Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. almMcLean Callum the analysis Fourier in elliptical complexity using shape pedipalp of amblypygid drivers and patterns the Assessing 00.Saei lokont aycnieal costegop ihtepsto,nme,rltv length relative number, position, Weygoldt, the 1936; with Fage, group, & the (Simon across length considerably body vary their to of known that also length times is of four in Shape members lengths Harvey and whilst 2000). 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. eew pl ut ftos nldn litclFuiraayi,t siaetesaecmlxt fthe pedipalp of interspecific complexity Hypotheses of shape and comparison quantitative the Aims a estimate order. time, to important first analysis, this Fourier the across elliptical for correlates shape including facilitates, also tools, This good habitats of pedipalp. a suite spider amblypygid be a within can apply diversity 2012). example, it we Hartz, and For suggesting Here & 2019), birds, Rodrigues King, in concepts. 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No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. o utoe 5 fttlosre aito nbt emns hsP1P1 eecridfradit the into forward carried accounted were 10 PC1-PC10 through thus 1 segments, PC’s both = tibia in tibia. tibia variation the and analysis. 56%, observed femur clustering in total = the spines species of both (femur those 95% in segments variation over of both of just length PC2 in 15% for the variation roughly ( where contributed observed and species elongation, PC2 of spines Phrynichidae pedipalp same species. half longest separated over the to the higher mainly just within of correlate a and for position species 59.2%) to show accounted relative where PC1 appears to the femur femur. 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Data may be preliminary. eso hsseiswti h aecutraogien diinlseis -en lopae h two the placed also K-means species. additional no alongside cluster same the femur, within the Damon species In this results. classify of different only markedly mens could provided clustering methods MANOVA k-means and unsupervised k-means using clustering Species = coronatus thophrynus iue3-PAo amnccecet,bakotie iuls ulnso ersnaiepdpls grey pedipalps, representative = Exo of species. 20 outlines within diadema the variation visualise intraspecific at outlines the represent (calculated black hulls convex morphs coefficients, hypothetical harmonic are of outline PCA - 3 Figure aahyu aztecus Paraphrynus pce oehrwti iceegnscutrwihcnandn te pce,btwsual to unable was but species, other no contained which cluster genus discrete a within together species e = med , ao medius Damon e = ger , i = wil , hru gervasii Phyrnus aahyu williamsi Paraphrynus o = lon , eeohyu longicornis Heterophrynus i = vir , h exophalmus Ph. 7 aahyu viridiceps Paraphrynus th amnc rmeteepsto nteP axes, PC the on position extreme from harmonic) ih10 cuay lcn l speci- all placing accuracy, 100% with h = whi , hyihsexophalmus Phrynichus o = lop , hyu whitei Phrynus hyu longipies Phrynus i = dia , o = cor , Damon Acan- azt , Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. .lniis5.%42.9% 80.0% 100% 14.3% 50.0% 14.3% 16.7% 28.6% 12.5% 14.3% 25.0% 16.7% 12.5% 16.7% 50.0% 57.1% 57.1% 42.9% 14.3% 20.0% 100% williamsi Pa. 85.7% whitei P. viridiceps Pa. medius D. longipies 28.6% P. longicornis H. 100% gervasii P. exophalmus Ph. diadema D. coronauts A. aztecus Pa. t eeietfidi ohtba n eoa emns(al 3). (Table segments femoral and complex- tibial shape both size-independent in and identified length were pedipalp ity between length relationships negative pedipalp significant absolute Statistically and complexity no shape However, between species. 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. h oprtv nlsspeetdhr ouet ohita n neseicdffrne nsaecom- frequent shows shape harmonics in Fourier of differences analysis interspecific component no principal and A intra- However, pedipalp. both amblypygid documents the 13.89). within here plexity = presented analysis F comparative 0.037, The = p (ANOVA, femur. tibia the Discussion in female found the were differences in significant higher statistically femur statistically and was mean. tibia the Complexity the complexity around in error shape length, standard in pedipalp show differences and bars Sexual complexity error means, shape species of represent measure Points between segments. 