00-0 ILeg one (Figure 35A) exopodal setaebearing lateral serratedmembranousflange, endopodal setae bilaterally denticulated.Exopod two (Figure35B) modifiedand devoid of cuticular flaps, first segment greatlyenlarged with one distalseta, secondsegment smaller bearing singlestout seta and conical protuberance, thirdsegmentarmed with two small setae, andone truncate seta. Leg two endopod (Figure35B) withtwo smooth setae. Legs threeand four similar; exopods three and four (Figure35C)with denticulated styliform setae, terminalsegment of endopods three and four (Figure35C) bearing single denticulated seta. Leg five(Figure 35D) longer than wide, distallybearing three slender setae plus one setaarising dorsally from the base; lateral surfacesparsely covered with cuticular flaps.Male: UnknownComments: Eudactylina nykterimyzon seems to bespecific to the California batray Myliobatis californica Gill, 1865.Other species possessing a distinctly 4-segmentedfirst antenna are: E. hornbosteli, E. indivisa, E. myliobatidos, E. pristiophori, E.squamosa, E. turgipes, and E.urolophi. E. acuta, E. aphiloxenous, andE. tuberifera all possess an indistinctly 4- or5-segmented first antenna, as segments three and fourshow incomplete fusion. Six ofthe aforementioned species possess serrated membraneson the setae of the first exopod, these are: E. hornbosteli, E. indivisa, E. myliobatidos, E. pristiophori,E. turgipes,and E. urolophi. Five of these speciespossess only four setae on their caudal rami,they are: E. hornbosteli, E. mdivisa,E. myliobatidos, E. pristiophori, and E. urolophi.Only three of these species, E. hornbosteli,E. mdivisa, and E. myliobatidos possesselongate spiniform processes on segments two and three of thesecond antennae, asdoes E. nykterimyzon. E. nykterimyzon can be distinguished from this small group ofallies by being the only member with three cuticular flaps on segment three of thesecond antenna; the others all have six. The extremely elongate components of thesecond antenna of E. hornbosteli set that species apart fromthe rest of this group. Thepossession of a denticulated seta on the second segment of the second exopod andcuticular flaps on the auxiliary spine of E. mdivisa and E. myliobatidos additionally guarantee the uniqueness of this new species.67Eudactylina oliveri Laubier, 1968(Figures 36-39)Syn: Eudactylina olivieri, of Pillai (1985)Material examined. Severalfemales and males from the branchiallamellae ofthe spinetail mobula,Mobula japanica (Muller andHenle, 1841) collected from PuntaArena de Ia Ventana (Gulf ofCalifornia), Mexico and from the same host speciesfromnear Anacapa Island(southern California ChannelIslands); several females.from thesmoothtail mobula Mobula thurstoni(Lloyd, 1908) (=Mobula lucasana Beebe andTee-Van, 1938) collected from Punta Arenade Ia Ventana, Mexico; several females fromMobula sp. collected from Nosy Be,Madagascar from the personal collectionof Dr.Roger Cressey, Smithsonian Institution,Washington, D.C.DescriptionFemale (Figure 36A)Overall length in lateral view approximately 1.9 mm. Cephalothorax longer thanwide, lateral margin notched accomodatinglacertus of second maxillae. Lateral surfaces of cephalothorax covered withcuticular flaps. Cuticle of first, second, and thirdfree thoracic somites laterally coveredwith cuticular flaps. Fourth free thoracic somitesmaller than previous three with cuticularflaps on ventral surface and bearing leg five.Genital segment smaller than preceedingsomite,with cuticular flaps on ventral surface.Abdomen two-segmented, ventral surface bearingcuticular flaps. Caudal ramus(Figure 36B) longer than wide, bearingfour terminal small stout setae, one subterminalsmall stout seta, plus one dorsal slenderseta; ventral surface covered with posteriorlydirected triangular cuticular flaps. Egg string(Figure 36C) uniseriate.First antenna (Figure 36E) six-segmented, armature (proximalto distal) as follows: first segment with triangularand rectangular cuticular flaps along lateral marginbearing one stout seta; second segment witheight naked setae of various sizes plusone large smooth, curving (prehensile) claw; thirdsegment with four slender setae plusone small papilliform seta; fourth segment with twolarge seta, one small stout seta, andone large distal spine; fifth segmentbearing one tiny seta, terminal segment with 14setae plus one aesthete. Second antenna (Figure 36F) five-segmented,prehensile.Basal segment short, second segmentelongate bearing long rectangular cuticularflaps, third segment bearing rectangularcuticular flaps, two slender setae arising fromnear base of very reduced spinousprocess. Fourth segment elongate with many rectangular cuticular flaps (only twovisible in illustrated view), fifth segment an elongate68unciform terminal claw bearing two slendersetae and one longslender spine. Mouthtube (Figure 36D) siphonostome andsimilar to that of otherspecies; labrum withsub-rectangular flaps, distal edgeof frons labri bearing membrane;lateral surface oflabiumwith elongate triangular cuticularflaps, ventrodistal surfacewith smali prickles.Mandible (Figure 36G)of two parts, dentiferous marginwith six teeth. Firstmaxilla(Figure 36H) biramous;sympod with small patchesof spinules, endopod bearing twostout spinulated setae;exopod longer surmountedby two small naked setae and onelonger denticulatedseta. Second maxilla (Figure361) brachiform, lacertus larger thanbrachium, with small triangularcuticular flaps, brachium withrectangular and triangularcuticular flaps, a distal patch ofprickles a tiny seta anda single tuft of fine setae at baseof terminal claw(calamus). Distal region of claw bearingtwo rows of denticles, proximal region with three pairsof serrated membranes. Maxilliped(Figure 37A) chelate,indistinctly segmented, proximalsegment pedunculate; corpusmaxillipedis robust bearing small stout spine on distalmargin, small semicircular cuticularflaps and small triangular flaps; myxal area expandedinto large receptacle to accommodateclaw of opposable segment. Shaftbearing two setae: one near midpointon lateral convex marginand the other adjacentto a strip of membrane along concave distalmargin. Claw(Figure 37B) unciform with reduced quadrangularcuticular expansion producinga verysmall lateral shield.First four pairs of legs biramouswith three-segmented endopods, three-segmented exopods and two-segmentedsympods. All basipods with lateral seta,(basipods two, three and four with spatulatesetae); first basipod bears additional distomedial seta. Ventral surfaces of all fourlegs bearing triangular and semicircularcuticular flaps. Armature of ramias follows:Leg one Exopod 1-01-0 III Endopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 IllEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IllEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IExopodal setae of leg one(Figure 37C) small and denticulated on the most proximal two segments, endopodalsetae unilaterally denticulated. Exopod two (Figure37D) somewhat modified, firstsegment elongate with a proximal patch of triangularcuticular flaps along lateral margin, onedistal naked seta, second segment smaller withsmall, naked seta, thirdsegment armed with two small sub-apical nakedsetae, plusone longer unilaterally denticulated(two teeth) seta. Leg two endopod (Figure 37D) with69two finely spinulated/denticulatedsetae on terminal segment.Setae of legs three andfour (Figure 37E) stout withlargest endopodal and exopodalseta bearing well developed spinules. Leg five (Figure 37F) elongatedistally bearing threespatulate setaeplus one seta arising dorsallyfrom the base; lateral surface withrectangular and triangular cuticular flaps along ventralsurface.Male: (Figure 38A)Overall length in lateral viewapproximately 1.6 mm. More slender thanfemale,abdomen four-segmented, leg fivearising from fourth free thoracic somiteand leg sixarising from genital segment.Caudal ramus (Figure 38B) bearing four robustsetae(three semipinnate), one dorsalpinnate seta and one small lateral nakedseta.First antenna (Figure 38C) indistinctlyeight- or nine-segmented; armature (proximal to distal) as follows:first segment bearing one smallseta and triangular cuticularflaps along lateral margin, secondsegment with eleven slender setaeplus one largecurving (prehensile) claw, thirdsegment armed with nine slender setae,fourth segmentwith four distal setae and one proximalslender seta, fifth segment with two slendersetae, sixth segment with two slendersetae, seventh segment armed with two setae,terminal segment bearinga single aesthete plus 13 slender setae of various heights.Second antenna (Figure 38D)five-segmented, prehensile; proximal segment short,second segment elongate with proximally directedrectangular cuticular flaps, third segment with proximally directed cuticularflaps and two slender setae arising from nearbase of small spiniform process, fourth segmentelongate and naked, terminal segmentan elongate claw with onesmall auxiliary spine and two proximal slendersetae.Mandible (Figure 38E) similarto that of female. First maxilla (Figure 38F) biramous;endopod bearing small triangular cuticular flapsand two apical setae, one smooth andone unilaterally denticulated;exopod with triangular cuticular flaps, two small setae andone long seta. Second maxilla (Figure38G) similar to that of female. Maxilliped (Figure38H) subchelate; corpus bearingmany small rectangular cuticular flaps, myxal areabearing a well produced denticulated,styliform process, shaft bearing two small slendersetae, claw produced into a large curved processproximally producing a robust spinealong the concave margin.First four pair of legs (Figures 39A-39E) biramousand trimerite. Sympods twosegmented. Variously shaped cuticularflaps on ventral surfaces of legs. Armature oframi as follows:Leg one Exopod 1-0 1-0 IV Endopod0-0 0-0 IILeg two Exopod I-I I-I VI Endopod0-I 0-I V70Leg three Exopod I-I I-I VIIEndopod 0-I 0-I IVLeg four Exopod I-I I-I VIIEndopod 0-I 0-I IIIExopod one (Figure 39A) three-segmented;first segment with single smoothseta, second segment bearing single shortseta, terminal segment with four apicalnaked setae. Endopod one (Figure 39A)tipped with two short setae. Leg two (Figure39B) lateral exopodal setae with membranesalong lateral margins, medial setae pinnate, lateralmost long seta semipinnatewith membrane along lateral margin; endopodal setae pinnate. Exopod three and four(Figure 39C) with stout setae aloog lateralmargin and pinnate setae along medial margin.endopodal setae pinnate along medialmargin, apical setae naked. Terminalsegment of endopod three (Figure 39D) with onenaked plus three pinnate setae. Terminalsegment of endopod four (Figure 39E) withone naked and two pinnate setae. Leg five(Figure 39F) composed of three pinnatesetae on distal edge of leg with one similar seta at base. Leg six (Figure39G) consistsof two pinnate setae on distal edge of smalllobe.Comments: E. 0//yen was originally describedby Laubier, 1968 from gills ofMobula mobu/ar (Bonnaterre, 1788) captured nearNarbonne Beach, along the FrenchMediterrranean. The next record of this parasite was from gills of MobuladiabolusSmith, 1943 (=Mobula kuhili (Valenciennes in Muller and Henle, 1841)) fromCapeComorin, India (Pillai, 1985). This redescriptionadds to the list Mobula japanica (MUllerand Henle, 1841) from the southern Sea of Cortez (Gulfof California) and from off theChannel Islands, southern California, Mobula thurstoni(Lloyd, 1908) (previously knownas M. lucasana Beebe and Tee-Van, 1938) from the southern Sea of Cortez and,Mobula sp. from Nosy Be, Madagascar.This host list is interesting in light of the recent revision of the genus MobulabyNotarbartolo-di-Sciara (1987) in which he mentions the possible synonymy of Mobulamobular and Mobula japanica and, the possibility of some records of M. kuhili being inerror and probably representing M. thurstoni. In absence of more reliable host idenetification one can speculate that E. oilyen maybe specific only to the spine-tail mobula,Mobula japan/ca and the smooth-tail mobula, Mobula thurstoni.Recently, another new species, Eudactyilna mobuli was described by Hameedet al, (1990) from Mobula diabolus from off the coast of Kerala in the Indian Ocean. Thehideous cartoon-like abstractions used for illustrations add nothing but pain and confusion to the researcher from that literature. Key attributes suchas the unique spatulatesetae found in E. oliveri are not shown or compared with features found on E. mobull.Hameed et al, (1990) claim their new species is morphologically similar to E. oliveri,71but differ by an unarmed claw of thesecond antenna (this does notoccur anywhere inthe genus that I am aware of),and a two-segmented exopod onthe first leg. The highlyabstracted illustrations of thesecharacters coupled with the lack ofso much detailmake these claims difficultto agree with. The general appearance of theillustrations(the elongate exopods of legsthree and four resulting in an ant-like appearance)andthe possibility of the hostbeing M. thurstoni, suggest to me thatE. mobuli may be synonymous with E. oilyen.Eudactylina oilyen is easily distinguished from allother congeners by the spatulate spine-like setae locatedon the basipods of legs two, three,and four and onreduced leg live.Eudactylina papillosa Kabata, 1970(Figures 40-41)Material examined. One paratype female (BMNH 1968.1.5.2)from the branchiallamellae of the stingray, Dasyatus kuhil (Mullerand Henle, 1838) collected fromMoreton Bay, Queensland, Australia.DescriptionFemale (Figure 40A)Overall length in lateral view approximately1.3 mm. Cephalothorax longer thanwide, lateral margin slightly notched accommodatinglacertus of second maxillae.Dorsolateral surfaces of cephalothorax covered with wavycuticular flaps. Cuticle offirst, second, third and fourth free thoracic somitescovered with cuticular flaps. Fourthfree thoracic somite smaller thanprevious three with cuticular flaps on ventral surfaceand bearing leg five. Genital segment smallerthan preceeding somite,with cuticularflaps on ventral surface. Abdomen two-segmented,ventral surface bearing cuticularflaps. Caudal ramus (Figure 40B) longerthan wide, bearing three stout terminal setae(one very small, two well produced) plus one dorsomedial slenderseta and, one lateralslender seta; ventral surface covered with posteriorlydirected cuticular flaps.First antenna (Figure 40C) five-segmented; armature (proximalto distal) as follows: first segment bearing one slender seta;second segment with six naked setae,three short setae with proximal denticulations/ serrations plus one large curving, denticulated (prehensile) claw; third segment with nineslender setae plus one large distalspine; fourth segment with one slenderseta; fifth (terminal) segment bearing 14 slen72der setae plus one aesthete. Secondantenna (Figure40D) five-segmented,prehensile. Basal segment short, second segmentwith short spiniformprocess and proximallydirected subtriangular cuticular flaps,third segment bearingrectangular cuticular flapsand two slender setae arising frombase of well developed spiniformprocess. Fourthsegment elongate with many smalltriangular cuticular flaps along convex margin,fifthsegment an elongate unciformterminal claw bearing twoslender setae and a very long,slender auxiliary spine. Mandible(not shown) of two parts,dentiferous margin witheight teeth. First maxilla (Figure40E) biramous; sympodnaked, endopod bearing twoapical setae (one smooth and one unilaterallydenticulated); exopod longersurmountedby two small setae and one longerslender seta. Second maxilla(Figure 40F) brachiform, lacertus larger thanbrachium, with semicircular cuticular flaps,brachium withsemicircular cuticular flaps,a distal patch of prickles and twotufts of setae at base ofterminal claw (calamus). Clawbearing two pairs of serrated membranes.Maxilliped(Figure 40G) chelate, indistinctlysegmented, proximal segment pedunculate;corpusmaxillipedis robust bearing smallspiniform seta on distal margin,and rectangular cuticular flaps; myxal area expandedinto large receptacle to accommodateclaw of opposable segment. Shaft bearingtwo setae: one near midpoint onlateral convex marginand the other adjacentto a strip of membrane along concave distalmargin. Claw(Figure 40G detail) unciformwith reduced subquadrangular cuticular expansionproducing a small lateral shield.First four pairs of legs biramouswith three-segmented endopods,three-segmented exopods and two-segmentedsympods. All basipods with lateral slenderseta,first basipod bears additionaldistomedial slender seta. Ventral surfacesof all four legsbearing various shaped cuticularflaps. Armature of rami as follows:Leg one Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 IIIEndopod 0-I 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-01-0 III Endopod 0-0 0-0 ILeg one (Figure 41A) exopod three-segmented;exopodal setae with serratedmembrane along lateral margins,largest seta on terminal segment additionally bearsdenticulations along the distalportion of the medial edge, endopodal setae bilaterallydenticulated. Exopod two (Figure 41C)modified, first segment elongate with four rectangular cuticular flaps near proximalregion and one distal papiiliform seta, secondsegment smaller, naked and bearingone papilliform seta, third segment armed with onesmall sub-apical seta plustwo papilliform setae. Leg two endopod (Figure41B) with73two naked setae on terminalsegment. Legs threeand four (Figure41 D) setaestoutwith largest exopodal setabearing denticles.Leg five (Figure 41 E)subcircular, distallybearing three small pinnatesetae plus one similarseta arising dorsally fromthe base;lateral surface withsubquadrangular andsubtriangular cuticular flaps alonglateral surface.Comments: Eudactylinapapillosa has not beenreported since its discoveryanddescription by Kabata (1970)from gills of the stingray,Dasyatus kuhil.E. papillosa is easilydistinguished from all othercongeners by the papilliform(nipple-like) setae locatedon the modified exopodof leg two.Eudactylina peruensisLuque and Farfan, 1991.(Figures 42-43)Material examined. Sixfemales (USNM 251 289) from thebranchial lamellae ofthe shovelnose guitarfishRhinobatos planiceps Garman,1880 collected from inshorewaters near Chorrillos,Peru.DescriptionFemale (Figure 42A)Overall length in lateralview approximately 1.4 mm.Cephalothorax longer thanwide, lateral margin slightlynotched accommodatinglacertus of second maxillae.Anterolateral anddorsal surfaces of cephalothoraxcovered with cuticular flaps. Dorsaland ventrolateral surfacesof first, second, and thirdfree thoracic somites covered withcuticular flaps; firstand second free thoracic somitesaliform. Fourth free thoracicsomite smaller thanprevious three, bearing leg five.Genital segment smaller than preceeding somite. Abdomentwo-segmented, ventral surface bearinga pair of semicircular cuticular flaps on theposterior segment. Caudal ramus (Figure42B) modified into adigitiform structure bearingtwo strongly sclerotized processes(fused modified setae?),one tiny proximal seta (sensifla?),one dorsomedial slenderseta, one lateral slenderseta plus two ventrodistal shortnaked setae.First antenna (Figure42C) five-segmented, armature(proximal to distal) as follows: first segmentbearing one seta; secondsegment with five small slender setae,two large naked setae, onelarge bilaterally denticulatedseta plus one large denticulated, curving (prehensile) claw;third segment with seven slendersetae, two large stout74naked spines, plus one large uncinate denticulatedspine; fourth segment with oneslender seta; terminal segment with one tiny seta, 11 long slendersetae plus one aesthete. Second antenna (Figure 42D) five-segmented, prehensile.Basal segment short(not shown), second segment with long unciform process and triangularcuticular flaps,third segment bearing large hook-like process, three cuticular flaps andtwo slendersetae arising from near base of well produced spinous process. Fourthsegment elongate and unarmed; fifth segment an unciformterminal claw bearing two slender setaeand one stout uncinate auxiliary spine. Mouthtube siphonostome and similar to that ofother species. Mandible (Figure 42E) oftwo parts, dentiferous margin with five teeth.First maxilla (Figure 42F) biramous; endopod bearingtiny triangular cuticular flaps andtwo apical setae, the longer with spinules or minutedenticles; exopod longer surmounted by two small setae and one longer seta. Second maxilla (Figure42G) brachiform,lacertus with large semicircular cuticular flaps, brachium with crescenticand subquadarangular cuticular flaps and one patch of prickles and one tuft ofrope-like setaeat base of terminal claw (calamus). Claw bearing two rows of serrated membranesparallelling the claw plus two strips of membrane along the convex margin of other side,small basal process at base of lacertus. Maxilliped (Figure 42H) chelate, indistinctlysegmented, proximal segment pedunculate; corpus maxillipedis robust bearing slenderseta on distal margin and narrow cuticular flaps; myxal area expanded into large receptacle to accommodate claw of opposable segment. Shaft bearing two setae: one withpedunculate proximal region near midpoint on lateral convex margin and the other adjacent to a strip of membrane along concave distal margin. Claw complex, cuticularexpansions producing a fused complex of claw and cuticle.First four pairs of legs biramous, leg four with two-segmented endopod, three-segmented exopods and endopods on remaining rami, sympods two-segmented. Allbasipods with lateral slender seta; first basipod bears additional distomedial nakedseta. Ventral surfaces of all four legs bearing semicircular to subtriangular shapedcuticular flaps. Armature of rami as follows:Leg one Exopod 1-0 1-0 III Endopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 III Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 III Endopod 0-0 0-0 ILeg four Exopod 1-0 1-0 III Endopod 0-0 - ILeg one (Figure 43A) exopodal setae unilaterally bearing serrated membranes,largest endopodal seta unilaterally bearing serrated flange, smaller naked. Exopod two75(Figure 43B) modified, first segmentgreatly elongated with tiny triangularcuticularflaps along the lateral margin and onedistal naked seta, second segmentsmaller withnaked seta, third segment armed withone short naked seta, one longercurved nakedseta, plus one large curved unilaterally denticulatedseta. Leg two endopod (Figure43B) with one small naked seta and one largebilaterally spinulated seta on terminalsegment. Scierite bar between leg two and legthree (Figure 43C) giving riseto a single, large medial stylet.Exopodal setae of legs three and four (Figures 43D and43E)large and unilaterally denticulated, endopodalseta unilaterally bearing serrated membranous flange. Leg five (Figure 43F)longer than wide; lateral surface smooth, probably bearing three pinnatesetae (only one intact in my specimens) plus one similarsetaarising dorsally from the base.Male: UnknownComments: E. peruensis can be readily identifiedby the uniquely modified caudal ramus. The similarities of thisstructure with the caudal ramus of the new speciesEudactylina dactylocerca, described herein, plus the two-segmentedendopod of legfour, suggest a close phylogeneticrelationship. Interestingly, these two species andEudactylina rhinobati Essafi and Raibaut, 1979 share the posteriorlyextended terga offree thoracic somites one and two, overlappingfree thoracic somites two and three,respectively. All three infect Rhinobatos, from theirrespective waters.Eudactylina pollex Cressey, 1967(Figures 44-45)Material examined. Several females (USNM 271635) from the branchial lamellae of the great hammerhead, Sphyrna mokarran(Rüppell, 1837) collected fromSarasota, Florida, and several females (USNM 262099) from the same host speciescollected from the Caribbean Sea during a shark tagging expedition.DescriptionFemale (Figure 44A)Overall length in lateral view approximately3.5 mm. Cephalothorax longer thanwide, lateral margin slightly notched accommodating lacertusof second maxillae.76Dorsolateral surfaces of cephalothorax coveredwith tiny triangular cuticularflaps.Cuticle of all body somites covered with patches of tinytriangular cuticular flaps. Fourthfree thoracic somite smaller than previousthree with cuticular flaps on ventralsurfaceand bearing leg five. Genitalsegment smaller than preceeding somite,with cuticularflaps on ventral surface. Abdomen(Figure 44B) two-segmented, ventral surface covered with tiny triangular cuticularflaps. Caudal ramus (Figure 44B) longer than wide,bearing two very large bilaterallydenticulate setae plus one or two (too small for certainty) very tiny setae tipped with a tiny setule,one lateral slender seta plus one dorsomedial slender seta; ventral surface coveredwith posteriorly directed tiny triangularcuticular flaps.First antenna (Figure 44C) five-segmented,armature (proximal to distal) as follows: first segment bearing onestout seta; second segment with nine stout setae; thirdsegment with ten stout setae, plus one very robustseta (only eight total in figure); fourthsegment with one short stout seta; fifth (terminal) segmentbearing 13 slender setaeplus one aesthete. Second antenna (Figure 44D)five-segmented, prehensile. Basalsegment short, second segment without spiniform process,bearing triangular cuticularflaps, third segment bearing a patch ofrectangular cuticular flaps and a patch of tiny triangular flaps along convex margin, two slendersetae arising from distal end of concaveedge. Fourth segment elongate with many small triangularcuticular flaps along convexmargin, fifth segment forminga stout unciform terminal claw bearing two short nakedsetae and a slender auxiliary spine. Mandible (Figure 44E) of two parts,dentiferousmargin with five teeth. First maxilla (Figure 44F) biramous;sympod naked or with spinules, endopod bearing two apicalsetae (one smooth and one sparsely semipinnate ordenticulated); exopod longer with or without spinulessurmounted by two small stoutsetae and one longer seta with or without tiny spinules. Secondmaxilla (Figure 44G)brachiform, lacertus larger than brachium, with triangular cuticular flaps, brachiumalsowith triangular cuticular flaps, a distal patch of densely packedtriangular flaps and fourlarge elongate triangular flaps (modified setal tuft?) at base of terminal claw (calamus).Claw bearing two rows of teeth. Maxilliped (Figure 44H) chelate,indistinctly segmented, proximal segment pedunculate; corpus maxillipedis robust bearing smallseta ondistal margin, and two patches of tiny triangular cuticular flaps;myxal area expandedinto large receptacle to accommodate claw of opposable segment.Shaft bearing twoshort stout setae: one near midpoint on lateral convex margin and the other adjacenttoa strip of membrane along concave distal margin. Claw (Figure 44H detail)unciformwith amorphous cuticular expansion bearing some very fine spinules along outeredgeproducing lateral shield.First four pairs of legs biramous with two-segmented endopodone, modified77endopod four and indistinctly three-segmentedendopods two and three,three-segmented exopods and two-segmented sympods.All basipods withlateral slender seta,first basipod bears additional distomedial denticulatedseta. Ventral surfaces ofall fourlegs bearing triangular shaped cuticular flaps.Armature of rami as follows:Leg one Exopod1-0 1-0 III Endopod 0-0 - IILeg two Exopod 1-01-0 III Endopod 0-0 0-0 IIILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0III Endopod0-0 -Leg one (Figure 45A) exopodal setae powerful clawswith largest seta on terminal segment additionally bilaterally denticulate,endopodal setae bilaterally denticulated.Exopod two (Figure 45B) not modified, allexopodal setae manifest as powerful clawswith largest seta on terminal segmentbilaterally denticulated, leg two endopod (Figure45B) with two denticulated stoutsetae and one seta on terminal segment. Legs threeand four (Figure45C) with powerful claw-like setae, largest exopodal seta bilaterallydenticulate, endopod indistinctly segmented of legthree and partially fused in leg four(Figure 45D) into a two-segmented, bilaterallydenticulated claw. Leg five (Figure 45E)ovoid; lateral surface witha few triangular cuticular flaps along lateral surface, distallybearing three small slender setae plus one similarseta arising dorsally from the base.Comments: Eudactylina pollex was reportedby Cressey (1967 and 1970) fromthe great hammerhead, Sphyrna mokarran (Ruppell,1837), collected from Madagascarand the West coast of Florida(Sarasota), and from the scalloped hammerhead,Sphyrna lewini (Griffith and Smith, 1834) fromoff the west coast of Florida. Thisredescription extends the known geographic range of E. pollextothe Caribbean.E. pollex is easily distinguished from all other congeners by the fused uncinateclaw-like nature of the third and especially the fourth endopod, and the lateral expansion of the proximal segment of endopods two, three,and four. Although the unusualabsence of an uncinate claw on the second segmentof the first antenna appears to beunique to this species, it is probable that the largestout spine on the proximal portionon the third segment represents thisstructure. It is also possible this apparent displacement is more apparent than real. The highly sclerotized condition of the firstantenna makes it very difficult to accurately follow segmentalboundaries.78Eudactylina pristiophorisp. nov.