Literature DB >> 25317062

The deep phylogeny of jumping spiders (Araneae, Salticidae).

Wayne P Maddison1, Daiqin Li2, Melissa Bodner3, Junxia Zhang3, Qingqing Liu4, Fengxiang Liu4.   

Abstract

In order to resolve better the deep relationships among salticid spiders, we compiled and analyzed a molecular dataset of 169 salticid taxa (and 7 outgroups) and 8 gene regions. This dataset adds many new taxa to previous analyses, especially among the non-salticoid salticids, as well as two new genes - wingless and myosin heavy chain. Both of these genes, and especially the better sampled wingless, confirm many of the relationships indicated by other genes. The cocalodines are placed as sister to lapsiines, in a broader clade with the spartaeines. Cocalodines, lapsiines, and spartaeines are each supported as monophyletic, though the first two have no known morphological synapomorphies. The lyssomanines appear to be non-monophyletic, of three separate groups: (1) Lyssomanes plus Chinoscopus, (2) Onomastus, and (3) the remainder of Old World species. Several previously-inferred relationships continue to be supported: hisponines as sister to the Salticoida, Amycoida as sister to the remaining Salticoida, and Saltafresia as monophyletic. The relationship of Salticus with Philaeus and relatives is now considered well enough corroborated to move the latter into the subfamily Salticinae. A new clade consisting of the Plexippoida + Aelurilloida + Leptorchesteae + Salticinae is recognized. Nungia is found to be an astioid, and Echeclus, Gedea and Diplocanthopoda to be hasariines. The euophryines are corroborated as monophyletic. The agoriines Agorius and Synagelides are salticoids, within the sister group to amycoids, but their further placement is problematical, perhaps because of their nuclear ribosomal genes' high GC bias, as also seen in the similarly problematic Eupoa.

Entities:  

Keywords:  Jumping spiders; Salticidae; phylogeny; systematics

Year:  2014        PMID: 25317062      PMCID: PMC4195940          DOI: 10.3897/zookeys.440.7891

Source DB:  PubMed          Journal:  Zookeys        ISSN: 1313-2970            Impact factor:   1.546


Introduction

Salticid spiders, remarkable for their excellent vision (Land 1969, Blest et al. 1990), include more than 5000 species (Platnick 2014) with a great diversity of body forms and behaviours. While this diversity has long resisted phylogenetic organization, recent molecular studies (Maddison and Hedin 2003, Su et al. 2007, Maddison et al. 2008, Bodner and Maddison 2012, Zhang and Maddison 2013), aided by compilations of morphological taxonomic knowledge (Prószyński 2013) have resolved much of the phylogenetic structure of the family. One of the best-supported clades is the Salticoida, recognized by both morphological and molecular data (Maddison 1996, Maddison and Hedin 2003) and containing about 95% of the known species in the family. Within the Salticoida, large groups such as the Amycoida, Astioida, Marpissoida and Plexippoida are well-corroborated (Maddison and Hedin 2003, Maddison et al. 2008). However, many of the deeper relationships of salticoids have been poorly resolved. Outside the Salticoida are the spartaeines, lyssomanines, and hisponines, showing ancestral features like limited tracheal systems, complex palpi, and the retention of a tarsal claw on the female palp. These non-salticoids (often called “basal salticids”) have been studied phylogenetically (Su et al. 2007), but with limited taxon sampling. In this work we attempt to resolve more firmly the basic structure of the family by increasing the taxon sampling, especially among non-salticoid salticids, and by using additional genes. Two of the genes, wingless and myosin heavy chain, are new to salticid molecular phylogenetics. By building a dataset that has a greater number of genes among selected species, we hoped to obtain a phylogenetic resolution with stronger confidence.

Methods

Taxon sampling

Taxa included in the analysis are 169 species of salticids and representatives of four dionychan families as outgroups (Table 1, Suppl. material 1). Based on previous phylogenetic work (Maddison and Hedin 2003, Maddison et al. 2008, Bodner and Maddison 2012, Zhang and Maddison 2013, in press), about 70 species of salticids from the major clade Salticoida were selected because they would represent most known major lineages, and because several genes are available for each (Table 1, Suppl. material 1). In addition, a few salticoids were added because their placement was unclear: , , , , , , and .
Table 1.

Specimens and sequences used in phylogenetic analyses, with GenBank numbers indicated. * marks previously published sequences. Specimen localities given in Suppl. material 1.

