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Intercontinental ecomorph convergence in jumping spiders (Araneae: Salticidae) Piascik, Edyta Katherine 2014

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     INTERCONTINENTAL ECOMORPH CONVERGENCE IN JUMPING SPIDERS (ARANEAE: SALTICIDAE)  by Edyta Katherine Piascik  B.A., Wilfrid Laurier University, 2005 B.Sc. (Honours), University of Toronto, 2009  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF  MASTER OF SCIENCE  in The Faculty of Graduate Studies (Zoology)  THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver)  July 2014   © Edyta Katherine Piascik, 2014 ii  Abstract  The ecomorph convergence hypothesis states that the relationship between morphology and ecology will be similar in independently evolved communities.  Molecular phylogenetic studies show that most major groups of jumping spiders (Salticidae) are primarily restricted to one continental region, indicating independent replicate radiations.  This study asks whether phylogenetically independent communities of salticids predictably converge in morphospace and how likely are they to be influenced by historical contingencies.  To test the ecomorph convergence hypothesis, 281 salticid species were collected in two comparable tropical rainforests in Ecuador and Borneo while recording habitat data.  Parsimony methods indicate that there are a minimum of 17 evolutionary transitions that have occurred between foliage, ground, and trunk microhabitats.  The transitions among all three microhabitats have occurred independently in a major clade of Euophryines, within the Freyines, within the Marpissoids, and within the Amycines.  Most of the diversification occurred within continents as the major clades are largely restricted to continental regions.  Strict criteria ensuring sufficient sample size, habitat data, and phylogenetic independence in species comparisons resulted in 36 species for all morphometric analyses.  Ecomorphs show signs of clustering in multivariate morphological space and there are significant differences in morphology between microhabitats.  Trunk salticids have flatter carapaces with shorter legs than ground salticids while foliage salticids have raised carapaces and longer legs when compared to trunk salticids.  The relationship between foliage and ground salticids is not as clear.  Foliage salticids exhibit the widest range of body forms when compared to the other two ecomorphs.  By combining morphometric measurements with molecular phylogenetic data, I show that independent origins of microhabitat use lead to similar body forms providing evidence for the ecomorph convergence hypothesis.  However, I also find differences in convergence patterns between continents in terms of the number of shifts between ecomorph origins and species composition between the three microhabitats. These results provide a rare example of a large scale study of intercontinental community-wide patterns that show a mix between convergence and contingency using a comparative phylogenetic framework while demonstrating quantitatively that large scale continental diversifications can behave as predictably as smaller diversifications.    iii  Preface  The field work was conducted as a team in which I was the co-leader in design and execution along with W. Maddison.  All specimens collected were sorted to species by W. Maddison with the help of M. Vega.  I took all morphometric photographs and measurements.  I designed the morphometric analyses with help from W. Maddison.  I performed all analyses, except the modified sign test, which was designed and conducted by W. Maddison. The phylogenetic MANOVA was conducted with help from L. Harmon.  I prepared the manuscript with input from W. Maddison.   iv  Table of Contents  Abstract ........................................................................................................................................... ii Preface............................................................................................................................................ iii Table of Contents…………………………………………………………………………………iv List of Tables .................................................................................................................................. v List of Figures ................................................................................................................................ vi Acknowledgements ....................................................................................................................... vii Introduction ..................................................................................................................................... 1 Materials and Methods .................................................................................................................... 4 Data Collection ............................................................................................................................ 4 Morphological Analyses ............................................................................................................. 5 Phylogenetic Analysis ................................................................................................................. 8 Statistical Analyses ..................................................................................................................... 9 Results ........................................................................................................................................... 12 Phylogenetic Analysis ............................................................................................................... 12 Statistical Analyses ................................................................................................................... 13 Discussion ..................................................................................................................................... 17 Bibliography ................................................................................................................................. 45 Appendices .................................................................................................................................... 50 A: List of all salticid species collected and habitat data from both Ecuador and Borneo.50 B: Variable loadings and cumulative percentage of variation from the PCA……………61 C: Plots of reconstructed specimens of small, large and average PC values for PC1…...64 D: Plots of reconstructed specimens of small, large and average PC values for PC2…...65 E: Plots of reconstructed specimens of small, large and average PC values for PC3……66 F: Plots of reconstructed specimens of small, large and average PC values for PC4……67 G: Plots of reconstructed specimens of small, large and average PC values for PC5…...68 H: Plots of reconstructed specimens of small, large and average PC values for PC6…...69 I: Plots of reconstructed specimens of small, large and average PC values for PC7…….70  v  List of Tables  Table 1. The total number and proportion of salticid species and specimens collected from each  microhabitat from Ecuador and Borneo………………………………………………….23   Table 2. List of morphometric study species and general microhabitat data…………………….24  Table 3. List of phylogenetic pairwise comparisons for trunk, foliage and ground salticids…...25 Table 4. Classification results for the discriminant function analysis…………………………...26  Table 5. Phylogenetic MANOVA results………………………………………………………..27  Table 6. List of values indicating which phylogenetic pair scored highest along principal  component scores 1-7……………………………………………………………………28 Table 7. Results of the modified sign test………………………………………………………..29    vi  List of Figures  Figure 1. Map of salticid distribution and sampling localities in Ecuador and Borneo…………30  Figure 2. Photograph of a jumping spider with body parts labelled……………………………..31 Figure 3. Photograph of geometric landmarks and linear leg measurements……………………32 Figure 4. Examples of several salticid species used in the morphometric analyses……………..33 Figure 5. Abundance plots showing how many species were represented by how many specimens  for each of the microhabitats for both Ecuador and Borneo……………………………..34 Figure 6. Phylogeny of salticid species collected from Ecuador and Borneo showing the number  of microhabitat transitions and placement of morphometric species…………………….35 Figure 7. Phylogeny of salticid species collected from Ecuador and Borneo showing  phylogenetic clustering by continent…………………………………………………….36   Figure 8. Discriminant function analysis of morphological data versus microhabitat…………..37 Figure 9. PCA for all 271 specimens from both Ecuador and Borneo…………………………..38  Figure 10. PCA for centroids of species from both Ecuador and Borneo……………………….39 Figure 11. PCA for 36 centroids of species separated by region………………………………...40  Figure 12.  X,Y coordinates for reconstructed specimens showing small, large and average PC  values for geometric landmarks on the lateral view of the carapace and linear leg measurements for PC3 and PC4…………………………………………………………41 Figure 13. Vector plots between phylogenetically paired species of ground and trunk salticids..42  Figure 14. Vector plots between phylogenetically paired species of foliage and trunk salticids..43  Figure 15. Vector plots between phylogenetically paired species of foliage and ground  salticids………………………………………………………………………………......44    vii  Acknowledgements  I would like to express my gratitude to my supervisor Wayne Maddison, for his comments, remarks and engagement throughout the learning process of this master thesis: he continually conveyed a sense of passion and excitement towards this project and towards the world of jumping spiders.  Without his guidance this dissertation would not have been possible.   I would also like to thank the members of my committee for their invaluable suggestions and guidance over the last few years.  Thank you to Luke Harmon, Dolph Schluter and Diane Srivastava for all their help.  Fieldwork in Ecuador and Borneo would not have been possible without the help of many people.  