(Figures 46-47)Material examined.Three females from the branchial lamellae ofthe longnosesawshark, Pristiophorus cirratus (Latham,1794) captured from near Green’s Beachatthe Tamar River mouth,northern Tasmania. Specimen (USNM 205516)examined atthe Smithsonian’s SupportCenter, National Museumof Natural History, SmithsonianInstitution, Washington, D.C. Femaleholotype (USNM 266529)and 1 female paratype(USNM 266530) depositedat the United States National Museumof Natural History.Etymology: The specific namepristiophori refers to the generic name ofthe host.DescriptionFemale (Figure 46A)Overall length in lateral view approximately1.8 mm. Cephalothorax longer thanwide, lateral margin slightlyconcave. Surface of cephalothorax, first,second, thirdand, fourth free thoracicsomites covered with cuticular flaps. Fourthtree thoracicsomite, genital segment and, two-segmentedabdomen all bearing cuticular flaps onventral surface. Caudal ramus (Figure46B) bearing terminally two verysquat curvedsetae, plus two elongate slender setae(one dorsomedial, one lateral).First antenna (Figure 46C) apparentlyfour-segmented, armature (proximaltodistal) as follows: first segmentbearing one slender seta; second segment withfourdenticulated setae (3 small, one large),four naked setae (two small, one large, onepapilliform), and one large denticulated,curving (prehensile) claw; third segment witheight naked setae, one stout nakedspine, plus one large denticulated spine; fourthsegment with 14 slender setae plus one aesthete.Second antenna (Figure 46D) five-segmented, prehensile. Basal segmentshort, second segment devoid of cuticular flapswith short spinous process, third segment bearingproximally directed irregular rectangular cuticular flaps and two slendersetae arising from near base of curving spinousprocess. Fourth segment elongateand unarmed; fifth segment an unciform terminalclaw bearing two slendersetae and one curving auxiliary spine. Mouth tube siphonostome and similar to that of other species.Mandible (Figure 46E) of two parts, dentiferous margin with eight teeth. Firstmaxilla (Figure 46F) biramous; endopod more robustbearing two unilaterally denticulatedapical setae; exopod longer and surmountedbytwo small setae and one longer, unilaterallydenticulated seta. Second maxilla (Figure46G) brachiform, lacertus larger thanbrachium bearing semicircular cuticular flaps andsmall basal process,brachium with semicircular and rectangular cuticularflaps, a distal79patch of triangular prickles and twotufts of setae (one coarse or rope-like (possiblyfused setae) and one fine or hair-likeat base of terminal claw (calamus)). Claw bearing four pairs of serrated membranes. Maxilliped(Figure 46H) chelate, indistinctlysegmented, proximal segment pedunculate; corpusmaxillipedis robust bearing smallsetaon distal margin and small rectangular cuticularflaps; myxal area expandedinto areceptacle to accommodate claw of opposable segment.Shaft bearing two setae: onenear midpoint and the other along concavedistal margin near membranous strip(seedetail). Claw (Figure 46H detail) unciformbearing cuticular flange and a quadrate lateral shield.First four pairs of legs biramous and trimerite,sympods two-segmented. Allbasipods with lateral slenderseta; first basipod bears additional distomedial slenderseta. Ventral surfaces of legs two, three, and fourbearing cuticular flaps. Armature oframi as follows:Leg one Exopod 1-0 1-0 IVEndopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg one (Figure 47A) lateral margins of proximal segmentswith single row offine triangular cuticular flaps; lateral exopodalsetae unilaterally bearing serrated membranous membrane, largest apical exopodal seta bilaterally denticulated, smallestslender and naked, endopodal setae bilaterally denticulated.Leg two exopod (Figure 47C)modified, first segment greatly enlarged with thickseta, second segment with stoutseta, terminal segment with one small naked setae,one small recurving seta plus onetruncate seta, endopod (Figure 47B) tipped with two bilaterallydenticulated setae.Legs three and four (Figure 47D) similar;all exopodal setae denticulated, endopodstipped by single finely denticulatedseta. Leg five (Figure 47E) slightly longer than wide,distally bearing three spiniform setae plus oneseta arising dorsally from the base; lateral surface with small subquadrangularcuticular flaps.Male: UnknownComments: Eudactylina pristiophori is the first record ofa parasitic copepodinhabiting the Pristiophoriformes (sawsharks).This species is readily identifiedby the very short, stout, and apically curving terminal setae on the caudal rami.Additionally, it is the only species in the genus with a80stout papilliform seta on segment two of the first antenna.Also, no other species bearsthe unique combination of two slender setae with lateralserrated membranes, an elongate bilaterally denticulated (not semipinnate) seta, anda medial small slender seta onthe terminal segment of the first exopod.Eudactylina pusilla Cressey, 1967(Figures 48-49)Material examined. Several females (USNM 153628)from the branchial lamelIae of the tiger shark, Galeocerdo cuvier (Peronand LeSueur, 1822) collected fromSarasota, Florida.DescriptionFemale (Figure 48A)Overall length in lateral view approximately 4.4 mm.Cephalothorax longer thanwide, lateral margin deeply notched accommodatinglacertus of second maxillae.Dorsolateral surfaces of cephalothorax covered with cuticularflaps. Cuticle of free thoracic somites one-four covered with cuticular flaps.Fourth free thoracic somite smallerthan previous three with cuticular flaps on ventralsurface and bearing leg five. Genitalsegment smaller than preceeding somite,with cuticular flaps on ventral surface.Abdomen two-segmented, ventral surface bearing cuticular flaps. Caudal ramus(Figure 48B) longer than wide, bearing three terminal large blunt (nearly amorphous)setae (the two lateralmost with a row of fine denticles), one lateral slender seta plus onedorsomedial slender seta; ventral surface with a few posteriorly directed tiny triangularcuticular flaps.First antenna (Figure 48C) five-segmented, armature (proximal to distal) as follows: first segment bearing one stout seta; second segment with six thick naked setaetwo slender naked setae, plus one finely denticulated curving (prehensile) claw; thirdsegment with nine thick naked setae and one large spine; fourth segment with oneslender seta; fifth (terminal) segment bearing 13 slender setae plus one aesthete.Second antenna (Figure 48D) five-segmented, prehensile. Basal segment short, second segment without spinous process, bearing subrectangular cuticular flaps, thirdsegment bearing a patch of subrectangular cuticular flapsand two slender setäe arisingfrom base of small spinous extension. Fourthsegment elongate with many small triangular cuticular flaps along convex margin, fifthsegment forming a stout unciform termi81nat claw bearing two slender setae and a single thick auxiliaryspine. Mandible (Figure48E) of two parts, dentiferous margin with five teeth. First maxilla(Figure 48F) biramous; endopod with triangular cuticular flaps bearing two denticulatedsetae; exopodlonger with subtriangular flaps surmounted by two small denticulatedsetae and onelonger denticulated seta. Second maxilla (Figure48G) brachiform, lacertus larger thanbrachium with semicircular cuticular flaps and basal process,brachium also with semicircular cuticular flaps, and distal patch of coarsesetae at base of terminal claw (calamus). Claw bearing two rows of teeth distally andtwo strips of serrated membranesproximally. Maxilliped (Figure 49A) chelate, indistinctly segmented, proximal segmentpedunculate; corpus maxillipedis robust bearing small seta on distal margin,and patches of tiny triangular cuticular flaps; myxal area expanded into large receptacletoaccommodate claw of opposable segment. Shaft bearing two stout setae: one nearmidpoint on lateral convex margin and the other adjacentto a strip of membrane alongconcave distal margin. Claw (Figure 49A detail) unciform with small cuticular expansion producing lateral shield.First four pairs of legs biramous and trimerite. All basipods with lateral pinnate tosemipinnate setae, first basipod bears additional distomedial denticulatedseta. Ventralsurfaces of all four legs bearing triangular shaped cuticular flaps. Armature of rami asfollows:Leg one Exopod 1-0 1-0 III Endopod 0-0 0-0 IILeg two Exopod 1-0 1-0 Ill Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 III Endopod 0-0 0-0 ILeg four Exopod 1-0 1-0 III Endopod 0-0 0-0 ILeg one exopodal setae (Figure 49B) denticulated, endopodal setae (Figure49C) bilaterally denticulated. Exopod two (Figure 49D) modified, all exopodal setaenaked except largest seta on terminal segment unilaterally denticulated, terminal segment atypically hemispherical in shape with setae displaced to medial margin, leg twoendopod (Figure 49E) with two small naked setae on terminal segment. Setae of legsthree and four (Figure 49F) claw-like with largest exopodal seta unilaterally denticulated, endopod tipped with a single bilaterally denticulated seta. Leg five (Figure 49G)wider than long, distally bearing three sparsely pinnate setae plus one seta arising dorsally from the base; lateral surface covered with subtriangular cuticular flaps along lateral surface.82Comments: Eudactylina pusillawas reported by Cressey (1967,1970) from gillsof the tiger shark, Galeocerdocuvier (Peron and LeSueur,1822), collected fromMadagascar and the Westcoast of Florida near Sarasota.E. pusilla is easilydistinguished from all other congenersby the large blunt(nearly amorphous) denticulatedsetae on the caudal rami and by the unusualshape ofthe terminal segment onthe second (modified) exopod and thelateral displacement ofthe three setae which areusually found terminally.Eudactylina similis T.Scott, 1902(Figures 50-51)Syn: Eudactylina rachelaeGreen,1958, (see Kabata, 1979)Material examined. Numerousfemales from the branchial lamellaeof thePacificelectric ray, Torpedo ca/ifornica (Ayres),and the big skate Raja binoculataGirard, captured near Palos Verdes,southern California.DescriptionFemale (Figure 50A)Overall length in lateral viewapproximately 4.5 mm. Cephalothorax longer thanwide, lateral margin slightly concave.Dorsolateral surface of cephalothorax, first,second, third and, fourth freethoracic somites covered with cuticular flaps.Fourth freethoracic somite, genital segmentand, two-segmented abdomen all bearingcuticularflaps on ventral surface.Caudal ramus (Figure 50B) with two terminalstout setae, onelateral slender seta, and one dorsomedialslender seta.First antenna (Figure500) apparently four-segmented, but could be interpretedas seven-segmented; armature (proximalto distal) as follows: first segment bearing oneshort seta; second segment withthree denticulated slender setae (twosmall, onelarge), five naked setae and onelarge denticulated, curving (prehensile) claw; thirdsegment with a proximal group offive slender setae (one denticulated) and a distalgroup of three setae (two slender,one stout) plus one large denticulated spine; fourthsegment elongate with one proximal slenderseta, and either 13 or 14 setae with eitherone or two aesthetes respectively.Second antenna (Figure 50D) five-segmented, prehensile. Basal segment short,second segment devoid of cuticular flaps with smallspinous process, third segmentbearing proximally directed subrectangular cuticular83flaps, distally directed subtriangular flapsalong convex margin, and two slendersetaearising from near base of large spinousprocess. Fourth segment elongate withcuticular flaps along convex margin; fifth segmentan unciform terminal claw bearing twoslender setae and one curving auxiliary spine. Mouthtube siphonostome and similar to thatof other species. Mandible (Figure 50E) of two parts,dentiferous margin with seventeeth. First maxilla (Figure 50F) biramous; endopod withtriangular cuticular flaps bearing two denticulated apical setae; exopodlonger and surmounted by two small nakedsetae and one longer, bilaterally denticulated thick seta.Second maxilla (Figure 50G)brachiform, lacertus slightly larger than brachium bearingsemicircular cuticular flaps,brachium with semicircular cuticular flaps,a distal patch of triangular prickles and twotufts of setae and possibly a small spine atbase of claw (calamus). Claw bearing multiple strips of serrated membranes. Maxilliped (Figure50H) chelate, indistinctly segmented, proximal segment pedunculate; corpus maxillipedisrobust bearing minute setaon distal margin and small rectangular cuticular flaps; myxal area expandedinto areceptacle to accommodate claw of opposable segment. Shaft bearingtwo setae: onenear midpoint and the other along concave distalmargin near membranous strip (seedetail). Claw (Figure 50H detail) unciform bearing cuticular flangeand a subquadratelateral shield.First four pairs of legs biramous and trimerite, sympods two-segmented. AHbasipods with lateral slender seta; first basipod bearsadditional distomedial slenderseta. Ventral surfaces of legs two, three, and four bearing cuticular flaps. Armature oframi as follows:Leg one Exopod 1-0 1-0 IV Endopod 0-0 0-0 IILeg two (modified) Exopod 1-0 1-0 III Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 III Endopod 0-0 0-0 ILeg four Exopod 1-0 1-0 Ill Endopod 0-0 0-0 ILeg one (Figure 51A) lateral margins of proximal segments with singie row offine triangular cuticular flaps; lateral exopodal setae unilaterally bearing serrated membranous flange, largest apical exopodal seta bilaterally denticulated, smallest with distalsetules, endopodal setae slender and bilaterally denticulated. Leg two exopod (Figure51C) modified, first segment greatly enlarged with stout seta, second segment withsquat papilliform seta, terminal segment with one small naked seta, one small recurvingseta plus one truncate seta, endopod (Figure 51 B) tipped with two slender bilaterallyspinulated setae. Legs three and four (Figure 51D) similar; all exopodal setae denticulated, endopod tipped by single finely denticulated slender seta. Leg five (Figure 51E)84longer than wide, distally bearingthree slender setae plusone seta arising dorsallyfrom the base; lateral surfacewith small cuticular flaps.Male: Not obtainedComments: Eudactylinas/mills was first describedby Scott (1902) in Britishwaters from gills of thethorny skate, Raja radiata Donovan,and the skate Raja fulloniCa. Green (1958) discovered it on theAtlantic torpedo, Torpedo nobiliana Bonaparte,and erroneously establisheda new species E. rachelae, forthat record. Also, DelamareDeboutteville and Nunes-Ruivo (1958)discovered E. s/mills on Raja asteriasin theMeditteranean. Boxshall (1974)found it on Raja montaguiand Raja naevus in theNorth Sea, and Kabata (1979)found this copepod on the longnoseskate, Raja rh/naJordan and Gilbert, and on thestarry skate, Raja stellulata Jordanand Gilbert, in theeastern North Pacificnear Vancouver Island. This reportadds the big skate, Rajabinoculata Girard and the Pacific electricray, Torpedo californica Ayres both from nearPalos Verdes in southern Californiato the list. The finding of E. s/mills onTorpedo cal/torn/ca supports the synonymyof E. rachelae and E. s/mills. This copepod is apparently a common parasite of bothRaja and Torpedo worldwide.This species canbe readily identified by the many small denticlesscattered overthe convex surface of the largeprehensile claw on the second segment ofthe firstantenna, and the apparently unique possession(in the genus) of two aesthetes on thedistal segment (s) of the firstantenna.Eudactylina squamosa Bere, 1936(Figures 52-53)Mater/al examined. Several females fromthe branchial lamellae of the cow noseray Rhinoptera bonasus (Mitchill,1815) collected from the West Coast ofFlorida (Gulfof Mexico), donatedto me by Dr. Roger Cressey (personal collection), SmithsonianInstitution; and two females from thegills of the cow nose ray, Rhinoptera ste/ndacheriEvermann and Jenkins, 1891 collectedfrom Punta Arena de Ia Ventana, in the southern Sea of Cortez (Gulf of California).DescriptionFemale (Figures 52A, B)Overall length in lateral view approximately 1 .2 mm. Cepalothoraxlonger than85wide, lateral margin notched accomodatinglacertus of second maxillae. Entiresurfaceof cephalothorax covered with cuticularflaps. Cuticular flapspresent on surfaces offirst, second, third and fourth free thoracic somites. Fourth freethoracic somite bearingleg five. Fourth free thoracic somite, genital segment, and abdomenbearing posteriorlydirected cuticular flaps on ventral surface. Genital segmentsmaller than preceedingsomite. Abdomen two-segmented.Caudal ramus (Figure 52C) longer than wide, bearing two terminal stoutsetae, one very small medialseta, one dorsomedial slender seta,one lateral slenderseta, and one proximal seta near the lateral edge (possibly homologous to the typical setule seen in this area in otherspecies); ventral surface armed withposteriorly directed su btriangu lar cuticularflaps.First antenna (Figure 52D) apparentlyfour-segmented, armature (proximal todistal) as follows: first segmentbearing one seta; second segment with five smallnaked setae, two large setae plus onecurving, denticulated (prehensile) claw; thirdsegment bearing nine naked setae, terminalsegment armed with 15 smooth slendersetae plus one aesthete. Second antenna (Figure 52E)five-segmented, prehensile.Basal segment short, second segmentarmed with blunt spinous process, third segmentbearing three cuticular flaps and twosetae (one of them nearly the size of spinousprocess) arising from base of well produced spinousprocess. Fourth segment elongateand unarmed; fifth segment an unciform terminal claw bearingtwo slender setae andone stout auxiliary spine. Mouthtube siphonostome and similar to that of otherspecies. Mandible (Figure 52F) of two parts, dentiferous marginwith seven teeth. Firstmaxilla (Figure 52G) biramous; endopod largerbearing two elongate setae, longer setaunilaterally denticulated, exopod shorter surmountedby two small naked setae and onelonger slender seta. Second maxilla (Figure 52H) brachiform, lacertuslarger thanbrachium with basal process and bearing cuticularflaps, brachium with irregular triangular cuticular flaps, a distal patch ofcoarse setae and apparently a small spine atbase of terminal claw (calamus). Claw bearing three rows of serrated membranesdistally, plus one longer serrated strip proximally.Maxilliped (Figure 521) chelate, indistinctly segmented, proximal segment pedunculate; corpus maxillipedisrobust bearingsmall stout spine on distal margin, and devoid of cuticular flaps; myxalarea expandedinto large receptacle to accommodate claw ofopposable segment. Shaft bearing twosetae: one near midpoint on lateral convex margin and the other near concave distalmargin. Claw unciform with small lateralshield.First four pairs of legs biramousand trimerite, two-segmented sympods. Allbasipods with lateral slender seta; first basipod bears additional distomedialseta tippedwith two setules. Ventral surfacesof all four legs bearing subtriangular cuticular flaps.Armature of rami as follows:86Leg one Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg two Exopod 1-0 1-0 IIIEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0III Endopod 0-0 0-0 ILeg four Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg one (Figure 53A) exopodal setae thickbearing lateral denticulations, endopod with one slender seta and one larger seta bilaterallydenticulated. Leg two (Figure53B) exopod not modified and bearing subtriangularcuticular flaps, first segmentlargest with one distal stout seta, second segment smaller bearingsingle stout seta,third segment armed with three small stout setae. Leg twoendopod (Figure 53B) withtwo bilaterally denticulated setae (one small, one large). Exopodsof legs three andfour (Figure 53C) with claw-likesetae, largest seta on terminal segment with denticles,terminal segment of endopods three and four (Figure53C) bearing single thick bilaterallydenticulated seta. Leg five (Figure 53D) longer than wide, distally bearing three slender setae plus one similar seta arising dorsally from thebase; lateral surface sparselycovered with triangular cuticular flaps.Male: UnknownComments: E. squamosa has not been reported since its discovery by Bere(1936) on Rhinoptera bonasus from Lemon Bay, Florida. This reportfrom Rhinopterasteindacheri extends the host spectrum and geographic range of the parasite. Thespecimen dissected and studied from the Gulf of California from A. steindacheridifferedfrom its counterpart from the Gulf of Mexicoby possessing cuticular flaps on the maxilliped, cuticular flaps on the fourth segment of the second antenna,cuticular flaps on thepenultimate segment of the first antennae, and terminal elements on the caudal ramiappearing a bit more amorphous and claw-like. Until more specimens are looked at Ifeel it is best to treat these differences as intraspecific. In either case, this (or these)species seem to be specific to the rhinopte rids.The squat habitus, the relatively short expansion of the corpus maxillipedis coupled with the transverse palm of the corpus (resulting in a reduced aperture) readily distinguish this species. Additionally, the thick setae approximating the dimensions of thespiniform process of the third segment of the second antenna are uniqueto thespecies. The relative lengths of the terminal setae on the third segment of the unmodified second exopod decrease from the lateralto the medial margins, in sharp contrastto what occurs in the other species.87Eudactylina tuberiferaCastro and Baeza, 1987(Figures 54-55)Material examined. Severalfemales (USNM 213114) from the branchiallamellae of the southern angelote,Squat/na armata (Philippi, 1887), collected fromthe eastern South Pacific nearAntofagasta, Chile.DescriptionFemale (Figure 54A)Overall length in lateral view approximately1.8 mm. Cephalothorax longer thanwide, lateral margin notched accommodatinglacertus of second maxillae. Entire surface of cephalothorax coveredwith cuticular flaps. Cuticular flapspresent on surfacesof first, second, third andfourth free thoracic somites. Fourth freethoracic somite bearing leg five. Fourth free thoracicsomite, genital segment, and abdomenbearing posteriorly directed cuticular flapson ventral surface. Genital segmentsmaller than preceeding somite. Abdomentwo-segmented. Caudal ramus (Figure 54B)longer than wide,bearing two terminal apically curvedstout setae, one dorsomedial slender seta, andone lateral slender seta; ventralsurface armed with posteriorly directedcuticular flaps.First antenna (Figure 54C) indistinctlyfour-segmented, armature (proximaltodistal) as follows: first segmentbearing one naked seta; secondsegment with fivesmall naked setae, twolarge naked setae (one elongate, onestout), one elongatebilaterally denticulated seta plus one curving,denticulated (prehensile) claw; third segment bearing six slendersetae, one short stout seta, one large naked spine, andonelarge denticulated spine, terminalsegment armed with 14 naked slendersetae, onelong unilaterally denticulatedseta plus one aesthete. Second antenna (Figure 54D)five-segmented, prehensile. Basalsegment short second segment armed with spiniform process, third segmentbearing proximally directed cuticular flaps and two slendersetae arising from base of well produced spinousprocess. Fourth segment elongateand unarmed; fifth segment an unciformterminal claw bearing two slender setae andone well developed auxiliaryspine. Mouth tube siphonostome and similarto that ofother species. Mandible (Figure54E) of two parts, dentiferous margin with eight teeth.First maxilla (Figure 54E) biramous;endopod with triangular cuticular flaps and bearingtwo slender setae, longer setadenticulated, exopod longer surmounted by two smallsetae and one longer bilaterally denticulatedseta. Second maxilla (Figure 54F) brachiform, lacertus larger than brachium andbearing cuticular flaps, brachium with irregularcuticular flaps, and two tufts ofsetae (one coarse or rope-like (possibly fused setae)and one fine or setule-like atbase of terminal claw (calamus)). Claw bearing four rows88of serrated membranes distally, plusone longer serrated stripproximally, two strips ofserrated membranes flankingconcave surface on oppositeside. Maxilliped (Figure55A) chelate, indistinctly segmented,proximal segment pedunculate; corpusmaxillipedis robust bearing smallstout spine on distal margin, and large patchof cuticularflaps; myxal area bearinglarge transverse cuticular flap along anteroventralmargin andexpanded into large receptacleto accommodate claw of opposable segment.Shaftbearing two setae: onenear midpoint on lateral convex marginand the other adjacentto membranous strip alongconcave distal margin. Claw (Figure55A, detail) unciformwith subrectangular lateral shield.First four pairs of legs biramousand trimerite, two-segmented sympods. Allbasipods with lateral slenderseta; first basipod bears additional distomedialslenderseta. Ventral surfaces of all fourlegs bearing variously shaped cuticular flaps.Armature of rami as follows:Leg one Exopod 1-0 1-0 IllEndopod 0-0 0-0 IILeg two Exopod 1-01-0 Ill Endopod 0-0 0-0 IILeg three Exopod 1-01-0 Ill Endopod 0-0 0-0 1Leg four Exopod 1-01-0 1W Endopod 0-0 0-0 ILeg one (Figure 55B) exopodal setae bearinglateral serrated cuticular expansions, endopodal setae and bilaterallydenticulated. Leg two exopod (Figure 55C)modified and bearing subtriangular cuticular flaps alonglateral margin, first segmentgreatly enlarged with one distal, toothedstout seta, second segment smaller. bearingsingle stout, naked seta, third segmentarmed with one small slender seta, one truncateseta, and one recurving seta. Leg twoendopod (Figure 55D) with two bilaterally denticulated slender setae. Exopods of legs three andfour (Figure 55E) with stout toothedsetae, terminal segment of endopods three and four (Figure55F) bearing single thickbilaterally denticulated seta. Leg five (Figure55G) longer than wide, distally bearingthree slender naked setae plus onepinnate seta arising dorsally from thebase; lateralsurface with cuticular flaps.Male: Not obtainedComments: E. tuberifera, the parasite of thesouthern angelote Squat/na armata, is very similar to E. acuta, a parasiteof the Mediterranean angeishark, Squat/nasquat/na and to the new species E. aphiloxenos,parasitizing the Pacific angelshark,Squat/na cailforn/ca off the southern California coast.Castro and Baeza (1987) claim89the swollen third segment of thesecond antenna of the female isa diagnostic characterof this species. However, this is notso; the second antenna is quite typical oftheentire genus and the specimens examined didnot display this accentuated trait.This species, and its two aforemetioned allies,E. acuta and E. aphiloxenous,share the large tines found on thesetae of exopods three and four, and the difficulttodistinguish four- or five-segmentedfirst antenna. The serrated membranes on the lateral edge of the setae on exopod one,possessing three setae on the terminal segmentof exopod one (not four), and the rowof cuticular flaps along the lateral edge of themodified exopod of leg two, separate this speciesfrom the two remaining squatinid parasitizing copepods.Eudactylina turgipes Bere, 1936(Figures 56-57)Material examined. Several females (USNM 155196)from the branchial lamellae of the butterfly ray Gymnurasp. collected from Lemon Bay, Florida (Gulf ofMexico), additional specimens fromsame host species and locality donated to me byDr. Roger Cressey (personal collection),Smithsonian Institution.DescriptionFemale (Figure 56A)Overall length in lateral view approximately 1.7mm. Cephalothorax longer thanwide, lateral margin notched accomodating lacertusof second maxillae. Entire surfaceof cephalothorax covered with cuticular flaps.Cuticular flaps present on surfaces offirst, second, third and fourth freethoracic somites. Fourth free thoracic somite bearingleg five. Fourth free thoracic somite, genitalsegment, and abdomen bearing posteriorlydirected cuticular flaps on ventral surface. Singlemedian intersomitic stylet presentbetween leg three and four (Figure57A). Genital segment smaller than preceedingsomite. Abdomen two-segmented. Caudal ramus (Figure 56B)longer than wide, bearing four terminal setae (two smallunilaterally denticulated setae, one small naked seta,and one elongate bilaterally denticulatedseta), one dorsomedial slender seta, and onelateral slender seta; ventral surfacearmed with posteriorly directed subtriangular cuticular flaps.First antenna (Figure 56C) indistinctly four-segmented, armature (proximaltodistal) as follows: first segmentbearing one small slender seta; second segment with90four small naked setae, onesmall unilaterally denticulatedseta, two elongatenakedsetae, one large stout seta plus onecurving, denticulated(prehensile) claw;third segment bearing eight slendernaked setae, one stout nakedspine, and one large denticulated spine, terminalsegment armed with 15 slendersetae plus one aesthete. Secondantenna (Figure 56D) five-segmented,prehensile. Basal segment short,second segment armed with spinousprocess, third segment bearingtwo rows of proximally directed subrectangular cuticularflaps and two slendersetae arising from base of long spinous process. Fourth segmentelongate and unarmed; fifthsegment an elongate, unciform terminal claw bearingtwo slender setae andone elongate auxiliary spine.Mouthtube siphonostome and similarto that of other species. Mandible (Figure56E) of twoparts, dentiferous marginwith eight teeth. First maxilla(Figure 56F) biramous; endopod bearing one unilaterallydenticulated seta and one longerbilaterally denticulatedseta, exopod longer surmountedby two small naked setae and one longerbilaterallyspinulated (sparsely pinnate)slender seta. Second maxilla (Figure56G) brachiform,lacertus larger than brachiumand bearing a proximal patchof triangular cuticular flapsadjacent to basal processand wavy cuticular flaps on otherside, brachium with wavy(sinusoidal) cuticularflaps, a distal patch of triangularflaps (prickles) and one tuft ofcoarse setae at base ofterminal claw (calamus). Claw bearingone pair of serratedmembranes distally and anotherpair proximally, two strips of serratedmembranesflanking concave surface onopposite side. Maxilliped (Figure 56H) chelate,indistinctlysegmented, proximal segmentpedunculate; corpus maxillipedisrobust bearing smallstout spine on distal margin,and large patch of sinusoidal cuticularflaps; myxal areaexpanded into large receptacleto accommodate claw of opposable segment.Shaftbearing two setae: one largestout seta near midpoint on lateral convex marginand theother adjacentto elongate membranous strip along concavedistal margin. Claw(Figure 56H, detail) complexwith ovoid lateral shield.First leg biramous and bimerite, legstwo three and four biramous and trimerite,two-segmented sympods. Allbasipods with lateral pinnate seta; first basipodbearsadditional distomedial slender seta.Ventral surfaces of all four legs bearing semicircular, triangular, and rectangularshaped cuticular flaps. Armature of ramias follows:Leg one Exopod1-0 - 1111 Endopod 0-0 - IILeg two Exopod1-0 1-0 III Endopod 0-0 0-0 IILeg three Exopod1-0 1-0 III Endopod 0-0 0-0 ILeg four Exopod1-0 1-0 III Endopod 0-0 0-0 ILeg one (Figure 57A) exopodalsetae distally bearing serrated cuticular expan91sions, longest seta on terminal segmentbilaterally bearing tiny spinules, endopod withunusual array of semicircular cuticular flapswrapping around lateral margin of elongated terminal segment, setae bilaterally spinulated. Legtwo exopod (Figure 57B) modified, bearing minute triangular cuticular flapsalong lateral margin, first segment greatlyenlarged, second and third segmentscompressed with a few tiny triangular flaps, allsetae robust and papilliform. Leg two endopod(Figure 57C) with relatively large proximal segment and rectangular cuticularflaps, terminal segment apically bears two pinnate slender setae. Exopodsof legs three and four (Figure 57D) with apically toothedsetae, largest seta on terminal segment pinnate,terminal segment of endopods threeand four bearing single pinnate seta. Leg five(Figure 57E) longer than wide, distallybearing three pinnate setae plus one similarseta arising dorsally from the base; lateralsurface with triangular cuticular flaps.lntersomitic stylet (Figure 57F) plunging downbetween pediger three and pediger four.Male: UnknownComments: E. turgipes was first recordedby Bere (1936) from gills of the butterfly ray Pteroplatea maclura(=Gymnura maclura); it was subsequently found byPearse (1952a) from the smooth butterfly ray, Aetoplatea micrura(=Gymnura micrura,(Bloch and Schneider)), and most recently it turnedup again (Raibaut et al., 1971) inTunisian waters (Medittereanean Sea) from the gills of the spiny butterfly ray, Gymnuraaltevela (Linnaeus). This additional record fromGymnura sp. from Lemon Bay, Floridastrongly suggests E. turgipes is specificto butterfy rays of the genus Gymnura.This species is easily distinguished by the peculiarly swollen modified exopod ofleg two, the papilliform setae of exopod two, the unique combination of setalcharacteristics on the caudal ramus, the bimerite condition of the exopod and endopodof legone, the apparently bifid ventral stylet, and the four semicircular flaps wrapping aroundthe lateral margin of the elongated terminal segment of theendopod of leg one.Eudactylina urolophi sp. nov.(Figures 58-59)Material examined. Several females from the branchial lamellae of the roundstingray Urolophus halleri Cooper collected from Los Angeles Harbor and Sel Beach,California. Female holotype (USNM 266566) and 1 female paratype (USNM 266567)92deposited at the United StatesNational Museum of NaturalHistory.Etymology: The specific name urolophirefers to the generic name of the host.DescriptionFemale (Figure 58A)Overall length in lateral view approximately2.3 mm. Cephalothorax longer thanwide, lateral margin notched accommodatinglacertus of second maxillae. Dorsolateralsurface of cephalothorax covered with cuticularflaps. Cuticular flaps present on surfaces of first, second, third and fourth freethoracic somites. Fourthfree thoracic somitebearing leg five. Fourth free thoracicsomite, genital segment, and abdomen bearingposteriorly directed cuticular flapson ventral surface. Genital segment smaller thanpreceeding somite. Abdomen two-segmented.Caudal ramus (Figure 58B) longer thanwide, bearing two stout setae, onedorsomedial slender seta (not illustrated), andonelateral slender seta; ventral surfacearmed with posteriorly directed triangular cuticularflaps.First antenna (Figure 58C) apparently four-segmented,armature (proximal todistal) as follows: first segment bearing one smallslender seta; second segment withonly four slender setae observed(certainly more exist but were missed) plus onecurving, denticulated (prehensile) claw;third segment bearing three small setae andoneelongate slender seta (again, more are certainto exist but were missed) plus one stoutseta and one large denticulated spine, terminalsegment with only 10 slender setaeobserved plus one aesthete. Second antenna (Figure58D) five-segmented, prehensile. Basal segment short, secondsegment armed with spinous process and singlesubrectangular cuticular flap, thirdsegment bearing proximally directed subrectangularcuticular flaps with two slender setae arising frombase of spinous process. Fourth segment elongate and unarmed; fifth segment anelongate, unciform terminal claw bearingtwo slender setae and one elongate auxiliaryspine. Mouth tube siphonostome andsimilar to that of other species. Mandible (Figure58E) of two parts, dentiferous marginwith six teeth. First maxilla (Figure58F) biramous; endopod bearing two naked setae,exopod longer surmounted by two small nakedsetae and one longer naked seta.Second maxilla (Figure 58G) brachiform,lacertus larger than brachium, brachium withrectangular cuticular flaps, a distal patch of triangular flaps (prickles)and two tufts ofsetae (one coarse and one fine) at base of terminal claw (calamus). Claw bearingserrated membrane distally and anotherstrip along concave margin. Maxilliped (Figure58H) chelate, indistinctly segmented, proximalsegment pedunculate; corpus maxillipedis robust bearing small stoutseta on distal margin, and five large rectangular cuticu93lar flaps; myxaf area expanded into largereceptacle to accommodate clawof opposable segment. Shaft bearing two setae: onelarge naked seta near midpoint onlateralconvex margin and the other adjacent toelongate membranous strip along concavedistal margin. Claw with subquadrangular lateralshield.First leg biramous and bimerite, legs two threeand four biramous and trimerite,two-segmented sympods. All basipodswith lateral slender seta; first basipod bearsadditional distomedial seta (not shown).Ventral surfaces of all four legs bearing triangular and rectangular shaped cuticular flaps.Armature of rami as follows:Leg one Exopod 1-0 - 1111Endopod 0-0 - IILeg two Exopod 1-01-0 III Endopod 0-0 0-0 IILeg three Exopod 1-0 1-0 IIIEndopod 0-0 0-0 ILeg four Exopod 1-0 1-0 Ill Endopod0-0 0-0 ILeg one (Figure 59A) exopodal setae laterally bearing serrated cuticularexpansions (membranes), longest seta on terminalsegment slender and naked, endopodwith elongated terminal segment, setae slenderand naked. Leg two exopod (Figure59B) modified, bearing minute cuticular flapsalong lateral margin and small triangularflaps along medial edge, first segment greatlyenlarged, second and third segmentssmaller, terminal segment with truncate seta, oneshort slender seta, and one recurvingseta. Leg two endopod (Figure 59B) with relatively large lateral protuberanceon proximal segment, terminal segment apically bears two naked slender setae.Exopods oflegs three and four (Figure 59C) with unilaterally denticulated setae,seta on proximalsegment of endopods with serrated membrane. Leg five (Figure 59D) longerthan wide,distally bearing three slender setae plus oneseta arising dorsally from the base; lateralsurface with rectangular and triangular cuticular flaps.Male: UnknownComments: E. urolophi is the first eudactylinid to be reported from the genusUrolophus. Only three species possess both a two-segmented endopod and two-segmented exopod on the first leg. These species are E. epaktoIampter E. corrugata, andE. turgipes. Of these three species, both E. epaktolampter and E. turgipes possesscaudal rami bearing six setae, E. urolophi bears only four setae on the caudal ramus.This simple combination sets E. urolophi apart from the remaining species in thegenus.94Eudactylina vaquetillaesp. nov(Figures 60-61)Material examined. Several femalesbranchial lamellae of the