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Data may be preliminary. nte osblt sta htsielnt scntandb eetv eie rdvlpetlfactors, developmental or regimes selective by constrained good is a length be spine not may that thus that work. and this is & intraspecific mixed in Fernandes possibility all provides report Silva are arachnids Another we da examples other trends these from 2009; interspecific that evidence al., the stressed possible, for et be be analogue and (Willemart must may longer and it females trade-off also However, & in conspecifics are a Girard contest, support. though seen female 1990; male-male Thus, not than in Kronestedt, 2014). apophyses used 1889; longer arachnids. opiliones, Willemart, coxal Peckham, in are male & elaborated of complex display (Peckham larger legs more courtship setae fourth have and of the in Furthermore, larger adults ridges used often 2014). are as of are Endler, is such selection that chelicerae it elaborations sexual However, legs also large possess of 1982). spines spider often influence the Bali, pedipalp male the & Some in under example, (Rajashekhar 2015). decreased For structures pedipalps (Bird, wear relatively large that cheliceral with case and of thelyphonids the function juveniles male a in dentition in be cheliceral reduced part example, pronounced appear in For may more arachnids. although this in arachnids, though be complexity within (Solifugae), to shape size we of increasing known with trends yet complex interspecific is limited as less about therefore though known becoming are cost, is structures inferences for little energetic such unit evidence lower and some per amblypygids, at is produce in There pedipalps to development cost. longer costly of developmental energetically achieve length, genetics lower some less to of pedipalp at be with nothing individuals achieved increased may know allow exists, be spines for may to complexity shorter allowing which contest, with and spines, form and length, length shorter complex selection less relatively sexual pedipalp a of with between Hypothetically, pressures form, trade-off the complex a under resource that less developing that that suggests a and possible This taking choice, be species mate 2016). therefore via length. Hebets, may selection pedipalp & sexual It with Chapin the of correlated during influence 2000; be and may the (Weygoldt, 2016) potential under group Hill-Lindsay, holding likely amblypygid & the of are Chapin across number pedipalps 2013; a courtship amblypygid Peixoto, in of & contests Porto stages territorial 2006; in first observed Hebets, conflicts been & of (Fowler-Finn also 80% has Roughly, species display has morphology. work Pedipalp pedipalp Previous 2017). of Reed-Guy, size. evolution centroid the to in normalised display revealed in once of was spines importance pattern ‘major’ the similar shorter demonstrated a predominantly relatively Interestingly, have complexity to 2). that found (Figure suggests therefore spines was outlines pedipalp length in Fourier major pedipalp GMM of of and using inspection length complexity relative between Visual the relationship traits. reflects negative segments. pedipalp pedipalp significant both overall using a any investigates in species 2, of identified analysis related Hypothesis use our closely of the between (as invalidate support differentiating not species In when do define caution results to urge our Prendini, characters do Although 2000; femur we (Weygoldt, 2012). segments shape), and pedipalp Armas, tibia tibia de and pedipalp & femur Joya specific the of a 2005; on within species spines Wheeler, placement specimen of identified & specimen’s length a have Weygoldt relative of of studies knowledge the placement of prior previous the basis no Notably, for have the speci- would analogue species. algorithm, species better k-means a which a the of within arguably like knowledge taxonomist, is prior a K-means as no not. species, has could which to, methods, belong k-means ‘super- mens a species unsupervised either MANOVA, each the to in characters. 