Thank you to Mauricio Vega for his involvement in collecting and sorting species from Ecuador.  Thank you to Alex Ang for his help in collecting salticids in Borneo.  To Ch’ien Lee and the staff at Mulu National Park, thank you for sharing your expertise and passion for the surrounding flora and fauna.  A special thank you is due to Jerzego corticicola, whose discovery on the majestic trees of Borneo will always bring a smile to my face.   I would further like to thank former and current members of the Maddison Lab and the Aviles Lab for their comments, suggestions and assistance along the way.  Last, but certainly not least, I thank my family and all my friends for their continued support that kept me harmonious, and who have continually incented me to strive towards my goal.   This work was funded by an NSERC grant to W.P. Maddison.1   Introduction  Phylogenetically independent adaptive radiations such as those seen in desert lizards (Melville et al. 2006), ranid frogs (Bossuyt & Milankovitch, 2000) and cichlid fishes (Young et al. 2009) allow evolutionary biologists to examine emergent evolutionary patterns.  Convergence implies that species will evolve similarities in form and function over time in response to similar environmental pressures.  On the other hand, historical contingencies suggest that repeated instances of evolutionary diversification will lead to different outcomes, regardless of whether starting conditions are similar or different (Gould 1989).  The extent to which species in similar environments but different geographic regions converge has been shown with relatively few species and primarily with island biota (Losos et al. 1998; Gillespie 2004).  Studies on mainland biota often show cases of non-convergence (Ricklefs & Schluter 1993; Price et al. 2000) or they show a mix between convergence and contingency (Melville et al. 2006; Harmon & Gibson 2006; Pinto et al. 2008).  The hypothesis of ecomorph convergence focuses on the relationship between morphology and ecology and states that this relationship will be similar in independently evolved communities (Williams 1972). To what extent do ecological communities converge through predictable evolutionary processes and how likely are they to be influenced by historical contingencies or chance? Jumping spiders (Salticidae), the most diverse spider family with 5678 described species (Platnick 2014), provide an opportunity to study the hypothesis of ecomorph convergence on a large continental scale.  Salticids are found in almost all habitats from rainforests to deserts yet are most diverse in tropical regions where they partition the environment by using many different microhabitats such as foliage, tree trunks, leaf litter, vines and suspended litter.  Initial non-2   quantitative observations demonstrate that salticids show consistent patterns in body form correlated to these microhabitats.  Salticids found on the foliage have raised carapaces with longer, thinner and often translucent legs; salticids found on tree trunks have flatter carapaces and shorter legs; salticids found on leaf litter have plump bodies with wider raised carapaces.   Extensive data involving molecular phylogenetic analyses (Maddison & Hedin 2003; Maddison & Needham 2006; Maddison et al. 2007; Maddison et al. 2008; Bodner & Maddison 2012; Zhang & Maddison 2013) provides evidence that each continental region contains salticid clades that have diversified in morphology and into many hundreds of species, largely independently from those in other regions.  This study focuses on the communities in Ecuador and Borneo to represent two of these continental regions and test for community-wide patterns of morphological convergence as it relates to microhabitat.  Ecuador and Borneo contain ideal sites to study salticid communities as they both have comparatively stable, undisturbed mature forests with high rainfall and high canopy. As there have not yet been any quantitative comparative studies linking morphology with functional traits in salticids, the aim of this study is to focus on the statistical relationship between overall body shape and microhabitat use.  This study tests the hypothesis of ecomorph convergence by examining whether community-wide patterns of convergence have occurred in assemblages of salticids within Ecuador and Borneo and how likely they are to be shaped by contingencies.  I will provide evidence for ecomorph convergence by comparing species means in morphological traits between microhabitats and also by examining whether species cluster together in multivariate morphological space.  Using habitat and combined geometric and traditional morphometric data paired with molecular phylogenetic analyses, I investigate whether 3   independent radiations of jumping spiders have similar patterns in morphology correlated to microhabitat.   4   Materials and Methods  Data Collection  Salticids were intensely sampled from a single locality in each of the two continental regions (Fig. 1).  In the Americas, the locality was Yasuní National Park in Ecuador (1.08° S, 75.91° W).  In Southeast Asia, the locality was Gunung Mulu National Park in Sarawak, Malaysia, on the island of Borneo (4.13° N, 114.91° E).  The sampling strategy was structured by microhabitat to be able to standardize person hours of effort in each microhabitat sampled which allowed for a quantification of each species’ preference for microhabitat.   “Microhabitat” refers to the subset of the habitat used by the species (i.e., the trunk of a tree).  For example, a salticid was considered a trunk specialist if we found more specimens of that species per hour effort on tree trunks than we did on any other microhabitat. On an average day, salticids were sampled from 3 sampling sites in the morning and from 3 different sampling sites in the afternoon where each individual collecting was randomly assigned a single microhabitat during the sampling session to ensure a non-biased collection.  Salticids were collected from many different microhabitats throughout the forest in order to gain a complete picture of salticid fauna from each continental region.  However for the purpose of the morphometric study, we only included specimens sampled from the three main microhabitats; foliage (all leaf sizes), tree trunks (trunks with a minimum of 10 cm diameter at breast height) and ground (including leaf litter).  The microhabitat designated as “foliage” consisted not only of leaves, but unavoidably an occasional branch and some suspended litter; however the latter was avoided whenever possible.  Specimens were collected from foliage by shaking vegetation onto a sheet, from tree trunks by sweeping trunks with a paintbrush directing material onto a sheet as well as visual searching, and from the ground by visual searching.  On an average day, each microhabitat was sampled six 5   times throughout the day for 40 minutes along 20 m wide transects approximately 50 meters long.  These transects ran along designated park trails that varied in altitude, elevation, vegetation density, proximity to a water source, and proximity to forest edge to ensure that microhabitats were sampled from a diversity of environmental conditions.      In order to obtain a record of each species collected and to help with species identification, a male and a female of each species was photographed alive in the field and then all samples collected were preserved in 95 % ethanol.  Specimens were sorted to species by morphology, with male-female matching aided by 16sND1 sequences (W. Maddison & M. Vega, unpublished).  Microhabitat data was recorded based on two different criteria for all species collected.  Criterion 1 listed species that had a minimum of 80 % of specimens collected from that particular microhabitat.  Criterion 2 was more conservative and listed species that had a minimum of 80 % of specimens collected from that particular microhabitat and also had a minimum of 4 specimens collected in total from any microhabitat.  Specimens are now stored in the Spencer Entomological Collection at the Beaty Biodiversity Museum, University of British Columbia (UBC-SEM). Morphological Analyses A species was included in the morphometric analyses only if a minimum of 80 % of its specimens collected came from one microhabitat, if there was a minimum of 4 specimens per sex per species, and if it contained a phylogenetic pairwise comparison to another morphometric species.  Because species share part of their evolutionary history, they cannot be considered independent data points (Felsenstein 1985).  Therefore pairs of phylogenetically independent species belonging to different microhabitats were chosen for analyses to ensure that any comparisons made between morphology and microhabitat were independent of any phylogenetic 6   relationships.  This phylogenetic independence was ensured by choosing species comparisons where the path between members of a pair, along the branches of the tree, did not touch the path of any other species pair (Maddison 2000).  All ant mimic species were excluded from the morphometric analyses because of the expectation of radically different selective pressures on mimics.     To summarize the variation observed in overall body shape, geometric morphometric analyses of the carapace and traditional linear measurements of the legs were combined.  A single investigator (EKP) photographed all specimens and obtained all measurements from both adult males and females.  Preserved specimens were positioned and photographed in a petri dish with white sand filled with 95 % ethanol.   The issue of projecting a three-dimensional organism, such as a jumping spider, into a two-dimensional plane must be considered to avoid the possibility of distortion.  To avoid this problem, specimens were consistently oriented in the sand under the camera using one particular plane for each of those orientations (i.e., dorsal carapace view, lateral carapace view, frontal carapace view, and sternum view).  Refer to figure 2 for a labelled photograph of a jumping spider.  For example, the dorsal carapace view was captured by positioning the specimen along the anteroposterior direction so that the posterior end of the outline of the anterior medial eye (AME) lenses was straight with only slight convexity while simultaneously ensuring the outline of the posterior end of the carapace was visible.  