vaquetilla,Mobulatarapacana (Philippi, 1892)collected from Punta Arena de Ia Ventana inthe .southernSea of Cortez (Gulf of California).Female holotype (USNM 266531) and5 femaleparatypes (USNM 266532) depositedat the United States National Museum ofNaturalHistory.Etymology: The specific namevaquetillae refers to the local vernacularname ascribedto this very large mobulid.DescriptionFemale (Figure 60A)Overall length in lateral view approximately3.3 mm. Cephalothorax longer thanwide, lateral margin notchedaccomodating lacertus of second maxillae.Lateral anddorsal surfaces of cephalothoraxcovered with cuticular flaps. Cuticle of first, second,and third free thoracic somitescovered with cuticular flaps. Fourth free thoracicsomitesmaller than previous threewith cuticular flaps on ventral surface and bearing leg five.Genital segment smaller than preceedingsomite,with cuticular flaps on dorsal.and ventral surfaces. Abdomen two-segmented,ventral surface bearing cuticular flaps. Caudalramus (Figure 60B) ellipsoid, bearingfour terminal short stout setae (only three illustrated), one dorsomedial slender seta and one ventrolateralslender seta; ventral surfacecovered with variously directed semicircular cuticularflaps.First antenna (Figure 60C) indistinctly five-segmented,armature (proximal to distal) as follows: first segment withcuticular flaps along lateral margin bearing one tinystout seta; second segment with eight naked setaeplus one large, curving (prehensile)claw; third segment witha proximal cluster of six naked setae (only four illustrated)plus a distal cluster of three nakedsetae and a stout spine; fourth segment bearingone slender seta, terminal segmentwith 14 slender setae plus one aesthete. Secondantenna (Figure 61A) five-segmented, prehensile.Basal segment short, second segment elongate bearing many smallsemicircular cuticular flaps, third segment bearingproximally directed rectangular cuticularflaps, two slender setae arising from near baseof very reduced spinous (possiblyabsent) process. Fourth segment elongate withmany semicircular cuticular flaps,fifth segment an elongate unciform terminal clawbearing two slender setaeand one similar in size, slender auxiliary spine. Mouth tube95siphonostome and similar to that of otherspecies. Mandible (Figure60D) of one part,dentiferous margin with six teeth.First maxilla (Figure 60D) biramous;sympod withsmall patches of spinules, endopod bearingtwo stout setae, one unilaterallydenticulated; exopod longer surmounted by two smallnaked setae and one longerbilaterallydenticulated seta. Second maxilla (Figure 60E) brachiform,lacertus larger than brachium, with small triangular and largersemicircular cuticular flaps, brachium withsemicircular cuticular flaps, a distal patch ofprickles a tiny naked seta and a singletuft of finesetae at base of terminal claw(calamus). Distal region of claw bearing two rows ofdenticles, proximal region with three pairsof serrated membranes. Maxilliped (Figure60F) chelate, indistinctly segmented,proximal segment pedunculate; corpus maxillipedis robust bearing smallstout spine on distal margin and small triangular cuticularflaps; myxal area expanded into large receptacleto accommodate claw of opposablesegment. Shaft bearing two setae: one near midpoint onlateral convex margin and theother adjacent to a strip of membrane alongconcave distal margin. Claw (Figure 60Fdetail) unciform with reduced subquadrangular cuticularexpansion producing a verysmall lateral shield.First four pairs of legs biramous with three-segmentedendopods, three-segmented exopods and two-segmented sympods. Allbasipods with short lateral slenderseta; first basipod bears additional distomedialsshort slender seta. Ventral surfaces ofall four legs bearing triangular and semicircular cuticularflaps. Armature of rami as follows:Leg one Exopod 1-0 1-0 III Endopod0-0 0-0 IILeg two Exopod 1-0 1-0 IllEndopod 0-0 0-0 IILeg three Exopod 1-0 1-0 III Endopod0-0 0-0 ILeg four Exopod 1-0 1-0 III Endopod0-0 0-0 ILeg one (Figure 61 B) exopod reduced; exopodalsetae small, stout and denticulated, endopodal setae unilaterally denticulated with elongatetriangular cuticular flapsnear bases. Exopod two (Figure 61 C)elongate; first segment long with triangular cuticular flaps along lateral margin and one enlarged, distalpapilliform seta, second segment smaller with papilliform seta, third segmentarmed with three slightly more elongated papilliform setae. Leg two endopod (Figure 61C) with two unilaterally denticulated setae on terminal segment and three elongate, triangularcuticular flaps near bases.Legs three and four (Figure 61 D);setae short and denticulated on terminal segments.Terminal segments of exopod of legfour (Figure 61E) and endopod of leg four (Figure61F) not as elongate as leg three withsmoother setae on exopod of leg four. Leg five96(Figure 61 G) elongate, distally bearing three slendersetae plus one seta arising dorsally from the base; lateral surface with semicircular andtriangular cuticular flaps. Leg six(Figure 61 H) represented by three short stout spines nearoviducal opening.Male: UnknownComments: Predictably, Eudactyilna vaquetillaeshares many characteristicswith other mobu lid-infecting species, Eudactylinadiabolophila and Eudactylina oliveri.The long legs, giving these species an ant-likeappearance, the reduced lateral shieldon the claw of the maxilliped, the relatively reducedexopods and elongated éndopodsof legs one and two, the relatively straight five- or six-segmentedfirst antenna with relatively reduced auxiliary spines on the thirdor fourth segment, the relatively elongatedcaudal rami bearing six setae, the two rows of thickdenticles found on the distal half ofthe second maxilla, the transverse palm of themaxilliped coupled to the short extensionof the myxa resulting in a reduced aperture, separatethese species from their remaining congeners. Undoubtedly, close phylogenetic relationshipsexist among species ofthis complex.Eudactylina vaquetillae is distinguished from other species of this complex andall other congeners by the globose, mammiform setaelocated on the exopod of leg two,the unusual and irregular orientation of the very largesemicircular cuticular flaps on thecaudal rami and the very elongate leg five.REMAINING UNOBTAINABLE NOMINAL SPECIESAs previously stated, Eudactylina now consists of 38 nominal species. Five ofthem, (E. carchariaeglauci Hesse, 1884, E. mustelilaevis Hesse, 1884, E.sqautinaeangeli Hesse, 1884, E. puriensis Tripathi, 1956, and E. mobuli Hameedet al, 1990)have not been sufficiently described to be recognized and are considered speciesinquirendae. Yamaguti (1963) includes E. versicolor Wilson, 1913 in the genus.Strangely enough, the only eudactylinid featured in Wilson (1913) is Nemesis versicolor. Thus, E. versicolor is presumably a lapsus calamidue to Yamaguti’s transcriptionalerror. E. rachelae and E. complexa have been relegatedto synonomy (see Kabata,1979), as has E. doilfusi and E. spinhfera herein, bringing the number ofaccepted taxawithin the genus to 30.For the sake of completeness and cohesiveness for future reference, this section97is a brief (alphabetical) accountof the females of the uncertainspecies above and theremaining species that werenot available for this revisionof Eudactylina.Eudactylina alata Pillal, 1968was described from Rhynchobatussp. fromnear Kerala, India. This species isrelatively well described andexhibits many unique ifnot bizarre characteristics. The first freethoracic somite immediatelyposterior to thecephalothorax is greatly enlarged withlateral aliform expansions overlappingthe majority of the followingsomite. The second segmentof the second antenna bears threestout spines or processes, andthe prehensile claw of thefirst antenna appears to bebifid.Eudactylina bicornis Hameedet al, 1990 was described from the smooth hammerhead, Sphyrna zygaena(Linnaeus,1 758) near Trivandrum, India.The illustrationsare recognizeable as Eudactylina,but the fine detail neededfor specific identification islacking.Eudactylina breviabdomina Pearse,1952 was originally reported from theblack-tip shark, Carcharhinus limbatus(Valenciennes, 1839) fromthe Gulf of Mexico nearTexas. I examined the old,colorless, flattened specimen preservedon a slide andcould not use it for the current purposes.The original description also lacks sufficientdetail.Eudactylina brevicauda Hameedet al, 1990 described from gills ofRhynchobatus djiddensis (Forsskal, 1775)from near Cape Comorin, India. Despite thespecific name bestowed upon it, thisspecies stands out amongst its congenersdue tothe presence of very longposteriorly directed extensions of the cephalothorax.Illustrations are insufficient for comparisonof systematically important fine detail.Eudactylina chelata Hameedet al, 1990 was collected from gills of thewhitecheek shark, Carcharhinus dussumieri(Valenciennes, 1 839). The authors claimthis species is unique within thegenus due to the modification of the terminal segmentof the endopod into a chela onlegs three and four. Again, the ilustrations make it difficult to distinguish the exact nature oftheir claim, however it appears there is a largecurving spine on that segment.Eudactylina carchariaeglauci Hesse,1883 was reported from the blue shark,Prionace glauca (Linnaeus, 1758) fromthe French Atlantic. Inadequate description relegates this to a species inquirendum(see Kabata, 1979).Eudactylina lancifera Pillai, 1968 was reportedfrom gills of the sawfish, Pristissp. and Rhynchobatus sp. from Kerala,India. This species possesses a long medianinterpodal stylet extending fromthe basipod of leg three, hence the specific name. Italso exhibits what appearsto be two instead of the typical single seta near the midpoint of the shaft on the maxillipedand along the distal medial margin of the myxal area98of the maxilliped. These latter twotraits are unique within the genus if they trulydoexist.Eudactylina minuta T. Scott, 1904 was recentlyredescribed by Kabata (1979)from the gills of the stingray Dasyatis pastinaca(Linnaeus). The fine illustrations allowthe researcher to easily identify this species andas always keen attention is paid to thefine detail. Because I was not ableto obtain specimens of this parasite, this specieswas not included in thisanalysis. This species has also been reported from therough-tail stingray Dasyatis centroura (Mitchill),and from Dasyatis pastinaca (Linnaeus,1758), both from Tunisian waters (Mediterrranean)(Essafi and Raibaut, 1977).Eudactylina mobuli Hameed et al, 1990 wasdescribed from MobuladiabolusSmith, 1943 (=Mobula kuhlii (Valenciennes in Mullerand Henle, 1841)). Although theillustrations appear unrealistic, the elongated legsunmistakeably alert one to the similarity of this species to the other mobulid-infectingeudactylinids. The two-segmentedmodified exopod of leg two is very atypical forthis genus. Unfortunately, the fine detailis lacking and since correct identification of mobulidsis not an easy task, this specieswas not included in the analyses.Eudactylina musteilaevis Hesse, 1884 wasconsidered a species inquirendumby Kabata (1979) due to insufficient description. This parasite was originallydescribedfrom gills of the smooth-hound shark, Muste/usIaevis Linck, 1790 (=Mustelus mustelus(Linnaeus, 1758)). One wonders if subsequent records of E. insolens(resdescribedherein) from this host and from the blackspotted smooth-hound,Mustelus mediterraneus (Quignard and Capape, 1972) (=Musteluspunctulatus Risso, 1826) from thesame geographic locality represent the same parasitespecies.Eudactyilna parva Castro and Baeza, 1991 was recently described fromthe gills of the skate Sympterygia brevicaudataCope, 1877 from Antofagasta, Chile.This appears to be a rather non-descript species. Itdoes possess what appears to bea four-segmented first antenna, two large claw-likesetae at the distal margin of the thirdsegmet of the first antenna, a medium length spiniform process on both second andthird segment of the second antenna, four rows of serrated membranes along the distalportion of the claw of the second maxilla, a subquadrangular lateral shield on the clawof the maxilliped, denticulated setae of exopods three and four, setae on the first exopod bearing lateral serrated membranes, and a truncate medial seta on the terminalsegment of the modified second exopod. All these characteristics should place thisspecies in with the other species of Eudactylina infecting batoids (see following cladisticanalyses).Eudactylina puriensis Tripathi, 1962 was described from gills ofRhynchobatus djiddensis from Pun, India. This description allows oneto recognize the99organism as Eudactylina but little else.The general habitusdoes resemble E. lanciferathat Pillal (1 968) described from thesame host genus. Interestingly,male E. puriensispossess two posteriorly-directed cephalothoracicextensions. These same attributesarealso shown to be shared by the male ofE. alata described by Pillai(1968), again fromthe same genus of host. It would appearthis confused and insufficientdescriptionshould relegate this taxonto a species inquirendum.Eudactylina rhinobati wasdescribed by Raibaut and Essafi (1979) from gillsofthe shovenose guitarfish,Rhinobatos rhinobatos (L., 1758) and RhinobatoscemiculusGeoffrey, 1817 both from southern Tunisia.The description suggests this species isclosely related to theother rhinobatid-infecting eudactylinidsdescribed herein; allspecies show similarities in structuresof the caudal rami and in the two-segmentedendopods on some if not all thoracic legs.Eudactylina spinula Pearse, 1950 parasitizes thesand devil, Squat/na dumerilLesueur, 1 81 8. This speciescould not be used in this analysis due to itbeing curled upand mounted on a slide. Kabata (1979)mentions E. spinula is unmistakably differentthan E. acuta which parasitizessquatinids on the European side of the Atlantic.Eudactylina squatinaeangeli Hesse, 1883 existsas a species inquirenda due tothe insufficient description originallyprovided by Hesse (1883) (see Kabata, 1979).This parasite was originally reportedfrom the angelshark, Squatina angelus Blainville,1816 (=Squatina squat/na (Linnaeus,1758). One wonders if this does not representEudactyilna acuta, a common parasite of this host,found along European and northern African continental shelves.Eudactylina valei Nunes-Ruivo, 1956 was reportedfrom the dusky smoothhound, Mustelus can/s (Mitchill, 1815) fromAngola, and from the spiny dogfish,Squalus acanthias Linnaeus, 1758, from southwestAfrica.Eudactylina vile/al Nunes-Ruivo, 1956 was originally found onthe little gulpershark, Centrophorus uyato (Rafinesque, 1810),and on the longnose spurdog, Squalusfernandinus Molina, 1782 (=Squalus b/a/n v/I/el(Risso, 1826)) both members of theSqualea (see Shirai, 1992b), from off the coast of Angola.Essafi and Raibaut (1977)collected this species from the blackmouth catshark, Pristiurusmelanostomus (=Ga/eusmelastomus Rafinesque, 1810) from the westernMediterranean, and off the coast ofFrance. The latter two species of Eudactylina,though both adequately described wereleft out of the analysis to maintainthe standardized interpretations generated by a single illustrator/author.100PHYLOGENETIC ANALYSISCLADOGRAM CONSTRUCTIONA phylogenetic analysiswas conducted for the 28 species of Eudactylinarevised herein. Deets and Ho (1988) generateda cladogram (Figure 62) and host-summary cladogram (63) of the Eudactylinidae.The outgroup used herein was composedof the supposed sister taxonEudactylinodes and at times members of their siSter dadenamely, Eudactylinella,Carnifossorius, and Eudactyilnopsis (Figure 62; see Deets andHo, 1988 for details). 75 characters(see Appendix A for data matrix and definition ofcharacters) were analyzed using the heuristic Tree-Bisection andReconnection (TBR)algorithm from the phylogenetic computer programPAUP (Phylogenetic Analysis UsingParsimony), version 3.OS (developed by Dr.David Swofford, Museum of NaturalHistory, Smithsonian Institution, Washington,D.C.). The most parsimonious tree forEudactylina (Figure 64) had a tree length of 239,a consistency index of 0.77 (maximum value = 1.00), a retention indexof 0.88 (maximum value = 1.00), and an F- ratioof 0.0589 (maximum value=0). Dueto the large number of characters, it became problematical mapping them back on the tree, therefore change and/orsynapomorphy listsmay be obtained from the author.The tree posits that one species E. aspera is the sister taxonto the remainingspecies in the genus. The next node separatesthe tree into two major lineages, onedade is composed of the E. diabolophila - E. insolens group, whilethe other dade iscomposed of the remaining 18 species.Since heuristic methods do not guarantee finding the most parsimonious tree(see Swofford, 1991, for a discussion on global versus local optima), ninespecies ofEudactylina were then selected in order to employ the Branch andBound algorithmwhich is capable of identifying all most parsimonioustrees in PAUP. This analysis waspursued as a check or test of the general phylogeneticframework revealed from theinitial heuristic TBR method. Hence, the selection of these nine speciesrequired thissubset to span across the cladogram, and to preferably represent parasites of majorsupraspecific host taxa (i.e., Triakidae, Carcharhinidae, squalids, squatinids, pristiophorids, and batoids in order to generate a general parasite-derived phylogeny forthese systematically more inclusive host taxa.The 9 species chosen to fulfill these systematic needs were: E. acanthii, E. chilensis, E. epaktoiampter, E. acuta, E. pristiophori,E. myliobatis, E. insolens, E.push/a and, E. pollex.101A phylogenetic analysis was conducted forthe 9 aforementioned species ofEudactylina. 55 characters (see Appendix B fordata matrix and definition of characters) were analyzed using the Branch and Bound algorithm.The most parsimonioustree for the subset of Eudacty//na on supraspecifichost taxa (Figure 65) hada treelength of 102, a consistency index of 0.88 (maximumvalue = 1.00), a retention index of0.88 (maximum value = 1.00), and an F-ratio of 0.0642 (maximum value=0). Again,change and/or synapomorphy lists maybe obtained from the author if the reader wishes to map character state changes back on tothe tree.The resultant cladogram is composed of two major lineages.The smaller dadecomposed of three species reveals E. insolensto be the sister taxon to the clde composed of E. pus//Ia and E. pollex. The otherdade composed of the remaining sixspecies places E. acanthll as the sister taxonto the remaining species which sortthemselves out into two distinct clades. The first groupis composed of E. chilensis andE. epaktolampter The remaining dade appearsquite closely related and derived relative to the other species is this analysis. Thisgroup is composed of E. acuta, which isthe sister taxon to the remaining two species,E. pristiophori and E. myliobatidos.PARASITE-DERIVED HOST CLADOG RAMSIn an attempt to reconstruct the history of this association betweenspecies ofEudactylina and their elasmobranch hosts and to infer possible phylogenetic relationships of the host taxa, the parasite cladogram was recodedby the additive binary coding technique (O’Grady and Deets,1987), and a host by parasite data matrix was created (Table I). Treating the parasites as characters and the phylogeny of thespecies as acharacter state tree was formally introduced by Brooks (1981). Accordingly, the most-parsimonious reconstruction from the data describes the evolution of the host-parasiteassociation.The phylogenetic analysis conducted from the recoded species of Eudactylinaby host matrix (Table I) resulted in a single most-parsimonious tree (Figure 66). TheTBR generated tree had a tree length of 55, anda consistency index with a maximumvalue of 1.00.The parasite-derived host cladogram postulates an unresolved group composedof Chiloscyllium, Sphyrna, Rhizoprionodon, and two Carcharhinus species (all sharingE. aspera) to be the sister group to the rest of the tree. The next node separates thetree into two major clades. The smaller complex enveloped by Manta birostris and102Mustelus asterias, is composed ofcarcharhiniforms and a closely relatedgroup of highlyderived batoids. Specifically, an unresolveddade composed of the generaGaleorhinus and Mustelus both members of the Triakidaeexist as the sister group to adade containing one lineage housing the generaCarcharhinus and Sphyrna, andanother dade with the tiger shark,Galeocerdo cuvier which in turn is the sister taxonto a group of highly derived epipelagic tropicalrays, the rhinopterids and mobulids.Although at first this appearsa very unlikely phylogeny for the hosts in question,a partial dismantling of the treesuggests otherwise. First, if one can apply a naivehypothesis to this chronicled pattern (see O’Hara, 1992 foran interesting discussion onthe use and misuse of evolutionarychronicle and narrative), two distinct (arid believable) patterns emerge. If a horizontaltransmission or colonization event occurred withthe emergence of the rhinopterids inthe lower Eocene (Eocene=35-55 MY) (Maisey,1984) from some carcharhinid parasitizingeudactylinid, as the cladogram suggests,then we are left with twoseparate or independent phylogenetic hypotheses(a carcharhinid dade and a batoid dade) to inspect. The carcharhiniddade now shows thetriakid dade composed of Galeorhinus andMustelus spp., as the sister group to thedade which now posits Galeocerdo cuvieras the sister taxon to the dade housing anunresolved polytomy of Carcharhinus plumbeus,C. acronotus, and C. obscurus whichin turn is the sister group to the Sphyrna (hammerheads)dade with these nested relationships (Sphyrna tiburo (Sphyrna lewini, Sphyrna mokarran)). These relationshipsbetween the Triakidae, Galeocerdo, Sphyrnidae, and Carcharhinidae are congruentwith those hypotheses presented by Maisey (1984b) and most recentlyby Compagno(1988). The relationships of the three sphyrnids revealedby their parasitic copepodsshows the same pattern as the dladogram of Sphyrnidae offeredby Compagno (1988)in his monumental treatise on the Carcharhiniformes. The colonized batoid dade positsthat the two species of Rhinoptera (cownose rays)is the sister group to the dade composed of Manta which itself is the sister group to the (Mobula tarapacana (Mobulajapanica, Mobula thurstoni)) dade. Again, these relationships are congruent with thephylogenetic relationships of this closely related subset of rays set foreward by Nishida(1990), except the parasites show more resolution between the three species of Mobulaand the single species of Manta. Nishida (1990) is left with an unresolved polytomybetween the taxa used in his analysis: Manta birostris, Mobula japan/ca, Mobulalucasana (=Mobula thurstoni), and Mobula diabolus (=M. kuhili), which is often amisidentified M. thurstoni (cf. Notarbartolo-di-Sciara, 1987).The remaining dade contains taxa spanning from Squalus acanthias toUrolophus hal/en. Fascinatingly, this portion of the parasite-generated host phylogenyproposes the monophyly of the squaloids, squatinids, pristiophorids and batoids. This103corroborates the revolutionary findings of Shirai(1992a, b) in which he proposeda radically new systematic framework for squaloidsand related taxa, the Squalea andHypnosqualea.The basal dade of this portion of the parasitegenerated host phylogeny is represented by the spiny dogfish, Squalus acanthias(Squalidae), the next branch containsthe lantern sharks (Etmopteridae). Thenext node up the tree mirrors the relationshipsof the supraspecific taxa comprisingShirai’s Hypnosqualea, which contains threespecies of angel sharks, Squatina, a singlesspecies of sawshark, Pristiophorus, and22 species of batoids or Rajiformes,hierarchically arranged as follows: (Squatina(Pristiophorus, Rajiformes)).The batoid dade is characterized by a majordichotomy. The first dade is composed primarily of members from the Rajoideiplus two species of electric rays,Torpedo. This rajoid dade contains onecompletely unresolved polytomy of eightspecies of Raja and two species of Torpedo. Since,these ten species of rays all possess the same parasite Eudactylina s/mills, anyadditional Eudactylina-derived hostresolution is impossible. This complex is the sister groupto a small group composed oftwo species of Rhinobatos which is the sistergroup to two species of Raja. It is surprising to see some species of Raja morerelated to Rhinobatos than to the other speciesof Raja. This incongruence maybe due to the author’s inability to find and/or discriminate more characters or character states in orderto resolve these inconsistencies. It isalso possible, as has been realizedby others that some minor incongruencies (such asthe aforementioned paraphyletic parasitederived host relationships of Raja versus themonophyletic relationships which supposedly distinguisha taxon) in host and associatecladograms might be more apparent than real. Recent advances (softwarenot available to the author at this point in time) comparing “hostand associate” trees (i.e. genetrees and species trees, host treesand parasite trees and, organism trees and areas)generate a single hypothesis called a reconciled tree. This hypothesis whichmaximizesthe amount of codivergence (shared history) between the associated,represents thecombined host and associate tree and makes explicit the cost ofa strict cospeciationhypothesis (Page, 1994). Future studies involving these rival methods with these datashould provide added insight regarding the historical association of this host - parasitesystem.The remaining and most apical dade of the cladogram is strictly composed ofmyliobatoids. The first subclade contains 4 species of Myliobatis. The other subcladecontains 3 myliobatoid genera that are hierarchically arranged as follows:(Urolophus(Dasyatis (Gymnura))).The phylogenetic analysis conducted from the recoded subset of nine species of104Eudactylina by host matrix (Table II) resultedin a single most-parsimonious tree (Figure67). The Branch and Bound generated tree had a tree length of 17and a consistencyindex with a maximum value of 1 .00 This cladogramsupports the relationships of thesupraspecific host taxa revealed in the initialanalysis containing all 28 recoded speciesrevised herein. Additionally, with the serendipitousremoval of the supposed colonization (horizontal transfer) event of the rhinopterid- mobulid dade, the putative relationships of the four carcharhiniform taxa used in thisanalysis break down into two distinctclades. One dade contains Galeorhinusgaleus and various species of Mustelus assister groups, both members of the Triakidae.The other dade contains Galeocerdocuvier and Sphyrna mokarran, two of the largest carcharhinoids. Theremainder of thetree once again shows the very interestingmonophyly (the Squalea of Shirai, 1992a, b)of the squaloids, squatinids, pristiophorids and,batoids. Again, Squalus acanthiasbranches off first becoming the sister taxon to theremainder of the squalean dade.The next dade composed of shark-like squalidscontains Etmopterus pusillus andAculeola nigra both members of the bioluminescent Etmopteridae. The relationships ofthe remaining three taxa confirm the relationships uncovered by the heuristic searchusing the TBR algorithm. This dade is composed ofSquat/na squat/na, the sister taxonto the final dade containing Pristiophorus and Myliobatis, further corroborating thevalidity of Shirai’s (1 992a, b) Hypnosqualea.HISTORICAL BIOGEOGRAPHYHistorical biogeography is completely reliant on phylogenetically accurate systematics. Describing the distributional relationships of a taxon is useless, unless itsmembers constitute a monophyletic group (Futuyma, 1986). Cladistics can provideoriginal hypotheses on the history of the continents by means of vicariance biogeography. This rests on the postulate that allopatric speciation is due to the origin of naturalbarriers (e.g. the breakup of Pangaea), so that the phylogeny of the monophyleticgroups under study is a reflection of the geological history of the areas they occupy.Vicariance biogeography offers the advantage of hypotheses about the history of areasthat can be independently tested with geological data, without noise from intervening,narrative, ad hoc hypotheses. Thus, it can suggest to geologists inconsistenciesbetween the reported biogeographical patterns and plate tectonic models (Janvier,1984).105Fortunately, two monophyletic groupsof hosts the angeisharks, Squatinaandthe bat rays, Myliobat/s appearto be parasitized by monophyletic subsetsofEudactylina (Figure 66).The 3 species of Squatina, are held togetheras a monophyletic unit by the parasites Eudactyl/na tuber/fera, Eudactyl/na aphiloxenous, andEudactyl/na acuta. Whatdoes the parasite-derived host and area summary-cladogram(Figure 68) tell us aboutthe possible evolutionaryhistory of this subset of angelsharks, and land masses?First, the summary-cladogram positsthat Squat/na armata (refer to discussion sectionof Eudactyilna aph/loxenous forcompeting ideas regarding the synonymy of S. armataand S. californica) is the sistertaxon to the dade housing S. californica and S. squatina.Nothing in the literature is available for comparisonregarding those specific relationships. However, the cladogramsuggests two possible vicariant events, representedbythe nodes in the cladogram, were responsiblefor the differentiation of ancestral stockinto these three descendants. Fossilsrecognizeable as Squatina have been found inJurassic period deposits (Maisey, 1984), pushing their divergencetime as far back as140-1 80 MY during the Mesozoic era. Assuming anintimate association was alreadyestablished between Eudactylina and Squat/na is interestingand I believe quite likely.Phylogenetically congruent host - parasite patterns foundbetween holocephalans andelasmobranchs with the Kroyeriidae (Deets, 1987) mayreflect a very ancient copepodchondrichthyan association and co-divergence event. This possiblyoccurred in the lateto post Devonian with the emergence of the Chondrichthyesapproximately 400 MY. Inany case, the first node separates the eastern South Pacific(S. armata) from the eastern North Pacific (S. californica) pluseastern North Atlantic and Mediterranean (S.squat/na). This apparent Gondwanaland - Laurasia distribution is likelythe result of theTethys Seaway separating North and South Americain the late Jurassic appràximately160 MY (Haq, 1984). Another concurrent mid-Jurassicevent, the opening of the NorthAtlantic began approximately 160-1 65 MY (Windley, 1984).Perhaps it is this latterevent that is represented by the node uniting the eastern NorthPacific and easternNorth Atlantic plus the Mediterranean. Thus,a possible explanation for the speciationpatterns of this subset of Squat/na interpreted from the summary cladogram would positan initial separation of eastern Pacific ancestralstock into southern and northern cornponets with the opening of Central America. The Tethys Seaway must have allowedsome of the northern population to migrate far enough Eastto eventually find themselves rifted away with western Europe during the evolution ofthe North Atlantic.Future collections of Eudactylina from the remaining nine or tenspecies of Squat/nafound worldwide, would be a fascinating study andtest of the aforementioned scenario.The 4 species of Myl/obatis, are held together asa monophyletic unit by the par-106asites Eudactylina h ornbosteli,Eudactyilna indivisa, Eudactylina myliobatidos, andEudactylina nykterimyzon. Whatdoes the parasite-derived host and area summarycladogram (Figure 69) tell us about the possibleevolutionary history of thissubset ofbat rays, and their associated coastal land masses? Thesummary-cladogram positsthat Myliobatis sp. is the sister taxon to the dadehousing M. californicus which is inturn the sister taxon to M.chilensis and M. peruvianus. The literature is devoid ofdataregarding those specificmyliobatid relationships. However, the cladogram posits threepossible speciation events representedby the nodes in the dladogram, responsible forthe differentiation of ancestral stock into these fourdescendants. The earliest fossilsrecognizeable as Myliobatis are from Tertiary deposits(Maisey, 1984b), postulatingtheir latest time of emergence occurred during the Eoceneepoch 35-55 MY. This veryrecent emergence would place all the continentsnearly in their present position (seeWindley, 1984; Figure 11 .3 a-h), and would makea vicariant interpretation of the relationships of the myliobatid hosts and their respectiveassociated land masses impossible. Is their any way to infer a much earlier emergencetime, independent of the relationships revealed by the parasite-derived summary-cladogram? I believeso. IfNishida’s (1990) Dasyaticloida (Urolophidae plus Dasyatididae) is the sister taxonto hisMyliobatidoidea (Gym nuridae plus Myliobatididae), then these groups sharea commonnode or a common point of divergence in space and time. Although,paleontologicaldata is lacking for the Gymnuridae (butterfly rays), there is a record of a few lateCretaceous teeth which have been referredto dasyatids (Case, 1978). Additionally,there appears to have been a rapid diversification of stingrays towards the end of theCretaceous; the group may primitively be represented by the extinct Cyclobatis (upperCretaceous. Lebanon; Capetta, 1980) which approaches recent dasyatid myliobatoidsbut still maintains uncertain systematic status (Maisey, 1 984b). Hence, it seems likelythe Dasyatidoidea and Myliobatidoidea diverged in the Cretaceous, andMyliobatis ispossibly much more ancient than what the current paleontological data base hasuncovered. Working with these assumptions, telling the tree with an accomodating tectonic history is now possible. The first branch leads to an unidentified sp of Myliobatisliving in the Indian Ocean (Mozambique Channel) off Madagascar. This node may represent the incipience of the South Atlantic with the separation of South America andAfrica during the mid- Jurassic approximately 160 MY. Some reconstructions suggestMadagascar might have been located