Reference28s18swinglessmyosin HCactin 5chistone 3CO116sND1
Outgroups
Anyphaenidae: Hibana sp.s318AY297295*KM033091KM032961KM032929AY297422*AY297295 / AY297358*
Gnaphosidae: Cesonia sp.s319AY297293; EF201663*KM032996EU522700*DQ665720*AY297420*AY296711 / AY297356*
Miturgidae: Cheiracanthium sp.s321AY297294; EF201664*KM032997KM032928AY297421*AY296712 / AY297357*
Oxyopidae: Oxyopes birmanicus Thorell, 1887Su et al. 2007EF419032 / EF419065*EF418998*EF419126*EF419097*EF418969 / EF419150*
Philodromidae: Philodromus alascensis Keyserling, 1884GR011KM033130KM033092KM032998KM032962
Thomisidae: Misumenops nepenthicola (Pocock, 1898)Su et al. 2007EF419029 / EF419062*EF418996*EF419123*EF419094*EF418967 / EF419148*
Thomisidae: Xysticus sp.s316AY297296; EF201665*KM033093EU522701*DQ665704*AY297296*AY296714 / AY297359*
Lyssomanines
Asemonea sichuanensis Song & Chai, 1992SC-03-0055EF418986*EF419082*
Asemonea sichuanensis Song & Chai, 1992MRB084KM033131KM032931
Asemonea cf. stella Wanless, 1980MRB083JX145767*KM033094KM032930JX145686*
Asemonea tenuipes (O. P.-Cambridge, 1869)d186KM033132KM033095KM032999KM032963KM032932
Chinoscopus cf. flavus (Peckham, Peckham & Wheeler, 1889)d273KM033133KM033096KM032888
Goleba lyra Maddison & Zhang, 2006d051DQ665768*KM033097KM033000EU522709*DQ665707*DQ665755*
Lyssomanes amazonicus Peckham & Wheeler, 1889ECU11-6112KM033134KM032889
Lyssomanes antillanus Peckham & Wheeler, 1889d298KM033135KM033001
Lyssomanes cf. benderi Logunov, 2002ECU11-5402KM033136KM032890
Lyssomanes cf. jemineus Peckham & Wheeler, 1889ECU11-5682KM033137KM032891
Lyssomanes longipes (Taczanowski, 1871)MRB086KM033138KM032933KM033208KM032892
Lyssomanes pauper Mello-Leitão, 1945d297KM033139KM033002
Lyssomanes taczanowskii Galiano, 1980ECU11-4193KM033141KM032894
Lyssomanes tenuis Peckham & Wheeler, 1889ECU11-4869KM033142KM032895
Lyssomanes viridis (Walckenaer, 1837)s160AY297231*AY297360*AY296652 / AY297297*
Lyssomanes viridis (Walckenaer, 1837)d129KM033098KM033003EU522715*DQ665715*
Lyssomanes sp. [Esmeraldas]d408KM033140KM032893
Onomastus nigrimaculatus Zhang & Li, 2005Su et al. 2007EF419031 / EF419064*EF418997*EF419125*EF419096*EF418968 / EF419149*
Onomastus sp. [Guangxi]MRB085JX145768*KM033099KM033004KM032964KM032934JX145687*JX145910*
Pandisus cf. decorus Wanless, 1980d303KM033143KM033005
Cocalodines
Allococalodes madidus Maddison, 2009d236KM033144KM033006KM032896
Cocalodes longicornis Wanless, 1982d291KM033145KM033007KM032935KM032897
Cocalodes macellus (Thorell, 1878)d230KM033146KM033100KM033008KM032936KM033209
Cucudeta gahavisuka Maddison, 2009d234KM033147KM033009KM032898
Cucudeta zabkai Maddison, 2009d235KM033148KM033010KM032965KM032899
Tabuina aff. baiteta Maddison, 2009d313KM033149KM033011
Tabuina rufa Maddison, 2009d232KM033151KM033013KM032900
Tabuina aff. rufa Maddison, 2009d312KM033150KM033012
Tabuina varirata Maddison, 2009d233KM033152KM033014KM032901
Yamangalea frewana Maddison, 2009d231KM033153KM033015KM032902
Spartaeines
Brettus cf. adonis Simon, 1900SWK12-4323KM033154
Brettus sp. [Yunnan]LiD-026-053-05KM033155SKM033101SKM033195S
cf. Phaeacius sp. [Sarawak]SWK12-3728KM033156
Cocalus murinus Simon, 1899LiD-013-027-05EF419019 / EF419053*EF418988*EF419116*EF419084*EF418959 / EF419140*
Cyrba algerina (Lucas, 1846)Su et al. 2007EF419021 / EF419054*EF418989*EF419086*EF418961 / EF419142*
Cyrba lineata Wanless, 1984MRB106JX145792*KM033016KM032966KM032937JX145704*
Cyrba ocellata (Kroneberg, 1875)Su et al. 2007EF418990*EF419087*EF418962 / EF419143*
Cyrba ocellata (Kroneberg, 1875)MRB104KM033157
Cyrba sp. [Kenya]Su et al. 2007EF419023 / EF419056*EF418991*EF419088*
Gelotia cf. bimaculata Thorell, 1890d250KM033158KM033017KM032938
Gelotia syringopalpis Wanless, 1984Su et al. 2007EF419024 / EF419057*EF419118*
Gelotia syringopalpis Wanless, 1984MRB105KM033019KM033212KM032903
Gelotia sp. [Guangxi]MRB199KM033018KM032939KM033210
Gelotia sp. [Yunnan]LiD002-053-05KM033102SKM033196SKM033211S
Holcolaetis vellerea Simon, 1910Su et al. 2007EF419025 / EF419058*EF418992*EF419119*EF419090*EF418963 / EF419144*
Holcolaetis cf. zuluensis Lawrence, 1937d036DQ665770*KM033103EU522711*DQ665721*DQ665757*
Meleon aff. kenti (Lessert, 1925)d287KM033159KM032940
Mintonia mackiei Wanless, 1984SWK12-4202KM033161
Mintonia cf. melinauensis Wanless, 1984d441KM033160
Mintonia ramipalpis (Thorell, 1890)SWK12-1442KM033162
Mintonia silvicola Wanless, 1987d104KM033020KM032904
Mintonia silvicola Wanless, 1987SWK12-1653KM033163
Mintonia silvicola Wanless, 1987Su et al. 2007EF418995*EF419122*EF419093*
Mintonia tauricornis Wanless, 1984d249KM033164KM033021KM032941KM032905
Neobrettus tibialis (Prószyński, 1978)LiD-001-055-05EF419030 / EF419063*EF419124*EF419095*
Neobrettus sp. [Sarawak]SWK12-1040KM033165
Paracyrba wanlessi Zabka & Kovac, 1996Su et al. 2007EF419033 / EF419066*EF418999*EF419098*
Phaeacius lancearius (Thorell, 1895)d111DQ665775*KM033022DQ665759*
Phaeacius malayensis Wanless, 1981Su et al. 2007EF419034 / EF419067*EF419000*EF419099*EF418970 / EF419151*
Phaeacius sp. [Guangxi]LQ-24-06KM033166SKM033104SKM033213SKM032906S
Phaeacius sp. [Hainan]Su et al. 2007EF419035 / EF419068*EF419001*EF418971 / EF419152*
Phaeacius sp. [Sarawak]SWK12-4541KM033167
Portia africana (Simon, 1886)Su et al. 2007EF419037 / EF419069*EF419003*EF419128*EF419101*
Portia crassipalpis (Peckham & Peckham, 1907)SWK12-2354KM033168
Portia fimbriata (Doleschall, 1859)LiD-001-04EF419038 / EF419070*EF419004*EF419129*EF419102*EF418973 / EF419154*
Portia heteroidea Xie & Yin, 1991Su et al. 2007EF419039 / EF419071*EF419005*EF419130*EF419103*EF418974 / EF419155*
Portia jianfeng Song & Zhu, 1998Su et al. 2007EF419040 / EF419072*EF419006*EF419104*EF418975 / EF419156*
Portia labiata (Thorell, 1887)S206AY297232*AY297361*AY296653 / AY297298*
Portia cf. schultzi Karsch, 1878d131DQ665776*KM033105KM033023KM032967EU522718*DQ665708*
Portia quei Zabka, 1985Su et al. 2007EF419042 / EF419074*EF419008*EF419132*EF419106*EF418977 / EF419158*
Portia taiwanica Zhang & Li, 2005MRB103KM033169KM032942KM033214KM032907
Portia sp. [Sichuan]SC-03-0011EF419043 / EF419075*EF419009*EF419133*EF418978 / EF419159*
Sonoita lightfooti Peckham & Peckham, 1903d226KM033170KM033215
Sonoita aff. lightfooti Peckham & Peckham, 1903MRB200JX145791*JX145705*JX145927*
Sparbambus gombakensis Zhang, Woon & Li, 2006d251KM033171KM033024KM032943
Spartaeus jianfengensis Song & Chai, 1991Su et al. 2007EF419045 / EF419076*EF419011*EF419109*EF418980 / EF419161*