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Of not k-means iden- accurately. unsupervised could does However, clusters MANOVA , species study. species. this most define in between to taxon differences struggles each shape representing species, pairwise individuals between significant of differences statistically numbers shape tify low exceeding relatively often the variation in shape even intraspecific with species, between overlap .longipes P. .variegatus D. r eie i ipa ntefvu fteidvda ihlne eiap Cai & (Chapin pedipalps longer with individual the of favour the in display via decided are ihlne eiap aigsotrsie.Lne mlpgdpdplsare pedipalps amblypygid Longer spines. shorter having pedipalps longer with , Paraphrynus ol o edffrnitdo h ai fsaecomplexity shape of basis the on differentiated be not could 10 Damon priori a Damon species, ol en ieecsbetween differences define could , and .diadema D. Paraphyrnus and atal on partially .medius D. . Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. tde htlo oadestepuiyo aao mlpgdlf itr,eooyadsca yaiswill dynamics system. social unique Future and this ecology in partitioning. history, morphology life niche functional again amblypygid Acknowledgements trophic the once on understanding and data studies, of further other courtship, paucity in in and the valuable intraspecifically address contest be seen to complex- based patterns look gross similar display in that follows dimorphism of studies and Sexual pressures tibia territorial capture). the the and potentially prey in selection highlighting pedi- in length, sexual present use of in pedipalp also for basis use is increasing (primarily the (for ity exceed length with length on spine occasionally total decreases species pedipalp pedipalp complexity can and defining in contest) gross when investment pedipalps between that taken in trade-off find be a variation to also uncovering complexity We needs shape caution characters. thus pedi- within-species palp amblypygid terms and that across in species, relationships variation find between shape interspecific differences we undocumented and previously Here intra- of considering number palps. of a highlighting importance by the shape, highlight of results our conclusion, history In life amblypygid morphology. into pedipalp work of this more drivers suggesting However, Conclusion the species, understand 2000). amblypygid and (Weygoldt, fully across contest to pressure behaviour Interestingly, needed selection forces in group. is driving common similar occur- the these very a across The that are be pervasive suggest and could may are 2020). display group complexity Brassey, involve the and & both across size courtship Garwood species in of numerous (McLean, dimorphism in pedipalps of display sexual dimorphism the pressure behind of pedipalp the of form for under analysis this optimise develops of GMM female which rence males using the pedipalp, while identified in male capture been larger the already prey relatively of has are length which pattern increased spines, similar variegatus the major A to the com- due selection. of discussed, sexual be length previously may the As This by counterparts. tibiae influenced male Signifi- female tibia. their be with time. than tibia, to first complexity pedipalp appears the greater amblypygid plexity by the for of characterised in amblypygids identified be across importance is to quantified complexity relative More found was shape dimorphism the in capture. length. shape dimorphism determine prey sexual sexual pedipalp study, cant to via to this selection dynamics of relative natural social part and isometry As and contest, than territorial history lower via direct life rates selection most amblypygid at selection, McCormick, the into scale sexual under & needed hypothetically also (Polis are similar may is prey a (which If capture) work sized spines chelae. prey species, similarly sized amblypygid equivalently for on across producing selection feed This