more southerly than its current position, therebybeing in relatively close proximity to the tip of South America. Additionally, the southerntip of South America may have been slightly tucked around the southern tip of southernAfrica (see Windley, 1984). Perhaps continued separation of these two continents andconcurrent expansion of the South Atlantic carried away and separated those parts of a107formerly contiguous population of Myliobatisin that southern region. Thenext majorevent represented by the following nodeon the cladogram appears to once again represent the separation of the northernand southern hemispheres (eastern North Pacificand eastern South Pacific) dueto the opening of Central America by the TethyanSeaway approximately 160 MY. The finalnode separates two eastern South Pacificcountries, Chile and Peru from one another, sharingMyliobatis chilensis but withMyliobatis peruvianus exclusive to Chile. This finalnode is difficult to assess withoutany obvious potential past vicariant events orany strong geological landmarks to referto. Note that E. mdivisa and E. myliobatidos maybe synonymous. Although small differences were found between the two parasitespecies it is possible a larger samplemay find synonymy a better choice inthe future. The author prudently leaves them withtheir own taxonomic integrity in this revision,but wonders how unique their two myliobatid hosts actually are.108KRO YERIAEXTERNAL MORPHOLOGYGENERAL HABITUSThe body of Kroyeria is typically elongate and cylindrical with six distincttagmata (Figure 70 and 71A): the cephalothorax (consisting of the segments bearingthe firstantenna, second antenna, mandible, first maxilla, secondmaxilla, maxilliped, and thefirst pedigerous somite bearing leg one), the well developedpedigerous somites two,three, and four (bearing biramous, trimerite legs),a greatly elongated tube-like genitalcomplex housing the gonads, and bearing the reduced fifthleg, and a multi-segmentedabdomen (posteriorly giving rise to the caudal rami).The cephalothorax comprises the somites bearing the principal appendages ofattachment for Kroyeria. The main appendages of attachment are the remarkablechelate second antenna, the slender subchelate maxilliped, andthe dagger-like dorsalstylets arising dorsolaterally along the posterior margin of the cephalothorax, from within the posterior sinuses. Additionally, large paired interpodal stylets arisingfrom swimming legs two, three, and four (stylets usually reduced on leg one) (Figures 72A-72D)undoubtably function as brakes with regards to the upstream orientation of this parasiteon carcharhiniform gills (see Benz and Dupre, 1987).The major articulation of the body exists between the third free thoracic somiteand the genital complex.The genital complex of the female bears the reduced fifth leg (leg six is neverfound), and the genital complex of the male bears reduced legs five and six. Theabdomen of the male is composed of three segments, while the abdomen of thefemale exists in various states of indistinctness ranging from one to three segments.Again, this seems to indicate that the “missing” abdominal segments are not beingincorporated into the genital complex but are being suppressed or becoming less distinct due to vague segmental boundary areas (sutures less scieritized?).Many of the previous functional and anatomical comments mentioned in the section regarding the appendages of Eudactylina apply to Kroyeria and need not berepeated in the following discourse.109CAUDAL RAMUSThe caudal rami (Figure 71 C) originate terminallyfrom the posteriormostabdominal somite. They are rather generalized in their appearance. Thesix setaearising from the distal regionare similarly rather generalized in their structure, thoughall homologous elementshave the ability to exhibit the slender pinnate state toa nakedstout state with intermediate characterstates exhibited by different species. The flatlamelliform shape of the ramus might allowit to function as a rudder as suggested byKabata and Hewitt (1971), for the Caligidae.Additionally, the more derived claw-likesetae when present on certain species of theupstream oriented Kroyeria, may functionto brake the parasite from slipping back and off the gillfilament.DORSAL AND INTER PODAL STYLETSThe dorsal stylet (Figures 70, 71A and 71B) isa novel structure in parasiticcopepods. Other parasitic copepods possess posteriorly-directedcephalothoracicexpansions such as in female Eudactylina brevicaudaHameed et al, 1990, and maleEudactylina alata Plllai, 1968. In the monotypic Jusheyus Deetsand Benz, 1987 dorsal spines arise from a thick, dorsal scierotizedbar along the posterior margin of thecephalothorax. However, Kroyeria is the only genus in which thesestyliform processes(arising from within the posterior sinuses of thecephalothorax) are articulated. Thearticulation of this structure appears to be dueto a complex ball and socket joint.Observations on living specimens reveal that the stylets have the abilityto rotate freelyin all directions. Undoubtably,these structures are wedged into the secondary lamellae of the host’s gills securing the parasite in itsupstream orientation (Figure 70). It hasbeen found that for Kroyeria carchariaeglauciHesse, 1879, on the blue shark, Prionaceglauca (Linnaeus, 1758), 80 percent of individuals were attached to the secondarylamellae and the remaining 20 percent werefound in the excurrent water channelsclinging to the interbranchialsepta (Benz and Dupre, 1987).Dorsal stylets differ between the species. They may occur as sweeping,elongate, lissome stylets as in Kroyeria sphyrnae Rangnekar, 1957,as stout, bluntprocesses as in Kroyeria dispar Wilson, 1932, as slightly curved spines vitha flangedterminus as in Kroyeria branchioetes sp. nov., or possessing a deeply incised bifid tipas seen in Kroyeria longicauda Cressey,1970. Hence, they are helpful characters foridentification.With the exception of Lewis’ (1 966b) discussion onthe possibility of the interpo110dal stylets giving rise to the sternal furca ofDissonidae, Trebiidae and Caligidae,theorigins and homologies of dorsal and interpodalstylets have never been critically discussed. During this revision, an interesting event recurred witheach dissection of theavailable species in this genus, which may shed someinsight on the homology of theseunique structures. Whenever themaxillipeds were carefully dissected from thecephalothorax, the dorsal stylets becamedisjoined from the cephalothorax and wereremoved in conjunction with the maxillipeds. Thisstructural complex of maxillipeds,scierites, and dorsal stylets is illustrated in Figure 83A. Themaxillipeds are united by aheavily sclerotized interpodal bar. Dorsally, the base of the maxillipedsappear to articulate with a complex scleritized ringdorsolaterally giving rise to the articulated dorsalstylets.Maxillipeds are appendages of the first or anteriormost thoracic somitethat havebeen incorporated into the cephalosome thus becominga cephalothorax during theprocess of cephalization (see Huys and Boxshall 1991, Kabata, 1979, andSchram1986). Crustaceans are metameric organisms(see Barnes, 1984), therefore differentstructures having the same segmental origin are saidto be serially homologous. Thus,the maxilliped is serially homologous not only to the remaining thoracic appendagesbutalso to the cephalic appendages, for all evolved fromoriginally similar segmentalappendages. Interestingly, all non-reduced thoracic appendages (maxillipeds and legs)possess paired scleritized styliform processes.Not unlike the external cuticular structures such as setae, appendages, processes, and the like, other cuticular structures such as sclerites or scierotized bars or rodshave serial homologues. Hence, we should be ableto use this type of information intracing morphological homologies.The thoracic legs are connected by an interpodal bar which gives rise to thepaired interpodal stylets (Figures 82B, 83A, and 83B). Although the interpodal styletsare not articulated, there are some soft, joint-like areas at the junction of the leg and theinterpodal bar. The interpodal bar anteriorly butts up against a large sclerite ring notassociated with an appendage. Anterior to this is a small gap before the pattern repeatsitself with legs united by an interpodal bar giving rise to paired interpodal stylets. Thisbar anteriorly butting up against a large non-appendage associated sclerite ring and soforth. Upon inspecting the complex of sclerites associated with the maxilliped (Figure83B), one can see the interpodal bar of the first leg with its typically reduced interpodalstylets anteriorly butting up against a non-appendage associated ring (as with the legsmentioned above), albeit not complete. This is followed anteriorly by a large gap, andthen another interpodal bar uniting the maxilipeds. This serially repeating pattern ofsclerites and appendages suggests that the interpodal stylets and the dorsal stylets111found in Kroyeria are one and the same. This is aneconomical explanation forthe origination of these novel structures. One must onlyhypothesize a single evolutionary eventgiving rise to paired, styliform, interpodal thoracicprocesses for their presence to beexplained on all thoracic appendages on non-reducedthoracomeres (thoracic legs fiveand six are often absent).Perhaps a homeotic mutation, known in arthropods, whichcauses all or part of a segment to develop ina manner inappropriate to itself but inapproximate conformity to the normal developmentof some other segment (see Arthur,1984), could explain the manifestation of an interpodalstylet associated with a maxilliped (the dorsal stylet).FIRST ANTENNAThe first antenna of Kroyeria (Figure 71 D) is indistinctly seven- oreight-segmented. Assuming similar (homologous) innervation exists hereas seen in other siphonostomes (see Kabata, 1979), this appendage haschemosensory and tactile functions.The apical segment bears 13 setae (12 terminal, 1sub-terminal) and one aesthete. Dueto differential fusion of segments amongst species within the genus,the details regarding specific armature on each segment will be covered in taxonomic descriptions.SECOND ANTENNAThe principle attachment organ for Kroyeria is the extraordinary four-segmented,chelate second antenna (Figure 71 E). The heavily scleritized, indistinctly divided firstand second segments form a base allowing great freedom of movement (Kabata, 1979,Deets, unpublished observations of living specimens). The third segment forms the corpus of the chela, its distal end greatly produced and terminally expanded formsa receptacle to receive tip of the terminal segment or claw. The receptacle can be quite expansive when possessing membranous extensions of the cuticle as in Kroyeria longicaudaCressey, 1970 (Figure 93E), or simply a small indentation barely large enough toaccommodate the tip of the opposable claw as in Kroyeria dispar Wilson, 1935 (Figure85F). Likewise, the tip of the claw as in Kroyeria longicauda Cressey,1970 (Figure93E) may possess membranous cuticular expansions giving it a cup-like appearancesimilar to the receptacle which typically houses it, alternatively, the tip mayexist smoothand unornamented as in Kroyeria dispar Wilson, 1935 (Figure 85F). Additionally, theproximal area of the claw is typically armed with a pair of spines as in Kroyeria cresseyiDeets, sp.nov. (Figure 79F), ora set of three spines as in Kroyeria longicauda Cressey,1970 (Figure 93E). Different combinations andstates of these characters make the112second antenna a valuable taxonomicand phylogenetic discriminant.MANDIBLE AND ORALCONEThe mandible, witha dentiferous distal end(Figure 74G), uniramous, subcylindrical, is typically siphonostome.It appears to be oftwo parts. Their boundaries demarcated by a proximal suture.The dentiferous marginbears from seven to ten teeth, withsome species exhibiting teeth uniformin size (Figure 74G)and other species with different-sized teeth (Figure93F), reminiscent ofthe primary and secondary teeth intheLernaeopodidae/Sphyriidae/Tanypleuridaecomplex.The oral cone (Figures81 H and 891) consisting ofanterior labrum and posteriorlabium house the mandibles.The labrum typicallybears distolateral patches of pricklesand terminal membranes,the labium is equipped withtwo rows of prickles along the lateral surface and a terminalmembrane. This structure beingquite uniform throughoutthe genus was not utilized fortaxonomic purposes.FIRST MAXILLAThe first maxilla (Figure71G) located adjacent to the oral cone andmandible is abiramous appendage composedof a relatively elongateendopod and smaller exopod.Both the exopod and endopodare tipped with two setae. Setae mayoccur as naked orpinnate depending onthe species.SECOND MAXILLAThe second maxilla of Kroyeriais a large brachiform appendage (Figure71 H),consisting proximally ofa heavily scleritized lacertus and distally ofa robust brachium.The orifice of the maxillarygland is present near the base of the lacertusas is the previously undescribed (probably unnoticed)basal process. The basal process in somespecies may approach half thelength of the second maxilla. The brachiumis typicallyarmed with two contiguous patchesof densely packed prickles. Distalto these patchesat the base of the claw isa tuft of long fine setae. The most terminal componentis theclaw which may or maynot represent a third segment. It isa robust, curving structure,typically armed alongthe lateral surfaces with lamelliformmembranes and a pricklymembrane along the convexsurface. No specific function for thesecond maxitla ofKroyeria has yet been observed.113MAXILLIPEDThe maxilliped (Figure 71 I) is a subchelatestructure. The corpus appears to betwo-segmented. The subchela is not dividedinto a proximal shaft and distalclaw as inmany siphonostomes. A minute slenderseta is present near the distal end of the subchela. A membranous flangeis present along the distal margin of the corpus in somespecies. Additionally, a seriesof three transverse cuticular flanges occur on the corpusof Kroyeria dispar Wilson, 1935 (Figure 86A)and Kroyeria longicauda Cressey, 1970(Figure 931).LEG ONEThe first thoracic leg (Figure 72A) pair is connectedby an interpodal bar. As previously mentioned, interpodal stylets arise from theinterpodal bar. Interpoda! stylets ofthe first thoracic leg are always reduced in relationto those of thoracic legs two, three,and four. The sympod is composed of the proximalcoxa and distal basis. The coxausually bears two strips of membrane,although in a few species only one or none wereobserved. The basis similarly bears two strips of membrane,one lateral seta, and onedistomedial seta. The first leg is always biramous, composedof a lateral exopod andmedial endopod. Both the exopod and endopodare trimerite throughout the genus.The first (proximal) segment of the exopodbears a row of setules along themedial margin, a distomedial pinnateseta, a distolateral seta, and a smooth and / orpectinate membrane along the lateral edge. The second segmentpossesses the samecharacteristics except the distolateral seta is absent in some species.The third segment shares similar characteristicsto the first two segments and bears one small lateralmost seta, one elongate slender seta occurring in various states(see detailedspecies descriptions), and four long pinnate setae.The first endopodal segment bears a single distomedial pinnateseta, a distolateral membrane, and a row of setules along thelateral margin.The second endopodal segment in most species hasa row of setules along thelateral edge and in some speciesa series of triangular denticulations connected by awebbing of membrane. Under both light microscopyand scanning electron microscopy(unpublished data and Oldewage unpublisheddata), these denticulations or teeth foundon some endopodal segments appearto be thickened, fused, or fortified regions of themembrane that typically run along that lateral margin. Under close examination, themembranous webbing running along these teeth appearto fuse with the teeth orendopodal denticulations. Hence, both the membrane and the teeth or endopodal den114ticulations appear to be one in the same.Additionally, in cases where these endopodaldenticulations or teeth are absent, thetypical membrane is present. Typically thissegment is devoid of pinnate distomedial setaebut in Kroyeria dispar Wilson, 1935 fromthe tiger shark, Galeocerdo cuvier (Peron and LeSeuer,1822), two pinnatesetae arepresent.The third or terminal endopodal segment from the firstleg simUarly exhibitsmembranous denticulations or a smoothmembrane and the fringing row of setules wasnot always observed. Six long, pinnatesetae are found on this segment throughout thegenus. Very minute pectinate membranesreminiscent of those present in the Caligidae(see Kabata, 1979), are presentat the setal bases (of some leg pairs) of some species(possibly most) but are clearly seen onlywith scanning electron microscopy.LEG TWOThe second thoracic legs (Figure 72B) are unitedby an interpodal bar whichgives rise in the majority of species to well developedinterpodal stylets. The exceptionsare Kroyeria caseyl Benz and Deets,1987 on the night shark, Carcharhinus signatus(Poey, 1868), Kroyeria dispar Wilson, 1935, and Kroyeriapapillipes Wilson, 1932 bothparasitic on the tiger shark, Galeocerdo cuvier (Peronand LeSeuer, 1822). The coxaand basis are well delimited with the basis distally bearing two welldeveloped membranes, and a single lateral seta.The first segment of the second exopod typically bears a medialrow of setules,pinnate seta, a distolateral slender seta, and a lateral smooth or pectinatemembrane.The second segment is similarto the first with the lateral slender seta absent in somespecies. The third segment similarly bears a smooth or pectinate membrane along thelateral margin and seven pinnate and variously modifiedslender setae.The first segment of the endopod bears a distomedial pinnate seta,a lateralmembrane, and a lateral fringe of setules. The second segment generally bearsa lateral fringe of setules, membranous denticulations in somespecies, and the atypical presence of pinnate setae on the distomedial edge in Kroyeria dispar Wilson, 1935. The terminal segment harbors six pinnate setae.LEG THREEThe third thoracic legs (Figure 720) are connectedby an interpodal bar possessing large interpodal stylets withthe exception of K. caseyl, K. dispat and K. papillipes.The coxa is unarmed, buta lateral seta and two distal membranes are found on the115basis.The proximal segment of the exopod typicallybears a medial fringe of setules,adistomedial pinnate seta, a lateral slenderseta, and a smooth and / or pectinate membrane. The second segment issimilar to the first, with the lateral slender seta absent incertain species. The terminalsegment similar to the previous two segments bears fourelongate pinnate setaeand three variously modified slender setae.The first segment of the endopod isequipped with a lateral fringe of setules, adistolateral membrane, anda single distomedial pinnate seta. The second segmentmay possess membranousdenticulations, a medial pinnate seta, ora medial fringe ofsetules depending on the species.The third, or terminal, segment shares similar attributes with the preceding segments andgenerally bears four pinnate setae. However, inK. caseyl and K. dispar, there are threepinnate setae, one slender seta, and anotherpinnate seta for a total of five setae.LEG FOURThe fourth thoracic legs (Figure 72D) as in the previousthree are connected byan interpodal bar which gives riseto interpodal stylets which are well-developed exceptin K. caseyl, K. dispar and K.papillipes. The sympod is composed of an unarmedcoxa and a basis with a single lateralseta and two membranes along the distal edge.The first exopodal segment bears a medial fringe of setules,a single distomedial pinnate seta, a single distolateral slenderseta, and a smooth and / or pectinate membrane flanking the lateral margin. The second segmentis similar to the first with exception of the absence of both the fringingsetules on the medial edge and the distolateralslender seta in some species. The terminal segment differsfrom these two segmentsby possessing four pinnate setae plus three variously modifiedsetae.The first segment of the endopod bears a lateral membrane,a lateral fringe ofsetules, and a single distomedial pinnate seta.The second segment bears a single distomedial pinnate seta and may or may not possess lateral membranous denticulationsand fringing setules. The terminal segment generallybears two pinnate setae plus onevariously modified seta. K. disparis an exception bearing two pinnatesetae, one slender seta, and one more pinnate seta,as is K. papillipes bearing three pinnate setae.LEGS FIVE AND SIXThe fifth leg (Figure 81J) is located approximately 1/2 to 2/3 the length ofthegenital complex down the lateral side of the genital complex and is rarelyfound.116However, in species where it is known itis a uniramous appendageconsisting of foursetae. A sixth leg does not appearto exist.In the males, the fifth and sixth legs(Figure 73A) are found on the genital complex. The fifth and sixth legboth setiform are representedby four and two setae respective ly.LIFE HISTORYGENERAL DESCRIPTIONThere have been few published attempts ofculturing Kroyeria, or descriptiveaccounts of life history stages other than theadult and first nauplius.The first report of the nauplius accompanied the first descriptionof this genus byvan Beneden (1853) for Kroyeria lineata,the type species. Carli and Bruzzone (1973)were able to hatch first-stage of the nauplius60 minutes after the egg was shed fromthe egg strand of Kroyeria carchariaeglauciHesse, 1878. The most recent descriptionof the first nauplius was offeredby Benz and Deets (1986), and illustrated the threenaupliar appendages, the uniramous first antenna,the biramous second antenna, andthe biramous mandible. Also shown were thetwo dorsal ocelli of the tripartite naupliareye (the ventral ocellus was not shown), and the two well developedfiliform balancerson the posterior end.It would not be surprising if Kroyeria exhibiteda holoxenous life-cycle similar tothat demonstrated by Cabral, et al (1984) inLernanthropus kroyeri, from the percichthyid (temperate bass) Dicentrarchus labrax(see Raibaut, 1985 for a general life history review of different parasitic copepod taxa).REPRODUCTIONCopulation and reproduction probably occurs ina manner similar to that inEudactylina, but no reports or observations have been published.I have not seen members of this genus in the presumed copulatoryembrace.117HOST-PARASITE RELATIONSHIPSDELETERIOUS EFFECTS / FEEDINGAs previously stated, Kroyeria attach themselvesto the secondary lamellae ofthe gill primarily with their clasping chelate secondantenna, and secondarily with theirarticulate posterolaterally oriented dorsal stylets and the ventroposteriorlydirected interpodal stylets (Figure 70). Benz and Dupre (1987), have shownthat 80 percent of all K.carchariaeglauci Hesse, 1879, attach themselves in anupstream-oriented fashion tothe secondary lamellae on the gills of its favoredhost, the blue shark, Prionace glauca(Linnaeus, 1758). The remaining 20 percent were found clipped onto the soft tissue inthe underlying excurrent water channels. The dorsal stylets andinterpodal stylets withtheir supposed braking function probably allowKroyeria increased mobility in its environment by not having to rely entirely on the second antennae for itssecurity. It is speculated that the second antennae may be used to reachout and align itself upstreamand possibly crawl with them, assuming the stylets have the ability to hold the parasitein place, as suggested by Figure 70.I have not witnessed gill tissue pathologies induced by Kroyeria inthe carcharhinform hosts I have inspected over the years. During microscopic examination,many specimens of Kroyeria still had tissue from the secondary lamellaegrasped intheir chelate second antennae. Hence, some damage is occuringto the respiratory surfaces of the host. In situ, the copepods are reddish, suggesting these parasites feed atleast in part on blood. Blood would be easily accessible from the secondary lamellaesince respiratory blood sinuses are only one epithelial cell thick inthis region (Benz,1984). With up to 1,250 individuals per hostas reported for K. carchariaeglauci onPrionace glauca (Benz and Dupre, 1987) some physiological effects on the host sharkwould be expected.The specific feeding mechanics have yet to be observed for any gill dwellingcopepod, but morphological evidence (from scanning electron microscopy, Oldewage,unpublished) suggests that it is similar to that of caligids as described by Kabata (1974,1979).A dentiferous ridge, the strigil, appears to be present along the inner edge of thelabium (Oldewage, unpublished SEM micrographs,). The strigil possibly saws away atthe epithelial cells, while the mandibles equipped with their dentiferous margins, maycontinue the maceration process and then convey the host tissue into the buccal cavity.Patches of prickles along the distolateral region of the labrum and the two rows of spin118ules flanking the lateral surfaces of the labiummay anchor the oral coneto the feedingsite.SPECIFICITYAlthough the literature suggestsa few species such as K. ilneata and K. carchariaeglauci have the abilityto infect a broad range of hosts, my personal collectionsover the last 1 3 years, suggest thathost specificity is the generalrule for the species inthis genus. One problem thatundoubtably beleaguers this host-parasite systemis themorphological conservatismthat both Kroyeria and its hosts, theCarcharhiniformesexhibit. Hence, host and/orparasite may have been misidentifiedand the .apparentbroad host range of these species maybe artifactual.With the exception of the mesoparasiticKroyeria caseyl Benz and Deets, 1986,which embeds approximately80% of its body into the interbranchial septa of itshostCarcharhinus signatus (Poey, 1868), Kroyeriatypically attach by the chelate secondantenna to the gills (secondary lamellae).Microniche specificity and the functional morphologicalrequirements associatedwith it has only recently beenexamined in parasitic copepods of elasmobranchs(seeBenz 1980, 1992; Benz andAdamson, 1990; Benz and Dupre, 1987). Reports ofmicroniche have typically not beengiven in sufficient detail (e.g., skin, gill, or branchiallamellae) to assess the ecological or morphologicalsignificance of the association.When collecting eudactylinids or kroyeriids from branchiallamellae, I havenoticed that genera that possess fully chelateattachment organs, such as the secondantenna of Kroyeria or the maxilliped ofEudactylina or Eudactylinodes, secure themselves primarily to the secondary lamellae of theirhost. On the other hand, Nemesis,attaches itself by surrounding the efferentbranchial arterioles near the gill filaments’free distal tips with its largesubchelate maxillipeds (Benz and Adamson, 1990), amicrofiche quite unlike that occupied by its fully chelate allies.SYSTEMATIC ACCOUNTGenus Kroyeria van Beneden, 1853Kroyeriidae: Female. Cephalothorax covered by well demarcated dorsal shieldDorsal stylets arising from posteriorsinuses of cephalothorax. Three pedigerous segments between cephalothorax and genital complex.Genital complex composed of pedi119gerous somites five and six, togetherconstituting more than 50% of length ofbody.Abdomen one- to three-segmented.Caudal ramus lamelliform bearing six setae distally.First antenna indistinctly seven-or eight-segmented, terminal segment bearingone aesthete and thirteen slendersetae. Second antenna four-segmented and chelate;third segment distally produced into a receptacleto accommodate claw of terminalsegment. Mouth tube siphonostome. Mandible oftwo parts, distal end dentiferous bearingfrom seven to ten teeth. First maxillabiramous with both endopod and exopod bearingtwo apical setae. Second maxilla brachiform, two-possibly three-segmented (includingclaw). Maxilliped subchelate,subchela not divided into shaft and claw. Legs onethrough four biramous and trimerite.Leg five when found represented by four setae.Male: Similar to female; abdomen three-segmented.Legs five and six setiformand represented by four and two setaerespectively.TYPE-SPECIES: Kroyeria lineata van Beneden,1853COMMENTS: Previous to this account, the literaturecontained references to 19nominal species (see Kabata, 1979 for review).An additional description of anunnamed male by Capart (1953) is devoid ofillustrations making correct identificationdifficult. The description suggests affinity with K.papillipes, and this possibility is furthersuggested by the fact that Capart’s material came from thetiger shark, Galeocerdocuvier(Peron and LeSueur, 1822), the principal hostto K. papillipes. Three species (K.acanthiasvulgaris Hesse, 1879, K. galeivulgaris Hesse,1884, and K. scyliicaniculaeHesse, 1879) have been insufficiently describedto be recognized and are treated asspecies inquirendae. K. trecai Delamare Deboutteville andNunes-Ruivo, 1953 wasnever fully described and remainsa nomen nudum. K. aculeata (Gerstaecker, 1854)and K. sublineata Yamaguti and Yamasu, 1959have been relegated to junior synonymsof K. lineata van Beneden, 1853. This revisionresurrects K. elongata Pillai, 1967 fromsynonymy, and treats it as a valid species. Five species newto science, K. branchiocetes, K. cresseyi, K. decepta, K. procerobscena, and K. rhophemophaga aredescribed herein.Kroyeria consists of the 16 species illustrated and phylogenetically analyzedherein, plus two unobtainable species for thiscurrent effort bringing the total number ofaccepted species to 18. As in the previous section, the unobtained specieswill bereviewed at the end of the following taxonomicaccount.The systematics of Kroyeria has suffered in the past from insufficient attentiontomorphological detail that can beused as specific discriminants (Kabata, 1979).Additionally, this genus is superficiallymorphologically conservative, and also difficult to120distinguish.The major host taxon for Kroyeria, is theCarcharhiniformes (see Raibaut, 1982).The latest revision of this host group was by Compagno(1988). He estimates thatthislargest group of living sharks comprises nearly60 percent or 200 species of the approximately 350 known shark species. This information coupledto the apparently highdegree of host fidelity exhibited by Kroyeria,suggests (as is true of Eudactylina) thereare many species yet to bediscovered in the genus.Kroyeria branchiocetessp. nov(Figures 71-73)Material examined. Several females (on loanfrom Dr. Z. Kabata, Pacific BiologicalStation, Nanaimo, Canada) collectedby Dr. Paperna, University of Israel, from the gillsof the grey reef shark, Carcharhinusamblyrhynchos (Bleeker, 1856) from the Red Sea.Female holotype (USNM 266533) and3 female paratypes (USNM 266534) depositedat the United States National Museum of Natural History.Etymology: The specific name branchiocetes is derivedfrom the greek Branch/a,meaning gill and oecetes an inhabitant, referringto the typical gill-dwelling nature ofthis rather attractive, albeit nondescript species.DescriptionFemale (Figure 71A)Overall length in dorsal view approximately 3.8 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures 71 A and 71 B) extending posteriorlyto nearly 30 % down the length of the thirdfree thoracic somite, stylets curving slightly inward and distolaterally bearinga flange-like cuticular expansion. Three free thoracic somites withoverlapping terga. Genitalcomplex cylindrical, constituting 60 % of totalbody length. Posterolateral corners of latter bearing oviducal openings. Abdomen indistinctly three-segmented. Caudalramus(Figure 71C) lamelliform, longer than wide with medial fringe of setules, distally bearingfour pinnate and two semipinnatesetae.First antenna (Figure 71 D) indistinctly eight-segmented, armature (base to apex)as follows: 10, 1, 5, 2, 3, 1 1, 1 2+1 aesthete. Second antenna (Figure 71 E) chelateand prehensile, apparently four-segmented.Proximal two segments heavily sclerotized121in such a way as to suggest relativelyunrestricted movementcapabilities. Third segment forming corpus of chela, extending intoa rigid arm distallyexpanded into a membranous receptacle to accommodate tip of fourthsegment. Latter forming heavily scierotized robust claw, bearing two prominent setae proximallyand similarly expanded intoa membranous receptacle distally. Mandible (Figure71 F) of two parts (only distal portion illustrated), dentiferousmargin with nine teeth, tiny apical tooth followedby onelarge, two small, two large, and threesmall. First maxilla (Figure 71G) biramous; endopod longer bearing two apical elongate,naked setae; exopod shorter bearing twoshort, naked setae. Second maxilla(Figure 71H) brachiform; lacertus heavily sclerotized with elongate basal process arising fromnear base. Brachium with two largepatches of prickles and a tuft of fine, longsetae near base of claw; claw bearing lateralmembranous lamellae with small pricklesscattered upon convex surface. Maxilliped(Figure 711) subchelate; corpus two-segmented;subchela not divided into shaft andclaw, distally uncinate and bearinga single small, slender seta.All four legs biramous and trimerite. Sympodstwo-segmented. All basipods withlateral pinnate seta and one or two distomedialmembranes: first basipod bears additional distomedial pinnate seta; first coxopod bearsadditional distolateral membrane.All four interpodal bars bearing interpodal stylets;interpodal stylets of leg one verysmall. Lateral fringe of setules oneach endopodal segment, medial fringe of setules oneach of first exopodal segment. Armatureof rami as follows (Arabic numerals denotefully pinnate setae, Roman numerals conditionsdiverging from that state):Leg one Exopod 1-1 0-1 11,4 Endopod 0-1 0-0 6Leg two Exopod I-i 1-1 111,4Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4 Endopod 0-1 0-0 4Leg four Exopod I-i 1-1 111,4 Endopod 0-1 0-1 1,2Exopod of leg one (Figure 72A) bearing distolateral membrane onfirst (proximal)segment; pectinate lateral