Spartaeus platnicki Song, Chen & Gong, 1991SC-03-069EF419046 / EF419077*EF419012*EF419135*EF419110*EF418981 / EF419162*
Spartaeus spinimanus (Thorell, 1878)S199KM033216KM032908
Spartaeus thailandicus Wanless, 1984BV-004EF419047 / EF419078*EF419013*EF419136*EF419111*EF418982 / EF419163*
Spartaeus uplandicus Barrion & Litsinger, 1995S185/S186AY297233*AY297363*AY296655*
Spartaeus wildtrackii Wanless, 1987Su et al. 2007EF419048 / EF419079*EF419014*EF419137*EF419112*EF418983 / EF419164*
Taraxella sp. [Johor]d246KM033172KM032944KM032909
Taraxella sp. [Pahang]d248KM033173KM032945KM033197
Taraxella sp. [Pahang]LiD-001-003-06KM033106SKM033217SKM032910S
Yaginumanis wanlessi Zhang & Li, 2005Su et al. 2007EF419050 / EF419081*EF419016*EF419139*EF419114*EF418985 / EF419166*
Lapsiines
Galianora bryicola Maddison, 2006d124DQ665771*DQ665741*KM033025EU522706*DQ665717*DQ665758*DQ665727*
Galianora sacha Maddison, 2006d116DQ665766*DQ665734*KM033026KM032968EU522707*DQ665716*DQ665754*
Lapsias canandea Maddison, 2012d442KM033174
Lapsias guamani Maddison, 2012UBC-SEM AR00191KM033175KM033027
Lapsias lorax Maddison, 2012UBC-SEM AR00194KM033176KM033028
Soesiladeepakius lyra Ruiz & Maddison, 2012GR130JQ312077KM033029JQ312074*JQ312079*
Thrandina bellavista Maddison, 2012d396KM033177KM033030
Thrandina cosanga Maddison, 2012d395KM033178
Thrandina parocula Maddison, 2006d123DQ665779*KM033107EU522720*DQ665718*DQ665761*DQ665726*
Thrandina parocula Maddison, 2006d394KM033031KM032969
Eupoa
Eupoa nezha Maddison & Zhang, 2007d220/MRB102EF201648*EF201666*KM033032EF201668*EF201667*
Hisponines
cf. Tomocyrba sp. [Madagascar]d305KM032881*
Hispo macfarlanei Wanless, 1981d404KM032882*KM032970
Hispo sp. [Madagascar]d309KM032883*
Jerzego cf. alboguttatus Simon, 1903SWK12-4787KM032884*
Jerzego corticicola Maddison, 2014SWK12-2900KM032885*KM032887*
Massagris contortuplicata Wesolowska & Haddad, 2013d082DQ665772*KM033108KM033033DQ665705*DQ665722*
Massagris schisma Maddison & Zhang, 2006d081DQ665762*KM033109KM033034DQ665728*
Tomobella andasibe (Maddison & Zhang, 2006)d127DQ665780*DQ665752*KM033035KM033198DQ665725*
Tomocyrba sp. [Madagascar]d306KM032886*
Tomomingi sp. [Gabon]MRB243JX145764*KM033110KM033036KM032971JX145850*JX145684*
Salticoida
Agoriines
Agorius constrictus Simon, 1901d172KM032953
Agorius constrictus Simon, 1901d213KM033119KM033072KM032921
Agorius sp. [Selangor]d299KM033189KM033073
Synagelides cf. lushanensis Xie & Yin, 1990d214KM033074
Synagelides cf. palpalis Zabka, 1985MRB050KM032922
Synagelides cf. palpalis Zabka, 1985d225KM033190KM033226
Amycoids
Cotinusa sp. [Ecuador]MRB024JX145746*KM033120KM033075KM032987JX145832*JX145671*JX145896*
Hurius vulpinus Simon, 1901S213AY297239*AY297368*AY296662 / AY297306*
Hurius cf. vulpinus Simon, 1901d156KM033076EU522712*KM033203
Hypaeus aff. miles Simon, 1900 [Ecuador]d130EU815499*KM033121KM033077KM032988EU522702*KM032923
Sarinda cutleri (Richman, 1965)MRB193JX145744*KM033078KM032954JX145669*JX145895*
Sitticus floricola palustris (Peckham & Peckham, 1883)d030DQ665778*KM033122KM033079KM032989KM033204DQ665760*DQ665729*
Astioids
Arasia mollicoma (L. Koch, 1880)d046EU815483*EU815532*KM032990JX145834*KM033205EU815598*EU815550*
Helpis minitabunda (L. Koch, 1880)d265KM033123KM033080KM032991KM032955KM033227
Ligurra latidens (Doleschall, 1859)d175JX145749*KM033081JX145835*JX145898*
Ligurra latidens (Doleschall, 1859)LiD-001-027-05EF418993*EF419120*EF419091*
Mopsus mormon Karsch, 1878d018EU815470*EU815529*KM033082JX145836*KM033206EU815586*
Myrmarachne sp. [Pahang]d162EU815507*KM033124KM033083KM032992JX145837*EU815616*EU815565*
Neon reticulatus (Blackwall, 1853)d283KM033191KM033125KM033084KM032993KM032956
Nungia epigynalis Zabka, 1985d221KM033192KM032924
Simaetha sp.d027EU815477*KM033126KM033085JX145839*EU815592*EU815546*
Trite pennata Simon, 1885d035EU815478*KM033086KM032957KM033207EU815593*EU815547*
Baviines
Bavia aff. aericeps Simon, 1877 [Sabah]d079EU815490*KM033127KM032958EU815603*KM032925
Stagetilus sp. [Selangor]MRB079KM033193KM033087KM032959KM032926
Marpissoids
Afromarengo sp. [Gabon]MRB262JX145758*KM033128KM033088KM032994JX145842*JX145682*JX145905*
Dendryphantes hastatus (Clerck, 1757)d043EF201646*KM033129KM033089KM033228KM032927
Platycryptus californicus (Peckham & Peckham, 1888)d316KM033194KM033090KM032995KM032960KM033229
Rhene sp. [Pahang]LiD-001-021-05EF419044*EF419010*EF419134*EF419108*EF418979 / EF419160*
Tisaniba mulu Zhang & Maddison, 2014SWK12-1244KM032876*KM032880*
Saltafresians
Aelurillus cf. ater (Kroneberg, 1875)d140EU815504*EU815536*KM033037KM032972JX145831*KM033199EU815615*EU815564*
Amphidraus complexus Zhang & Maddison, 2012JXZ035KC615380*KM033038KC616069*KC615640*KC615806*
Athamas cf. whitmeei O. P.-Cambridge, 1877JXZ345KC616286*KC615649*KC615822*
Bacelarella pavida Szüts & Jocqué, 2001d195EU815511*EU815538*KM033039KM032973KM032946EU815618*EU815569*
Bathippus macrognathus (Thorell, 1881)JXZ372KC615407*KM033040KC616305*KC615835*
Bianor maculatus (Keyserling, 1883)d017EU815469*KM033041KM033200EU815585*EU815542*
Bristowia afra Szüts, 2004JXZ363KC615409*KC616301*
Bristowia afra Szüts, 2004MRB230KM033042KM033218
Cheliceroides longipalpis Zabka, 1985d222KM033111KM033043JX145830*KM033219EU815579*
Cheliceroides cf. longipalpis Zabka, 1985d415KM033179
Chinattus parvulus (Banks, 1895)d009EU815464*EU815525*KM033044JX145848*KM033201EU815581*
Chinophrys pengi Zhang & Maddison, 2012JXZ145KC615416*KM033045KC616146*KC615843*
Corythalia locuples (Simon, 1888)JXZ315KC615390*KM033046KC616260*KC615645*KC615816*
Cosmophasis umbratica Simon, 1903Su et al. 2007EF419020*EF419117*EF419085*EF418960 / EF419141*
Cytaea nimbata (Thorell, 1881)JXZ229KC615474*KM033047KC616197*KC615693*KC615899*
Diolenius varicus Gardzińska & Zabka, 2006JXZ349KC615480*KM033048KC616290*KC615695*KC615905*
Diplocanthopoda marina Abraham, 1925d209KM033180KM032947KM033220KM032911
Eburneana sp. [Gabon]MRB231KM033181KM033049JX145858*KM033221KM032912
Echeclus sp. [Selangor]MRB089KM033182KM032948KM033222KM032913
Euophrys frontalis (Walckenaer, 1802)JXZ137KC615536*KM033050KC616139*KC615960*
Evarcha proszynskii Marusik & Logunov, 1998d096DQ665765*KM033112EU522704*DQ665723*
Evarcha proszynskii Marusik & Logunov, 1998d323KM033051KM032974
Freya decorata (C. L. Koch, 1846)d211EU815521*EU815539*KM032975EU522705*JX145908*
Gedea cf. tibialis Zabka, 1985MRB090KM033183KM032949KM033223KM032914
Habrocestum cf. albimanum Simon, 1901d132EU815500*EU815611*EU815562*
Habronattus borealis (Banks, 1895)d207KM033184KM033052KM032976KM032950KM033224KM032915
Hasarius adansoni (Audouin, 1826)d295KM033113KM033053KM032977