exists species at size larger sizes 2010). to prey size Summers, body benefit in in & added overlap different increase Herrel little vastly which Meijden, is chelae, der there with (Van suggesting pedipalp size species 1986), scorpion body because in to found be relation is with may Amblypygi isometry paucity than in lower current form much the selection palpal to rates the addressing sexual leading to by of species investigated analogue amblypygids. pressures across An properly in to similar be diet due remain to only prey pedipalps related also can should longer data may hypothesis Therefore, attained in this size have size. However, spine pedipalp that contest. group, than species indeed and size is the in spines prey across spines of with equal smaller function closely primary relatively or especially more the similar scale arthropods, If may remain consumer 2016). size primary size Hebets, pedi- about spine of & known capture, forming composed (Chapin is prey species largely Blattodea Little in and of be items. Orthoptera number to prey orders thought secure a the and are with of capture they capture, help but prey to diets, hypothesised length. in display-based amblypygid are pedipalp function in which to important primarily relative baskets’, spines isometry ‘catching not to with longer palp are thought increasing with absolutely length and are spine species that capture, devel- spines to advantage possible case, prey by Pedipalp little limited to is this is be benefits there it In meaning could functional Additionally, contest, length additional species. length. spine limited across example, provide pedipalp size For may to absolute spines. relative similar shorter constraints relatively a opmental have at would remained pedipalps longer simply have spines meaning ihteosre eulsaedmrhs en huh oaiedet eaeotmsn for optimising female to due arise to thought being dimorphism shape sexual observed the with , 11 D. Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. awo J,Dno A,Keh J,HgaT.21.Tepyoeyo oslwi spiders. whip fossil of spiders. phylogeny Peacock The 2014. JA. across 2017. Endler complexity MB., TA. Girard shape Hegna 3D BJ., Knecht Determining biology JA., evolutionary shapes: Dunlop Alpha RJ., Garwood 2018. CA. Brassey structures. Pro- diverse J., of morphologically Behnsen analysis Fourier JD., elliptic Gardiner using deformation Anthropologists Physical cranial of artificial Association Exploring spider whip outlines. 2003. aligned the crustes M. in Baylac interactions M., agonistic Friess (Orcinus of whale examination killer An in 2006. variation marginemaculatus EA. Quantifying Hebets KD., 2019. Fowler-Finn JM. analysis. Fourier Waite elliptical ME., using Dahlheim morphology JJ., orca) Hard CK., Emmons Amblypygi the in cannibalism size-symmetric atypical longipes spider mediates Territoriality whip Phrynus 2017. the S. in Reed-Guy value KJ., Chapin resource asymmetric by evidenced Territoriality longipes 2016. Phrynus amblypygids. S. of Hill-Lindsay ecology KJ., behavioral Chapin In: The Complexity. 2016. Shape EA. 2D Hebets KJ., Exploring Chapin 2018. K. Leonard for C., value Analysis Grimm and Shape R. T., applications, Emerson using fundamentals, E., analysis analysis: Chambers Fourier outline Elliptical 2017. Momocs: CN. anthropology. Stephan forensic J., 2014. Byrd J. J., Claude Caple C., Gaucherel S., Software Statistical Picq char- and V., terminology, Thesis. homology, Bonhomme Primary PhD University. (Arachnida): State Solifugae Colorado of survey. morphology acter Cheliceral effects. 2015. spatial TL. fragmented and a Bird environmental in of web-spiders of importance composition relative Metacommunity forest: 2012. SM. neotropical Hartz ENL., Rodrigues R., Baldissera of behavior and Biology badensis 1962. this in AJ. used Alexander specimens all 10.6084/m9.figshare.c.5289733 of at lengths Figshare References pedipalp via and available made body be and will outlines study binarized raw photographs, Pedipalp Society Arachnological statement American Data the by supported was Michael work and This Horweg Christoph Christoph Prendini, Spiegel, Lorenzo der Funding van Colmenares, Natural USA; Didier Pio York, The thank New Beccaloni, we Belgium; History, Specifically, Tervuren, Jan Seiter. Natural Museum, Austria. Jocque, of African Vienna, Rudy Musuem Central in Allard, American Royal Museum The History the Kingdom; Natural of The United staff London, the Museum, thank History to like would We ooko rndd I(rcnd,Pedipalpi). (Arachnida, WI Trinidad, of Pocock pigr 61–83. Springer, . . . Aahia Amblypygi). (Arachnida, Ethology Processes Behavioural 56:1–24. 