membrane on segmentstwo and three. Segment three withfour pinnate setae, one elongate slender setabearing a lateral membrane, and onesmall naked seta. Endopod of leg one (Figure72A) with distolateral membrane on firstsegment; segment two with four to five (only four shown) endopodal denticulations;segment three with four to six (only four shown)endopodal denticulations. Exopod ofleg two (Figure 72B) similar to leg one, exceptsecond segment bears additional seta(bilaterally bearing membranes), and third segment withfour pinnate setae, one semipinnate seta with setules along the medial edge anda membrane along the lateral, oneslender seta bearing a membrane alongthe lateral edge, and one seta bearing mem122branes on both lateral and medial edges.Endopod of leg two similar to leg one, exceptfive to six endopodal denticulations (fiveshown) are present on the second segment;five endopodal denticulations on the thirdsegment. Exopod of leg three (Figure 72C)as in leg two, except seta adjacent to lateralmostseta bilaterally bears membranes(unilaterally in leg two). Endopod of leg three(Figure 71C) with six and five endopodaldenticulations on segments two and three respectively;segment three with four pinnatesetae. Exopod of leg four (Figure 72D) as in leg three.Endopod of leg four (Figure 72D)with seven and five endopodal denticulationson segments two and three respectively;segment three with two pinnate setae and onebilaterally bearing serrated membranes.Fifth leg (not shown) representedby four setae.Male: (Figure 73A)Overall length in lateral view approximately2.8 mm. Cephalothoracicappendages and swimming legs similar to those of female.Genital complex bearingfifth and sixth legs representedby four and two setae respectively. Dorsal stylet (Figure73B) shorter and more stout than that of female witha hyaline flange along the proximalmedial margin. Caudal ramus (Figure73C) more elongate than that of female, setulesfringing medial margin; six distal setae (twosemipinnate, four pinnate).Comments: Kroyeria branchiocetes is the first kroyerid reported from the greyreef shark, Carcharhinus amblyrhynchos(Bleeker, 1856). This non-descript parasiteresembles K. cresseyl, K. ilneata, K. rhophemophagus, and K.triakisae, in possessingonly two slender setae on the claw of thesecond antenna. It can be distinguished fromthese species by the presence of endopodal denticulationson segments two and threeof all four swimming legs. Interestingly, all four ofthe aforementioned parasites lackingthe complete set of endopodal denticulations and possessingonly two (not the morecommon condition of three) slender setae onthe claw of the second antennae arefound on various hosts of the family Triakidae. Kroyeria branchiocetes isfound onCarcharhinus amblyrhynchos, a member of Carcharhinidaeas opposed to Triakidae.Kroyeria carchariaeglauci Hesse, 1879(Figures 74-76)Syn: Kroyeria gracilis Wilson, 1932,(see Delamare-Deboutteville and Nunes Ruivo)123Material examined. Several males and femalescollected by George Benz, TennesseeAquarium, Chatanooga, Tennessee, U.S.A., fromthe gills of the blue shark, Prionaceglauca (Linnaeus, 1758) from the western NorthAtlantic; numerous males and femalescollected from the same host species from the southernCalifornia bight near theChannel Islands and from the southern Sea of Cortez (Gulf ofCalifornia) near IslaCerralvo; a few specimens were collected from the silky shark, Carcharhinusfalciformis(Bibron, 1839) near Punta Arena de Ia Ventanaand Isla Cerralvo in the southern Sea ofCortez; a few specimens collected from the pelagicwhite tip shark, Carcharhinus longimanus (Poey, 1861) near the RevillagigedosIslands in the tropical eastern NorthPacific, Mexico.DescriptionIllustrated specimen from blue shark, Prionace glauca (Linnaeus, 1758).Female (Figure 74A, B)Overall length in dorsal view approximately5.5 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures 74A-C) extending posteriorly to approximately60 % down the length of thesecond free thoracic somite, stylets curving slightly inward and distallybifurcating.Three free thoracic somites with overlapping terga. Genital complex cylindrical,constituting approximately 65 % of total body length. Posterolateral cornersof latter bearingoviducal openings; egg strand bearing 44 eggs. Abdomen indistinctlythree-segmented.Caudal ramus (Figure 74D) lamelliform, longer than wide with medial fringe of setules,distally bearing six pinnate setae (one short, one short and pyriform,and four elongate).First antenna (Figure 74E) indistinctly seven- or eight-segmented, armature(base to apex) as follows: 11 (only nine shown), 5, 2, 3, 1, 1, 13 +1 aesthete. Secondantenna (Figure 74F) chelate and prehensile, apparently four-segmented. Proximal twosegments heavily scierotized in such a way as to suggest relatively unrestricted movement capabilities. Third segment forming corpus of chela, extending into a rigid armdistally expanded into a receptacle to accommodate tip of fourth segment. Latter forming heavily sclerotized robust claw, bearing threeprominent slender setae proximally.Mandible (Figure 74G) of two parts, dentiferous margin with nine teeth. First maxiMa(Figure 74H) biramous; endopod longer bearing two apical elongate, naked setae; exopod shorter bearing two short, naked setae. Second maxiMa (Figure 741) brachiform;lacertus heavily scierotized with elongate basal process arising from near base.Brachium with two large patches of prickles and a tuft of fine, long setae near base ofclaw; claw bearing pairedlateral membranous lamellae with small prickles scatteredupon convex surface. Maxilliped (Figure 75A) subchelate; corpus two-segmented,124proximal segment bearing two conicalprocesses, and proximal end ofadjacent segment bearing single small, conicalprocess; subchela not divided into shaftand claw,distally uncinate and bearing a single small, slenderseta.All four legs biramous and trimerite.Sympods two-segmented. All basipods withlateral pinnate seta and one or twodistomedial membranes; first basipod bears additional distomedial pinnate seta; firstcoxopod bear additional distolateral membranes.All four interpodal bars bearing interpodalstylets; interpodal stylets of leg one relativelysmall. Lateral fringe of setules on eachendopodal segment, medial fringe of setules oneach of first exopodal segment.Armature of rami as follows (Arabic numeralsdenotefully pinnate setae, Roman numeralsdenote conditions diverging from that state):Leg one Exopod 1-10-1 11,4 Endopod 0-1 0-0 6Leg two Exopod I-i 1-1111,4 Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4Endopod 0-1 0-0 4Leg four Exopod I-i 1-1 111,4Endopod 0-1 0-1 1,2Exopod of leg one (Figure 75B)bearing distolateral membranes on segmentsone, two and three. Segment three with fourpinnate setae, one elongate, slender setabearing a finely serrated, lateralmembrane, and one small naked seta. Endopod of legone (Figure 75B) with distolateral membraneon first segment; segment two with six toeight (only six shown) endopodal denticulations;segment three with six to seven (onlysix shown) endopodal denticulations(specimens collected from the pelagic white tip,Carcharhinus longimanus (Poey, 1861), bear nine). Exopod of leg two (Figure75C)similar to leg one, except secondsegment bears additional lateral, slender seta, andthird segment with four pinnatesetae, one semipinnate seta with setules along themedial edge and a membrane (smooth or finelyserrated, too small for certainty) alongthe lateral edge, one slender seta bearinga membrane along the lateral edge, and onenaked, slender seta. Endopod of leg two similarto leg one, except seven endopodaldenticulations (eight in the specimens collected from the pelagic white tip) are presenton the second segment and eightto ten (ten shown) endopodal denticulations on thethird segment. Exopod of leg three (Figure 75D) as in legtwo, except seta adjacent tolateralmost seta appears to be devoid of lateral membrane(unilaterally in leg two).Endopod of leg three (Figure 75D) with seven and ten endopodal denticulations onsegments two and three respectively (eight arepresent on both segments from specimenscollected from the pelagic white tip); segment three with four pinnatesetae. Exopod ofleg four (Figure 75E) as in leg three. Endopodof leg four (Figure 75E) with seven andnine endopodal denticulations on segments two and three respectively (specimens ccl-125lected from the pelagic white tip bear sevento eight on segment three);segment threewith two pinnate setae and one bilaterally bearingserrated membranes; bases of setaebearing small pectinate membranes (reminiscent ofthose in Caligidae). Fifth leg (seeFigures 74A and 74B) representedby four setae.Male: (Figure 76A)Overall length in lateral view approximately4.4 mm. Cephalothoracicappendages and swimming legs similar to those of female.Genital complex bearingfifth and sixth legs representedby four and two setae respectively. Dorsal stylet shorterand more stout than that of female. Caudalramus (Figure 76B) more elongate than thatof female, setules fringing medial margin;six distal setae (two semipinnate, four pinnate).Comments: Kroyeria carchariaeglauci Hesse, 1879 was originallydescribedfrom gills of the blue shark, Carcharhinus glaucus(=Prionace glauca (Linnaeus, 1758))near Brest, France in the eastern North Atlantic. It wassubsequently redescribed fromfrom the same host in the Mediterranean (DelamareDebouteville & Nunes Ruivo,1953), from Japanese waters (Shiino, 1957),from off Valaparaiso, Chile (Stuardo andFagetti, 1961), from the West coast of South Africa (Kensley and Grindley,1973), fromthe western North Atlantic and Mediterranean (Carliand Bruzzone, 1972; 1973), fromTunisian waters (Essafi and Raibaut, 1977), from the western North Atlantic (Benz,1986; Benz and Dupre, 1987), and herein from the eastern North Pacific from thesouthern Sea of Cortez, Mexico, Revillagigedos Islands, Mexico, and Channel Islands,southern California Bight.K. carchariaeglauci has also been reported from carcharhiniform hosts otherthan the blue shark. The first report of this occurred with the description of K. gracilisby Wilson (1932). He reported K. grad/is from the blue shark and brown shark,Carcharhinus milberti (Valenciennes,inMuller and Henle, 1839)(=the sandbar shark,Carcharhinus plumbeus, Nardo, 1827). Additionally, K. grad//is is considered a juniorsynonym of K. carchariaeglauci by Delamare-Deboutteville and Nunes-Ruivo (1953)and Shiino (1957). Kabata and Gusev (1966) report K. carchar/aeglauci from Eu/am/asp. from near Cocos Island, Indian Ocean but provided no illustrations. Finally, Essafiand Raibaut (1977) add questionable records of K. carchariaeg/auci inhabiting the starry smooth-hound, Mustelus aster/as Cloquet, 1821, and the smooth-hound Mustelusmustelus (Linnaeus, 1758) from France. Equally suspect, is their report of K. carchariaeg/auci from gills of the longnose spurdog, Squalus blainvillei (Risso, 1826).I have examined specimens from Dr. Roger Cressey’s personal collection from126elasmobranchs from both the IndianOcean and Florida coast,(see Cressey, 1967 and1970) and conclude that the many unillustratedrecords of K. gracilis(=K. carchariaeglauci) on hosts other than blue shark, Prionaceglauca (Linnaeus, 1758),pelagicwhite tip, Carcharhinus longimanus,(Poey, 1861), and silky shark, Carcharhinusfalciform/s (Bibron, 1839) should beviewed with caution (see following newspeciesdescriptions of K. decepta and K.procerobscena).The combined characteristicsof the bifid dorsal stylet,the relative length to widthratio of the males’ caudal rami (seeK. decepta), and thestout, pyriform, pinnate setaadjacent to the two elongate pinnatesetae of the caudal rami, readily distinguish thisspecies from its congeners.Kroyeria Casey! Benz and Deets, 1986(Figures 77-78)Material examined. Two females and one malecollected from the interbranchial septaof the night shark, Carcharhinus signatus(Poey,1 868) from the western North Atlantic.DescriptionFemale (Figure 77A-C)Overall length in dorsal view approximately60 mm. Cephalothoracic suturesarising anterolaterally and unitingposteromedially. Eyes not evident. Dorsal stylets(Figures 77A-D) very short androbust, extending posteriorly to approximately 50 %down the length of the firstfree thoracic somite, stylets curving inward slightly and terminating somewhat bluntly. Three free thoracicsomites with overlapping terga. Genitalcomplex cylindrical, inflating distally and constitutingapproximately 95 % of total bodylength. Posterolateral corners of latterbearing oviducal openings. Abdomen swollen,one-segmented bearing tiny posteroventrally directedspinules. Caudal ramus (Figure77E) lamelliform, longer than widedevoid of typical medial fringe of setules, distallybearing six stout, naked,setae.First antenna (Figure 77F) indistinctly seven- oreight-segmented, armature(base to apex) as follows: 7, 2, 5, 2,3, 1, 1, 13 +1 aesthete. Second antenna (Figure77G) chelate and prehensile, apparently four-segmented. Proximaltwo segments heavily sclerotized in such a wayas to suggest the capablity of relatively unrestricted movement. Third segment forming corpusof chela, extending into a rigid arm distallyexpanded into a receptacleto accommodate tip of fourth segment. Latter forming heavi127ly sclerotized robust claw, bearingone elongate, somewhat bluntseta near midpoint,one tiny seta along concave margin inaperture of chela, and one proximal seta.Mandible (Figure 77H) of two parts, dentiferousmargin with nine teeth. First maxilla(Figure 771) biramous; endopod longer bearingtwo apical elongate, nakedsetae; exopod shorter bearing two short, naked setae. Second maxilla(Figure 77J) brachiform;lacertus heavily sclerotized with stout basal process arisingfrom near base. Brachiumwith two large patches of prickles anda tuft of fine, long setae near base of claw;clawvery elongate, bearing paired lateralmembranous lamellae. Maxilliped (Figure 77K)subchelate; corpus two-segmented, proximalsegment bearing two conical processes,and proximal end of adjacentsegment bearing single small, conical process; subchelanot divided into shaft and claw, distally uncinateand bearing a single small, slenderseta.All four legs biramous and trimerite. Sympodstwo-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta; first coxopod bears additionalmembranes. All four interpodalbars bearing small interpodal stylets; interpodalstylets of leg one smaller than others.Lateral fringe of setules on each endopodal segment; medialfringe of setules on theproximal segment of each exopod. Armature of ramias follows (Arabic numeralsdenote fully pinnate setae, Roman numerals denoteconditions diverging from thatstate):Leg one Exopod I-i 1-1 11,4 Endopod0-1 0-1 6Leg two Exopod I-i 1-1 111,4 Endopod 0-10-1 6Leg three Exopod I-i 1-1 111,4 Endopod 0-10-1 1 ,I,3Leg four Exopod I-i 1-1 111,4 Endopod 0-10-1 1,2Exopod of leg one (Figure 78A) bearing distolateral membranes on segmentsone, two and three. Segment three with four pinnatesetae, plus two naked setae.Endopod of leg one (Figure 78A) with distolateral membrane on first segment; segmenttwo with three to eight (only three shown) endopodal denticulations;segment three withthree to six (four shown) endopodal denticulations. Exopod of legtwo (Figure 78B) similar to leg one, except third lateralmost seta is semipinnate with setules along the medial edge and a finely serrated membrane alongthe lateral edge, and two lateralmostseta bilaterally bearing smooth or finely serrated membranes(too small for certainty).Endopod of leg two similar to leg one, with threeto six endopodal denticulations (threeshown) present on the second segmentand four to six (four shown) endopodal denticulations on the third segment. Exopod of leg three (Figure 77C)as in leg two. Endopod128of leg three (Figure 78C) with three tofive and four to five endopodaldenticulations onsegments two and three respectively (threeand four are shown,respectively); segmentthree bears one medial pinnate seta followedlaterally by one stout seta, andthree elongate pinnate setae. Exopod of leg four (Figure78D) as in leg three. Endopod of leg four(Figure 78D) with three to five(three shown) endopodal denticulations on segmenttwo,and two to seven (three shown)endopodal denticulations on segment three.Fifth legnot found.Male: (Figure 78E)Overall length in dorsal view approximately3.0 mm. Cephalothoracicappendages and swimming legs similarto those of female. Genital complex bearingfifth and sixth legs representedby four and two setae, respectively. Dorsal styletslightlymore stout than that of female.Caudal ramus (Figure 78F) more elongate than that offemale, with setules fringing medial margin: sixelongate distal setae (two pinnate).Comments: Kroyeria caseyl has been reportedonly once from the night shark,Carcharhinus signatus (Poey, 1868)by Benz and Deets (1986) from the western NorthAtlantic. Kroyeria caseyi is perhapsthe most remarkable member of the entireKroyeriidae. It not only is the largest member ofthe family at approximately 60 mm, butit is also the only known memberto date of this family to be mesoparasitic, embeddedup to 80% of its body (anteriorly) into the interbranchialsepta. The extremely elongategenital complex (comprising approximately95% of the total body length) coupled to thevery elongate claw of the second maxilla, thereduced seta in the aperture of the second antenna, the 1,1 3 formula of the terminalsegment of endopod three, the inflatedone-segmented abdomen, and the very derived caudalrami devoid of the typical medialfringe of setules, and bearingstout naked setae, distinguish this species from all otherkroyeriids.Kroyeria cresseyi sp. nov.(Figures 79-80)Material examined. Several females collected from thesecondary lamellae of the leopard shark, Triakis semifasciataGirard, 1854 from inshore waters off El Segundo, SealBeach, and Palos Verdes, California,U.S.A. Female holotype (USNM 266535) and 5female paratypes (USNM 266536)deposited at the United States National Museum of129Natural History.Etymology: The specific name honors mygood friend and pioneer in the studies of theparasitic copepods infectingelasmobranchs. Dr. Roger Cressey, Curator of Crustacea,USNM, Smithsonian Institution, WashingtonD.C.DescriptionFemale (Figure 79A)Overall length in dorsal view approximately5.0 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures 79A and 79B) extending posteriorlyto approximately 50 % down the length ofthe second free thoracic somite, stylets curvinginward slightly with a bifid terminus.Three free thoracic somites with overlapping terga.Genital complex cylindrical, constituting approximately 66% of total body length. Posterolateral corners of latter bearingoviducal openings. Abdomen indistinctly three-segmented. Caudalramus (Figure 79C)lamelliform, longer than wide bearing the typicalmedial fringe of setules, distally bearing four elongate pinnate setae and two shortersemipinnate setae. Egg strands (Figure79D) containing six eggs.First antenna (Figure 79E) indistinctly seven- or eight-segmented, armature(base to apex) as follows: 11, 1,5, 2, 3, 1, 1, 13 (only 12 shown) +1aesthete. Secondantenna (Figure 79F) chelate and prehensile, apparently four-segmented.Proximal twosegments heavily sclerotized in such a wayas to suggest the capablity of relativelyunrestricted movement. Third segment forming corpus of chela, extendinginto a rigidarm distally expanded into a receptacle toaccommodate tip of fourth segment. Latterforming heavily sclerotized robust claw, bearing one elongate seta along concave margin in aperture of chela, and one proximal elongateseta. Mandible (Figure 79G) of twoparts, dentiferous margin with nine (1 apical, 2 large, 2 small, 2 large, and 2 small)teeth. First maxilla (Figure 79H) biramous;endopod longer bearing two apical elongate, naked setae; exopod shorter bearing two short, naked setae. Second maxilla(Figure 80A) brachiform; lacertus heavily sclerotized with large basal process arisingfrom near base. Brachium with two large patches of prickles and atuft of fine, longsetae (not shown) near base of claw. Claw bearing paired lateral membranous lamellae, with a unilaterally serrated membrane enveloping the latter.Maxilliped (Figure80B) subchelate; corpus two-segmented, proximal segmentbearing two conicalprocesses, and proximal end of adjacent segment bearing single small, conicalprocess; subchela not divided into shaft and claw, distally uncinate and bearinga single small slender seta.130All four legs biramous and trimerite.Sympods two-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta; first coxopod bears twoadditional membranes. All four interpodal bars bearing elongate interpodal stylets; interpodal stylets ofleg one smaller thanothers. Lateral fringe of setules on each endopodal segment,medial fringe of setuleson the proximal segment of each exopod. Armatureof rami as follows (Arabicnumeralsdenote fully pinnate setae, Romannumerals denote conditions diverging from thatstate):Leg one Exopod 1-1 0-1 11,4Endopod 0-1 0-0 6Leg two Exopod 1-1 0-1 111,4Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 11,4Endopod 0-1 0-0 4Leg four Exopod I-i 1-1 111,4 Endopod0-1 0-1 1,2Exopod of leg one (Figure 80C) bearinglateral membranes on segments one,two and three. Segment three with four pinnatesetae, plus two slender setae; longer ofthe two with membrane along lateral edge. Endopod ofleg one (Figure 80C) with distolateral membrane on first segment; segmenttwo with six endopodal denticulations; segment three with distolateral membrane, and sixpinnate setae. Exopod of leg two(Figure 80D) similar to leg one, except thirdlateral most seta is semipinnate withsetules along the medial edge and a finely serrated membrane alongthe lateral edge,the two lateralmost setae slender and apparently naked.Endopod of leg two similar toleg one, with three to six endopodal denticulations (six shown)present on the secondsegment, and a distolateral membrane and six pinnate setae on the thirdsegment.Exopod of leg three (Figure 80E) similar to leg two except secondsegment bears asmall lateral seta, and the terminal segmentbears only 2 lateral setae laterally bearingserrated membranes; longer of the two semipinnate. Endopod of leg three(Figure 80E)with three to seven endopodal denticulations onsegment two (four shown); segmentthree bears four pinnate setae and one distolateral membrane. Exopodof leg four(Figure 80F) as in leg two. Endopod of leg four (Figure 80F) with sixto thirteen (thirteenshown) endopodal denticulations on segment two; one lateral stoutseta bilaterallybearing serrated membranes, and two pinnatesetae tip segment three. Fifth leg notfound.Comments: Kroyeria cresseyl along with K. branchiocetes, K. lineata, K.rhophemophagus, and K. triakos all possess second antennae with the claw armed withonly two slender setae, asopposed to three found amongst the remaining congeners.131K. cresseyl differs most notably form K.branchiocetes and K. ilneataby not possessingthe large membranous expansions founddistally on the claw and the corpus ofthe second antennae of these two species. K. cresseyl iseasily distinguished from K. triakosby possessing only four pinnate setae (plus two slendersetae (one with serrated membrane and one semipinnate)) on the terminalsegment of exopod three, while K. triakosbears five pinnate setae(plus one naked seta). Finally, the orbicular cephalothorax ofK. cresseyl is quite dissimilarfrom the distinctly subquadrangular cephalothorax of K.rhophemophagus. Additionally, K. cresseyistands apart from this species complex inbeing the only member witha bifid dorsal stylet.As noted above, K. branchiocetes differs from thesespecies in that it occurs inthe Carcharhinidae not theTriakidae.Kroyeria decepta sp. nov(Figures 81-84)Material examined. Several males and females collectedby Dr. Roger Cressey, fromthe gills of the dusky shark, Carcharhinus obscurus (LeSueur, 1818) fromthe Westcoast of Florida; numerous males and females collected from the same host speciesfrom near the Revillagigedos Islands in the tropical northeastern Pacific,Mexico.Female holotype (USNM 266537) and 7 female paratypes (USNM 266538) depositedat the United States National Museum of Natural History.Etymology: The specific name decepta is derived from the Latin dec/plo, to deceive, inreference to the subtle morphological differences possessed by this species, relative toK. carchar/aeglauci, making specific identification difficu It.DescriptionFemale (Figure 81A, B)Overall length in dorsal view approximately 7.6 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially. Eyes not evident. Dorsal stylets(Figures 81 A, B, and 81 D) extending posteriorly to approximately 60 % down the lengthof the second free thoracic somite, stylets curving slightly inward and distally bifurcating. Three free thoracic somites with non-overlapping terga. Genital complex cylindrical,constituting approximately 66 % of total body length. Posterolateral corners of latterbearing oviducal openings. Abdomen indistinctly three-segmented. Caudal ramus132(Figure 81C) lamelliform, longer than widewith medial fringe of setules, distallybearingsix pinnate setae (two stout and fourelongate).First antenna (Figure 81 E) indistinctly seven-or eight-segmented, armature(base to apex) as follows: 10, 5, 2, 3, 1, 1, 13 +1 aesthete.Second antenna (Figure81 F) chelate and prehensile, apparently four-segmented.Proximal two segments heavilyscierotized in such a way asto suggest relatively unrestricted movement capabilities.Third segment forming corpus of chela,extending into a rigid arm distally expandedinto a receptacle to accommodate tip offourth segment. Latter forming heavily sclerotized robust claw, bearing threeprominent slender setae proximally. Mandible(Figure81 G) of two parts, dentiferousmargin with nine teeth (2 large, 2 small, 2 large,3 slightlysmaller). Labrum of oral cone (Figure81H) typical of the genus. First maxilla (Figure81 I) biramous; endopod longer bearingtwo apical elongate pinnate setae; exopodshorter bearing two short, naked setae.Second maxilla (only claw detailed) (Figure82A) brachiform; lacertus heavily scierotizedwith elongate basal process arising fromnear base. Brachium with two largepatches of prickles and a tuft of fine, long setaenear base of claw; claw bearing pairedlateral membranous lamellae with small pricklesscattered upon convex surface. Maxilliped(Figures 82B, 83A, 83B) subchelate; corpus two-segmented, proximal segment bearing twoconical processes; subchela notdivided into shaft and claw, distally uncinate and bearinga single small slender seta.[The possible homologous relationshipsbetween the interpodal and dorsal stylets’ sclerites illustrated in Figures 823, 83A, B have been discussedin the previous dorsal andinterpodal stylet section dealing with this hypothesis].All four legs biramous and trimerite. Sympodstwo-segmented. All basipods withlateral pinnate seta and one or two distomedialmembranes; first basipod bears additional distomedial pinnate seta; first coxopod bears twoadditional distolateral membranes, and a small patch of spinules. All four interpodalbars bearing interpodalstylets; interpodal stylets of leg one relatively small. Lateral fringe of setuleson eachendopodal segment, medial fringe of setules on the first exopodalsegment of .each leg.Armature of rami as follows (Arabic numeralsdenote fully pinnate setae, Romannumerals denote conditions diverging from thatstate):Leg one Exopod 1-1 0-1 11,4 Endopod 0-10-0 6Leg two Exopod I-i 1-1 111,4 Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4 Endopod 0-10-0 4Leg four Exopod I-i 1-1 111,4 Endopod 0-1 0-1 1,2Exopod of leg one (Figure 82C) bearing distolateral membranes onsegments133one, two and three (membranes pectinateon segments two and three).Segment threewith four pinnate setae, plus two lateralsetae bearing finely serrated, lateralmembranes. Endopod of leg one (Figure 820) withdistolateral membrane on first segment;segment two with eight endopodal denticulations;segment three with eightto nine (nineshown) endopodal denticulations (specimensof K. carchariaeglauci collectedfrom thepelagic white tip, Carcharhinus longimanus(Poey, 1861), similarly bear nine)..Exopodof leg two (Figure 82D) similarto leg one, except second segment bearsadditional lateral seta, and third segment withfour pinnate setae, plus one semipinnateseta withsetules along the medial edge anda membrane along the lateral edge, and two slendersetae bearing membranes along their lateraledge. Pectinate membranes are presentalong the lateral margin ofsegments two and three. A strip of pectin is also presentparalleling the typical membrane along thelateral margin if the first segment of bothexopod one and two. Endopod of leg two(Figure 82D) similar to leg one, except eightto nine (nine shown) endopodal denticulations(eight in the specimens of K. carchariaeglauci collected from the pelagic white tip)are present on the second segment andnine to ten (ten shown) endopodal denticulationson the third segment. Exopod of legthree (Figure 82E) as in leg two, except the twolateralmost setae appear to be devoidof lateral membrane (unilaterally in leg two).Endopod of leg three (Figure 82E) witheight to nine endopodal denticulations onsegments two and nine to ten endopodal denticulations on segment three (eight are present on bothsegments from specimens of K.carchariaeglauci collected from the pelagic white tip);segment three with four pinnatesetae. Exopod of leg four (Figure 82F) as in legthree. Endopod of leg four (Figure 82F)with seven to eight and eightto ten endopodal denticulations on segments two andthree respectively (specimens of K. carchariaeglaucicollected from the pelagic white tipbear seven to eight on segment three); segmentthree with two pinnate setae and onebilaterally bearing serrated membranes. Fifth leg(Figure 81J) represented by four setae(three pinnate and one naked).Male: (Figure 84A and 84B)Overall length in lateral view approximately 4.6 mm. Cephalothoracicappendages and swimming legs similarto those of female. Genital complex bearingfifth and sixth legs representedby four and two pinnate setae respectively. Dorsal styletshorter and more stout than that of female. Caudal ramus (Figure84C) more elongatethan that of female, setules fringing medial margin;six distal setae (one stout andsemipinnate, one stout and naked, two large and two small pinnate). Leg one endopod(two most distal segments illustrated) (Figure 84D)similar to female, bearIng’ four andfive endopodal denticulations on segments twoand three respectively. Endopods of134legs two, three andfour similar to leg one, all bearingfour to five endopodal denticulations on segmentstwo and three.Comments: Kroyeria deceptais very similar to K. carchariaeglauci,and thisundoubtedly has led to confusionin my mind and in the literature.Having examinedspecimens of Kroyeria fromCarcharhinus obscurus from both theWest coast of Floridaand off the RevillagegedosIslands in the tropical easternNorth Pacific, I believethisrepresents a distinctform and have treatedit herein as such. Many specificdifferences, albeit subtle, contribute tothis decision. First, thecaudal rami of male K. decepta are greatly elongated relative tothat of male K. carchariaeglauci,being approximately9.5 times the width, comparedwith about 6.5 times thewidth in male K. carchariaeglauci. The lengthof the genital complex of female K.decepta is approximately10.25 times its width, comparedwith that of C. carchariaeglauciwhich is only about 7.6times its width. The endopodof the first maxillaof K. decepta bears pinnatesetae,these setae are nakedin K. carchariaeglauci. Thelateral membranes on the secondand third segments of theexopod from leg oneare pectinate in K. decepta,while thoseof C. carchariaeglauc/are thin and smooth. The twospecies are differentin size; K.decepta is approximately7.6 mm in total length,while K. carcharaieglauciis about 5.5mm long, or about 72% thelength of the former. Thelength of the dorsalstylet of K.decepta is approximately6.5 times its width, comparedto that of K. carecharchiaeglauci, which is nearly 8.8 times its width.And finally, theteeth on the mandiblefrom K.decepta exhibit alternatingsizes; teeth are moreuniform in size in K.carchariaeglauci.K. decepta appears to beconfined to the duskyshark, Carcharhinus obscurus.Interestingly, Kabata and Gusev(1966) report K.carchar/aeglauci fromEu/am/a sp. inthe Indian Ocean. The copepodwas reported to be 6.72mm, large for that species,anda genital complex length approximately9 times thatof its width. Thesemeasurementssuggest that copepodmay be K. decepta, butwithout examining thespecimen or beingmore certain what Eulamia sp.represents, no conclusionshould be drawn.Kroyeria dispar Wilson,1932(Figures 85-86)Material examined.Several females andmales (USNM 153870)collected byDr.Roger Cressey, fromthe gills of the tiger shark,Galeocerdo cuvier(Peron & LeSueur,1822) from the West coast of Florida;numerous malesand females (USNM153864)135collected from the same host species fromthe Indian Ocean near Madagascar.DescriptionFemale (Figure 85B)Overall length in dorsal view approximately13.1 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Cephalothorax laterally extended relative to congeners.Dorsal stylets (Figures 85B and85C) extendingto just below posterior margin of first free thoracicsomite, stylets curving slightly inwardwith blunt tips. Three freethoracic somites with non-overlapping terga. Genital complexcylindrical, constitutingapproximately 65 % of total body length. Posterolateral cornersof latter bearing oviducalopenings. Abdomen indistinctly one- or two-segmented.Caudal ramus (Figure 85D) lamelliform, longerthan wide with medial fringe of setules,distally bearing two stout semipinnate setae,two long thick pinnate setae, and twosmall slender pinnate setae;two duct-like openings (see detail) are present ventrodistally.First antenna (Figure 85E) indistinctlynine-segmented, armature (base to apex)as follows: 9, 1, 1 5, 2, 3, 1, 1, 13 +1 aesthete. Secondantenna (Figure 85F) chelateand