Hasarius adansoni (Audouin, 1826)S130/S131/S324AY297281*AY297409*
Heliophanus cupreus (Walckenaer, 1802)d044DQ665769*KM033114EU522710*DQ665710*DQ665756*KM032916
Idastrandia cf. orientalis (Szombathy, 1915)d108EU815535; EU815496*EU815535*JX145852*EU815608*EU815560*
Langerra aff. longicymbium Song & Chai, 1991d182KM033185KM033054KM032917
Leptorchestes berolinensis (C. L. Koch, 1846)d086EU815491*EU815534*KM033055EU815604*EU815556*
Longarenus brachycephalus Simon, 1903MRB258JX145798*KM033056KM032978KM032951JX145707*KM032918
Nannenus sp. [Pahang]d105EU815493*KM033057KM032979JX145853*EU815558*
Naphrys pulex (Hentz, 1846)JXZ081JX145760*KM033115KM032980JX145844*KC615749*JX145907*
Omoedus orbiculatus (Keyserling, 1881)d008KC615792*
Omoedus orbiculatus (Keyserling, 1881)JXZ136JX145762*KM033116KM033058JX145846*KM033202
Omoedus papuanus Zhang & Maddison, 2012JXZ286KC615619*KM033059KC616234*KC615790*KC616042*
Pellenes peninsularis Emerton, 1925d057DQ665774*KM033117KM033060JX145864*DQ665712*
Pellenes peninsularis Emerton, 1925d400KM032981
Phaulostylus grammicus Simon, 1902d304KM033186KM033061
Philaeus chrysops (Poda, 1761)d025EU815475*EU815530*KM033062JX145855*EU815590*EU815545*
Phintella sp. [Gabon]d402KM033187KM033063KM032982
Plexippus paykulli (Audouin, 1826)LiD-001-029-05EF419002*EF419127*
Plexippus paykulli (Audouin, 1826)MRB016JX145784*KM033064EU522713*
Plexippus paykulli (Audouin, 1826)S73AY297384*AY296674 / AY297317*
Pochyta cf. pannosa Simon, 1903MRB257JX145806*KM033065KM032983KM032952JX145715*KM032919
Saitis barbipes (Simon, 1868)JXZ147KC615589*KM033066KC616147*KC615767*KC616011*
Salticus scenicus (Clerck, 1757)d003DQ665777*KM033118KM033067KM032984EU522719*DQ665713*JX145663*AY296707 / AY297352*
Thiania bhamoensis Thorell, 1887LiD-001-028-05EF419049 / EF419080*EF419015*EF419138*EF419113*EF418984 / EF419165*
Trydarssus cf. nobilitatus (Nicolet, 1849)MRB270KM033188KM033068KM032985JX145847*KM033225KM032920
Tusitala lyrata (Simon, 1903)MRB226JX145771*KM033069JX145856*JX145689*JX145912*
Yllenus arenarius Menge, 1868d013EU815527*EU815583*EU815541*
Yllenus arenarius Menge, 1868JXZ173JX145766*KM033070KM032986JX145851*
Zabkattus furcatus Zhang & Maddison, 2012JXZ218KC615503*KM033071KC616190*KC615928*
Specimens and sequences used in phylogenetic analyses, with GenBank numbers indicated. * marks previously published sequences. Specimen localities given in Suppl. material 1. Our sample targeted especially the non-salticoid salticids, those that lie outside the major clade of familiar salticids (Maddison and Needham 2006). We included PageBreakmost available data from non-salticoid salticids, both new data and data previously published by Su et al. (2007) and others (Maddison and Hedin 2003, Maddison and Needham 2006, Maddison et al. 2007, Bodner and Maddison 2012, Ruiz and Maddison 2012, Zhang and Maddison 2013, Maddison and Piascik, in press). Included for the first time in a molecular phylogeny are the cocalodines (Wanless 1982, Maddison 2009), which are Australasian non-salticoid salticids. Also analyzed for the first time are the lyssomanine genera and , the lapsiine , and the spartaeines , , , and . Some previously-published data from non-salticoid salticids was either excluded or represented under a different species name here. Excluded are sequences of cf. frenata, because its limited data made it unstable in the analyses (see Maddison and Piascik 2014), “” from Su et al. (2007), because its identification is in doubt and no voucher specimen is available, and the actin 5C sequence of sp. voucher d243, which we discovered to have been a contaminant from the euophryine . The species labeled as by Su et al. (2007) is included here as “ sp. [Hainan]”, because the specimen was a juvenile female and thus identified with doubt; by its DNA we suspect it is . The specimen labeled as by Su et al. (2007) is actually a female . This misidentification arose because of an error in male-female matching by Wanless (1984), whose female “” is actually the female of . The correct match of male and female is evident by intimate co-collecting in a recent expedition to Sarawak (Maddison and Piascik, unpublished) and in DNA sequence comparison. We have therefore blended data from Su et al.’s female with that from our males to represent . Some of the species studied appear to be undescribed, or are doubtfully the same as described species. Following the usual convention, the names of some of our specimens includes “cf.” to indicate that they may be the same as the mentioned species, “aff.” to indicate that they are close to, but distinctly different from, the mentioned species. Figures 1–13 give illustrations of some of the undescribed species, in order to facilitate future association of our data with a species name. The species we refer to as “cf. PageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreak [Sarawak]” (Figs 1, 2) is known from a single female and juvenile from Lambir Hills, Sarawak. It resembles but the legs are shorter, and the epigynum is distinctively different. sp. [Sarawak] (Figs 3, 4) is a fairly typical whose epigynum resembles that of Wijesinghe, 1991, but with the atria elongated posteriorly. sp. [Guangxi] is shown in Fig. 5. aff. lightfooti (Fig. 6) has longer grooves for the openings of the epigynum than , and is distinctive in gene sequences as well. sp. [Guangxi] (Fig. 7) has a palp resembling , but the tibial apophyses are much shorter. sp. [Selangor] (Figs 8, 9) was identified as an by the distinctive form of the palp tibia, and the embolus hidden behind a ledge of the tegulum, through which several dark sclerites can be seen (Prószyński 1987). It might equally well have been identified, by the same features, as a species (Zabka 1988). Indeed, the two genera are likely synonyms. “” is used as that is the older name. sp. [Johor] (Figs 10, 11) and sp. [Pahang] (Figs 12, 13) are typical species of . The specimen MRB024 identified as sp. is the same as that named “unidentified thiodinine” by Bodner and Maddison (2012). The specimen (d130) was formerly identified as sp. (Bodner and Maddison 2012). The specimen d105 labeled as “” by Maddison et al. (2008) is not , but another apparently undescribed species of . The data for comes from two specimens, d222 which is clearly , and d415 which may be a different but very closely related species. Notes on the undescribed hisponines are given by Maddison and Piascik (2014), whose data we use.
Figures 1–13.