17:105. nentoa ora fLglMedicine Legal of Journal International mrcnJunlo hsclAtrplg:TeOca ulcto fteAmerican the of Publication Official The Anthropology: Physical of Journal American 123:772–777. M vltoayBiology Evolutionary BMC 122:11–22. 122:110–115. ora fArachnology of Journal ao variegatus Damon urn Biology Current aieMma Science Mammal Marine 12 Zoologica 131:1675–1690. 18:184. 24:R588–R590. et fSuhArc and Africa South of Perty 47:25–37. 34:62–76. h ora fArachnology of Journal The lsone Plos 35:5–21. (0,p.e48099 7(10), deu bar- Admetus eerhin Research ora of Journal Phrynus 44:1–15. BMC Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. oeL,Anvs .21.Sxa eeto n h vlto fGntlSaeAdCmlxt In Complexity And spider Shape whip the Genital by Of capture Arachnology Prey Evolution of 2009. The EA. Hebets And RD., Santer Selection Sexual abun- 2012. by Striders. G. revealed Water not Arnqvist associations L., habitat Rowe avian reflects plumage in Variation manip- genetic 2019. dance. by DI. performance King HP., flight Roberts Enhanced Drosophila. in 2016. shape RJ. wing Bomphrey P., of Henningsson ulation T., Nakata RP., Ray genus whipscorpion, a the in dimorphism of Uropygi). Sexual nida, spiders 1982. G. Whip Bali KP., Rajashekhar 2010. J-I. Island. Kojima Borneo from shape. MS., Charinidae) tooth Harvey paleoeco- of of measure C., evaluation free Rahmadi an homology a extinction: anthropology using primate physical primates major stem of first American journal The North 2016. of MT. dynamics Silcox logical DM., Boyer KA., Prufrock DNA. and morphology the behaviour, of from Systematics Evolution evidence 2005. Amblypygi): WC. (Chelicerata: Wheeler spiders P., Weygoldt L., Amazonian the Prendini in territoriality and selection habitat of spider evidence scorpion. Experimental whip desert 2013. of PEC. species Peixoto among TJ., structure Porto age Ecology and Animal use Attidae. resource of Spiders of Journal Patterns of The Family 1986. the SJ. McCormick in Soc. GA., Selection Polis Hist. saurians. Sexual Nat. dentigerous on Wisc. Observations living . in 1889. Pap complexity E. Peckham tooth scorpions; G., and Peckham in diet force between pincer relationship Morphology and of The size rap- Journal chela the 2017. of of Comparison KM. 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Joya French from spider whip 2012. inhabiting APL. bromeliad Giupponi M., Jocque h isnJunlo Ornithology of Journal Wilson The :395–402. 5:203–236. olgc Scripta Zoologica eeohyu longicornis Heterophrynus olgshrAnzeiger Zoologischer 37:109–112. ultnBiihAahooia Society Arachnological Bulletin-British Evolution ora fZoology of Journal 7:0–2.DI 10.1002/jmor.20645. DOI: 278:500–522. 64–4 O:10.1111/j.1558-5646.2011.01411.x. DOI: 66:40–54. 1:3–60. 19:203–225. :59–73. Zootaxa 159:683–697. 273:56–64. hrnsbromeliaea Charinus auecommunications Nature Aahia Amblypygi). (Arachnida, 280:319–325. 2612:1–21. 131:339–347. 13 () pp.330-331 5(7), . Zootaxa p .Abyyi hrnde;anwseisof species new a Charinidae); n.(Amblypygi: sp. lpcs aculeata Alopecosa 7:1–8. hyu marginemaculatus Phrynus 3158:53–59. ora fethology of Journal ao variegatus Damon hlpou indicus Thelyphonus ora fZoology of Journal Sarax Cec)b morphological, by (Clerck) io 82(Amblypygi: 1892 Simon raim iest & Diversity Organisms 31:299–304. ru fArcnwhip African of group LKoch. CL 1:55.DOI: 310:45–54. tlck (Arach- Stoliczka PeerJ oei de Boletin American 6:e5751. Journal c . Occ Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. avrdcp 33 66.7% 25.0% 50.0% 12.5% 100% 11 100% 10 50.0% 14.3% 33.3% 9 8 25.0% 85.7% 75.0% 7 6 25.0% 14.3% 5 42.9% 87.5% 42.9% 4 Pa.williamsi P.whitei 3 Pa.viridiceps 50.0% 