The same view was positioned along the mediolateral direction by ensuring that a similar proportion of the carapace was seen on either side of the posterior medial eyes (PME).  The lateral carapace view was photographed from the left side.  This view was captured by positioning the specimen along the anteroposterior direction by ensuring that only the left PME was visible.  The specimens were positioned along the 7   mediolateral direction so that both AMEs eclipse and only the left AME was visible.  The frontal carapace view was captured by positioning the specimen along the anteroposterior direction so that the AMEs and anterior lateral eyes (ALE) were not tilted in either direction and that the PMEs or posterior lateral eyes (PLE) were not visible.  The sternum was positioned along both the anteroposterior and mediolateral direction to capture an equal amount of convexity from the sternum.  Specimens were photographed under a Leica MZ16 dissecting microscope and a Leica IC D camera (images with 2088 x 1550 pixels) with Leica Application Suite version 3.1.0.  Geometric Morphometrics: Sets of aerial (n=10), lateral (nL=9, nSL=20), frontal (n=12) and ventral (n=4) anatomical landmarks (L) and semilandmarks (SL) were used to represent the carapace shape (Fig. 3A-D).  Only two-dimensional type 1 landmarks were used; this type of landmark represents a locally defined homologous point on a specimen, or in some cases it is defined in terms of structures close to that point (Bookstein 1991).  Semilandmarks are defined as points whose position along a curve is arbitrary but which provide information about curvature (Bookstein 1991).  Landmarks and semilandmarks were digitized using TPSDig 2.16 (Rohlf 2006) which converts points marked on the photographs into Cartesian x,y, coordinates.  The coordinates from each view were then superimposed by a full Procrustes fit using MorphoJ version 1.05d (Klingenberg 2011) to remove the effects of translation, rotation, and scale.   To account for the issue of missing legs or partially damaged specimens, missing values were estimated using the Bayesian Principal Component Analysis (BPCA) method with the R package bpca (Faria & Demétrio 2013) and pcaMethods (Stacklies et al. 2007).  BPCA uses an expectation maximization (EM) approach combined with a Bayesian estimation method to 8   calculate the likelihood of an estimated value for a missing data point (Stacklies et al. 2007).  The resulting landmarks were used in the proceeding analyses.  Traditional Morphometrics: Linear measurements were made on the first, third, and fourth legs of each specimen (Fig. 3 E-G) and were taken using ImageJ version 1.44 (Abràmoff, et al. 2004). The first leg had three measurements; tibia depth at the base, tibia length, and length of longest tibial spine.  The third and fourth legs had four measurements each; femur length, patella length, tibia length, and tarsus plus metatarsus length.  Legs were only measured from the left side unless they were missing or damaged, in which case they were measured from the right side of the specimen. All four geometric landmark datasets were combined with the linear leg measurements into one dataset to summarize variation in overall body shape.  However since the geometric landmark datasets had undergone a Procrustes fit, all the specimens had to be re-scaled by dividing each coordinate by the centroid size.  The resulting combined coordinates and linear measurements were then treated as variables in all analyses. Phylogenetic Analysis  A phylogeny based on two gene regions, nuclear 28s and mitochondrial 16sND1 (W. Maddison unpublished), was used to examine the continental distribution and microhabitat of sampled species in relation to phylogenetic relationships.  The placement of Agorius is not to be trusted as previous work with more genes shows that its position on the tree is not stable (Maddison et al. submitted).  Therefore it was excluded from the following phylogenetic analyses.  Parsimony methods were used on the phylogeny to determine the minimum number of evolutionary 9   transitions that have occurred from one microhabitat specialist to another as well as to determine the minimum number of evolutionary transitions that have occurred between continents.     Statistical Analyses  Several tests were performed to determine whether morphology differs between spider species in different microhabitats (i.e., microhabitat specialists).  First, a discriminant function analysis (DFA) was applied using the R package MASS (Venables & Ripley 2002) to determine whether the a priori microhabitat assemblages are identifiable morphologically.  To determine whether microhabitat specialists form discrete clusters in morphological space, a principal component analysis (PCA) was conducted.   An exploration analysis was performed to visualize the changes in shape that accompany variation along each PC axis one at a time.  This was achieved by computationally manipulating the original PC scores matrix to determine what shape a specimen would have with different values of PC scores along each of the PC axes.  The PC scores are linear transformations of the original landmark coordinates.  For each PC axis I generated the average shape and then two new specimens varying only in the single PC dimension (by adding and subtracting a single unit).  These new specimens were then back-transformed to the original landmark coordinates.  The difference between the back-transformed shapes indicates the influence of the given PC in isolation of overall shape.  The shapes of these new experimental specimens were visualized against the average shape using Excel 2010 (Microsoft 2010).    Next, mean PC scores were calculated for each species from the BPCA analysis.  A phylogenetic multivariate analysis of variance (MANOVA) was conducted on the first 7 PCs to test the hypothesis that microhabitat specialists differ in morphology using the species mean PC scores.  This test, as implemented in the R package Geiger (Harmon et al. 2008), takes into 10   account phylogenetic non independence among species.  First, the standard F-statistic and Wilk’s lambda are calculated and then the null distribution was obtained by running 10,000 simulations on the phylogenetic tree, trimmed to include only the morphometrics taxa, under a Brownian motion model. A table of values and vector plots were created to determine whether there is consistency in which microhabitat specialist of each phylogenetic pair scored highest along each of the PC axes (PCs 1-7).  If a representative of a habitat within a group consisted of a set of closely related species rather than a single species, as with Laufeia trunk salticids, then reconstructed ancestral states were used for the set of species.  If the set consisted of two species, then the average was used.  If the set consisted of more than two species, then ancestral states were reconstructed using square change parsimony on the small clade isolated from the rest of the phylogeny, using Mesquite (Maddison & Maddison 2013). To examine whether microhabitat specialists had consistent differences in their values of principal components, we conducted a modified sign test of the null expectation that for a given PC axis the average difference between spiders from two microhabitat types is in either direction with equal frequency — e.g., a ground salticid is as likely to have a smaller as a larger value of a principal component than its related trunk salticid.   Thus, the 7 phylogenetically independent pairwise comparisons between ground and trunk salticids should be distributed binomially in their number in which the ground's value in a PC is larger than the trunk's.  However, as a sample size of 7 is too small to test for individual effects of ecomorph on a single PC, the data were pooled across principal components.  Two test statistics were compared to their null distributions simulated by a Python program written by W. Maddison.  The first test statistic ("sum smaller") was the sum of the smaller numbers in the comparisons; e.g., if PC1 had ground greater than 11   trunk in 2 of the pairs and trunk greater in 5 of the pairs, and PC2 had ground greater than trunk in 6 of the pairs and trunk greater in 1 of the pairs, then the sum of smaller numbers is 2 + 1 = 3.  The second test statistic ("sum P") summed the two tailed P values for each PC comparison from a sign test.  For the combined analysis, a similar method was used except that instead of finding the average difference between spiders from just two microhabitats, the sum smaller or sum P was calculated over all 21 comparisons – 7 PCs for ground and trunk salticids, 7 PCs for foliage and trunk salticids, 7 PCs for ground and foliage salticids.  The p-values were generated by examining the number of simulated cases when the test statistic was small or smaller than that calculated for our data.  The simulations simulated coin flips to construct comparisons of the same number and sample sizes as the observed against which they are comparing.  Simulation sample size is 10,000 for analyses involving specific comparisons of two microhabitat specialists and a simulation sample size is 100,000 for a combined analysis of all three microhabitat specialists.   12   Results  A total of 281 species were collected from two different continents: 168 species from Yasuní National Park, Ecuador and 113 species from Gunung Mulu National Park, Borneo.  A list of all species collected with microhabitat data is provided in appendix 1.  Figure 4 shows examples of salticid species used in morphometric analyses, organized by clade.  Abundance plots demonstrate how many species were represented by how many specimens for each of the microhabitats and for each of the continents (Fig.5).  Of the 281 species collected in total, 207 species had clear microhabitat data (at least 80 % of all specimens were collected from a single microhabitat).  When the number of specimens available was considered (minimum 4 specimens collected from any microhabitat), 90 species had clear microhabitat data with a sufficient sample of measureable specimens.  