Specimens of undescribed species. 1, 3, 5, 6 are of epigyna, ventral view; 8, 11, 13 of left palps, ventral view; 7 of the right palp tibia, retrolateral view. Scale bar 0.1 mm. 1–2 Female cf. [Sarawak], voucher SWK12–3728 3–4 Female sp. [Sarawak], voucher SWK12–4541 5 female sp. [Guangxi], voucher MRB085 6 Female aff. lightfooti, voucher MRB200. 7 male sp. [Guangxi], voucher MRB199. The drawing is reversed so as to appear to be the left palpus 8–9 Male sp. [Selangor], voucher MRB089 10–11 Male sp.[Johor], voucher d246 for the palpus. The photo of the living male may or may not be of the same specimen 12–13 Male sp. [Pahang], voucher d248. The photo of the living male may or may not be of the same specimen. Figures 1–13 are copyright ©2014 W.P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license.

Specimens of undescribed species. 1, 3, 5, 6 are of epigyna, ventral view; 8, 11, 13 of left palps, ventral view; 7 of the right palp tibia, retrolateral view. Scale bar 0.1 mm. 1–2 Female cf. [Sarawak], voucher SWK12–3728 3–4 Female sp. [Sarawak], voucher SWK12–4541 5 female sp. [Guangxi], voucher MRB085 6 Female aff. lightfooti, voucher MRB200. 7 male sp. [Guangxi], voucher MRB199. The drawing is reversed so as to appear to be the left palpus 8–9 Male sp. [Selangor], voucher MRB089 10–11 Male sp.[Johor], voucher d246 for the palpus. The photo of the living male may or may not be of the same specimen 12–13 Male sp. [Pahang], voucher d248. The photo of the living male may or may not be of the same specimen. Figures 1–13 are copyright ©2014 W.P. Maddison, released under a Creative Commons Attribution (CC-BY) 3.0 license. Specimens whose voucher ID’s (Table 1, Suppl. material 1) are of the form S###, d###, MRB###, or JXZ###, SWK12-####, or ECU11-####, where # is a digit, are deposited in the Spencer Entomological Collection of the Beaty Biodiversity Museum, University of British Columbia. The remaining vouchers are in the Lee Kong Chian Natural History Museum (formerly Raffles Museum for Biodiversity Research or RMBR), National University of Singapore. In addition to analyses done on all 176 sampled taxa (“Complete”), subsets of taxa were analyzed alone. A first subset (“Salticoida”) of 78 taxa highlighted the Salticoida, with just 7 non-salticoid outgroup taxa (4 hisponines, 1 spartaeine, 1 cocalodine, 1 lapsiine), in order to obtain an alignment that was less perturbed by highly divergent non-salticoids. A second subset highlighted the non-salticoids (“Non-salticoid”, 120 taxa), to obtain an alignment primarily for non-salticoid salticids, and also to be able to explore their relationships in more detail.