prehensile, apparently four-segmented. Proximaltwo segments heavily sclerotizedin such a way as to suggest relatively unrestrictedmovement capabilities. Third segment forming corpus of chela, extending into a rigidarm distally expanded into a smallreceptacle to accommodate tip of fourth segment.Latter forming heavily sclerotizedrobust claw, proximally bearing three prominentsetae (one seta nearly half the lengthof claw and very blunt). Aperture of chelanearly circular due to strongly curving claw.Mandible (Figure 85G) of two or three parts assuggested by divisions of the sclerites,dentiferous margin with nine teeth. First maxilla (Figure 85H) biramous;endopodlonger bearing two apical elongate setae with rowsof denticles; exopod shorter bearingtwo short, naked setae. Second maxilla (Figure 851) brachiform;lacertus heavily sclerotized with elongate basal process arising from nearbase. Brachium with two largepatches of prickles and a tuft of fine, longsetae near base of claw; the latter bearingpaired lateral membranous lamellae with small prickles scattered uponconvex surface.Maxilliped (Figure 86A) subchelate; corpus two-segmented, proximal segmentbearingtwo conical processes; myxal region of corpus withlarge protuberance bearing threecuticular expansions reminiscent of the cuticular flaps in Eudactylina; subchela notdivided into shaft and claw, distally uncinate and bearinga single minute seta.All four legs biramous and trimerite. Sympods two-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta; first coxopod bears two additional membranes. All four interpo136dal bars with poorly developed (absent?)interpodal stylets. Lateral fringe ofsetules oneach endopoda! segment, medial fringe of setules on thefirst exopodal segment ofeach leg. Armature of rami as follows (Arabic numeralsdenote fully pinnate setae,Roman numerals denote conditions divergingfrom that state):Leg one Exopod I-i 1-1 11,4Endopod 0-1 0-2 6Leg two Exopod I-i I-i 111,4Endopod 0-1 0-2 6Leg three Exopod I-i 1-1 111,4 Endopod0-1 0-1 1,1,3Leg four Exopod I-i 1-1 111,4Endopod 0-1 0-1 1,1,2Exopod of leg one (Figure 86B) bearing distolateralmembranes on segmentsone, two and three (additional pectinatestrips on segments two and three). Segmentthree with four pinnate setae, plus two lateralsetae bilaterally bearing membranes.Lateral short setae on all exopods with flagellate tips.Endopod of leg one (Figure 86B)with distolateral membranes on eachsegment. Exopod of leg two (Figure 86C) similarto leg one, except third segment with four pinnate setae, plus one semipinnateseta withsetules along the medial edge and a membrane along the lateraledge, and the two lateralmost setae bilaterally bearing membranes. Pectinate membranes paralleling thetypical membranes are present along the lateral margin ofsegments one, two, andthree. Endopod of leg two (Figure 86C) similar to leg one. Exopod of leg three (Figure86D) as in leg two, except pectinate membranes not observed. Endopodof leg three(Figure 86D) with only one pinnate seta on segment two and segment three bears (laterally to medially) one pinnate seta, one short seta bearingmembranes followed bythree pinnate setae. Exopod of leg four (Figure 86E)as in leg three. Endopod of legfour (Figure 86E) similar to leg three except terminal segment bears one pinnate setafollowed by one short seta bearing membranes, followedby only two pinnate setae.Fifth leg (not illustrated) represented by four setae (three pinnate and one naked).Male: (Figure 85A)Overall length in lateral view approximately 7.2 mm. Cephafothoracicappendages and swimming legs similar to those of female. Genital complex bearingfifth and sixth legs represented by four and two pinnate setae respectively. Dorsal styletshorter and more stout than that of female. Caudal ramus more elongate than that offemale.Comments: Kroyeria dispar was originally described by Wilson (1935) from anunnamed shark. Since then, (see Cressey, 1967, 1970) all subsequent records have137been from gills of the tiger shark, Galeocerdo cuvier(Peron & LeSueur, 1 822).The unusually wide cephalothorax, the absenceof endopodal denticulations,plus the presence of two elongate, pinnate setae arisingfrom the medial margin of legsone and two, the spinulated / denticulated endopod ofthe first maxilla, and the peculiarcuticular flaps found on the myxal area of the maxillipeddistinguish this species from allothers in the genus. This speciesshares several attributes with K. papillipesalso fromGaleocerdo cuvier: short dorsal stylets, veryreduced interpodal stylets, circular aperture of the second antenna, and an indistinctlynine-segmented first antenna.Kroyeria elongata Pillai, 1967Syn: KroyeriaspatulataPearse, 1948 of Pillal, 1985nec Kroyeria spatulata Pearse, 1948nec Kroyeria elongatus Fukui, 1965(Figures 87-88)Material examined. Several females (USNM 154002) collectedby Dr. Roger Cressey,from the gills of the spot-tail shark, Carcharhinus sorrah(Valenciennes, 1 839) from theIndian Ocean near Madagascar.DescriptionFemale (Figure 87A)Overall length in dorsal view approximately 3.0 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures 87A, C) extending into anterior quarter of free thoracic somite three, styletscurving inward with expanded distolateral portion forminga cuticular flange. Three freethoracic somites with non-overlapping terga. Genital complex cylindrical,constitutingapproximately 63 % of total body length. Posterolateral corners of latter bearing oviducal openings. Abdomen indistinctly three-segmented. Caudal ramus (Figure87B) lamelliform, longer than wide with medial fringe of setules, distally bearing twostout setae(the longer distally pinnate, the shorter semipinnate), two thick longer pinnatesetae,and two slender pinnate setae.First antenna (Figure 87D) indistinctly seven-to eight--segmented, armature(base to apex) as follows: 10, 1, 5, 2, 3, 1, 1, 13 +1 aesthete. Second antenna (Figure87E) chelate and prehensile, apparently four-segmented. Proximal twosegments heavily sclerotized in such a as wayto suggest relatively unrestricted movement capabilities.138Third segment forming corpus of chela,extending into a rigid arm distallyexpandedinto a large receptacle to accommodate tip offourth segment. Latter forming heavilyscierotized robust claw, distally bearing membranousexpansions forminga large cuticular receptacle fitting into the large receptacleof the corpus. Claw additionallybeañngthree prominent slender setae proximally (middleseta shorter and blunt). Aperture ofchela elliptical due to elongatedcorpus and claw, reminiscent of the maxillipedinEudactylina. Mandible (Figure 87F) oftwo or three parts as suggested by divisions ofthe sclerites, dentiferous margin with nineteeth (1 apical, 1 large, 2 small, 2 large, and3 small decreasing in size). First maxilla (Figure87G) biramous; endopod longer bearing two apical elongate nakedsetae; exopod shorter bearing two short, naked setae.Second maxilla (Figure 87H) brachiform;lacertus heavily sclerotized with elongatebasal process arising from near base. Brachium withtwo large patches of prickles anda tuft of fine, long setae near base of claw;the latter bearing paired lateral membranouslamellae with small prickles scattered uponconvex surface. Maxilliped (Figure 871)subchelate; corpus two-segmented, proximalsegment bearing three conical processes; subchela not divided into shaft andclaw, distally uncinate and bearing, a singlesmall slender seta.All four legs biramous and trimerite.Sympods two-segmented. All basipods withlateral pinnate seta and twodistomedial membranes; first basipod bears additional clistomedial pinnate seta; first coxopod bears two additional membranes.Interpodal barswith interpodal stylets (leg one interpodalstylet very small or absent). Lateral fringe ofsetules probably present on each endopodalsegment, though not seen on all segmentsfrom specimens inspected,medial fringe of setules on the first exopodal segment ofeach leg. Armature of ramias follows (Arabic numerals denote fully pinnate setae,Roman numerals denote conditions diverging from thatstate):Leg one Exopod 1-1 0-1 11,4 Endopod0-1 0-0 6Leg two Exopod I-i 1-1 111,4 Endopod0-1 0-0 6Leg three Exopod I-i 1-1 111,4 Endopod0-1 0-0 4Leg four Exopod I-i 1-1 111,4 Endopod0-1 0-1 1,2Exopod of leg one (Figure 88A) bearing distolateral membraneson segmentsone, two and three (membranes pectinate on segments twoand three). Segment threewith four pinnate setae, plus twolateral slender setae (the lateralmost seta naked, theadjacent seta bearing a membrane along lateraledge). Endopod of leg one (Figure88A) with distolateral membrane on first segment, five to sixendopodal denticulationson the second segment, and threeto five endopodal denticulations on the terminal seg139ment. Exopod of leg two (Figure 88B) similarto leg one, except second segment bearslateral spiniform seta, and third segment bearsfour pinnate setae, plus one semipinnate seta with setules along the medial edge anda membrane along the lateral edge,and the two lateralmost setae laterally bearing membranes.Endopod of leg two (Figure88B) similar to leg one. Exopod of leg three (Figure88C) as in leg two. Endopod of legthree (Figure 88C) similar to leg two but segment three bears onlyfour pinnate setae.Exopod of leg four (Figure 88D) as in leg three. Endopod of leg four(Figure 88D) similar to leg three except second segment mediallybears pinnate seta, and terminal segment bears one stout seta bilaterally bearing shortsetules plus two pinnate setae.Male: not acquiredComments: Kroyeria elongata was originally describedby Pillai (1967) frommilk shark, Scoliodon sorrokowah (Bleeker, 1853)(=Rhizoprionodon acutus (Ruppel,1837). PiIlai (1985) synonomized this species withK. spatulataPearse, 1948. No reaSons were given for the action, but the two speciesare quite distinct from another.K. elongata can be distinguished from its congeners easilyby the structure of theSecond antenna alone. No other Kroyeria has such anelongated claw and corpus ofthe second antenna. These attributes give the second antenna a veryelongate ellipticalaperture resulting in a general morphology that is reminiscient of that seen inthechelate maxilliped of Eudactylina. K. elongata is further distinguished from K.spatulatain that the terminus of the dorsal stylet of K.elongata is an inward curving flange, whilethat of K. spatulata is bifid (see following species description ofK. spatulata). Thecephalothorax of K. elongata is subtriangular not subquadrangularas in K. spatulata.This report adds the spot-tail shark, Carcharhinussorrah, as the second known host forthis species. Both records come from the Indian Ocean.Kroyeria gemursa Cressey, 1967(Figures 89-90)Material examined. Two paratype females (USNM 113296) collected fromthe gills ofthe great hammerhead, Sphyrna mokarran (Ruppel, 1837) from the Indian Ocean nearMadagascar; numerous females (USNM 153855) from the same host species fromnear Sarasota, Florida.140DescriptionFemale (Figure 89A)Overall length in dorsal view approximately 14.4 mm.Cephalothoracic suturesarising anterolaterally and unitingposteromedially. Eyes not evident. Ventral surfaceofrostrum (Figure 890) with twosmall spinous processes. Dorsal stylets (Figures 89A-C,E) extending near anterior quarterof free thoracic somite three, stylets relativelystraight, with a tiny bifid terminus. Threefree thoracic somites with non-overlappingterga. Genital complex cylindrical, constitutingapproximately 73 % of total body length.Posterolateral corners of latter bearing oviducalopenings. Abdomen indistinctly threesegmented. Caudal ramus (Figure 89D)lamelliform, longer than wide with medialfringe of setules along proximalhalf, bearing four pinnate setae distally (2large, twosmall), and distolaterally bearingtwo stout semipinnate setae each witha ventromedialrow of tiny denticles.First antenna (Figure 89F) indistinctlyseven-segmented, armature (base toapex) as follows: 11, 3, 1, 3, 1, 1, 1 2+1 aesthete. Second antenna (Figure 89G) chelateand prehensile, apparently four-segmented.Proximal two segments heavily sclerotizedin such a way as to suggest relativelyunrestricted movement capabilities. Third segment forming corpus of chela, extending intoa rigid arm distally expanded into a smallreceptacle to accommodate tip of fourth segment.Latter forming heavily scierotizedrobust claw, proximally bearing three setae. Apertureof chela small due to stout clawand stout pollex of corpus. Mandible (Figure89H) of two parts, dentiferous margin withnine teeth (1 apical, 1 large, 2 small,2 large, and 3 small decreasing in size). Oralcone (Figure 891) typical kroyeriidtype but with lateral patches of spinules on thelabrum much larger than those of congeners. Firstmaxilla (Figure 89J) biramous; endopod longer bearing two apical elongate, pinnatesetae; exopod shorter bearing twoshort, naked setae. Second maxilla (Figure 90A)brachiform; lacertus heavily sclerotized with elongate basal process arising from nearbase. Brachium with two largepatches of prickles and a tuft of fine,long setae near base of claw; the latter bearingpaired lateral membranous lamellaewith small prickles scattered upon convex surface.Maxilliped (Figure 90B) subchelate;corpus two-segmented, proximal segment bearingtwo conical processes, one conicalprocess on proximal part of main corpus; subchelaindistinctly divided into shaft and claw, distallyuncinate and bearing a single smallslender seta.All four legs biramous and trimerite. Sympods two-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta. Interpodalbars with interpodal stylets (leg one interpodal styletvery small and blunt). Lateralfringe of setules present on segment one of the first141endopod and on the first and second segments ofendopods two, three, and four, medial fringe of setules on the first exopodal segment ofeach leg. Armature of ramias follows (Arabic numerals denote fully pinnate setae, Roman numeralsdenote conditionsdiverging from that state):Leg one Exopod 1-1 0-1 11,4Endopod 0-1 0-0 6Leg two Exopod I-i 1-1 111,4Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4Endopod 0-1 0-0 4Leg four Exopod I-i 1-1 111,4Endopod 0-1 0-1 1,2Exopod of leg one (Figure 90C) bearing distolateral membraneson segmentsone, two and three. Segment three with fourpinnate setae, plus two shorter lateralsetae (lateralmost seta with membrane along medialedge, the adjacent seta bearing afinely serrated membrane along lateral edge).Endopod of leg one (Figure 9CC) withdistolateral membrane on first segment, 28to 30 endopodal denticulations on the second segment, and 29 to 31 endopodal denticulationson the terminal segment. Exopodof leg two (Figure 90D) similar to leg one, exceptsecond segment bears lateral seta,and third segment bears four pinnate setae, plus one semipinnateseta with setulesalong the medial edge and a finely serrated membrane along the lateraledge, onespiniform seta bilaterally bearing finely serrated membranes anda small naked seta.Endopod of leg two (Figure 90D) similar to endopod of legone, except second segmentbears 25-32 endopodal denticulations, and segment three bears 28-33endopodal denticulations. Exopod of leg three (Figure 90E) as in leg two, except lateralseta on second segment bears a finely serrated lateral membrane. Endopod of legthree (Figure90E) similar to leg two but segment three bears only four pinnate setae, segment twobears 10-12 endopodal denticulations, and segment three bears 15 endopodal denticulations. Exopod of leg four (Figure 90F) as in leg three. Endopod of leg four (Figure90F) similar to leg three except second segment medially bears pinnateseta, and terminal segment bears one stout seta bilaterally bearing serratedmembrane plus twopinnate setae. Second and third segments bear 10-12 and 11-13 endopodal denticulations respectively.Male: not acquiredComments: Kroyeria gemursa was originally described by Cressey (1967) fromthe great hammerhead, Sphyrna mokarranfrom off Madagascar. Since then it has beenreported from the same host from the West coast of Florida (Cressey, 1970) and from142the Indian Ocean near Trivandrum, India(Pillai, 1985).The laterally bulging, heavily sclerotizeddistal region of the last segment of theabdomen, the thickened claw and thickenedextension of the corpuswith the resultantreduced aperture of the second antenna, the largepatches of spinules on the distolateral surfaces of the labrum, and the numerous (25-33)endopodal denticulationsfoundonthe second and third segments of leg one andleg two, readily distinguish thisspecies.Kroyeria lineata van Beneden, 1853Syn: Lonchidium aculeatumGerstaecker, 1854Lonchidium Iineatum;of Bassett-Smith (1899)Kroyeria lineata Beneden; of Brian (1906)Kroyeria aculeata Gerstaeker; of Brian (1906)Krøyeria lineata Beneden; of Wilson (1932)Krøyeria sublineataYamaguti and Yamasu,1959(Figures 91-92)Material examined. One female (BMNH1928.6.11.6-15); one female (BMNH191 3.9.18.250-259) both collected by Andrew Scottfrom the gills of the tope shark,Galeus canis Bonaparte, 1834(=Galeorhinus galeus (Linnaeus, 1758) from the IrishSea; one female (BMNH 23.5.1975)collected by G.A. Boxshall from the gills of thesmooth-hound, Mustelus mustelus (Linnaeus, 1758).DescriptionFemale (Figure 91A)Overall length in dorsal view approximately 4.9mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures 91A and 91B) extending near hind margin of first free thoracicsomite, styletscurve sharply inward, forming an uncinateterminus. Three free thoracic somites withnon-overlapping terga. Genital complexcylindrical, constituting approximately 68 % oftotal body length. Posterolateral corners of latter bearing oviducal openings.Abdomenindistinctly three-segmented. Caudal ramus (Figure 91C) lamelliform, longerthan widewith medial fringe of setules, bearing two pinnatesetae distally, one distomedial pinnate seta, one distal thick semipinnate seta, and distolaterallybearing one stput semi-pinnate setae, and one slender pinnate seta.First antenna (Figure 91D) indistinctlyseven- to eight-segmented, armature(base to apex) as follows: 10, 1, 5, 2,3, 1, 1, 12 +1 aesthete. Second antenna (Figure14391 E) chelate and prehensile, apparentlyfour-segmented. Proximal twosegments heavily scierotized in such a way as tosuggest relatively unrestricted movementcapabilities.Third segment forming corpus of chela, extendinginto a rigid arm distallyexpandedinto a large membranous receptacle to accommodatetip of fourth segment. Latter forming heavily sclerotized robust claw, bearing twoprominent setae proximally, and distallyexpanded into a large membranous receptacle.Mandible (Figure 91 F) of two parts,dentiferous margin with nine teeth (1apical, 1 large, 2 small, 2 large, and 3 smalldecreasing in size). First maxilla(Figure 91G) biramous; endopod longer bearing twoapical elongate, pinnatesetae; exopod shorter bearing two short, naked setae.Second maxilla (Figure 91H) brachiform; lacertus heavilyscierotized with short basalprocess arising from near base. Brachium with two largepatches of prickles and a tuftof fine, long setae near base of claw; thelatter bearing paired lateral membranouslamellae with small prickles scattered upon convexsurface, and atypically, a large distal, subquadrangular membranous flap.Maxilliped (Figure 901) subchelate; corpustwo-segmented; subchela not divided into shaft and claw,distally uncinate and bearinga single small, slender seta.All four legs biramous and trimerite. Sympods two-segmented. All basipodswithlateral pinnate seta and two distomedial membranes; first basipodbears additional distomedial pinnate seta. Interpodal bars with interpodalstylets (leg one interpodal styletvery small). Lateral fringe of setules present on allendopodal segments, medial fringeof setules on the first exopodal segment of each leg. Armatureof rami as follows(Arabic numerals denote fully pinnate setae, Roman numeralsdenote conditionsdiverging from that state):Leg one Exopod 1-1 0-1 11,4 Endopod 0-1 0-0 6Leg two Exopod 1-1 0-1 111,4 Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4 Endopod 0-10-0 4Leg four Exopod I-i I-i 111,4 Endopod 0-1 0-1 1,2Exopod of leg one (Figure 92A) bearing lateral membranes on segments one,two and three. Segment three with four pinnatesetae, plus two lateral slender setae(lateralmost seta smaller and naked, adjacent seta bearing membrane along lateraledge). Endopod of leg one (Figure 92A) with distolateral membranes on all threesegments, and devoid of endopodal denticulations. Exopod of leg two (Figure 92B) similarto leg one, except third segment bears four pinnate setae, plus one semipinnatesetawith setules along the medial edge anda membrane along the lateral edge, one setalaterally bearing a membrane, anda small naked seta. Endopod of leg twa (Figure14492B) similar to endopod of leg one.Exopod of leg three (Figure 92C)as in leg two,except second segment bears a lateralslender seta. Endopod of leg three (Figure92C)similar to leg two but segmentthree bears only four pinnate setae. Exopod ofleg four(Figure 92D) as in leg three. Endopod ofleg four (Figure 92D) similar to leg threeexcept second segment medially bears pinnateseta, and terminal segment bears onestout seta bilaterally bearing serrated membrane plustwo pinnate setae.Male: not acquiredComments: Kroyeria lineata van Beneden,1853, type species of the genus,was originally reported fromtope shark, Galeus canis Bonaparte, 1834(=Galeorhinusgaleus (Linnaeus, 1758)). Since then,a few questionable records come from hosts inthe family Carcharhinidae, suchas the smooth hammerhead, Sphyrna zygaena(Linnaeus, 1758) (Cf. Wilson, 1932),the blacktip shark, Carcharhinus limbatus(Valenciennes, 1839), and the lemon shark, Hypoprionbrevirostris (=Negaprion brevirostris (Poey, 1868)), (cf. Wilson,1936), and the blue shark, Prionace glauca(Linnaeus, 1758) (cf. Rokicki and Bychawska, 1991;Kabata, 1979), the species hasbeen reported principally from tope sharks, smooth-hounds orwhiskery sharks of thegenera Galeorhinus and Mustelus, of the familyTriakidae. K. lineata has been reported from the tope shark, Galeorhinus galeus (Linneaus, 1758) fromBelgian waters (vanBeneden, 1853 and 1861), from theMediterranean (Claus, 1858), (Brian, 1906); fromthe Irish Sea (Scott and Scott, 1913, A.Scott, 1929),and from off England (LeighSharpe, 1933). K. lineata has also been reportedfrom the smooth-hound Mustelusmustelus (Linnaeus, 1758),(=Mustelus equestrias, see Compagno, 1984b), from theAdriatic Sea (Valle, 1880), the NorthSea (Boxshall, 1974), and the Mediterranean offTunisia (Essafi and Raibaut, 1977). The specieshas also been found on the starrysmooth-hound, Mustelus aster/as Cloquet, 1821 and Musteluspunctulatus Risso, 1826(=M. mediterraneus accordingto Compagno (1984b)), and off Tunisia in theMediterranean (Essafi and Raibaut, 1979). Finally,the relegation of K. sublineataYamaguti and Yamasu, 1959 to synonymy with K. lineata by Kabata (1979) extendsthehost and geographic range to include the starspotted smooth-hound, MustelusManazoBleeker, 1854 from Japanese waters (InlandSea).Kabata (1979) suggested that K lineata may be a synonym of the incompletelydescribed Lonchidium aculeatum Gerstaeker, 1854,collected from Galeorhinus galeus,in the eastern North Atlantic. Brian(1906) and Delamare Deboutteville and NunesRuivo (1953) agreed with this synonymy.Evidence accumulated over the years indicates that G. galeus is one of the preferredhosts for K. lineata, and supports this synonymy.145K. lineata is the only species in thegenus that has endopodswithout endopodaldenticulations and a second antenna bearingonly two prominent setae on theproximalregion of the claw. Additionally,no other Kroyeria has the distal membranousextensions near the tip of the claw of the secondmaxilla. The relatively short, sharply incurving dorsal stylets coupledwith the angular cephalothorax, aid in specific identificationofthis parasite.Kroyeria longicauda Cressey, 1970(Figures 93-95)Material examined. One paratype female (USNM128497) collected from the gills of theblacktip shark, Carcharhinus limbatus(Valenciennes, 1839) from Sarasota, Florida;numerous females and a male (USNM 128496)collected by Dr. Roger Cressey, fromthe same host species from nearSarasota, Florida; and a few females (USNM154003) collected from the gills of the spinnershark, Carcharhinus brevipinna (Mullerand Henle, 1839) collected by Dr. RogerCressey from Nosy Be Island, nearMadagascar in the Mozambique Channel.DescriptionFemale (Figure 93A)Overall length in dorsal view approximately 3.5 mm.Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures 93A and 93B) extending near anterior quarterof third free thoracic somite,stylets slightly curving inward with a deeply bifid terminus,lateral tine approximately20% the length of the other. Three free thoracic somiteswith non-overlapping terga.Genital complex cylindrical, constituting approximately65 % of total bod’ length.Posterolateral corners of latter bearing oviducal openings. Abdomen indistinctly threesegmented. Caudal ramus (Figure 93C) lamelliform, longerthan wide without medialfringe of setules along proximal half, but with a lateralcuticular flange not a membraneas represented by Cressey (1970) or Pillai (1985), bearing five pinnatesetae and onestout, plumose seta.First antenna (Figure 93D) indistinctly seven- to eight-segmented, armature(base to apex) as follows: 10, 1,5,2,3, 1, 1, 13+1 aesthete. Second antenna (Figure93E) chelate and prehensile, apparentlyfour-segmented. Proximal two segments heavilysclerotized in such a way as to suggest relativelyunrestricted movement capabilities.146Third segment forming corpus of chela, extendinginto a rigid arm, distallyexpandedinto a large receptacle to accommodatetip of fourth segment. Latter formingheavilysclerotized robust claw, distallyexpanded into a large membranous receptaclesimilarto corpus, and bearing threeprominent setae proximally. Aperture of chelasmall dueto the large membranous expansionsof both claw and corpus. Mandible (Figure 93F)of two parts, dentiferousmargin with nine teeth (1 apical, 1 large,2 small, 2 large, and3 small decreasing in size). First maxilla (Figure93G) biramous; endopod longer bearing two apical elongate, nakedsetae; exopod shorter bearingtwo short, naked setae.Second maxilla (Figure 93H)brachiform; lacertus heavily sclerotizedwith basal processarising from near base.Brachium with two large patches ofprickles and a tuft of fine,long setae near base of claw; the latterbearing paired lateral membranous lamellaewith small prickles scattered upon aninflated convex surface. Maxilliped (Figure931)subchelate; corpus two-segmented, myxal areabearing three crescent-shaped, membranous, cuticular flaps reminiscent ofthose in Eudactylina; subchela not divided intoshaft and claw, distally uncinate andbearing a single small, slender seta.All four legs biramous and trimerite.Sympods two-segmented. All basipods withlateral pinnate seta and two distomedialmembranes; first basipod bears additional distomedial pinnate seta. Interpodalbars with interpodal stylets (leg one interpodal styletvery small and blunt). Lateral fringeof setules present on segments one and two of allendopods, medial fringe of setules on the firstexopodal segment of each leg. Armatureof rami as follows (Arabic numeralsdenote fully pinnate setae, Roman numerals denoteconditions diverging from that state):Leg one Exopod 1-1 0-1 11,4Endopod 0-1 0-0 6Leg two Exopod I-i 1-1 111,4 Endopod0-1 0-0 6Leg three Exopod I-i 1-1 111,4 Endopod 0-10-0 4Leg four Exopod I-i 1-1 111,4 Endopod 0-10-1 1,2Exopod of leg one (Figure 94A) bearing pectinate lateralmembranes on segments one, two and three. Segment three with four pinnatesetae, plus two lateral slender setae (lateralmost seta naked,the adjacent seta semipinnate and bearing a membrane along lateral edge). Endopod ofleg one (Figure 94A) with distolateral membraneon first segment, five to six endopodaldenticulations on the second segment, and fourendopodal denticulations on the terminal segment. Exopodof leg two (Figure 94B)similar to leg one, except secondsegment bears lateral seta , and third segment bearsfour pinnate setae, plus onesernipinnate seta with setules along the medial edge and afinely serrated membrane alongthe lateral edge, and two slender seta bearing a mem147brane alonge their lateral edge. Endopod of leg two (Figure94B) similar to endopod ofleg one, except second segment bears six to seven endopodaldenticulations, segmentthree bears four endopodal denticulations.Exopod of leg three (Figure 94C)as in legtwo, except lateral seta on second segmentbears a lateral membrane. Endopod of legthree (Figure 94C) similar to leg twobut segment three bears only four pinnate setae,segment two bears five endopodal denticulations,and segment three bears threeendopodal denticulations. Exopod of leg four (Figure94E) as in leg three. Endopoci ofleg four (Figure 94D) similar to leg three except secondsegment medially beas pinnateseta, and terminal segment bears one stout seta bilaterally bearingsmooth membranesplus two pinnate setae. Secondand third segments bear three to four and threeendopodal denticulations respectively. Leg fivecomposed of four setae.Male: (Figure 95A)Overall length in lateral view approximately 2.1 mm. Cephalothoracicappendages and swimming legs similar to those of female. Genital complex bearingfifth and sixth legs represented by four and two pinnatesetae respectively. Dorsal stylet(Figure 95B) more stout than that of female, and similarly bearinga small lateral tine.Caudal ramus (Figure 950) more elongate than that of female, lackingsetules fringingmedial margin; six distal setae (one stout and semipinnate, one stout and naked, andfour slender and pinnate).Comments: Kroyeria longicauda was originally described by Cressey (1970)from the blacktip shark, Carcharhinus limbatus from off Sarasota, Florida. It has beenreported from this host from the Indian Ocean near Kerala, India (Pillai, 1985). Thisreport adds the spinner shark, Carcharhinus brevipinna (Muller and Henle, 1839) fromthe Mozambique Channel to the host and geographic range of this parasite.Interestingly, both the spinner shark, Carcharinus brevipinna and the blacktip shark,Carcharhinus limbatus are members of Garrick’s (1967) “C. limbatus group” andGarrick’s (1982) “C. limbatus-amblyrhynchoides group” (see Compagno, 1988). Thepresence of K. longicauda is consistent with the hypothesized close relationshipbetween these species.The lateral tine on the deeply incised, bifid dorsal stylet, the lateral cuticularflange (the hyaline fringe of Cressey (1970)) on the caudal rami, and the small numberof unusually large endopodal denticulations are unique to this species. The three atypical cuticular flaps on the myxal area of the rnaxilliped are also found on K. dispat aspecies obviously unrelated.148The dorsal stylet of male K. elongata (Pillai,1967), is very similarto that of K.Iongicauda.Kroyeriapapillipes Wilson,1932(Figures 96-97)Material examined. Oneholotype female (USNM 56672) collectedfrom the tiger shark,Galeocerdo cuvier (Peron& LeSueur, 1822); several females anda male (USNM153884-1 53899) collectedby Dr. Roger Cressey, from the same hostspecies from theWest coast of Florida.DescriptionFemale (Figure 96A)Overall length in dorsal view approximately12.0 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Cephalothorax laterally extended relative to congeners.Dorsal stylets (Figures 96A, B) extending to midpoint of first free thoracic somite,stout, and curving slightly outward with blunt tips.Three free thoracic somites with non-overlappingterga. Genital complex cylindrical,constituting approximately 68%of total body length. Posterolateral cornersof latterbearing oviducal openings. Abdomen indistinctlyone- or two-segmented. Caudal ramus(Figure 96C) lamelliform, longer than wide, devoidof typical medial fringe of setules,distally bearing six pinnate setae.First antenna (Figure 96D) indistinctly nine-segmented,armature (base to apex)as follows: 9, 1, 1, 5, 2, 3, 1, 1, 13 +1 aesthete.Second antenna (Figure 96E) chelateand prehensile, apparently four-segmented. Proximaltwo segments heavily scierotizedin such a way as to suggest relativelyunrestricted movement capabilities. Third segment forming corpus of chela, extending intoa rigid arm distally expanded into a smallreceptacle to accommodate tip of fourth segment.Latter forming heavily sclerotizedrobust claw, bearing three smallsetae proximally. Aperture of chela nearly circular dueto strongly curving claw. Mandible (Figure96F) of two parts, dentiferous margin withten teeth somewhat uniform in size.First maxilla (Figure 96G) biramous; endopodlonger bearing two elongatepinnate (short pinnules) setae; exopod shorter bearing twoshort, naked setae. Second maxilla(Figure 96H) brachiform; lacertus (not illustrated)heavily sclerotized withelongate basal process arising from near base. Brachium withtwo large patches of prickles anda tuft of fine, long setae near base of claw; the latter149bearing paired lateral membranouslamellae with small prickles scatteredupon convexsurface. Maxilliped (Figure 961) subchelate;corpus two-segmented, proximalsegmentbearing two conical processes; myxalregion of corpus with a tiny conical process; subchela not divided into shaft and claw,distally uncinate and bearing a single small, slender seta.All four legs biramous and trimerite.Sympods two-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta. Interpodal bars with poorlydeveloped (absent?) interpodalstylets. Lateral fringe of setuleson each endopodal segment, medial fringe of setuleson the first exopodal segment of eachleg. Armature of rami as follows (Arabic numeralsdenote fully pinnate setae, Roman numeralsdenote conditions diverging from thatstate):Leg one Exopod I-i 0-1 11,4Endopod 0-1 0-0 6Leg two Exopod I-i I-i 111,4Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4Endopod 0-1 0-0 4Leg four Exopod I-i 1-1 111,4 Endopod0-1 0-1 3Exopod of leg one (Figure 97A) bearing lateral membranes on segments one,two and three (additional pectinate strips onsegments one, two and three). Segmentthree with four pinnate setae, plus two lateral nakedsetae. Endopod of leg one (Figure97A) with distolateral membranes on segments oneand three; second segment bearsfive endopodal denticulations. Exopod of leg two(Figure 97B) similar to leg one,except second segment bears a lateral seta, third segment with four pinnatesetae, plusone semipinnate seta with setules along the medialedge and a membrane along thelateral edge, and two naked lateralmost setae. Pectinatemembranes paralleling thetypical membranes are present along the lateral margin of segments one, two, andthree. Endopod of leg two (Figure 97B) similarto leg one, except second segmentbears seven endopodal denticulations. Exopod of leg three (Figure97C) as in leg two.Endopod of leg three (Figure 960) similar to endopod of leg two except seCondsegment bears five to eight endopodal denticulations,and terminal segment bears four pinnate setae. Exopod of leg four (Figure 96D)as in leg three. Endopod of leg four (Figure96D) similar to leg three except terminal segment bears three pinnateseta, secondsegment similarly bears five to eight endopodal denticulations. Fifth leg not observed.Male: (Figure 97E)Total length approximately 5.4 mm. Appendages similarto female. Caudal rami150moreelongatethan thatof female,and legfive andsix representedby fourand twosetaerespectively.Comments:Kroyeriapapillipeswas originallydescribedby Wilson(1932)fromthe tigershark,Galeocerdocuvier(Peron &LeSueur,1822).Therecordof thisspeciesparasitizingthe smoothhammerhead,Sphyrnazygaena(Linnaeus,1758)hasbeenfound tobe inerror(see Cressey,1970).All subsequentrecordshavebeenfromgills ofGaleocerdocuvier.This speciesis theonly memberof thegenuswithall sixsetaeon thecaudalrami elongateand pinnate,makingit readilyidentifiable.Additionally,the endopodaldenticulationsare restrictedto the secondsegmentof eachendopod,similar tothatinK. cresseyl.The shortdorsalstylets,reducedor absentinterpodalstylets,circularapertureofthe chelatesecondantenna,and theindistinctlynine-segmentedfirst antennaeareattributessharedwith K.dispar Wilson,1935,whichoccurson thesamehost •andgeographiclocality.Kroyeriaprocerobscenasp.nov.