Among these, the morphometric species were chosen after confirming that they had sufficient numbers of males and/or females and matched the phylogenetic criteria as discussed below.    A count of the total number and proportion of salticid species and specimens collected allows for comparisons in species composition between continents.  Aside from the ground microhabitat, there are differences between Ecuador and Borneo in the proportion of species and specimens collected from each of the three microhabitats (Table 1).  The proportion of the total species (74.8 %) and specimens (64.8 %) collected from the foliage microhabitat was greater in Ecuador than in Borneo.  The trunk microhabitat had twice as many species (32.4 %) and three times as many specimens (45.3 %) collected in Borneo than in Ecuador.        Phylogenetic Analysis  The phylogenetic distribution of species in various microhabitats indicates that there are a minimum of 17 evolutionary transitions that have occurred between microhabitats (Fig. 6).  The 13   transitions among all three microhabitats have occurred independently in a major clade of Southeast Asian Euophryines and several times within the remainder of the Euophryines, within the Freyines (New World), within the Marpissoids, and within the Amycines (New World).  We cannot say for certain the direction of these transitions, but according to parsimony reconstruction the foliage microhabitat is ancestral with 9 independent transitions to the ground microhabitat (6 are from Ecuador and 1 from Borneo), as well as 8 independent transitions to the trunk microhabitat (4 are from Ecuador and 3 are from Borneo).  As expected, most of the diversification occurred within continents (Fig. 7).  This is evident when examining the major clades that are restricted to continental regions such as with the Amycines and the Freyines.  Even those major clades that do contain species in both continents like the Euophryines show phylogenetic clustering by continent within the subgroups.  When considering continent as a character, parsimony methods show 14 evolutionary steps.  This number of steps would be far fewer than expected were you to assume no relationship with a random phylogenetic placement of the species because under randomization with 100,000 replicates, the lowest number of evolutionary steps was found to be 53.    A total of 36 species that fit the previously outlined criteria were used for morphometric analyses: 20 species from Yasuní National Park, Ecuador and 16 species from Gunung Mulu National Park, Borneo, as listed in Table 2.  The phylogenetically paired morphometric species are listed by clade in Table 3.      Statistical Analyses  The results of the non-phylogenetic discriminant function analysis reveal that the first discriminant function axis achieves 60.9 % of the separation between ecomorphs while the second discriminant function axis improves separation by another 39.1 % (Fig.8).  The re-14   classification results based on a cross-validation procedure correctly classified foliage ecomorphs by 77.1 %, ground ecomorphs by 71.4 % and trunk ecomorphs by 80.6 % (Table 4).   The first 7 axes of the PCA explained 92 % of the variation among specimens.  The loadings and cumulative proportion of variation for these axes are reported in appendix 2.  The foliage ecomorphs occupy the largest area and overlap with the trunk and ground ecomorphs in multivariate morphological space (Fig.9, Fig.10).  Although there is scatter, the trunk and ground ecomorphs tend to cluster in different positions in morphological space.  This pattern persists when species centroids are plotted separately for Ecuador and Borneo (Fig. 11).  Foliage ecomorphs overlap with the trunk and ground ecomorphs and also occupy the largest area in morphological space.  Trunk ecomorphs cluster in a smaller area and lie consistently in lower values along PC3 when compared to the ground ecomorphs.   The exploration analysis, showing the changes in shape that accompany variation along each PC axis, revealed that the first principal component is dominated by size.  The second principal component shows that a variation from small to large values of PC scores is associated with an increase in carapace width and increase in the length of the first and third legs.  The third and fourth principal components show that a variation from small to large values of PC scores is associated with an increase in carapace height as well as an increase in the first leg (tibia depth) and decrease in the length of the third (femur) and fourth leg (tibia and tarsus and metatarsus) lengths (Fig. 12).  Variation along the fifth principal component appears less clear, however the variation from small to large values of PC scores is associated with an increase in carapace height as well as an increase in the length of the third (tarsus and metatarsus) and fourth (femur) legs.  The sixth principal component is associated with an increase in carapace width, from the small to large PC score spectrum, as well as an increase in the tibial spine length on the first leg.  15   Finally the seventh principal component reflects an increase in carapace width from the small to large PC score spectrum, although not as clearly as the other components, and also describes an increase in the tibial depth of the first leg and overall length of the third leg.  Plots illustrating PC 1-7 for all views are shown in appendix 3-9. A phylogenetic MANOVA supports the hypothesis that there is a significant difference in morphology among ecomorphs (Wilks’ λ=0.37, F14,54=2.45, P=0.04) (Table 5).   In addition, ecomorphs have consistent patterns of variation along each PC axis (Table 6).  Plotting the phylogenetically paired species along PC3 and PC4 for each ecomorph comparison revealed interesting patterns between body shape and microhabitat.  When plotting phylogenetically paired species of ground and trunk ecomorphs, the trunk body form consistently moves from positive to negative values along PC3 and PC4 which implies that trunk ecomorphs have a flatter carapace and shorter first legs than the third and fourth legs when compared to the ground ecomorphs (Fig. 13).  A similar pattern occurs between phylogenetically paired species of foliage and trunk ecomorphs along PC3, which also implies that trunk ecomorphs have a flatter carapace and shorter first legs than the third and fourth legs when compared to foliage ecomorphs (Fig. 14).  The pattern between foliage and ground ecomorphs is not as consistent within all phylogenetic pairs however there still is a trend for foliage ecomorphs to have increased values along PC3 which implies that the foliage body form consists of a raised carapace with longer legs when compared to ground ecomorphs (Fig. 15).   The modified sign test did not show any significant patterns between pairs of ecomorphs (ground vs trunk, P=0.15; trunk vs foliage, P=0.17; foliage vs ground, P=0.16), but it did show significant patterns in PC values when comparing phylogenetic pairs between all the ecomorphs combined (Table 7).  When all three ecomorphs are combined to ask whether ecomorph 16   comparisons show a more uneven distribution of contrasts in PC values than you would expect by chance, the pattern is significant (P=0.039).     17   Discussion  There is evidence for ecomorph convergence within salticid assemblages in Ecuador and Borneo as ecomorphs do show signs of clustering in multivariate morphological space and there are significant differences in morphology correlated to microhabitat.  However, there is a fair bit of scatter in the data as the habitat groups in morphospace are not very distinct.  Additionally, there is inconsistency in the patterns between continents regarding the number of shifts between ecomorph origins and the number of species within the three microhabitats.  Ground and trunk ecomorphs show the least amount of overlap when all three ecomorphs are plotted in multivariate morphological space.  In addition, there is a general pattern for trunk ecomorphs to show the least amount of variation in morphological space.  Trunk ecomorphs have consistently lower values of PC3 relative to the ground ecomorphs, and this pattern is consistent in both Ecuador and Borneo.  However, foliage ecomorphs appear to occupy the largest area in morphological space showing little signs of clustering.  But why might this be the case for foliage ecomorphs?  Initial field observations show that salticids found on foliage (of all sizes) tend to display a wider diversity in body forms.  Some species living on foliage appear extremely flat with short legs and others have raised carapaces with long, thin, translucent legs.  One way to explain this pattern is through the microhabitat’s heterogeneity: the foliage microhabitat includes branches, leaves, and suspended litter.  Salticids may have evolved specific morphological traits for each of the microhabitats that are contained within what has been classified as ‘foliage’ habitat.        The phylogenetic MANOVA reveals a significant difference in morphology between ecomorphs.  Further, although the modified sign test did not reveal significant patterns in contrasts between PC values when making specific ecomorph comparisons (i.e., ground versus 18   trunk ecomorphs), there are significant patterns when all three ecomorphs are combined in the analysis.  The lack of significance in specific ecomorph comparisons may be because the sample size for specific ecomorph comparisons was too low to detect any significant patterns and future studies could account for this low power by increasing the number of phylogenetic pairwise comparisons.     When PC values are examined between the phylogenetic pairs of ecomorphs (table 6), interesting trends in body form patterns are revealed that tend to match initial observations in the field.  The lower values in PC3 for trunk ecomorphs (when compared to ground ecomorphs) indicate that trunk ecomorphs have flatter carapaces with shorter first legs when compared to the third and fourth legs.  These salticids live on tree trunks that range from flaky rough bark to smooth bark on trees in open and closed forest areas.  