Gene choice and sequencing

Eight genes were used for this analysis. Two are nuclear ribosomal genes, 28s and 18s (Maddison and Hedin 2003, Maddison et al. 2008). Four are nuclear protein coding genes: actin 5C (Vink et al. 2008, Bodner and Maddison 2012), wingless (Blackledge et al. 2009), myosin heavy chain (“myosin HC”, Blackledge and Hayashi, unpublished), and histone 3 (Su et al. 2007). Two mitochondrial regions were also used, CO1 and another region including 16s and NADH1 ("16sND1", Hedin and Maddison 2001, Maddison and Hedin 2003). Following Bodner and Maddison (2012), the intron region of actin 5C was deleted from the analyses as it is highly variable and difficult to align. The sequencing protocols for wingless and myosin HC are described below. For other genes, sequences marked “S” in Table 1 and Suppl. material 1 were obtained by the protocols of Su et al. (2007), all others by the protocols of Bodner and Maddison (2012) and Zhang and Maddison (2013). For most PageBreakwingless sequences, the forward and reverse primers used were respectively Spwgf1 and Spwgr1 (Blackledge et al. 2009). PCR amplification included a 2 min 94 °C denaturation and 35 cycles of 30 s at 94 °C, a 30 s annealing step at 52–57 °C, 30 s at 72 °C and one 3 min extension step at 72 °C. For some specimens this did not succeed in amplifying wingless, and for those we used a nested protocol starting with outer primers wg550F and wgABRz (Wild and Maddison 2008). The resulting product was then amplified using two internal primers, forward Wnt8MBf1 5’-TGTGCTACTCARACKTGYTGG-3’ and reverse Wnt8MBr3 5’-ACAAWGTTCTGCA ACTCATRCG-3’. For both the external and internal reactions amplification was done with 2 min 94 °C denaturation and 37 cycles of 20 s at 94 °C, a 20 s annealing step at 52 °C (wg550F/wgABRz) or 56 °C (wnt8MBf1/wnt8MBr3), and 2 min at 72 °C, and no final extension. The nested protocol obtained sequences for aff. aericeps (voucher d389), (d295), sp. (GR011), sp. (d027), and (JXZ173). In other specimens, the nested protocol often resulted in amplification of a different member of the wingless family (e.g. WNT-8), but these were readily detected (and excluded) by BLASTing them to other genes in the NCBI database (http://www.ncbi.nlm.nih.gov). The region of myosin HC sequenced corresponds mostly to an intron. Primers used are (forward) Myhc1f 5'-ACAACAATTCTTCAACCATCAC-3' and (reverse) Myhc5r 5'-CTTCCTCAAGGATGGACA-3' (Blackledge and Hayashi, unpublished). PCR amplification included a 2 min 95 °C denaturation and 35 cycles of 20–45 s at 95 °C, a 45 s annealing step at 52 °C, 1 min at 72 °C and one 10 min extension step at 72 °C. The boundary between the exon and intron was determined by aligning the salticid implicit amino acid translations against the known transcript for myosin HC in (Genbank accession AAM97635.1; Ruiz-Trillo et al. 2002). Two small single-nucleotide errors in the sequences were corrected after the analyses but before submission to Genbank. These are near the ends of CO1 of MRB199 ( sp. [Guangxi]) and MRB231 ( sp. [Gabon]). Given that CO1 had little resolution, these are unlikely to have affected the results.