(Figures98-99)Materialexamined.A few females(USNM153860,153858,153857)collectedby Dr.RogerCressey,from thegills ofthe bullshark,Carcharhinusleucas(Valenciennes,1839) fromnear NosyBe island,near Madagascarin theMozambiqueChannel.Femaleholotype(USNM266541)and 2femaleparatypes(USNM266542)depositedat the UnitedStatesNationalMuseumof NaturalHistory.Etymology:The specificnameprocerobscenaderivedfromthe Latinprocerus,meaning slender,long, stretchedout, andfrom theLatin obscenus,meaningindecent,or private part,in referenceto theextremelyelongatedgenitalcomplexof thisspecies.DescriptionFemale(Figure98A)Overalllengthin dorsalview approximately17.7 mm.Cephalothoracicsuturesarisinganterolaterallyand unitingposteromedially.Eyesnot evident.Dorsalstylets(Figures98A-C)extendingposteriorlyto anteriormarginof thesecondfreethoracic151somite, styletscurvingslightly inward.Dorsal styletsdistally bifurcatedwith a sharptine(Figure98B)or protuberance(Figure 98C)on the proximolateraledgeapproximately30%down thelength ofthe stylet.Three freethoracic somiteswith overlappingterga.Genitalcomplexcylindrical,constitutingapproximately80 % oftotal bodylength.Posterolateralcornersof latterbearingoviducalopenings.Egg strands(Figure98A)containing89 and97 eggs.Abdomen(Figures98A, D)indistinctlytwo- or three-segmented. Caudalramus (Figure98E) lamelliform,longer thanwide withoutmedialfringeof setules,distallybearingtwo largepinnate setae(setulesalong inflatedproximalhalves),two slenderpinnate setae,one robustsemipinnateseta,and one stoutplumoseseta (asmall setulearises froma raisedcrypt nearthe lateralborder).First antenna(Figure 98F)indistinctlyseven-segmented,armature(base toapex) asfollows:11, 5, 2, 3,1, 1, 13÷1 aesthete.Second antenna(Figure98G) chelateand prehensile,apparentlyfour-segmented.Proximaltwo segmentsheavilyscierotizedin such away as tosuggestrelativelyunrestrictedmovementcapabilities.Third segment formingcorpusof chela,extendinginto a rigidarm distallyexpandedinto a receptacle toaccommodatetip of fourthsegment.Latterforming heavilysclerotizedrobustclaw, bearingthreeprominentsetae (twoslenderand onetruncate)proximally.Mandible(Figure98H) oftwo parts,dentiferousmargin withnine teeth(2 large,2 small,2 large, 3slightlysmaller).First maxilla(Figure 98H)biramous;endopodlonger bearingtwo apicalelongate,pinnate setae;exopod shorterbearingtwo short,naked setae.Secondmaxilla(only distalportionillustrated)(Figure981) brachiform;lacertusheavilyscierotizedwith basalprocessarising fromnear base.Brachiumwith twolarge patchesof pricklesand a tuftof fine,long setaenear baseof claw;claw bearingpairedlateralmembranousIamellaewith smallpricklesscatteredupon convexsurface.Maxilliped(Figure99A) subchelate;corpustwo-segmented,proximalsegmentbearingtwo spinous processes;distalportionof corpusbearingsharp cuticularflange;subchelanotdividedinto shaftand claw,distally uncinateand bearinga singlesmall, slenderseta.All four legsbiramousand trimerite.Sympodstwo-segmented.All basipodswithlateral pinnateseta andone ortwo distomedialmembranes;first basipodbears additional distomedialpinnate seta;first coxopodbearstwo additionaldistolateralmembranes.All four interpodalbars bearinginterpodalstylets;interpodalstyletsof leg oneacute butrelativelysmall.Lateral fringeof setuleson eachendopodalsegment,medial fringeof setuleson the firstexopodalsegmentof eachleg. Armatureof ramias follows (Arabicnumeralsdenotefully pinnatesetae, Romannumeralsdenoteconditionsdivergingfrom thatstate):Leg oneExopod1-1 0-111,4Endopod0-1 0-06152Leg two Exopod I-i I-i 111,4Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4Endopod 0-1 0-0 4Leg four Exopod I-i I-i 111,4Endopod 0-1 0-1 1,2Exopod of leg one (Figure 99B) bearing distolateralmembranes on segmentsone, two and three(membranes finely pectinate on segments two andthree). Segmentthree with four pinnate setae, plustwo lateral slender setae, the largest bearsa membrane along its lateral edge andthe smaller and lateralmost seta is naked. Endopodofleg one (Figure 99B) withdistolateral membrane on first segment; segment twowith sixto seven (six shown) endopodal denticulations;segment three with eight to en (nineshown) endopodal denticulations.Exopod of leg two (Figure99C) similar to leg one,except second segment bears additional lateralseta, and third segment with four pinnate setae, plus one semipinnateseta with setules along the medial edge and a membrane along the lateral edge, and two slendersetae bearing membranes along their lateral edge. Pectinate membranes are present alongthe lateral margin of segments twoand three. Endopod of leg two (Figure99C) similar to leg one, except seven endopodaldenticulations are present on the secondsegment and nine to twelve (nine shown)endopodal denticulations on the third segment.Exopod of leg three (Figure 99D) as inleg two, except the lateral membranes of thesecond and third segments are not pectinate. Endopod of leg three (Figure 99D) withseven to eight endopodal denticulationson segment two and eleven endopodal denticulationson segment three; segment threewith four pinnate setae. Exopod of leg four (Figure99F) as in leg three, but pectinatemembranes on segments two and three and an additional pectinatestrip parallels thetypical membrane on segment one. Endopod of leg four(Figure 99E) with six and nineendopodal denticulations on segments two and threerespectively; segment three withtwo pinnate setae and one slender seta bilaterallybearing serrated membranes. Fifthleg not observed.Male: not acquiredComments: Kroyeria procerobscena is one of the giants in the genus. At nearly18 mm long its size is only rivalvedby K. papillipes, K. dispai and K. gemursa, atapproximately 12.0, 13.1, and 14.4 mm respectively. Of course the mesoparasitic K.caseyl at nearly 60 mm in length remains ina class of its own.K. procerobscena also has an unusually long genitalcomplex (hence its name),comprising 80% of the body length. The unusually long genital complex, the large size,the unique lateral tine on theproximal region of the bifid dorsal stylet, and the two elon153gate, proximally inflated, medially-pinched pinnatesetae on the caudal rami distinguishthis from any other species in this genus.Kroyeria rhophemophaga sp. nov.(Figures 100-1 02)Material examined. Several femalescollected from the secondary lamellae of theCalifornia soupfin shark, Galeorhinus galeus(Linnaeus, 1758) from inshore waters offEl Segundo and Newport Beach,California, U.S.A. Female holotype (USNM 266539)and 5 female paratypes (USNM 266540) depositedat the United States NationalMuseum of Natural History.Etymology: The specific name is derived fromthe greek rhomphema, meaning soup,and the greek phagein, to eat, referring to thisspecies’ predilection for feeding uponsoupfin sharks.DescriptionFemale (Figure 1 OOA)Overall length in dorsal view approximately 8.1 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially.Eyes not evident. Dorsal stylets(Figures bOA-C) extending posteriorlyto approximately 50 % down the length of thesecond free thoracic somite, stylets curving inward slightly with anacute terminus.Three free thoracic somites with non-overlapping terga. Genital complex cylindrical,constituting approximately 67 % of total body length. Posterolateral corners of latterbearing oviducal openings. Abdomen indistinctly three-segmented. Caudal ramus(Figure 100D) lamelliform, longer than wide bearing the typical medial fringe of setules,distally bearing four elongate pinnate setae and twostout, semipinnate setae. Eggstrands (Figure 100E) containing six eggs.First antenna (Figure 100E) indistinctly seven- or eight-segmented, rmature(base to apex) as follows: 11, 5, 1, 3, 1, 1 (missing in this specimen), 13+1 aesthete.Second antenna (Figure 100G) chelate and prehensile, apparently four-segmented.Proximal two segments heavily scierotized in such a way as to suggest the capablity ofrelatively unrestricted movement. Third segment forming corpus of chela, extendinginto a rigid arm distally expanded into a receptacle to accommodate tip of fourth segment. Latter forming heavily scierotized robust claw, bearing one elongate, slender seta154arisingfrom concavemarginin apertureof chela,and oneproximalslenderseta.Mandible(Figure 100H)of two parts,dentiferousmargin withnine (2large, 2small, 2large, and3 smalldescendingin size)teeth.First maxilla(Figure1001) biramous;endopodlonger bearingtwo apicalelongate,pinnate setae;exopodshorter bearingtwoshort,naked setae.Secondmaxilla(Figure100J) brachiform;lacertus heavilyscierotized withlarge basalprocessarisingfrom nearbase.Brachiumwith two largepatchesof pricklesand a tuftof fine,long setaenear baseof claw.Claw bearingpaired lateralmembranouslamellae,with membranebearing pricklesalong theconvex surface.Maxilliped(Figure lOlA)subchelate;corpus two-segmented;subchela notdividedintoshaft andclaw, distallyuncinateand bearinga singlesmall slenderseta.All four legsbiramousand trimerite.Sympodstwo-segmented.All basipodswithlateral pinnateseta andtwo distomedialmembranes;first basipodbearsadditionaldistomedia!pinnate seta.All four interpodalbars bearingelongateinterpodalstylets;interpodal styletsof leg onesmallerthan others.Lateral fringeof setuleson endopodalsegments oneand two,and on segmentthree onleg four.Medialfringe of setuleson theproximalsegmentof eachexopod.Armatureof rami asfollows(Arabicnumeralsdenotefully pinnatesetae, Romannumeralsdenote conditionsdivergingfrom thatstate):Leg oneExopod[-1 0-111,4 Endopod0-1 0-06Leg twoExopod1-1 0-1111,4Endopod0-1 0-06Leg threeExopodI-i (-1 111,4Endopod0-1 0-04Leg fourExopodI-i 1-1111,4Endopod0-1 0-11,2Exopodof leg one(Figure1O1B)bearingpectinatelateralmembraneson segments one,two andthree. Segmentthree withfour pinnatesetae,plus twoslendersetae; longerof the twowith membranealong lateraledge.Endopodof leg one(Figure1018) withdistolateralmembraneon firstsegment;segmenttwo withfive to sixendopodaldenticulations;segmentthree withlateralmembrane,and sixpinnate setae.Exopodof legtwo (Figure1O1C) similarto legone, exceptthree distolateralslendersetae(two bearinglateral membranesand lateralmostseta smallestand naked)arepresent.Endopod ofleg two (Figure1O1C) similarto legone, withfive tosevenendopodaldenticulationspresenton the secondsegment,a distolateralmembraneand sixpinnate setaeare presenton thethird segment.Exopod ofleg three(Figure101 D) similarto legtwo exceptsecondsegmentbears asmall lateralslenderseta, andthe largestslender onthe terminalsegment setais semipinnatewith a membranealongthe lateraledge. Endopodof leg three(Figure101 D) withthree toeight endopodalden-155ticulations onsegment two (eightshown); segment three bearsfour pinnate setaeandone lateral membrane.Exopod of leg four(Figure 101 E) similar toexopod of leg three.Endopod of legfour (Figure 101 E) withnine endopodal denticulationson segmenttwo;one lateralstout seta bilaterallybearing finelyserrated membranes,and two pinnatesetae tip segmentthree. Fifth legnot found.Male: (Figures102A, B)Overall lengthin dorsal view approximately2.1 mm. Cephalothoracicappendages andswimming legs similar tothose of female.Genital complexbearingfifth and sixth legsrepresented by four andtwo pinnate setaerespectively. Dorsalstylet(Figure 102A) shorterand more stoutthan that of female.Caudal ramus(Figure 102C)more elongatethan that of female,bearing six distalsetae (two shortand apicallyplumose; two longand pinnate, andtwo small slenderpinnate). Secondsegment ofendopods (Figure102D) bears threelarge endopodaldenticulations and arow of longsetules fringing thelateral margin.Comments: Kroyeriarhophemophagaresembles K. branchiocetes,K. lineata,K. cresseyl, andK. triakos in thatthe claw of thesecond antennaebears two ratherthan three elongateslender setae.K. rhophemophagadiffers mostnotably form K.branchiocetesand K. lineata bylacking the largemembranous expansionsfound distallyon the claw andcorpus of the secondantennae. K.rhophemophagais easily distinguished from K. triakosby possessingonly four pinnate setaeplus two lateralslendersetae on the terminalsegment of exopodthree; K. triakosbears five pinnatesetae plusone naked slenderseta. Finally, thecephalothorax ofK. cresseyl isorbicular ratherthan subquadrangular asin K. rhophemophaga.I was also able to examinefour specimensof Kroyeria fromGaleorhinus galeusfrom Kaikoura,New Zealand, collectedby Dr. Pilgrim.These specimensresembledK.rhophemophaga fromsouthern Californiaspecimens thatwere only about4.4 mm inlength. This is approximately54% the lengthof the specimensdescribed herein.Untilmore data are acquired,populations of Kroyeriafrom the California andNew Zealandsoupfin sharks,Galeorhinus galeus,will be consideredconspecific.Interestingly, K. lineatafrom the soupfinor tope shark,Galeorhinus galeusfromthe eastern Atlantic,is markedly differentfrom the K.rhophemophagafrom the easternNorth Pacific (California)and New Zealand.Many of species-specificelasmobranchinfecting copepods,havea geographicrange concomitantwith that of theirhosts.Perhaps the synonymyof the six speciesof Galeorhinusinto G. galeusby Compagno(1984b) is mistaken.The different parasitespecies andstriking differencein the size156between the aforementioned Pacific samples suggest distinct stocks ofthese soupfins.Kroyeria spatulata Pearse, 1948(Figuresi 03-104)Material examined. Several females (USNM 154009) collected by Dr. Cressey from thegills of the bull shark, Carcharhinus leucas (Valenciennes, 1839) from Sarasota, Florida;and a few females (USNM 271 635) collected by Dr. Roger Cressey, from the gills ofthe lemon shark, Negaprion brevirostris (Poey, 1868) from Sarasota, Florida.DescriptionFemale (Figure 103A)Overall length in dorsal view approximately 5.4 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially. Eyes not evident. Dorsal stylets(Figures 103A and 103B) extending near posterior margin of second free thoracicsomite, stylets slightly curving inward with a bifid terminus. Three free thoracic somiteswith slightly overlapping terga. Genital complex cylindrical, constituting approximately57 % of total body length. Posterolateral corners of latter bearing oviducal openings.Abdomen indistinctly three-segmented. Caudal ramus (Figure 103C) lamelliform, longerthan wide with fringe of setules along medial margin, bearing five pinnate setae (fourelongate and one stout) and one large, stout, semipinnate seta.First antenna (Figure 103D) indistinctly seven- to eight-segmented, armature(base to apex) as follows: 11, 5, 2, 3, 1, 1, 13 +1 aesthete. Second antenna (Figure103E) chelate and prehensile, apparently four-segmented. Proximal two segmentsheavily sclerotized in such a way as to suggest relatively unrestricted movement capabilities. Third segment forming corpus of chela, extending into a rigid arm, distallyexpanded into a receptacle to accommodate tip of fourth segment. Latter forming heavily sclerotized robust claw, bearing three prominent slender setae. Mandible (Figure103F) of two parts, dentiferous margin with nine teeth (1 apical, 2 large, 2 small, 2large, and 3 small decreasing in size). First maxilla (Figure 103G) biramous; endopodlonger bearing two apical elongate, unilaterally denticulated setae; exopod shorterbearing two short, naked setae. Second maxilla (Figure 103H) brachiform; lacertusheavily sclerotized with large basal process arising from near base. Brachium with twolarge patches of prickles and a tuft of fine, long setae near base of claw; the latter bearing paired lateral membranous lamellae with small prickles scattered upon convex sur157face. Maxilliped (Figure 104A) subchelate; corpustwo-segmented, proximal segmentwith three small spinous processes, distal regionof distal segment of corpus bearingatransverse cuticular flange; subchelanot divided into shaft and claw, distally uncinateand bearing a single small slenderseta.All four legs biramous and trimerite.Sympods two-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta, coxopod of leg one bears additionaldistolateral membrane.Interpodal bars with interpodal stylets (leg one interpodalstylet very small and blunt).Lateral fringe of setules present on segmentsone and two of all endopods, and on segment three of the endopod of leg four; medialfringe of setules on the first exopodalsegment of each leg. Armature of rami as follows(Arabic numerals denote fully pinnatesetae, Roman numerals denote conditions divergingfrom that state):Leg one Exopod 1-1 0-1 11,4 Endopod0-1 0-0 6Leg two Exopod I-i 1-1 111,4 Endopod 0-10-0 6Leg three Exopod I-i 1-1 111,4 Endopod 0-10-0 4Leg four Exopod I-i 1-1 111,4 Endopod 0-1 0-1 1,2Exopod of leg one (Figure 104B) bearing pectinate lateral membranes onsegments one, two and three, typical membrane paralleling pectinatemembrane on firstsegment. Segment three with four pinnate setae, plus two lateralslender setae (lateral-most seta naked, the adjacent seta bears a membrane along lateraledge). Endopod ofleg one (Figure 104B) with distolateral membrane on first segment,seven to nineendopodal denticulations on the second segment, and ten to eleven endopodal denticulations on the terminal segment. Exopod of leg two (Figure 104C) similar to leg one,except second segment bears lateral seta , and third segment bears four pinnatesetae,plus one semipinnate seta with setules along the medial edgeand a membrane alongthe lateral edge, and laterally, two slender seta, the larger bearinga membrane alongthe lateral edge. Endopod of leg two (Figure 104C) similarto endopod of leg one,except second segment bears nine to eleven endopodal denticulations, segment threebears thirteen endopodal denticulations. Exopod of leg three (Figure 104D)as in legtwo, except pectinate membrane paralleling typical membrane is present. Endopod ofleg three (Figure 104D) similarto leg two but segment three bears only four pinnatesetae, segment two bears nineto ten endopodal denticulations, and segment threebears thirteen to fifteen endopodal denticulations. Exopod of leg four (Figure 104F)with pectinate membranes arising medial to the distolateralsetae of segments one andtwo, and arising medial to the second most lateralseta, and all sinuously wrapping158down along lateralmargins of segments one, two,and three (obviouslythe diagnosticsaddlelike sclerotizationsnoted by Cressey(1 970)). Membranes present orrall threesegments. ndopodof leg four (Figure 104E)similar to leg three except secondsegment medially bears pinnateseta, and terminalsegment bears one stout setabilaterallybearing finely serrated membranesplus two pinnate setae. Secondand third segmentsbear ten and eleven endopodaldenticulations respectively.Leg five not observed.Male: not acquiredComments: Kroyeria spatulatawas originally described byPearse (1948) fromthe sharpnosed shark, Scoiodonterraenovae (Richardson,1836) (=Rhizoprionodonterraenovae (Richardson,1836)) from off Beaufort,North Carolina, U.S.A.Since then ithas been reportedfrom the sandtiger shark,Carcharias littoralis (=Eugomophodestaurus (Rafinesque, 1810)),and from the Bahamas(Pearse, 1951), fromthe blacktipshark, Carcharhinus limbatus(Valenciennes, 1 839)from the Gulf of Mexico(Pearse,1952b). Cressey (1967)reported it from the spinnershark, Carcharhinusmaculipinnis(Poey, 1865) (=Carcharhinusbrevipinna (Muller andHenle, 1839)), thespot-tail shark,Carcharhinus sorrah (Valenciennes,1839), and from themilk shark, Rhizoprionodonacutus (Ruppel, 1837)all from the IndianOcean. Following thesereports, K. spatulatawas reported by Cressey(1970) from the lemonshark, Negaprionbrevirostris (Poey,1868), and the bull shark,Carcharhinus leucas(Valenciennes, 1839)from off the Westcoast of Florida.Having been fortunateenough to examinemany of these specimenscollected bythe International IndianOcean ExpeditionI found the recordof K. spatulataon C. brevipinna to bein error. The correct identificationof the parasiteis K. longicauda.Furthermore, the recordof K. spatulata on C.sorrah should be changedto K. elongata(assuming host identificationis accurate).This species is problematicin that the originaldescription byPearse (1948) wasincomplete, andinconsistent with theillustrations. First,the corpus of thesecondantenna is not uniquelyspatulate as describedin the text. Second,Pearse statesthatthere are no papillae (theendopodal denticulationsherein) on the middleendopod segment of all legs, yet theillustrations of theapparently broken firstleg and the fourthleg(legs two and threewere not illustrated) distinctlyshow these to bepresent. Addingmisery, the hideously distortedtype specimenis distorted and compressed,and was ofno value in sorting these inconsistenciesout. The only illustrationof this speciessincethe original one by Pearsewas a single illustrationof the fourth leg byCressey (1970)detailing the diagnostic“saddlelike sclerotizationson the distolateralcorners on seg159ments two andthree of the fourthleg exopod”, which corresponds tothe sinuous pectinate membranes describedherein. Pillai (1985)sync nimized K. spatulata withK. elongata, but this is clearlyin error as outlined abovein the comments section underK.elongata.This species is easily distinguishedfrom all other congenersby the unique sinuous, pectinate membranesarising medial to the spiniform setaeand wrapping down thelateral margins of segmentstwo and three on the exopodof leg four.Kroyeria sphyrnae Rangnekar,1957(Figures 105-1 06)Syn: Kroyeria praelongaciculaLewis, 1966, syn.nov.Material examined.Several paratype femalesof K. praelongacicula(USNM 110800)collected ByDr. Alan Lewis, University ofBritish Columbia,from the gills of thescalloped hammerhead,Sphyrna lewini (Griffithand Smith, 1834) fromthe HawaiianIslands near KaneoheBay; numerous femalesfrom the same host speciesfrom PuntaArena de Ia Ventana, inthe southern Sea of Cortez(Gulf of California),Mexico; andnumerous femalesfrom the smooth hammerhead,Sphyrna zygaena(Linnaeus, 1758)from Punta Arena deIa Ventana, Mexico,and from near the ChannelIslands, in thesouthern California bight.DescriptionFemale (Figure 105-106)Overall length in dorsalview approximately8.7 mm. Cephalothoracicsuturesarising anterolaterallyand uniting posteromedially.Eyes not evident.Dorsal stylets(Figures 105A-C) extendingbeyond fourth freethoracic somite, styletsextremely elongate, lissome, with anacute terminus. Threefree thoracic somiteswith non-overlappingterga. Genital complex cylindrical,constituting approximately60 °k of total bodylength.Posterolateral cornersof latter bearing oviducalopenings. Abdomenindistinctly threesegmented. Caudalramus (Figure 105D)lamelliform, longerthan wide withmedialfringe of setules, bearingfour pinnate setae distally,and distolaterallybearing two stoutsemipinnate setae.First antenna (Figure105E and (terminalsegment detail)104F) indistinctlyseven- to eight-segmented,armature (base to apex) asfollows: 11,3, 1, 3,1, 1, 13+1aesthete. Second antenna(Figure 1 05G) chelateand prehensile,apparently .four-seg160mented. Proximal two segments heavily sclerotizedin such a way asto suggest relatively unrestricted movement capabilities. Thirdsegment forming corpus of chela,extending into a rigid arm distally expandedinto a small receptacle to accommodate tipof fourth segment. Latter forming heavily scierotizedrobust claw, bearing three prominent setae proximally. Aperture of chela smalldue to stout claw and extension ofcorpus. Mandible (Figure 105H) of two parts,dentiferous margin with seven teeth (1large, 2 small, 2 large, and 2 small decreasing in size). Firstmaxilla (Figure 1051) biramous; endopod longer bearing two apical elongate,setae (one naked and one pinnate); exopod shorter bearing two short, nakedsetae. Second maxilla (Figure 106A)brachiform; lacertus heavily sclerotized with elongatebasal process arising from nearbase. Brachium with two large patches of pricklesand a tuft of fine, long setae nearbase of claw; the latter (see detail) bearing paired lateral membranous lamellaewithsmall prickles scattered upon convex surface.Maxilliped (Figure 106B) subchelate;corpus two-segmented, proximal segment bearinga single conical process; subchelanot divided into shaft and claw, distally uncinateand bearing a single small slenderseta.All four legs biramous and trimerite. Sympods two-segmented. All basipodswithlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta. Interpodal bars with interpodal stylets (legone interpodal styletvery small and blunt), remaining three legs allpossess interpodal stylets relativelysmaller than the majority of the genus. Lateral fringe of setules present onthe first andsecond segments of endopods one through four, andon the third segment of leg four;medial fringe of setules on the first exopodal segment of each leg. Armature oframi asfollows (Arabic numerals denote fully pinnatesetae, Roman numerals denote conditions diverging from that state):Leg one Exopod 1-1 0-1 11,4 Endopod 0-10-0 6Leg two Exopod I-i 1-1 111,4 Endopod 0-1 0-0 6Leg three Exopod I-i 1-1 111,4 Endopod 0-1 0-0 4Leg four Exopod I-i 1-1 111,4 Endopod 0-1 0-1 1,2Exopod of leg one (Figure 106C) bearing lateral membranes on segments one,two and three. Segment three with four pinnatesetae, plus two lateral slender setae(lateralmost seta small and naked, the adjacentseta bears a membrane along lateraledge). Endopod of leg one (Figure 1060) with distolateral membrane on first segment,six to nine endopodal denticulationson the second segment, and eight to nine endopodal denticulations on the terminalsegment. Exopod of leg two (Figure 106D) similar to161leg one, except second segment bears lateral seta , and third segment bears four pinnate setae, plus one semipinnate seta with setules along the medialedge and a finelyserrated membrane along the lateral edge, one slender seta bearinga lateral membrane and a small naked lateralmost seta. Endopod of leg two (Figure 1 06D) similar toendopod of leg one, except second segment bears seven to eleven endopodal denticulations, and segment three bears eight to ten endopodal denticulations. Exopod of legthree (Figure 106E) as in leg two. Endopod of leg three (Figure 106E) similarto leg twobut segment three bears only four pinnate setae, segment two bears nine to tenendopodal denticulations, and segment three bears nine to eleven endopodal denticulations. Exopod of leg four (Figure 106F) as in leg three. Endopod of leg four (Figure106F) similar to leg three except second segment medially bears pinnate seta, and terminal segment bears one stout seta bilaterally bearing serrated membrane plus twopinnate setae. Second and third segments bear eight to nine and eight to elevenendopodal denticulations respectively.Male: not acquired.Comments: Kroyeria sphyrnae was originally described by Rangnekar (1957)from an unknown species of hammerhead, Sphyrna sp. in India, presumably in theBombay region. The next record of K. sphyrnae came from the gills of the smoothhammerhead, Sphyrna zygaena (Linnaeus, 1758) from Trivandrum, India (Pillai, 1967).Kabata (1970) presented a tentative find of K. sphyrnae from a single non-ovigerousspecimen collected from the brownbanded bamboo shark, Chiloscyllium punctatumMuller and Henle, 1838 (Orectolobiformes: Hemiscylliidae), from Moreton Bay,Queensland, Australia. Unfortunately, no illustrations or dimensions accompanied thatreport, and specific identification of the specimen remains open to question. Cressey(1970) claimed to have discovered it from the blacknose shark, Carcharhinus acronotus(Poey, 1860) from the West coast of Florida. Again, no illustrations accompanied thatreport, but Cressey did include total length of the specimens, which were reported to bevery small (2.2 mm). PilIai (1985) superficially describes the species and summarizesreported host associations of this parasite.Lewis (1 966a) described a new species, Kroyeria praelongacicula, from the scalloped hammerhead, Sphyrna lewini (Griffith and Smith, 1834) from Kaneohe Bay,Oahu, Hawaii. These specimens appear to be conspecific with K. sphyrnae. Thestrange irregular bifid tip of the dorsal stylet illustrated in Lewis’ description of K. praelongacicula, was not observed in the paratypes I examined. Based on the .paratypespecimens examined and specimens collected from the same host species in the162southern Sea of Cortez, and southern California Bightin the eastern North Pacific, andthe descriptions offered by Rangnekar (1957) andPillai (1967, 1985), K praelongacicuIa is considered a synonym of K. sphyrnae.Cressey’s (1 970) record of K. sphyrnae is suspect.Cressey reports the length ofK. sphyrnae as only 2.2 mm, which is quite small consideringthe original description byRangnekar measures this species at 4.7 mm. Lewis (1966a)reports a range of 6.58-8.12 mm, and a total length of approximately8.7 mm was found herein. It is unlikelyCressey’s (1970) record represents the samespecies. Cressey notes that the smallsize combined with the last two segmentsof the endopod of the fourth leg bearing onlytwo or three lateral spinules (the endopodaldenticulations herein) make specific identification easy. Neither of these characteristicsapply to K. sphyrnae. What speciesCressey’s (1970) record represents is uncertain.K. sphyrnae is a parasite of smooth and scallopedhammerheads, Sphyrnazygaena and Sphyrna lewini, respectively. My unpublishedfield observations from bothsouthern California and the southern Sea of Cortez suggestS. zygaena is the preferredhost as the parasite load is higher on this host.The long, acute, lissome dorsal stylets, the formula of theseven-toothedmandible, and the relatively short interpodal stylets which barely reach thedistal marginof the basipods of legs two, three, and four readily distinguish thisspecies.Kroyeria triakos Fukui, 1965 nom. emend.