Having a flatter body with shorter first legs could benefit a salticid in this environment for a variety of reasons including crypsis which would allow a spider to use the bark as shelter and be better capable of moving around and blending into a flat trunk surface to avoid being seen by predators.  Ground ecomorphs, conversely, have more of a plump body form with raised carapaces and longer legs.  Selective pressures of living in a highly dimensional habitat, such as leaf litter, may have resulted in a raised body form with longer legs to help with moving around in their environment.  Foliage ecomorphs exhibit greater values in PC3 when compared to trunk dwellers indicating raised carapaces and longer legs yet also exhibit lower values in PC3 when compared to ground dwellers indicating a flatter carapace and shorter legs.  This lack of clear pattern may be attributed to heterogeneity in the foliage microhabitat as previously mentioned.  Salticids living on large, broad leaves may have evolved longer legs to allow them to move quickly to find food or escape predators, whereas on smaller leaves or on branches, selection for longer legs is not as 19   strong.  Also, perhaps body form is also being driven by sexually selective pressures as males are more visible to females in a foliage habitat, whereas the signalling between sexes may fail under the ecological conditions of ground and trunk habitats.  Previous studies have shown that the extent and form in sexual dimorphism may vary among ecomorphs (Butler 2007) and so this may explain some of the variation in salticid foliage ecomorphs, however this idea may only explain differences in males and would be worth investigating.   These patterns provide evidence that there is convergence in morphology correlated with microhabitat in salticids within continents; however it is not as clear as to whether these patterns are convergent between continents.  Similarities and differences can be examined on two levels; patterns in morphology and patterns in species composition between Ecuador and Borneo.   In terms of morphological patterns, it appears that multiple lineages of salticids have invaded all three microhabitats in both Ecuador and Borneo (Fig. 6).  The phylogenetic distribution of species indicates that there have been a minimum of 17 evolutionary transitions that have occurred between the microhabitats and these transitions have occurred independently in major clades that are largely restricted to each of the continents.  However, there are inconsistencies when comparing the shifts in ecomorph origins between phylogenetically paired species in Ecuador and Borneo.  Trunk ecomorphs from Ecuador have consistently lower values of PC3 compared to ground ecomorphs whereas trunk and ground ecomorphs from Borneo show the opposite pattern.  A similar inconsistency is seen when comparing foliage to trunk ecomorphs as well as foliage to trunk ecomorphs.  However although shifts in habitat origin have occurred many times, there are only a few cases to compare the shifts in ecomorph origin between the continents which may make it difficult to interpret similarities and differences.   20   In terms of species composition, a count of the total number of salticid species collected in each of the continents reveals that species composition in different microhabitats is similar (table 1).  The largest number of species was collected from the foliage microhabitat, followed by the trunk microhabitat and the ground microhabitat, respectively, for both Ecuador and Borneo.  However, with the exception of the ground microhabitat, there are differences between Ecuador and Borneo in the proportion of species and specimens collected from each of the three microhabitats.  The proportion of the total species and specimens collected from the foliage microhabitat was greater in Ecuador than in Borneo.  However this difference in foliage species may be due, again, to habitat heterogeneity implying that in fact multiple microhabitats were sampled which would facilitate the collection of more species.  There were also twice as many species and specimens collected from the trunk microhabitat in Borneo than there were in Ecuador.  Perhaps this difference is due to a single radiation of Laufeia that dominate the trunk microhabitat leaving fewer unoccupied niches for other species to fill.  Another possibility is that it is simply due to differences within the trunk microhabitat that provides more unfilled niches.   In studies of convergent evolution, convergence indicates determinism whereby lack of convergence implies the role of contingency or chance, where taxa may respond differently to unique historical events and differences in environments in different areas.  As the idea of ecological opportunity - the origin of a key innovation or extinction of antagonists through the entry into a new environment (Schluter 2000; Losos & Mahler 2010) - appears to be one of the underlying mechanisms driving convergent evolution (Schluter 2000), it is expected that as ecological opportunity varies, so will the degree of selective pressures that drive convergent evolution.  Studies have shown that island versus mainland environments provide different selective pressures that may produce alternative evolutionary outcomes in lineages that have 21   evolved independently (Harmon & Gibson 2006; Schaad & Poe 2010).  Among the best studied island radiations are Anolis lizards that have evolved convergently multiple times in the Caribbean but have not evolved in parallel on the mainland (Schaad & Poe 2010).   Island ecosystems represent an environment with differences in competitors, predator abundance, dispersal ability, food resources, and greater selective pressure as a result of less geographic space which can provide explanations for greater competition (MacArthur & Wilson 1967; Schluter 1988, Irschick et al 1997; Losos & Ricklefs 2009).  Salticids represent one of the few examples of independent radiations deviating away from the common island biota.  The independent radiations in mainland communities of salticids provide evidence for convergence within continents yet also show some differences between continents.  This mix between convergence and contingency allows us to speculate as to the importance of competition in driving divergence in this system.  Resource-induced competition may be driving adaptive divergence of salticid body forms however due to the variable landscapes and larger geographic area of the continents, the selective pressures may not be as strong which in turn may produce different evolutionary outcomes in Ecuador and in Borneo. Future work might also link these morphology and microhabitat relationships to understand a wide variety of other important ecological and evolutionary factors such as size effects and divergence time. For example, although the current study reveals general patterns in body form correlated to microhabitat, preliminary non-quantitative observations suggest that these patterns in smaller salticid species are not as distinct as in larger species within each ecomorph category.  There has been some focus on size and allometry in other cases of ecomorph convergence (Butler & Losos 2002) and so it would be interesting to further examine 22   the adaptive radiation of jumping spiders and determine whether ecomorphological divergence is size-dependent.   In addition to potential size-dependency, it would be interesting to further study divergence time and how it affects convergence.  As previously mentioned, some salticid species have a weaker correlation between morphology and microhabitat than other species within the same ecomorph category.  Why have certain species among ecomorphs not evolved similar body forms related to microhabitat?  Studies have shown that divergence times may have important consequences for ecomorphological divergence (Kozak et al. 2005).  Perhaps the evolution of microhabitat-specific body forms requires time to reach equilibrium of adaptation to the environment.  Future studies could examine the large scale adaptive radiation in jumping spiders and determine whether divergence time can explain why certain species have a weak correlation between morphology and microhabitat.   Simpson (1953) stated that adaptive radiations might explain all of life’s diversity.  Examples of convergent evolution suggest that natural selection can often produce predictable evolutionary outcomes (Losos et al. 1998; Melville et al. 2006; Schluter 2000) which continue to make adaptive radiations a key component to the study of evolution.  The current study, demonstrating signs of convergent evolution, contributes to salticid taxonomy by providing a more complete picture of a tropical salticid fauna than any previous study has presented.  Previously there has been almost no data on habitat, ecology, or natural history of salticids.  By combining the habitat and morphometric data along with phylogenetic placement of these two communities, we can begin to understand the patterns and processes responsible for the evolutionary diversification in salticids.  23   Table 1. The total number and proportion (%) of salticid species and specimens collected from each of the microhabitats from Ecuador and Borneo.  A species was considered as part of a microhabitat if a minimum of 80 % of the specimens collected came from that one microhabitat.  ‘Other’ includes species that did not meet the 80 % criterion, species that were collected outside the main foliage, ground or trunk microhabitats as well as species collected outside of the timed sampling for foliage, ground, and trunk microhabitats.     Ecuador Borneo Habitat Species % Species Specimens % Specimens Species % Species Specimens % Specimens Foliage 98 74.8 569 64.8 41 55.4 231 37.1 Trunk 20 15.3 131 14.9 24 32.4 282 45.3 Ground 13 9.9 178 20.3 9 12.2 110 17.6 Other 37 - -   39  -   Total 168 100 878 100 113 100 623 100       24   Table 2. List of morphometric study species, number of specimens used in morphometric analyses and general habitat from which they were collected; trunk (T), ground (G), and foliage (F).   