100% D.medius 62.5% P.longipies 2 H.longicornis P.gervasii Ph.exophalmus 1 D.diadema A.coronauts tibia Pa.aztecus segment and tibia femur for the Cluster results of K-means analysis and the Manova in – used individuals 2 female Material and Supplementary male of number the of segments. sizes Sample – S1 Opil- (Arachnida, species. harvestmen 18 Phalangida 1 on of Material data biology Supplementary morphological new Sensory with 2009. review, P. a Gnaspini iones): J., Farine RH., Willemart (Chelicerata: in spiders system size phylogenetic 2000. a and P. towards Weygoldt shape spiders: genital whip of male Amblypygi). morphology on Arachnida: Evolutionary study 1996. P. association Weygoldt Genome-wide partie). (1re 2015. Araneae melanogaster. KH. et Drosophila Takahashi Solifuga Scorpiones, B., Pedipalpi, Takahara III. Arachnida. l’Omo sexually de use 1936. scientifique Opiliones) L. Fage (Arachnida, environment. E., harvestmen the Simon Neotropical in chemicals 2014. spread RH. to glands Willemart dimorphic N., Fernandes Silva da whipspider, a significance. of evolutionary anatomy its Muscular and 1999. JW. Shultz ikb ad emr:Aol Books. Apollo Denmark: Sand, Kirkeby . hpsies(hlcrt:Abyyi:terbooy opooyadsseais Whip systematics, and morphology biology, their Amblypygi): (Chelicerata: spiders Whip pce M F exophalmus Ph. whitei P. longipies P. M gervasii P. williamsi Pa. viridiceps Pa. aztecus Pa. longicornis F H. medius D. diadmea D. coronatus A. Species 3:293–340. ora fZooia ytmtc n vltoayResearch Evolutionary and Systematics Zoological of Journal LSOne PLoS olgclJunlo h ina Society Linnean the of Journal Zoological 07,p.e0132846. 10(7), 4 4 4 3 4 4 4 3 4 3 3 4 4 4 4 3 4 4 4 3 3 4 3 4 4 2 4 2 4 4 3 4 4 3 4 3 4 3 3 4 4 4 (n) Tibia 4 (n) Tibia 4 (n) Femur (n) Femur 14 hyu longipes Phrynus ope edsbiologies rendus Comptes caZoologica Acta Pcc)Aahia Amblypygi), (Pocock)(Arachnida: 126:81–116. 90:209–227. 337:269–275. 34:185–202. 37.5% Mission Posted on Authorea 16 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161349101.13487593/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. P.whitei on,3)Atclto on ewe eu n ii,4)Atclto ewe eu n trochanter. and femur between inflection end Articulation femur Distal necessary. 4.) where tibia, placement 2.) landmark and point, aid femur inflection to end between consulted were femur were point Landmarks photographs Proximal Articulation Specimen alignment. aid 1.) to 3.) positions locations 4 following point, in placed the were in Landmarks placed landmarks. femur of Positions – S3b were photographs Specimen Landmarks shaft. central alignment. pedipalp point aid of Articulation necessary. end to where 2.) at placement tibia, positions point landmark and 3 femur aid Inflection between in to 3.) point consulted placed Articulation tarsus, were and 1.) Landmarks tibia locations between following the landmarks. in tibia placed of were Positions – S3a segment femur and tibia for positions Landmark – are 3 differences Material significant Supplementary no show contain that tables MANOVA comparisons bold. species segment. in comparison, tibial highlighted pairwise to applied for p-values species magnitude between tests the MANOVA pairwise of Results S2b- Pa.viridiceps D.medius P.longipies H.longicornis P.gervasii Ph.exophalmus D.diadema of A.coronauts percentage the contain species. Pa.aztecus tables represent rows K-means K-means, by segment. defined clusters tibial represent to Columns applied cluster. each clustering into fall k-means that of specimens Results - S2a .ooat .idm heohlu .evsiHlnionsPlniisDmdu avrdcp .htiPa.williamsi P.whitei Pa.viridiceps D.medius P.longipies H.longicornis P.gervasii Ph.exophalmus D.diadema A.coronauts < 0.001 < < 0.001 0.001 < < < 0.001 0.001 0.001 15 < < < < 0.001 0.001 0.001 0.05 < < < < < 0.001 0.001 0.001 0.001 0.001 < < < < < < 0.01 0.001 0.001 0.001 0.01 0.01 < < < N.S. < < < 0.001 0.001 0.001 0.001 0.001 0.001 < < < < < < < N.S. 0.001 0.01 0.001 0.001 0.001 0.001 0.001 < < < < < < < < < 0.01 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 < < < < < < < < < < 0.001 0.01 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001