(Figures 107-108)Syn: Kroyeria elongatus Fukui, 1965Kroyeria triakisae Fukui, 1965nec Kroyeria elongata Pillai, 1967Material examined. Two females collected from the secondary lamellae of the bandedhoundshark, Triakis scyllium Muller and Henle, 1839 (USNM 22607) from inshorewaters off Awa, Japan.DescriptionFemale (Figure 1 07A)Overall length in dorsal view approximately 4.6 mm. Cephalothoracic suturesarising anterolaterally and uniting posteromedially. Eyes not evident. Dorsal stylets(Figures 1 07A, B) extending to the posterior margin of the second free thoracic somite,163stylets curving inward slightly witha narrow distolateral flange.Three free thoracicsomites with slightly overlapping terga. Genitalcomplex cylindrical, constitutingapproximately 66 % of total body length.Posterolateral corners oflatter bearing oviducal openings. Abdomen indistinctly three-segmented.Caudal ramus (Figure107C) lamelliform,longer than wide bearing the typical medialfringe of setules,distally bearing two elongate pinnate setae, two shorter pinnatesetae and two stout semipinnatesetae.First antenna (Figure 107D) indistinctlyseven- or eight-segmented, armature(base to apex) as follows: 9, 1,5, 1, 3, 1, 1, 13 +1 aesthete. Second antenna(Figure107E) chelate andprehensile, apparently four-segmented. Proximaltwo segmentsheavily scierotized in sucha way as to suggest the capablity of relativelyunrestrictedmovement. Third segment forming corpusof chela, extending into a rigid armdistallyexpanded into a receptacle to accommodate tip offourth segment. Latter forming heavilysclerotized robust claw, bearing one elongateslender seta along concave margin inaperture of chela, and proximally, oneelongate seta. Mandible (Figure 107F) of twoparts, dentiferous margin with nine (1apical, 2 large, 2 small, 2 large, and 2 small) teeth.First maxilla (Figure 1 07G) biramous;endopod longer bearing two apical elongate, pinnate setae; exopod shorter bearing two short,naked setae. Second maxilla (Figure107H) brachiform; lacertus heavilysclerotized with long basal process arising from nearbase. Brachium with two large patches of prickles anda tuft of fine, long setae nearbase of claw. Claw bearing membranous lamellae, specificsdifficult to ascertain due tosmall size. Maxilliped (Figure 1 071)subchelate; corpus two-segmented, proximal segment bearing one conical process, distolateral regionof main corpus bears transversecuticular flange; subchela not divided intoshaft and claw, distally uncinate and bearinga single small, slender seta.All four legs biramous and trimerite. Sympodstwo-segmented. All basipods withlateral pinnate seta and two distomedial membranes;first basipod bears additional distomedial pinnate seta; first coxopod bears two additionalmembranes. All four interpodalbars bearing elongate interpodal stylets; thoseof leg one smaller than others. Lateralfringe of setules on endopodal segments two and three,medial fringe of setules on thefirst segment of each exopod. Armature of ramias follows (Arabic numerals denote fullypinnate setae, Roman numerals denote conditions divergingfrom that state):Leg one Exopod 1-1 0-1 11,4 Endopod 0-1 0-06Leg two Exopod 1-1 0-1 111,4 Endopod 0-10-0 6Leg three Exopod I-i 1-1 1,5 Endopod 0-10-0 4Leg four Exopod 0-1 0-1 111,4 Endopod 0-1 0-1 1,2164Exopod of leg one (Figure 108A) bearing lateral membranes on segments one,two and three. Segment three with four pinnate setae, plus two slender naked setae.Endopod of leg one (Figure 108A) with distolateral membrane on first segment; segment two with seven endopodal denticulations; segment three bearing six pinnatesetae, and apparently devoid of typical distolateral membrane. Exopod of leg two(Figure 108B) similar to leg one, except third lateralmostseta is semipinnate withsetules along the medial edge, the two lateralmost setae slender and apparently naked.Enclopod of leg two (Figure 1080) similar to leg one, with six to eight endopodal denticulations (eight shown) present on the second segment, and a lateral membrane and sixpinnate setae on the third segment. Exopod of leg three (Figure 108D) similar to legtwo except second segment bears a small lateral seta, and the terminal segment bearson’y one naked, lateral setae and five pinnate setae. Endopod of leg three (Figure108E) with seven to ten endopodal denticulations on segment two (seven shown); segment three bears four pinnate setae and one distolateral membrane. Exopod of leg four(Figure 108F) without typical lateral setae on segments one and two, terminal segmentwith four pinnate setae, one semipinnate seta, one slender seta bearing a serratedmembrane, and the lateralmost slender seta naked. Endopod of leg four (Figure 108F)with indistinct membranous, endopodal denticulations forming the membrane, or transparent supports for the membrane (illustrated as a typical membrane) on segment two(this could be interpreted as devoid of endopodal denticulations or antithetically, possessing very many fused, membranous endopodal denticulations); one lateral stoutseta bilaterally bearing serrated membranes, and two pinnate setae tip segment three.Fifth leg not found.Comments: As noted above, Kroyeria triakos, K. branchiocetes, K. lineata, K.rhophemophaga, and K. cresseyi differ from their congeners in that the claw of the second antenna is armed with only two slender setae. Differences between species in thiscomplex have already been outlined in the comments section for K. cresseyl.K. triakos was originally described by Fukul (1 965) from gills of a young, bandedhoundshark, Triakis scyllium Muller and Henle, 1839, from Kurihama, in the Kanagawaprefecture collected in 1955. Although his description is incomplete, the unique shapeof the cephalothorax (almost arrowhead shaped; “the cobra with protrusions in the posterior corners” of Fukui (1965)), the similar dimensions, and the fact that both collections come from the same host species in Japanese waters leave no doubt that specimens examined here are conspecific with Fukui’s material. Fukui (1965) assigned thenew Japanese name Dochi-zame-hoso-yadori Ken-mizinko, (meaning, the elongatedKroyeria from Triakis) to this parasite. Strangely, he also assigns two binomens to this165copepod, keeping the Japanese name the samein both instances. Kroyeriaelongatusis the first name to show up in the publicationin the figure caption preceedingthe actualdescription. The name Kroyeria triakisae occurs inthe narrative, describing the species.Fukui’s K. elongatus should be K. elongata(Kroyer/a is feminine), making it ahomonym of Pillai’s (1967) K. elongata. Thiswould force one to correct thegender ofFukul’s species and rename Pillai’s. However,for the sake of nomenclatural stabilityand to minimize further taxonomic confusionin the genus, this species will retain thename K. triakisae. Since the name isa Greek noun, its genitive singular istriakos. Ipropose, therefore to amend Fukui’s K.triakisae to K. triakos. This leaves Pillai’s (1967)K. elongata, unchanged.K. triakos is easily recognized by beingthe only species in the genus that hasfive elongate, pinnate setae on the thirdsegment of the third leg. It is also the onlyspecies lacking the typical lateral setae onsegments one and two of the fourth exopod.The latter feature is suggested here only tentatively,for two reasons. Firstly, it is otherwise unknown in Kroyeriidae; secondly, materialavailable for examination was not adequate to determine it beyond a reasonable doubt.REMAINING UNOBTAINABLE NOMINAL SPECIESKroyeria consists of 18 nominal species. Three ofthem, (K. acanthiasvulgarisHesse, 1879, K. galeivulgaris Hesse, 1884, andK. scyliicaniculae Hesse, 1879) havenot been sufficiently well described to be recognizedand must be considered speciesinquirenda. K.trecai Delamare Deboutteville andNunes-Ruivo, 1953 was neverdescribed and remains a nomen nudum. K.aculeata (Gerstaecker, 1854) and K. sublineata Yamaguti and Yamasu, 1959 are consideredjunior synonyms of K. lineata vanBeneden, 1853. K. grad/is Wilson, 1 932 similarly isconsidered a junior synonym of K.carchariaeglauci Hesse, 1879, and K. praelongacicula Lewis 1966is a synonym of K.sphyrnae Rangnekar, 1957. This revision resurrects K. elongata Pillai,1967.This section is a brief (alphabetical) account ofthe females of the uncertainspecies above and the remaining species that werenot available for this revision ofKroyeria.Kroyeria acanthiasvulgaris Hesse, 1879 was described from the spinydogfish,Acanthias vulgar/s Risso, 1826 (=Squalus acanthiasSmith and Radcliffe, 1912), fromoff Brest, France. I haveexamined literally hundreds of this host species, and theyonly harbored Eudactylinaacanthil A. Scott, 1901. Since many triakids or smooth-hounds are morphologically similarto this reported squaloid host, host misidentification166is a likely explanation for this unlikely host-parasite association.Kroyeria echinata Rangnekar, 1956 was reported from the body surface of thesmooth hammerhead, Sphyrna zygaena (Linnaeus, 1758) from the Indian Ocean nearBombay, India. The description is too superficial for detailed comparisons. AlthoughPilIai’s (1985) redescription is helpful, some interpretations (setation on distal elementsof the caudal rami and legs, and the number of slender setae on the claw of the secondantennae) are suspect. Pillai (1985) reports this parasite from the smooth hammerheadfrom Kerala, India. The short stout dorsal stylets are reminiscent of those found in K.dispar and K. papillipes, but can be distinguished from these two by their small size(approximately 3.2-4.1 mm, compared to the 13.1 and 12.0 mm of K. dispar and K.papillipes respectively). The endopods of K. dispar are devoid of endopodal denticulations, and they are restricted to the second segment only of all four legs in K. papillipesand K. echinata. The orbicular cephalothorax of K. papillipes is quite distinct from thatof K. echinatus as is the armature of the setae on the caudal rami and the legs.Kroyeria gale/vulgar/s Hesse, 1884 was reported from the Tope shark, Galeusvulgar/s Fleming, 1828 (=Galeorhinus galeus (Linnaeus, 1758)). This may represent K.lineata, considering the host and geographic locality (Brest, France).Kroyer/a minuta Pillai, 1968 was described from gill filaments of the milk shark,Scoliodon sorrokowah (Bleeker, 1853), (=Rhizoprionodon acutus (Ruppel, 1837)), fromKerala, India. This is a small species, only about 3.0 mm in length. The long bifid dorsalstylets reaching the posterior margin of the fourth free thoracic somite, coupled with thesupposed and questionable serrated medial margins (I suspect these serrations are thetypical endopodal denticulations characteristically found throughout the genus), and theatypical patch of spinules on the lateral margin of the coxopod of leg two, plus the existence of six fully pinnate setae on the terminal segment of the exopod of leg two, readily distinguish this species.Kroyeria scyllicaniculae Hesse, 1879 was originally reported from the small-spotted catshark, Scyllium canicula (=Scyliorh/nus canicula (Linnaeus, 1758)), fromBrest, France. Unfortunately, K. scyllicaniculae a species /nquirenda comes from theScyliorhinidae, a host family with very little parasitic copepod information.Kroyeria trecai Delamare Deboutteville and Nunes-Ruivo, 1953 was reportedfrom the smooth hammerhead, Sphyrna zygaena (Linnaeus, 1758) and from the scalloped hammerhead, Sphyrna diplana Springer, 1941 (=Sphyrna lewini (Griffith andSmith, 1834)) from off the coast of Senegal. Although no description was ever published, they mention the species is characterized by its styliform processes projectingpast the level of the genital segment, by having a one-segmented abdomen and by itscaudal rami ending in 4 subequal setae, the 2 longer being ciliated. The long dorsal167styets coupled to the fact that this species isfound on the same sphyrnid hostsasKroyeria sphyrnae suggests that thesetwo species may be synonyms.PHYLOGENETIC ANALYSISCLADOGRAM CONSTRUCTIONA phylogenetic analysis was conducted ofthe 1 6 species of Kroyeria revisedherein. The outgroup was composed of thepresumed sister taxon, Kroeyerina, andProkroyeria (presumed sister to Kroyeria andKroeyerina) (Figure 109; see Deets,1987 for details). A total of 44 characters (seeAppendix C for data matrix and definition of characters) were analyzed using an exactsearch Branch and Bound algorithmof PAUP. The most parsimonious tree for Kroyeria(Figure 110) had a tree length of165, a consistency index of 0.75 (maximumvalue = 1.00), a retention index of 0.75(maximum value = 1.00), and an F- ratio of 0.0589(maximum value=0). Change and/orsynapomorphy lists may be obtained from the author.The first two branches arising from thebase of the cladogram are representedby K. dispar and K. papillipes respectively, both parasites of thetiger shark, Galeocerdocuvier. The next dade is composed of parasites specificto the Triakidae: the majordichotomy in this dade separates K. lineata (parasites ofGaleorhinus galeus andMustelus spp.) and K. rhophemophaga (parasite of CalifornianGaleorhinus galeus)from K. triakos and K. cresseyl, parasites of theJapanese and Californian ‘eopardsharks, Triakis scyllium and Triakis semifasciata, respectively. Thenext dade up thecladogram holds two species, K. sphyrnae and K. gemursa,parasites of the hammerhead genus Sphyrna. A paraphyletic triad follows consisting of the bizarre, mesoparasitic K. caseyl, parasitic on the night shark, Carcharhinus signatus,K. spatulata, foundon various species of Carcharhinus, Negaprion, and Rhizoprionodon, and K. procerobscenum a parasite of the bull shark,C. leucas. The final major bifurcation leads to onedade composed of K. branchiocetes, K. longicauda, and K. elongata, parasitesof various species of Carcharhinus, and a dade composed of K. decepta, specificto thedusky shark, C. obscurus, and K. carchariaeglauci parasitic on the epipelagicsilkyshark, C. falciformis, the pelagic whitetip, C. longimanus, and the blue shark Prionaceglauca.168PARASITE-DERIVEDHOST CLADOG RAMSThe Kroyeria cladogram wasrecoded by additive binary coding and a host byparasite data matrix wascreated (Table Ill).The phylogenetic analysis conductedfrom the recoded species of Kroyeriabyhost matrix (Table 3) resulted ina single most-parsimonioustree (Figure 111). TheBranch and Bound generated tree hada tree length of 30, and a consistency indexwith a maximum value of 1.00.The parasite-derived hostcladogram shows some unexpected patterns. Thefirst two branches at thebase of the tree posit a paraphyletic relationshipfor the presumably monophyletic tiger shark,Galeocerdo cuvier. This artifact isdue to this hostpossessing two different species ofKroyeria. Using parasite species as characters inthis case would be analogous to codingan organism for phylogenetic analysisthatpossessed two different character states ofa given character simultaneously. InclusiveOR’ing, a technique previously usedto code for the occurrence of morethan one parasite taxon per host could have beenused to force Galeocerdo into monophyly, howeverthis technique, if not restricted totreatment of parasite sister taxa, createsa chimeraout of unmodified data, resulting in the distortionof phylogenetic information (seeO’Grady and Deets, 1987 for details).The next dade contains members of the Triakidae.This dade consists of onegroup housing the leopard sharks Triakis semifasciatafrom the California coast ,and Tscyllium from Japanese waters. The remainingdade contains the soupfin sharkGaleorhinus galeus functioning asthe sister taxon to a dade composed of the G.galeus and various species ofMustelus from the western North Atlantic andMediterranean. The monophyly ofG. galeus with the many Mustelus species is due tothis host complex sharing the common parasite,K. lineata. The other branch leading toG. galeus stands alone as this soupfin shark from Californiawaters possesses its owncopepod K. rhophemophaga.The next dade contains all hammerheads ofthe genus Sphyrna. Thegreat hammerhead, Sphyrna mokarran is sistertaxon to the dade consisting of scalloped and smooth hammerheads,Sphyrna lewini and S. zygaena, respectively. Thenight shark, Carcharhinus signatusis the first Carcharhinus to appear on the cladogram followed by a dade consistingof the bull shark, C. leucas and the lemon shark,Negaprion brevirostris both possessing K. spatulata.C. leucas is unresolved with twoclades, one with five the other withfour taxa. The five taxon dade consists of the Greyreef shark, C. amblyrhynchos whichis the sister taxon to a dade composed of two169smaller groups. The first group contains the spinner shark,C. brevipinna and the black-tip, C. limbatus. The remaining group is composed of thespot-tail shark, C. sorrah andthe milk shark Rhizoprionodon acutus. The four taxon dadeon consists of the duskyshark, C. obscurus, the sister taxon to an unresolvedtrichotomy of pelagic tropical totemperate carcharhinids, the silky shark,C. falciformis, the pelagic whitetip, C. longimanus, and the blue shark, Prionaceglauca.COMBINING PARASITE CLADOG RAMSIf parasites can be used as characters,and if they possess symplesiomorphic,synapomorphic, or autapomorphic relationshipswith their hosts as do typical charactersin phylogenetic analyses, then we should gainresolution regarding host relationshipswith addition of parasite phylogenies (assuming anhypothesis of strict co-speciation).Additionally, Brooks and McClennan (1991, 1993)claim the methods appliedin singledade analysis can be used for multiple clades simultaneously.Kroeyerina Wilson, 1932 (Figure 112) is a genus ofcopepods dwelling in olfactory Iamellae of elasmobranchs. Deets (1987)generated a parasite cladogram and aparasite-derived host cladogram of the genus. During thecourse of this studyKroeyerina cortezensis Deets, 1987, originallyreported from the silky shark,Carcharhinus falciformis, was discovered from olfactory lamellae of the pelagicwhitetip,Carcharhinus longimanus, captured in a tuna seine off theRevillagegedos Islands,Mexico in September of 1988. Hence theparasite-derived host dladogram forKroeyerina is slightly different from the original with the incorporation of the pelagicwhitetip (Figure 113).The Kroeyerina cladogram from Deets (1987) was converted intoa binary code(Table Ill), and then added to the binary code for Kroyeria (Table II) resulting inthe combined matrix for both genera (Table IV). A phylogeneticanalysis was conducted for thishybridized data set using the exact search Branch and Bound algorithm inPAUP.Three equally parsimonious trees with a tree length of 49,a CI of 0.90 (maximijm value= 1.00), and an RI of 0.96 (maximum value = 1.00)resulted. One tree preserved theparaphyly of Galeocerdo cuvier, as in the previouslygenerated Kroyeria cladogram(Figure 110). One tree had the thresher shark,Alopias vulpinus and the mako, Isurusoxyrinchus both lamnids, tucked between the triakids and Galeocerdo cuvier resultingin the paraphyly of the carcharhinids and lamnids.The remaining tree that maintainedmonophyly of the aforementioned specific andsupraspecific taxa was chosen (Figure114).The resultant tree accommodates bothsets of parasite data (the additive binary170recoded cladograms from Kroyeria andKroeyerina) with little conflict.The holocephaIan dade represented by Callorhynchuscallorhynchus (Linnaeus, 1758) is thesistertaxon to the remaining elasmobranchs.The batoid dade composed of thegeneraMobula, Dasyatis, and Rhinobatusis the sister dade to the remaining componentsofthe tree composed of the twolamnids, A/op/as and Isurus plus the remaining carcharhinids. Ga/eocerdo, previouslyshowing a paraphyletic relationship with itselfdueto the sharing of Kroyeria dispar and K.papi//ipes becomes a monophyletic entity heldtogether by the presence of Kroeyerinaelongata Wilson, 1932.Also of interest is the placement of theblue shark, Prionaceg/auca. In theKroeyerinacladogram (Figure 113)Prionace is basally placed in the carcharhinid-lamnid lineage, in theKroyer/a-derived cladogram (Figure 111) Prionace wasat the top ofthe cladogram, and the combined tree(Figure 114) (in fact all three of thecombinedKroeyerina-Kroyeria trees) maintained thisplacement of Prionace. This placement ofPrionace is consistent with that of Compagno(1988) and Lavery (1992) in their morphologically- and molecular-based cladograms,respectively. The taxonomic or phylogenetic congruence (see Lanyon, 1993) amongstthree independent data sets suggeststhat the presence of Kroeyerina e/ongata,on Prionace is a result of colonization fromGaleocerdo.Not mentioned in other studies however, the methodused equally combinesparasite species from one genus with those fromanother genus to generate the hostcladogram. The more species a cladogram contains,the more nodes and branches orinformation the binary code must contain, resulting ina significantly longer binary stringin order to represent that topology. Having morecharacter states in a parsimony analysis will functionally weight that set of dataas graphically illustrated in this example.Serendipitously, this combination of data sets has almost no overlap. With allthe Kroyeria species restricted to the Carcharhiniformes,and Kroeyerina extendingover a broad range of host taxa, minimal characterconflict or competition occurs,Although the specific results are not included in this effort, combining this hybriddata matrix composed of Kroeyerina and Kroyeria withthe recoded Eudactyll;’7a cladogram resulted in thousands of rival trees with heavilyconflicting topologies (data setmay be obtained from the author). Surveyinghundreds of these trees revealed thateach tree possessed subsets of clades fromeach of the three data sets. Due to the differential possession (via sampling biasor extinction) of a given parasite group orgroups, members of putative host clades apparently well resolved in the independentanalyses would be ripped away from one another.The general problem here has beentouched upon superficially by Page (1994) and specifically hereinby me. The problemis in the treating of parasites strictlyas characters and not as lineages with their own171independent histories. Generating a host phylogeny withmultiple parasite lineages thatare differentially possessed by the host complex under analysisis somewhat analogousto generating a copepod parasite tree based on the maxilliped froma couple of speciesof one genus, the antennae of another,and on the legs from yet another with only someof these characteristics overlappingacross the taxa under study. The result will beuninterpretable noise and artifact at best. Ifcladists recognize that clades or lineages,have independent histories, then single parasitelineages are what cospeciation analyses should focus on. Despite this, informative analysesare possible if the parasitetaxa are restricted to: 1) a single lineage, 2) multipleparasite taxa equally distributedand weighted across the host axis, or3) parasite lineages that occur in different hostlineages minimizing character conflict. Thisis nothing more than following the sameprotocol one follows in using typical characterdata. Point one is analagous to character state analysis of a single character, pointtwo is the same as undifferentially usingall character data available from organisms, andpoint three is equivalent to synapomorphic or autapomorphic character datadistributions. Therefore, though combined analyses may consolidate information into a single tree under the aboveconditions, cladograms may offer more information separately than whencombined in other situations.Subsequent tree comparison or tree reconcilation methodologies can proceed from thatpoint. Of course tree comparison/reconciliationcan be conducted with the parasite orassociate phylogeny directly, a parasite-derived host phylogeny need notbe generateda priori.COMPETING HOST CLADOGRAMSThe several parasite-derived host cladograms generated herein function asindependent tests or hypotheses of other elasmobranch phylogenies previously reported in the literature.Two of the most comprehensive phylogenetic analyses of relationships of thesqualoids, squatinids, pristiophorids (Shirai, 1992a, b), and batoids (Nishida, 1990) arehybridized herein for comparative purposes (Figure 115), with the irrelevant or non-parasitized host taxa excluded. The topology is notas different as it appears compared tothat of Eudactylina-derived host trees (Figures66 and 67). As previously mentioned,the Eudactylina-derived cladogram contains two distinct estimates of the host cladogram. One (read from left to right) bracketted by Manta and Mustelus, is essentiallyacarcharhinid dade with a host capture to derived epipelagic myliobatids. The otherbracketted by Squalus and Urolophus is a dade infecting the Squalea. The parasite-derived host tree corroborates Shirai’s (1992b) postulate ofa paraphyletic assemblage172of shark-like squaloids, and the monophyly orexistence of the hypnosqualea, consisting of Squatina as the sister taxon to thepristiophorid plus batoid dade. Theparasitedata posits a more basal basal placementfor Squalus relative to theetmopteridsAculeola and Etmopterus than does Shirai(1 992b).The batoid relationships from Nishida(1990) show genera (in ascending order)Torpedo, Rhinobatus, and Raja existingas a paraphyletic grade. Theparasite-derivedhost tree suggests the possibility of monophyly forthese respective taxa. Ina somewhat complementary approach, Heemstra andSmith (1980) hypothesizea monophyletIc Raja plus Rhinobatus withTorpedo as the sister taxon to that dade.Nishida (1990)places both Dasyatis (Dasyatididae)and Urolophus (Urolophidae) as the sister grouptoGymnura (Gymnuridae) plus Myliobatis,Rhinoptera, Manta, and Mobula , hisMyliobatididae. The parasite-derived host tree is verysimilar except the group composed of ((Urolophus) (Gymnura, Dasyatis)) is thesister group to Myliobatis. The supposed horizontal transfer or colonizationof the eudactylinid-infecting rhinopterid-mobulid dade from the carcharhinid lineage is congruentwith the relationships proposed byNishida (1 990) with the exception that the parasite-derived hosttree yields greater specific resolution between the sampled species of Mobulaand Manta.With regard to carcharhiform relationships, a cladogramdeduced and hybridizedfrom Compagno’s (1988) many cladograms and text is presentedin Figure 116A. Anadditional cladogram derived from molecular data (Naylor, 1992)is presented in Figure11 6B. Again, non-parasitized host taxa have beenexcluded from the host trees. Keepin mind different host taxa are involved because many more squaloidsand batoids hostEudactylina than do carcharhinids, many more carcharhinids host Kroyeria thando thesqualoids and batoids, and different sets of hosts wereused from that herein, withCompagno’s (1988) and Naylor’s (1992) analyses. Host relationships derivedfromEudactylina postulate an unresolved group composed of Chiloscylliumpunctatum (ano recto Ic bid), Rhizoprionodon acutus, Carcharhinus brevipinna,Carcharhinus limbatus,and Sphyrna lewini to be the sister groupto the rest of the tree; these four all harborE. aspera. The next carcharhiniform group is composed oftriakids Mustelus andGaleorhinus. This is sister group to a dade composed of Galeocerdo which,with thedade of rhinopterid-mobulid colonizers removed, is sister taxonto a dade composed ofSphyrna which is the sister groupto a small Carcharhinus dade. Additionally, if thehost association of E. aspera on the orectolobid, Chiloscylilum represents host-parasiteco-divergence, and if the other records of E. aspera indicate colonization of thisspeciesto those four carcharhinid hosts, then the parasite-derived host phylogeny closely parallels the hybrid host tree of Compagno (1988) (Figure 116A). Theorectolobid is placedoutside the carcharhiniform group with the remaining galeomorphs. The triakidgenera173Triakis, Mustelus, and Galeorhinus though shown to be monophyleticwith the parasitetree, are hypothesized to be paraphyletic by Compagno (1988),even though heassigns them to Triakidae. Galeocerdo is the sister taxon to the sphyrnidsand carcharhinids as in the Eudactylina-derived host tree. Interestingly, the host andparasite-derived cladograms show Sphyrna tiburoas sister taxon to S. lewini and S. mokarran.Additionally, both host morphological and molecularcladograms show Carcharhinusacronotus, well separated from the more closely relatedC. plumbeus and C. obscurus,while the Eudactylina-derived host cladogram (Figure66) leaves the three as an unresolved polytomy. All three trees suggest C. obscurus and C.plumbeus are closelyrelated.The combined Kroyeria plus Kroeyerina-derived host cladogram(Figure 114)shows the holocephalan to be the sister group to the remaining membersof the tree, orto all extant elasmobranchs as Maisey (1984a) has shown. The parasite-derivedhostcladogram shows batoids to be removed from the next node and postulatessistergroup relationships of Carcharhiniformes and Lamniformes (Alopias andIsurus), asCompagno (1988) has suggested. The Kroyeria-Kroeyerina-derived host tree impliesthat the tiger shark, Galeocerdo cuvier is the most basally placed carcharhinid of thisdata set. Hence the Triakidae is situated between sphyrnids and Galeocerdo. Thisrelationship is not supported by the host morphology-based cladogram (Figure 116A) ofCompagno (1988), nor by the host molecular-based cladogram (Figure 116B) of Naylor(1992). This arises because of the many plesiomorphic character states found inKroyeria dispar and Kroyeria papillipes which are specific to Galeocerdo. Becausecladistic analyses form groups on shared derived characters (synapomorphies) there isa possibility of species or groups of species being split off near the base of a cladogrambecause plesiomorphic characters invoke no particular groupings (Lambshead andPaterson, 1986). Although there is much evidence placing Galeocerdo outside of themain body of Carcharhinus (see Compagno, 1988 and Naylor, 1992) it is currently recognized as a member of the Carcharhinidae.Lavery (1992) in his phylogenetic analysis of carcharhinids from Australia usingallozyme electrophoresis concluded Galeocerdo may be more closely related to sharksof the Hemigaleidae (weasle sharks), traditionally placed between the Triakidae andGaleocerdo (Compagno, 1988 and Maisey, 1984b). Compagno (1988) mentions thatlittle has been done to elucidate the relationships of Galeocerdo because of its distinctiveness and ubiquity which invites neglect. Applegate (1978) compared dental andexternal morphology of Galeocerdo with a few triakids. It also shares similarities withthe Hemeigaleidae with regards to the nasal fontanelles. Additionally, manyGaleocerdo characters (see Compagno, 1988 for a comprehensive list) are primitive174characters by virtue of comparison with hemigaleids, triakids,proscyllilds, scyliorhinids,and noncarcharhinoid sharks. Many of these characters makeGaleocerdo a transitional form between hemigaleids and the subfamilyCarcharhininae, and through hemigaleids the Triakidae (Compagno, 1988). Compagno (1988)adds several characters suchas the very short snout, serrated andanaulacorhizous teeth, caudal keels, basin-likerostrum, pits and keels in front of the anteriorfontanelle, great size, massive jaws,Carcharhinus-like arrangement of dorsal and anal fins,and high vertebral number areprobably derived characters, suggesting that it is specializedaway from its common origin with the rest of the carcharhinidsbut has evolved in parallel with the large macro-predatory species of Carcharhinus,Glyphis, Negaprion, and Prionace. Galeocerdo isprobably the most primitive living carcharhinid, and maybe closer to the commonancestry of all carcharhinids than any other living memberof its family. It is probablythe primitive sister group of all other carcharhinidsplus the Sphyrnidae. If theseCarcharhinus-like characteristics are in fact parallelisms orhomoplasies, and the aforementioned primitive characteristics are found in even more basallyplaced families(Proscylliidae and Scyliorhinidae) than the Triakidae, then perhapsthe KroyeriaKroeyerina-derived host cladogram placing Galeocerdo below theTriakidae describesan accurate relationship not readily embraced by currentichthyologists. The most parsimonious explanation for the basal placement of Kroyeria dispar Kroyeria papililpes,and Kroeyerina elongata, in their respective genera is one of host-parasite co-divergence, not three independent host captures by three plesiomorphic parasites, furthercorroborating the unorthodox placement of Galeocerdo in the Kroyeria-Kroeyerinaderived host cladogram. In any case, Galeocerdo appearsto remain somewhat of anenigma, basally placed within the Carcharhinidae or possibly even outsidethat family.The next dade of the Kroyeria-Kroeyerina-derived host cladogram (Figure 114)posits a monophyletic Triakidae with a monophyletic Triakis dadeas the sister group toa dade composed of a paraphyletic Galeorhinus plus Mustelus. Host morpholgy(Figure 116A) depicts a paraphyletic grade consisting of, in ascending order Triakis,Mustelus, and Galeorhinus. A dade composed of ((Sphyrna mokarran)(Sphyrna lewini,Sphyrna zygaena)) is found next in the parasite-derived tree (Figure 114).Relationships among these sphyrnids are unresolved in the morphological and molecular cladog rams.The parasite-based tree suggests that Carcharhinus is paraphyletic, includingwith it Prionace, Negaprion, and Rhizoprionodon. This placement of Prionace is ingeneral consistent with its placement in morphologically-based and molecular-based hosttrees. Furthermore, Negaprion is nested within Carcharhinus in the morphologicallybased tree, and Lavery (1992) reached a similar conclusion in an allozyme study of car-175charhinids. Morphologically-based andmolecular-based treestreat Rhizoprionodonasexternal to Carcharhinus. Thisdiscrepancy in the parasite-derivedhost tree maybe theresult of parasite colonization orhost misidentification.SUMMARY ANDCONCLUSIONSA systematic revision and phylogeneticanalysis was conductedfor 26 of the 38nominal species of Eudactyilna,and for 16 of the 18 nominalspecies of Kroyeria. Ninenew species of Eudactyilna andfive new species of Kroyeria aredescribed herein. Anew host record for Kroeyerinacortezensis from Carcharhinuslongimanus is included.Parasite-derived host phylogenies weregenerated from the recoded(additive-binary)Eudactylina and Kroyeria cladogramtopologies and combinedin part with the previouslyrevised and phylogeneticallyanalyzed Kroeyerina. Congruenthost and parasitetopologies from both holocephalanand elasmobranch hostssuggest the existence ofwell established host-parasiteassociations as early as the late Devonianapproximately400 MY.The parasite-derived host phylogeniesgenerated from Eudactylina hypothesizedthe monophyly of the squaloids,squatinids, pristiophoridsand batoids. This corroborates the revolutionary findingsof Shirai (1992a, b) in which he proposeda radicallynew systematic frameworkfor squaloids and relatedtaxa, the Squalea andHypnosqualea. Eudactylina-derivedbatoid relationships were similarto previoushypotheses of these host groups. Parasitecladograms resolved batoid species relationships previously not addressedin the literature. The parasite-derived hostphylogenies generated fromKroyeria and from Kroyeria combined with Kroeyerinaposits theparaphyly of the genera containedwithin the Carcharhinidae. Again withthe exceptance of the basal placementof Galeocerdo below the Triakidae, theparasite-derivedgrams are generally congruentwith the molecular and morphologicalhost cladogramspreviously generatedby other authors and presented herein.Parasite-derived phylogeneticrelationships of a subset of species fromSquat/naand Myiobatis indicatea speciation pattern consistent with majorvicariant events associated with the Pangaeanbreakup during the Jurassic period approximately160 MY.If taxonomic congruence (analysisof the congruence between topologies produced from independent datasets) is preferable to character congruence (analysis ofthe congruence between individualcharacters) for estimating the accuracy of phylogenetic hypotheses (Lanyon, 1993),then it appears that parasite-derived host phylogennies offer much information inconjunction with the conventionally-derived (morphologi176cal and molecular data) phylogenetic hypotheses. 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