Genus species N general habitat Ecuador        Amphidraus ADBRG 4 T   Amphidraus ADYEL 10 T   Amycine AABBT 4 F   Amycus AMRBG 4 G   Amycus AMSLV 4 G   Amycus AMSVS 8 F   Balmaceda BMSTN 4 T   Capidava CAPLG 11 G   Capidava CAPSM 9 G   Chira CRWHT 5 F   Corythalia COYLF 9 G   Encolpius ENBND 9 T   Encolpius ENDRK 5 T   Eustiromastix EUYHS 4 F   Freya FRTYF 5 T   Galianora sacha 10 F   Kalcerrytus KCBMP 9 G   Laufeia SOBAR 8 G   Metaphidippus MPBRY 4 F   Noegus NGPAL 10 F Borneo        Bavia BVBND 4 F   Chalcotropis EMGRB 9 G   Colyttus striatus 10 F   Epeus EPGRG 9 F   Idastrandia WHSPS 9 G   Laufeia BBLCK 4 T   Laufeia BKYEL 10 T   Laufeia BNBRK 4 T   Laufeia FZLIN 10 F   Laufeia PXGRY 10 T   Laufeia RHRLQ 10 T   Laufeia SMCOL 4 T   Mintonia WFNCH 10 T   Pristobaeus TRLNG 4 T   Pristobaeus TRSTR 5 T   Tisaniba TSBLG 5 G 25   Table 3. List of phylogenetic pairwise comparisons for trunk (T), foliage (F), and ground (G) salticids. Trunk vs Ground Foliage vs Ground Foliage vs Trunk Group Species  Ecomorph Group Species  Ecomorph Group Species  Ecomorph                Amycine Encolpius ENBND T Amycine Amycus AMSVS F Lapsiine Galianora sacha F Amycine Encolpius ENDRK T Amycine Amycus AMSLV G Spartaeine Mintonia WFNCH T Amycine Amycus AMRBG G           Amycine Amycus AMSLV G Amycine Amycine AABBT F Amycine Noegus NGPAL F      Amycine Amycus AMRBG G Amycine Encolpius ENBND T Marpissoid Balmaceda BMSTN T      Amycine Encolpius ENDRK T Marpissoid Tisaniba TSBLG G Freyine Chira CRWHT F           Freyine Kalcerrytus KCBMP G Marpissoid Metaphidippus MPBRY F Freyine Freya FRTYF T Freyine Capidava CAPLG G Marpissoid Balmaceda BMSTN T Freyine Kalcerrytus KCBMP G Freyine Capidava CAPSM G      Freyine Capidava CAPLG G      Freyine Eustiromastix EUYHS F Freyine Capidava CAPSM G Baviine Bavia BVBND F Freyine Freya FRTYF T      Marpissoid Tisaniba TSBLG G      Euophryine Amphidraus ADYEL T      Euophryine Laufeia FZLIN F Euophryine Amphidraus ADBRG T Euophryine Colyttus striatus F Euophryine Laufeia BBLCK T Euophryine Laufeia SOBAR G Euophryine Chalcotropis EMGRB G Euophryine Laufeia BKYEL T           Euophryine Laufeia BNBRK T Euophryine Laufeia BBLCK T Plexippoid Epeus EPGRG F Euophryine Laufeia PXGRY T Euophryine Laufeia BKYEL T Nannenine Idastrandia WHSPS G Euophryine Laufeia RHRLQ T Euophryine Laufeia BNBRK T      Euophryine Laufeia SMCOL T Euophryine Laufeia PXGRY T           Euophryine Laufeia RHRLQ T           Euophryine Laufeia SMCOL T           Euophryine Chalcotropis EMGRB G                          Euophryine Pristobaeus TRSTR T           Euophryine Pristobaeus TRLNG T           Euophryine Corythalia COYLF G                          Spartaeine Mintonia WFNCH T           Nannenine Idastrandia WHSPS G             26    Table 4. Classification results for the cross-validation testing procedure for the first two discriminant functions for three ecomorphs; foliage, ground and trunk.    Predicted group membership   Ecomorph Foliage Ground Trunk Total Count      Foliage 74 18 4 96 Ground 17 55 5 77 Trunk 15 4 79 98 Percent      Foliage 77.1 18.8 4.1 100.0 Ground 22.1 71.4 6.5 100.0 Trunk 15.3 4.1 80.6 100.0 27    Table 5. Results of a phylogenetic MANOVA with 10,000 simulations run on the phylogeny.      Df Wilk's approx-F num-Df den-Df Pr(>F) Pr(phy) Habitat 2 0.37 2.45 14 54 0.009 0.04* Residuals 33                28   Table 6. List of values indicating which phylogenetic pair scored highest along principal component scores (PC) 1-7 for trunk (T), ground (G) and foliage (F) salticids. Average vector is calculated by finding the difference in PC values between all ecomorph pairs for each PC and then averaging those values.   PC T>G G>T Avg. Vector F>G G>F Avg. Vector F>T T>F Avg. Vector 1 3 4 0.0973 2 4 -0.6131 1 4 -0.1611 2 3 4 -0.0697 2 4 -0.3615 4 1 -0.0478 3 2 5 -0.5058 1 5 -0.9008 4 1 0.6217 4 4 3 0.3436 3 3 0.0845 2 3 0.1600 5 5 2 0.3261 2 4 -0.2411 1 4 -0.4283 6 5 2 0.2745 2 4 -0.2794 1 4 -0.4802 7 7 0 1.1796 6 0 1.5361 2 3 0.1634 29   Table 7.  Results of the modified sign test of the null expectation that for a given principal component (PC) axis the average difference between spiders from different microhabitat types is in either direction with equal frequency for ground (G), trunk (T), and foliage (F) microhabitats.  X is the number of pairs for which one type is greater than the other.  P values were generated by comparing the observed sumX against sumX’s and sumP’s.    Observed SumX Observed SumP P value for SumX P value for SumP Simulation Sample Size PC G vs T vs F 36 12.25 0.045 0.039 100,000 1-7 G vs T vs F 30 10.19 0.036 0.032 100,000 2-7 G vs T 12 3.38 0.146 0.150 10,000 2-7 G vs F 10 3.31 0.165 0.158 10,000 2-7 T vs F 8 3.50 0.264 0.168 10,000 2-7 30    Figure 1. (A) Geographic distribution of salticids with each colour representing an independent radiation.  Geographic sampling localities denoted in dark green; Yasuní National Park, Ecuador (B) and Mulu National Park, Borneo (C).31    Figure 2. Photograph of a jumping spider.  Eyes are labelled as follows: anterior medial eyes (AME), anterior lateral eyes (ALE), posterior lateral eyes (PLE), and posterior medial eyes (PME).  Specimen shown is Parabathippus shelfordi.   32    Figure 3. Photograph of a salticid specimen denoting geometric landmarks and linear leg measurements.  A. Landmarks collected on the carapace in aerial view (N=10). B. Landmarks and semilandmarks collected on the carapace in lateral view (NL=9, LSM=20). C. Landmarks collected on carapace in frontal view (N=12).  D. Landmarks collected on the sternum (N=4).  E-G. Linear leg measurements collected on the first, third and fourth legs, respectively.   33    Figure 4. Examples of several salticid species used in the morphometric analyses.  Species are organized by habitat (foliage, ground, trunk) and by clade: (A) Amycines, (B) Freyines, (C) Spartaeine, (D) Euophryines.       34     Figure 5. Abundance plots showing how many species were represented by how many specimens for each of the microhabitats and for each of the continents, Ecuador (A) and Borneo (B).  Specimens were considered part of a habitat class if more than 80% of the total specimens collected came from that habitat. 35    Figure 6. Phylogeny of salticid species collected from Ecuador and Borneo showing the number of habitat transitions and placement of morphometric species (extended colour bars).  The phylogeny continues from left to right.  The phylogeny was constructed using all salticid species, based on two genes, 28s and 16sND1, and then trimmed to include only species collected from Ecuador and Borneo.  Phylogeny from W.P. Maddison (unpublished). 36    Figure 7. Phylogeny of salticid species collected from Ecuador and Borneo showing phylogenetic clustering by continent.  The phylogeny continues from left to right.  The phylogeny was constructed using all salticid species, based on two genes, 28s and 16sND1, and then trimmed to include only species collected from Ecuador and Borneo.  Phylogeny from W.P. Maddison (unpublished).37     Figure 8. Discriminant function analysis of the three ecomorphs, foliage (F), ground (G) and trunk (T) salticids, on the combined geometric landmark and linear leg measurement dataset for 36 species from both Ecuador and Borneo.   38    Figure 9.  Principal component analysis of the three ecomorphs, foliage (F), ground (G) and trunk (T) salticids, on the combined geometric landmark and linear leg measurement dataset for all 271 specimens from both Ecuador and Borneo.   39    Figure 10.  Principal component analysis of the three ecomorphs, foliage (F), ground (G) and trunk (T) salticids, on the combined geometric landmark and linear leg measurement dataset for centroids of 36 species from both Ecuador and Borneo.     40     Figure 11.  Principal component analysis of the three ecomorphs, foliage (F), ground (G) and trunk (T) salticids, on the combined geometric landmark and linear leg measurement dataset for 36 centroids of species separated by region; Borneo (BOR) and Ecuador (ECU).  41    Figure 12.  X,Y coordinates for reconstructed specimens showing small, large and average PC values for geometric landmarks on the lateral view of the carapace for PC3 (A) and PC4 (B) and linear leg measurements for PC3 (C) and PC4 (D).  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg.  42    Figure 13. Vector plots between phylogenetically paired species of ground (G) and trunk (T) salticids along PC3 and PC4 axes.   43    Figure 14. Vector plots between phylogenetically paired species of foliage (F) and trunk (T) salticids along PC3 and PC4 axes.   44     Figure 15. Vector plots between phylogenetically paired species of foliage (F) and ground (G) salticids along PC3 and PC4 axes.  45   Bibliography  Abràmoff, M.D., Magalhães, P.J. & Ram, S.J. (2004) Image Processing with ImageJ. Biophontics International, 11 (7), 36-42. Bodner, M.R. & Maddison, W.P.M. (2012) The biogeography and age of salticid spider radiations (Araneae: Salticidae). Molecular Phylogenetics and Evolution. 65, 213-240. Bookstein, F.L. (1991) Morphometric tools for landmark data: geometry and biology. Cambridge University Press, Cambridge. Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. 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Molecular Phylogenetics and Evolution. 68, 81-92.  50   Appendices Appendix A. List of all salticid species collected with habitat data from both Ecuador (ECU) and Borneo (BOR).  Criterion 1 includes species that had at least 80% of the specimens collected from one of the habitats.  Criterion 2 includes species that had at least 80% specimens collected come from one habitat with a minimum of 4 specimens collected in total.  Habitat denotes the main habitat from both criteria.  The number of specimens collected from the main habitat is listed as a percentage of the total number of specimens collected from all habitats.  Species that were collected from outside of the foliage, ground or trunk habitats or species that did not meet criterion 1 were denoted by ‘NA’. Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Amycine Acragas HY6EX NA NA NA NA ECU Amycine Acragas HY6SP foliage 100  ECU Amycine Acragas HY8SP NA NA NA NA BOR Agoriine Agorius AGBL1 ground 100  BOR Agoriine Agorius AGBL2 NA NA NA NA BOR Agoriine Agorius AGMTW foliage 100  BOR Agoriine Agorius AGSLT ground 100  ECU Freyine Akela AKGRA NA NA NA NA ECU Freyine Akela AKTHN foliage 100  ECU Euophryine Amphidraus ADBRG trunk 100  ECU Euophryine Amphidraus ADDKB trunk 89  ECU Euophryine Amphidraus ADYEL trunk 92  ECU Amycine amycine genus AABBT foliage 100  ECU Amycine amycine genus AABWH foliage 100  ECU Amycine amycine genus AASBW NA NA NA NA ECU Amycine amycine genus AASTR foliage 100  ECU Amycine Amycus AMDKS foliage 100  ECU Amycine Amycus AMFBP foliage 100  ECU Amycine Amycus AMRAP foliage 85  51   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Amycine Amycus AMRBG ground 100  ECU Amycine Amycus AMRBL ground 100  ECU Amycine Amycus AMRBP ground 100  ECU Amycine Amycus AMRED foliage 100  ECU Amycine Amycus AMSLV ground 100  ECU Amycine Amycus AMSVS foliage 96  BOR Plexippoid Anarrhotus AHOGL foliage 100  BOR Plexippoid Anarrhotus ANMUL foliage 100  BOR Plexippoid Anarrhotus sp. NA NA NA NA ECU Other Amycoida Arachnomura HTCHV trunk 100  ECU Other Amycoida Arachnomura HTLIN trunk 100  ECU Freyine Asaracus CRCOP foliage 100  ECU Marpissoid Ashtabula ASHTB foliage 100  ECU Marpissoid Balmaceda BMFZZ trunk 100  ECU Marpissoid Balmaceda BMLNG trunk 100  ECU Marpissoid Balmaceda BMSTN trunk 100  BOR Baviine Bavia BVBGB foliage 100  BOR Baviine Bavia BVBML NA NA NA NA BOR Baviine Bavia BVBND foliage 100  BOR Baviine Bavia BVDUC foliage 100  BOR Baviine Bavia BVMDB foliage 100  BOR Baviine Bavia BVMTT trunk 100  BOR Baviine Bavia BVSMB foliage 100  ECU Marpissoid Beata BEAYA foliage 100  ECU Euophryine Belliena BELLI foliage 91  ECU Marpissoid Bellota BGREY NA NA NA NA ECU Marpissoid Bellota BGROR foliage 100  BOR Plexippoid Bianor BIMUL ground 100  52   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Other Amycoida Breda BRGRY foliage 100  ECU Other Amycoida Breda BRORG NA NA NA NA BOR Plexippoid Burmattus pococki NA NA NA NA ECU Freyine Capidava CAPLG ground 98  ECU Freyine Capidava CAPSM ground 100  BOR Salticine Carrhotus CRBCH NA NA NA NA BOR Salticine Carrhotus CRFOR NA NA NA NA BOR Heliophanine cf. Heliophanus sp. NA NA NA NA ECU Euophryine cf. Laufeia NEOLF trunk 100  BOR Astioid cf. Porius GNBLK foliage 100  BOR Marpissoid cf. Psecas CFBAV trunk 100  BOR Euophryine Chalcotropis EMGRB ground 100  BOR Euophryine Chalcotropis EMSTR NA NA NA NA ECU Euophryine Chapoda CHAWS foliage 100  ECU Freyine Chira CR2SP NA NA NA NA ECU Freyine Chira CRRBE NA NA NA NA ECU Freyine Chira CRRST foliage 100  ECU Freyine Chira CRSDF foliage 100  ECU Freyine Chira CRSMP foliage 100  ECU Freyine Chira CRWHT foliage 100  ECU Marpissoid Chirothecia CHIRO NA NA NA NA BOR Euophryine Colyttus CLCSP foliage 100  BOR Euophryine Colyttus CLCSR foliage 100  ECU Euophryine Corythalia COBBW foliage 100  ECU Euophryine Corythalia COBFC NA NA NA NA ECU Euophryine Corythalia COBFL NA NA NA NA ECU Euophryine Corythalia COBWT foliage 100  ECU Euophryine Corythalia CODUS trunk 100  53   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Euophryine Corythalia COGLD trunk 100  ECU Euophryine Corythalia COTNY foliage 100  ECU Euophryine Corythalia COYBN foliage 100  ECU Euophryine Corythalia COYHQ trunk 100  ECU Euophryine Corythalia COYLF ground 100  ECU Other Amycoida Cotinusa CSBLK foliage 100  ECU Other Amycoida Cotinusa CSCLW NA NA NA NA ECU Other Amycoida Cotinusa CSTIP NA NA NA NA ECU Other Amycoida Cylistella CLGRY foliage 100  BOR Euophryine Cytaea CYTMU foliage 100  BOR Euophryine Cytaea CYTSM trunk 100  BOR Euophryine Emathis EMFLU foliage 100  BOR Euophryine Emathis EMRUS foliage 100  BOR Euophryine Emathis EMWHB foliage 100  ECU Amycine Encolpius ENBND trunk 95  ECU Amycine Encolpius ENDRK trunk 100  ECU Amycine Encolpius ENSDK trunk 100  ECU Amycine Encolpius ENSPG trunk 100  BOR Plexippoid Epeus EPERA foliage 100  BOR Plexippoid Epeus EPGRG foliage 100  BOR Plexippoid Epeus EPORR foliage 100  BOR Plexippoid Epeus EPPAI foliage 100  ECU Other Amycoida Erica EIDGO foliage 100  ECU Other Amycoida Erica EIHZO foliage 100  ECU Freyine Eustiromastix EUINT NA NA NA NA ECU Freyine Eustiromastix EUYHS foliage 100  BOR Plexippoid Evarcha EVBNZ NA NA NA NA BOR Plexippoid Evarcha EVCIR NA NA NA NA 54   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Other Amycoida Fluda FDEPL foliage 100  ECU Other Amycoida Fluda FDMKK foliage 100  ECU Other Amycoida Fluda FDMTT foliage 100  ECU Other Amycoida Fluda FDTGB foliage 100  ECU Freyine Freya FRBNZ NA NA NA NA ECU Freyine Freya FREUL trunk 100  ECU Freyine Freya FRLTF NA NA NA NA ECU Freyine Freya FRTRF NA NA NA NA ECU Freyine Freya FRTYF trunk 100  ECU Freyine Freya decorata NA NA NA NA ECU Freyine Frigga sp. foliage 100  ECU Marpissoid Fritzia FZYAS foliage 100  ECU Lapsiine Galianora sacha foliage 100  ECU Marpissoid Gastromicans GMBLD foliage 100  ECU Marpissoid Gastromicans GMSPE NA NA NA NA BOR Hasariine Gedea GDFRY trunk 100  BOR Hasariine Gedea GDMUL trunk 100  BOR Cyrbine Gelotia GEPUR ground 100  BOR Cyrbine Gelotia GEROO NA NA NA NA ECU Plexippoida Habronattus paratus ground 100  BOR Plexippoid Harmochirus HRMUL NA NA NA NA BOR Plexippoid Hyllus HYLAC NA NA NA NA BOR Plexippoid Hyllus HYMOT foliage 100  ECU Amycine Hypaeus AGBBR foliage 86  ECU Amycine Hypaeus AGGHO foliage 100  ECU Amycine Hypaeus AGGRN foliage 100  ECU Amycine Hypaeus AGSPB foliage 100  ECU Amycine Hypaeus AGYOR foliage 100  55   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 BOR Nannenine Idastrandia WHSPS ground 100  ECU Marpissoid Itata ITYAS foliage 100  BOR Hisponine Jerzego corticicola trunk 100  ECU Other Amycoida Jollas JLBLK NA NA NA NA ECU Other Amycoida Jollas JLCOP NA NA NA NA ECU Freyine Kalcerrytus KCBMP ground 97  ECU Freyine Kalcerrytus KCEMM ground 94  BOR Nannenine Langerra WHFCL NA NA NA NA BOR Euophryine Laufeia BBLCK trunk 90  BOR Euophryine Laufeia BKYEL trunk 100  BOR Euophryine Laufeia BNBRK trunk 100  BOR Euophryine Laufeia BRWHT trunk 80  BOR Euophryine Laufeia FZDUL trunk 100  BOR Euophryine Laufeia FZLIN foliage 90  BOR Euophryine Laufeia LFBWS foliage 100  BOR Euophryine Laufeia LFCAN trunk 100  BOR Euophryine Laufeia LFRSK NA NA NA NA BOR Euophryine Laufeia LIMST trunk 100  BOR Euophryine Laufeia PXGRY trunk 100  BOR Euophryine Laufeia PXORG trunk 100  BOR Euophryine Laufeia RHRLQ trunk 90  BOR Euophryine Laufeia SMCOL trunk 100  BOR Marpissoid Leikung LKMUL NA NA NA NA ECU Marpissoid Lurio LURYA foliage 100  ECU Lyssomanine Lyssomanes amazonicus foliage 100  ECU Lyssomanine Lyssomanes cf. benderi foliage 100  ECU Lyssomanine Lyssomanes cf. jemineus foliage 100  ECU Lyssomanine Lyssomanes robustus foliage 100  56   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Lyssomanine Lyssomanes taczanowskii foliage 100  ECU Lyssomanine Lyssomanes tenuis foliage 100  ECU Euophryine Maeota MAEGS NA NA NA NA ECU Euophryine Maeota MAEHT foliage 96  ECU Euophryine Maeota MAESR foliage 100  ECU Euophryine Maeota MAESW foliage 100  ECU Euophryine Maeota MAETF foliage 100  ECU Euophryine Maeota MAEVN NA NA NA NA ECU Euophryine Maeota MAEWS NA NA NA NA ECU Euophryine Maeota MAEYY trunk 100  ECU Amycine Mago MGBLK NA NA NA NA ECU Amycine Mago MGBOR foliage 100  ECU Amycine Mago MGCWF foliage 84  ECU Amycine Mago MGHON NA NA NA NA ECU Amycine Mago MGHQN foliage 90  ECU Marpissoid Metaphidippus MPBRY foliage 100  ECU Marpissoid Metaphidippus MPGBR foliage 100  ECU Marpissoid Metaphidippus MPMET foliage 100  BOR Cyrbine Mintonia BSOME trunk 100  BOR Cyrbine Mintonia WFNCH trunk 100  BOR Astioid Myrmarachne alticephala foliage 100  BOR Astioid Myrmarachne cornuta foliage 100  BOR Astioid Myrmarachne exasperans foliage 100  BOR Astioid Myrmarachne gedongensis NA NA NA NA BOR Astioid Myrmarachne malayana foliage 100  BOR Astioid Myrmarachne MYBCH NA NA NA NA ECU Astioid Myrmarachne MYECM foliage 100  ECU Astioid Myrmarachne MYECO NA NA NA NA 57   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Astioid Myrmarachne MYECS foliage 100  BOR Astioid Myrmarachne nigella NA NA NA NA BOR Astioid Myrmarachne subehna NA NA NA NA BOR Nannenine Nannenus PANDS ground 100  BOR Cyrbine Neobrettus NEOMU foliage 100  ECU Amycine Noegus NGHPY foliage 100  ECU Amycine Noegus NGHQN foliage 100  ECU Amycine Noegus NGOR2 NA NA NA NA ECU Amycine Noegus NGORN foliage 100  ECU Amycine Noegus NGPAL foliage 98  ECU Amycine Noegus NGSPK NA NA NA NA ECU Amycine Noegus NGSPP foliage 94  ECU Amycine Noegus NGWFB foliage 100  ECU Amycine Noegus NGWHH foliage 100  BOR Astioid Nungia NUBWH trunk 100  BOR Astioid Nungia NUMUL NA NA NA NA ECU Freyine Nycerella NYDRB foliage 100  ECU Freyine Nycerella NYWHS foliage 100  BOR Euophryine Omoedus sp. NA NA NA NA BOR Heliophanine Orsima ORSMU NA NA NA NA BOR Plexippoid Pancorius PNBRS foliage 90  BOR Plexippoid Pancorius PNBRZ foliage 100  BOR Plexippoid Pancorius PNEVA trunk 100  BOR Plexippoid Pancorius PNORL NA NA NA NA BOR Euophryine Parabathippus PBBRN foliage 100  BOR Euophryine Parabathippus PBDKB NA NA NA NA BOR Euophryine Parabathippus PBDKS NA NA NA NA BOR Euophryine Parabathippus PBMAG NA NA NA NA 58   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 BOR Euophryine Parabathippus PBSBT foliage 100  ECU Marpissoid Parnaenus PNAEN foliage 100  ECU Marpissoid Peckhamia PKGRA NA NA NA NA ECU Freyine Phiale bulbosa foliage 100  ECU Freyine Phiale cf. gratiosa NA NA NA NA BOR Heliophanine Phintella PHBAR foliage 100  BOR Heliophanine Phintella PHLGL foliage 100  BOR Heliophanine Phintella PHSML foliage 100  BOR Heliophanine Phintella vittata foliage 100  BOR Aelurilloid Phlegra PGMUL ground 100  ECU Plexippoida Plexippus paykulli NA NA NA NA BOR Plexippoid Plexippus petersi NA NA NA NA BOR Astioid Porius GNBLK foliage NA  BOR Cyrbine Portia crassipalpis trunk 100  BOR Euophryine Pristobaeus PBHBR NA NA NA NA BOR Euophryine Pristobaeus TRLNG trunk 100  BOR Euophryine Pristobaeus TRSTR trunk 100  ECU Marpissoid Psecas PSGIA NA NA NA NA ECU Marpissoid Psecas PSRBW foliage 100  BOR Heliophanine Pseudicius PSMUL trunk 100  ECU Marpissoid Ramboia RBNOR foliage 100  ECU Marpissoid Ramboia RBUNI NA NA NA NA ECU Marpissoid Ramboia RBYST NA NA NA NA ECU Marpissoid Rhetenor RHETY foliage 100  ECU Freyine Romitia UPCRV foliage 100  ECU Freyine Romitia UPMIN foliage 100  ECU Freyine Romitia