Sequence alignment

Automatic multiple sequence alignment was performed by MAFFT (Katoh et al. 2002, 2005), run via the align package of Mesquite (prerelease of version 3, Maddison and Maddison 2014), aided by Mesquite for manual corrections and for alignment by amino acid. Coding regions were easily aligned by hand according to amino acid translations. This was done starting with an initial automated nucleotide alignment, followed by hand correction in Mesquite using the Color Nucleotide By Amino Acid function to reveal amino acid translation. Non-coding regions (28s, 18s, noncoding region of 16sND1, myosin HC intron) were aligned by MAFFT using the L-INS-i option (--localpair --maxiterate 1000). Mesquite was used to color the matrix via the option ‘‘Highlight Apparently Slightly Misaligned Regions’’ so as to identify regions that needed correction. These were almost always near the ends of sequences. Alignment was done separately on the Complete, Non-salticoid and Salticoida datasets. Following the MAFFT alignment, the Salticoida dataset required 5 small realignments by hand in 18s. The first 60 positions in the initial alignment of 16s were also realigned locally, and in addition 8 minor shifts by one or two positions were made by hand. The Non-salticoid dataset required three simple hand fixes in 28s. The PageBreakfirst 24 positions of 16s in the initial alignment were realigned by MAFFT in isolation because of several obvious misalignments. The Complete dataset appeared poorly aligned in 28s from sites 375 to 489 in the initial alignment, which were therefore realigned by MAFFT in isolation. The first 60 positions in the initial alignment of 16s were also realigned locally, and in addition 8 minor shifts by one or two positions were made by hand. Five small shifts were performed by hand for 18s. Many analyses were done with different variants of the alignments as this study was progressing, and the phylogenetic trees remained substantially consistent.

Phylogenetic analysis

Phylogenetic analyses using maximum likelihood were run using RAxML version 7.2.8alpha (Stamatakis 2006a, 2006b). The protein coding genes and 16sND1 were each divided into partitions. Protein coding regions were divided into one partition for 1st and 2nd codon positions, and another partition for third codon positions. Introns and non-coding regions were treated as separate partitions. For the fused 8 gene analyses, there were 7 partitions total: (1) 1st + 2nd codon positions in nuclear genes, (2) 3rd codon position nuclear, (3) nuclear intron, (4) nuclear ribosomal, (5) 1st + 2nd codon positions mitochondrial, (6) 3rd codon position mitochondrial, (7) noncoding mitochondrial. Each partition was permitted to have its own model parameters. Analyses were done for each gene region separately with the Complete taxon set. In addition, analyses fusing all 8 genes were done for the Non-salticoid and Salticoida taxon sets. For all of these, RAxML runs assuming the GTRCAT model were used with 100 search replicates, to seek maximum likelihood trees. In addition, likelihood bootstrap analysis was performed with 500-1500 bootstrap replicates (as indicated in the figures), each involving a single search replicate. Phylogenetic analyses using GARLI version 1.0.699 (Zwickl 2006) under the model GTR+gamma+I were also done but are not reported; they resulted in substantially similar trees.

Data resources

The data underpinning the analyses reported in this paper are deposited in the Dryad Data Repository at doi: 10.5061/dryad.v53h1.

Results

Sequences obtained and used in analyses are indicated in Table 1 and Suppl. material 1, along with those sequences taken from the literature. Figure 14 summarizes the results of the phylogenetic analyses, which are given in more detail in Figures 15–27. Colors assigned to clades in Figure 14 are shown in the remaining figures. Figures 15–19 show the All Genes results for the Complete, Non-PageBreakPageBreakPageBreaksalticoid and Salticoida datasets. Figures 20–27 show the results for individual genes analyzed separately.
Figure 14.

Summary of phylogenetic results. Number above branch shows percentage of maximum likelihood bootstrap replicates with clade. For clades outside the Salticoida, these percentages come from the Non-salticoid dataset with 1500 replicates; within the Salticoida, these come from the Salticoida dataset with 1000 replicates; the Salticoida percentage comes from the Complete dataset with 1000 replicates. Long bar on branch shows same percentage graphically: black 91–100%; dark gray 81–90%; gray 71–80%; light gray 51–70%. Oval spots show presence of clade in maximum likelihood tree for individual genes, with exceptions noted by * and adjacent notes. The notes about wingless on the Spartaeinae node and actin on the Salticoida node are ambiguous in placement; they could equally well have been placed one node deeper because of missing data. Pale gray outline indicates no conclusion because of inadequate taxon sampling. All indications of support are from analyses excluding , agoriines, and “” uplandicus. Bars show colors used to highlight taxa in Figs 15–27.