UPSRD NA NA NA NA ECU Freyine Romitia UPTTA foliage 100  59   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 ECU Marpissoid Rudra cf. wagae foliage 100  ECU Other Amycoida Sarinda cf. nigra foliage 84  ECU Other Amycoida Sarinda SRCIR foliage 100  ECU Other Amycoida Sarinda SRCRS foliage 100  ECU Other Amycoida Sarinda SRSEP NA NA NA NA ECU Other Amycoida Sarinda SRSPK foliage 100  ECU Other Amycoida Sarinda SRSTA foliage 100  ECU Other Amycoida Scopocira SCORG foliage 84  ECU Other Amycoida Scopocira SCSTR foliage 100  ECU Euophryine Sidusa SDBRN foliage 100  ECU Euophryine Sidusa SDGRN foliage 100  BOR Heliophanine Siler semiglaucus NA NA NA NA ECU Euophryine Siloca SLOCA foliage 100  BOR Astioid Simaetha sp. NA NA NA NA ECU Euophryine Soesilarishius SOBAR ground 91  ECU Euophryine Soesilarishius SOLOO ground 100  BOR Cyrbine Spartaeus SPRTM NA NA NA NA ECU Other Amycoida Synemosyna SYORB foliage 100  ECU Other Amycoida Synemosyna SYSMB foliage 100  ECU Euophryine Tariona TRBUL foliage 100  BOR Plexippoid Telamonia TMCAN foliage 100  BOR Plexippoid Telamonia TMMUL foliage 100  BOR Euophryine Thiania sp (Thianitara) NA NA NA NA BOR Euophryine Thiania THIMU foliage 100  BOR Euophryine Thiania THIPA foliage 100  ECU Other Amycoida Thiodina THEAG foliage 100  ECU Other Amycoida Thiodina THOWL foliage 100  BOR Marpissoid Tisaniba dik NA NA NA NA 60   Region Group Genus Species Habitat Specimens Criterion 1 Criterion 2 BOR Marpissoid Tisaniba mulu ground 100  BOR Marpissoid Tisaniba selan NA NA NA NA BOR Marpissoid Tisaniba selasi NA NA NA NA ECU Other Amycoida Titanattus cf. paganus trunk 100  ECU Amycine Toloella sp. ground 100  ECU Euophryine Tylogonus TYGUI NA NA NA NA ECU Euophryine Tylogonus TYSPK foliage 100  BOR Plexippoid unknown genus YELMU foliage 100  BOR Astioid Uroballus UROBM NA NA NA NA BOR Astioid Viciria praemandibularis foliage 100  61   Appendix B. Variable loadings and cumulative percentage of variation in the first seven principal components for all 36 morphometric species of salticids listing geometric landmarks and linear leg measurements, respectively. Variable PC1 PC2 PC3 PC4 PC5 PC6 PC7 P1x1 -0.061 0.005 0.005 -0.002 0.000 -0.002 0.001 P1x5 0.060 -0.005 -0.005 0.002 0.000 0.002 -0.001 P1x2 -0.060 0.003 0.005 0.000 -0.002 0.000 0.002 P1x4 0.061 -0.003 -0.004 0.000 0.002 0.000 -0.003 P1x3 0.023 -0.008 0.010 -0.006 -0.011 -0.002 0.001 P1x6 -0.059 0.011 0.005 0.001 0.001 0.000 0.004 P1x8 0.059 -0.010 -0.006 0.000 -0.001 0.000 -0.004 P1x7 -0.059 0.010 0.005 0.001 0.001 0.000 0.004 P1x9 0.059 -0.010 -0.006 0.000 -0.001 0.000 -0.004 P1y1 -0.055 0.011 -0.001 -0.003 0.011 0.004 0.007 P1y5 -0.042 0.009 -0.005 0.026 0.002 -0.012 0.006 P1y2 -0.059 0.010 0.000 0.004 0.009 -0.001 0.003 P1y4 -0.049 0.008 -0.003 0.022 0.004 -0.010 0.003 P1y3 -0.057 0.007 -0.001 0.015 0.005 -0.005 0.003 P1y6 0.021 -0.014 0.003 -0.034 -0.003 0.012 -0.006 P1y8 0.031 -0.019 -0.005 0.010 -0.017 -0.014 -0.009 P1y7 0.042 -0.010 -0.001 -0.026 -0.002 0.010 -0.003 P1y9 0.054 -0.012 -0.005 0.000 -0.009 -0.006 -0.005 P1y10 0.060 -0.004 0.005 -0.005 -0.006 0.005 -0.003 P2x1 0.060 0.000 0.002 -0.008 0.000 0.004 -0.003 P2x2 0.060 0.000 0.001 -0.002 -0.001 0.001 -0.006 P2x3 0.061 -0.002 -0.001 -0.001 -0.002 -0.002 -0.005 P2x4 0.060 -0.005 0.003 0.000 -0.006 0.001 -0.005 P2x5 0.060 -0.007 0.002 0.001 -0.006 0.000 -0.004 P2x6 0.059 -0.003 0.008 0.000 -0.002 -0.006 -0.009 P2x7 0.059 -0.003 0.008 -0.001 -0.001 -0.006 -0.009 P2x8 0.047 -0.008 0.022 0.026 -0.018 0.007 0.004 P2x9 0.050 -0.006 0.021 0.026 -0.017 0.004 0.007 P2x10 0.060 0.000 0.001 -0.009 0.000 0.004 -0.003 P2x11 0.060 0.000 0.002 -0.015 0.000 0.009 -0.002 P2x12 0.059 0.000 0.003 -0.021 0.000 0.014 -0.001 P2x13 0.056 0.002 0.003 -0.029 -0.001 0.022 0.002 P2x14 0.042 0.006 0.003 -0.052 0.000 0.044 0.010 P2x15 -0.050 0.008 -0.003 -0.041 0.004 0.038 0.015 P2x16 -0.060 0.003 -0.004 -0.011 0.004 0.010 0.007 P2x17 -0.061 0.002 -0.004 -0.003 0.004 0.003 0.005 P2x18 -0.061 0.001 -0.004 0.001 0.004 -0.001 0.004 P2x19 -0.061 0.001 -0.004 0.003 0.003 -0.003 0.003 62   Variable PC1 PC2 PC3 PC4 PC5 PC6 PC7 P2x20 -0.060 0.001 -0.004 0.004 0.003 -0.004 0.003 P2x21 -0.060 0.000 -0.004 0.006 0.003 -0.005 0.003 P2x22 -0.060 0.001 -0.005 0.006 0.003 -0.006 0.003 P2x23 -0.060 0.002 -0.005 0.006 0.004 -0.007 0.003 P2x24 -0.060 0.004 -0.004 0.006 0.004 -0.007 0.002 P2x25 -0.059 0.005 -0.003 0.009 0.004 -0.007 0.003 P2x26 -0.059 0.006 0.000 0.015 0.000 -0.006 0.005 P2x27 -0.055 0.006 0.005 0.027 -0.007 -0.003 0.010 P2x28 -0.019 0.003 0.023 0.056 -0.025 0.003 0.022 P2x29 0.053 -0.005 0.019 0.019 -0.012 0.002 0.003 P2y1 0.057 0.006 -0.005 0.006 0.008 -0.019 0.005 P2y2 0.044 0.011 -0.022 -0.018 -0.008 0.011 0.003 P2y3 -0.057 -0.003 -0.004 -0.001 -0.009 0.004 0.006 P2y4 -0.045 -0.026 -0.011 -0.008 -0.001 0.025 0.006 P2y5 -0.060 -0.010 0.000 -0.005 0.002 0.009 0.000 P2y6 -0.060 -0.006 0.002 -0.006 0.002 0.012 -0.004 P2y7 -0.060 -0.002 0.002 -0.008 -0.001 0.013 -0.003 P2y8 -0.059 -0.002 0.001 -0.009 -0.002 0.016 -0.006 P2y9 -0.060 0.000 0.004 -0.006 -0.002 0.011 -0.006 P2y10 0.057 0.006 -0.005 0.005 0.009 -0.018 0.004 P2y11 0.059 0.003 -0.003 0.001 0.009 -0.014 0.004 P2y12 0.059 0.003 -0.002 -0.002 0.007 -0.010 0.005 P2y13 0.060 0.003 -0.002 -0.003 0.005 -0.007 0.007 P2y14 0.060 0.002 -0.001 -0.003 0.004 -0.005 0.007 P2y15 0.060 0.001 -0.001 -0.002 0.003 -0.004 0.008 P2y16 0.060 -0.001 -0.001 -0.001 0.001 -0.004 0.008 P2y17 0.060 -0.003 -0.001 0.001 0.000 -0.004 0.009 P2y18 0.060 -0.006 -0.001 0.005 -0.002 -0.006 0.010 P2y19 0.059 -0.010 0.000 0.012 -0.004 -0.008 0.009 P2y20 0.058 -0.013 0.001 0.020 -0.006 -0.012 0.009 P2y21 0.055 -0.016 0.002 0.030 -0.007 -0.018 0.006 P2y22 0.035 -0.017 0.006 0.053 -0.011 -0.037 -0.015 P2y23 -0.050 0.008 0.005 0.014 -0.002 -0.011 -0.026 P2y24 -0.055 0.012 0.004 0.002 0.000 -0.001 -0.019 P2y25 -0.057 0.012 0.004 0.001 -0.001 0.002 -0.016 P2y26 -0.058 0.010 0.003 -0.001 -0.002 0.005 -0.012 P2y27 -0.059 0.007 0.003 -0.002 -0.003 0.007 -0.009 P2y28 -0.060 0.004 0.003 -0.003 -0.003 0.009 -0.007 P2y29 -0.060 0.001 0.003 -0.004 -0.002 0.009 -0.005 P3x1 0.061 -0.005 -0.006 0.002 0.000 0.002 0.000 P3x12 -0.061 0.005 0.006 -0.002 0.000 -0.002 0.000 P3x2 0.061 -0.004 -0.005 0.001 0.001 0.002 -0.001 63   Variable PC1 PC2 PC3 PC4 PC5 PC6 PC7 P3x10 -0.061 0.004 0.006 -0.001 -0.001 -0.002 0.002 P3x3 0.061 -0.004 -0.005 0.001 0.001 0.002 -0.001 P3x11 -0.061 0.005 0.006 -0.002 0.000 -0.002 0.001 P3x5 0.061 0.002 -0.006 -0.002 0.001 -0.002 -0.003 P3x9 -0.060 -0.003 0.005 0.001 -0.002 0.002 0.002 P3x4 0.061 0.003 -0.005 -0.001 0.001 -0.001 -0.002 P3x8 -0.061 -0.003 0.005 0.002 -0.002 0.002 0.002 P3x6 -0.024 0.000 -0.025 0.013 -0.013 0.006 -0.011 P3x7 0.001 -0.002 -0.014 0.004 -0.010 -0.001 -0.004 P3y1 -0.047 -0.029 0.001 0.009 0.016 0.009 -0.008 P3y12 -0.051 -0.026 0.003 0.006 0.016 0.007 -0.005 P3y2 0.033 -0.040 -0.011 0.015 0.012 0.017 0.002 P3y10 0.030 -0.041 -0.010 0.015 0.014 0.017 0.001 P3y3 -0.058 -0.010 0.006 0.001 0.011 0.002 -0.007 P3y11 -0.058 -0.009 0.007 0.000 0.010 0.001 -0.006 P3y5 -0.059 0.009 0.005 0.000 -0.010 0.002 0.002 P3y9 -0.059 0.008 0.005 -0.001 -0.010 0.002 0.002 P3y4 0.058 0.015 -0.007 -0.003 -0.007 -0.004 -0.004 P3y8 0.058 0.015 -0.006 -0.004 -0.006 -0.004 -0.004 P3y6 0.057 0.008 0.001 -0.005 0.000 -0.009 0.012 P3y7 0.007 0.039 -0.007 -0.004 -0.036 -0.003 -0.003 P6x2 0.054 0.013 0.006 0.005 0.007 0.009 -0.003 P6x3 -0.054 -0.013 -0.006 -0.005 -0.006 -0.008 0.004 P6x1 0.006 0.008 -0.023 0.012 -0.009 0.004 -0.010 P6x4 -0.001 -0.002 0.020 -0.012 -0.004 -0.014 -0.002 P6y2 0.048 0.008 0.011 0.003 0.003 0.011 -0.002 P6y3 0.048 0.008 0.001 0.008 0.001 0.016 -0.005 P6y1 -0.055 -0.005 -0.010 -0.006 -0.006 -0.011 0.007 P6y4 0.055 0.005 0.010 0.006 0.008 0.011 -0.008 L1_TibL -0.051 -0.027 -0.002 -0.003 -0.008 -0.004 0.003 L1_TibD -0.057 -0.001 -0.008 -0.005 -0.001 -0.010 0.008 L1_SL -0.046 -0.026 0.015 -0.010 -0.017 -0.007 -0.004 L3_FemL -0.056 -0.020 0.004 -0.006 -0.007 -0.007 -0.008 L3_PatL -0.059 -0.006 -0.001 -0.006 -0.004 -0.008 -0.006 L3_TibL -0.054 -0.025 0.002 -0.005 -0.011 -0.005 -0.004 L3_MetaTar -0.052 -0.025 0.003 -0.009 -0.010 -0.010 0.001 L4_FemL -0.057 -0.015 -0.008 -0.007 -0.013 -0.009 0.004 L4_PatL -0.055 -0.004 -0.012 -0.007 -0.006 -0.013 0.008 L4_TibL -0.051 -0.022 -0.009 -0.007 -0.015 -0.007 0.010 L4_MetaTar -0.051 -0.021 -0.006 -0.010 -0.014 -0.011 0.007          Cum.Prop. (%) 78.0 82.1 83.9 86.2 88.1 90.9 92.2 64                Appendix C. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC1.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg. A B C D E i ii iii iv v vi vii viii ix x xi xii 65                 Appendix D. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC2.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg.   A B C D E i ii iii iv vi vii viii ix x xi xiii vi 66                 Appendix E. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC3.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg.   A C B D E i ii iii iv v vi vii viii ix x xi xii 67                Appendix F. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC4.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg.    A B C D E i ii iii iv v vi vii viii ix x xi xii 68   Appendix G. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC5.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg. 69                   Appendix H. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC6.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg.   A B C D E i ii iii iv v vi vii viii ix x xi xii 70                Appendix I. X,Y coordinates of geometric landmarks (A-D) and linear leg measurements (E) for reconstructed specimens of small, large and average PC values for PC7.  Leg measurements include: (i) reference point for the length of carapace in lateral view, (ii) tibia length first leg, (iii) tibia depth first leg, (iv) tibial spine length first leg, (v) femur length third leg, (vi) patella length third leg, (vii) tibia length third leg, (viii) tarsus and metatarsus length third leg, (ix) femur length fourth  leg, (x) patella length fourth  leg, (xi) tibia length fourth leg, and (xii) tarsus and metatarsus length fourth leg. A B C D E i ii iii iv v vi vii viii ix x xi xii 

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