Figure 15.

Phylogeny from complete taxon sample, All Genes analysis. Numbers beside branches show percentage of 1000 RAxML likelihood bootstrap replicates with clade in analysis with and agoriines excluded. In analyses with these taxa included (500 bootstrap replicates), bootstrap percentages are within 5 of those shown, except for branches with two values (e.g. “100/60”), in which case the first value is from an analysis with and agoriines excluded, the second value with them included. Colors of branches are the same as those highlighting taxa in Fig. 14.

Figures 23–27.

Phylogeny from gene regions analyzed alone, complete taxon sample. 23 myosin HC 24 actin 5C 25 Histone 3 26 CO1 27 16sND1. Colors of branches are the same as those highlighting taxa in Fig. 14.

Figures 18–19.

Phylogeny from Salticoida dataset, All Genes analysis. 18 Salticoida analysis with all taxa included 19 Salticoida analysis with and excluded. Numbers beside branches show percentage of 1000 RAxML likelihood bootstrap replicates with clade. Colors of branches are the same as those highlighting taxa in Fig. 14.

Figures 20–22.

Phylogeny from gene regions analyzed alone, complete taxon sample. 20 28s 21 18s 22 wingless. Colors of branches are the same as those highlighting taxa in Fig. 14.

Summary of phylogenetic results. Number above branch shows percentage of maximum likelihood bootstrap replicates with clade. For clades outside the Salticoida, these percentages come from the Non-salticoid dataset with 1500 replicates; within the Salticoida, these come from the Salticoida dataset with 1000 replicates; the Salticoida percentage comes from the Complete dataset with 1000 replicates. Long bar on branch shows same percentage graphically: black 91–100%; dark gray 81–90%; gray 71–80%; light gray 51–70%. Oval spots show presence of clade in maximum likelihood tree for individual genes, with exceptions noted by * and adjacent notes. The notes about wingless on the Spartaeinae node and actin on the Salticoida node are ambiguous in placement; they could equally well have been placed one node deeper because of missing data. Pale gray outline indicates no conclusion because of inadequate taxon sampling. All indications of support are from analyses excluding , agoriines, and “” uplandicus. Bars show colors used to highlight taxa in Figs 15–27. Phylogeny from complete taxon sample, All Genes analysis. Numbers beside branches show percentage of 1000 RAxML likelihood bootstrap replicates with clade in analysis with and agoriines excluded. In analyses with these taxa included (500 bootstrap replicates), bootstrap percentages are within 5 of those shown, except for branches with two values (e.g. “100/60”), in which case the first value is from an analysis with and agoriines excluded, the second value with them included. Colors of branches are the same as those highlighting taxa in Fig. 14. Phylogeny from Non-salticoid dataset, All Genes analysis. Numbers beside branches show percentage of RAxML likelihood bootstrap replicates with clade. 16 Non-salticoid analysis with all taxa included (1500 bootstrap replicates used) 17 Non-salticoid analysis with , , and “” uplandicus excluded (500 bootstrap replicates used). Colors of branches are the same as those highlighting taxa in Fig. 14. Phylogeny from Salticoida dataset, All Genes analysis. 18 Salticoida analysis with all taxa included 19 Salticoida analysis with and excluded. Numbers beside branches show percentage of 1000 RAxML likelihood bootstrap replicates with clade. Colors of branches are the same as those highlighting taxa in Fig. 14. Phylogeny from gene regions analyzed alone, complete taxon sample. 20 28s 21 18s 22 wingless. Colors of branches are the same as those highlighting taxa in Fig. 14. Phylogeny from gene regions analyzed alone, complete taxon sample. 23 myosin HC 24 actin 5C 25 Histone 3 26 CO1 27 16sND1. Colors of branches are the same as those highlighting taxa in Fig. 14. Several taxa stood out as being problematical, especially for nuclear ribosomal genes. PageBreakPageBreakPageBreakPageBreak was not only difficult to sequence (Maddison et al. 2007) but its 28s and 18s genes stand as outliers in alignments, remarkably different from other salticids. The same holds for the agoriines and and, in 28s, for the hasariine . These sequences do not appear to be contaminants, as they BLAST in the NCBI database to salticids. In analyses with just 28s or 18s, these taxa tend to appear on long branches, wandering to different parts of the salticid phylogeny in different analyses, attaching themselves together and to clearly inappropriate relatives (e.g. within the pellenines, Fig. 21). This instability and unexpected placement are likely artifacts due to long branch attraction (Felsenstein 1978), possibly related to compositional bias (Hasegawa and Hashimoto 1993). and the agoriines have the highest GC bias of the sample (0.72–0.78, compared to 0.60–0.69 for all other species) in 28s, and are similar outliers in 18s. With wingless, appears on a normal-length branch (Fig. 22). However, the agoriines with wingless are on a long branch in an unlikely place, within the euophryines (Fig. 22). Their placement is unstable: in slight variants of the analyses they come out in other places. There is nothing obviously unusual about the wingless sequences in agoriines, but whatever has shifted the GC bias in the nuclear ribosomal genes might also be affecting the rest of the genome. When and the agoriines are excluded from analyses, bootstrap percentages rise through much of the tree, suggesting their instability is adding noise to the other relationships in the tree. For this reason, the reported bootstrap percentages and other indications of support are generally those for analysis with and the agoriines excluded. was left in the bootstrap analyses, because CO1, actin 5C and 16sND1 all agree on a clear placement in the hasariines.

Discussion

Many of the salticid clades now recognized by molecular data had been previously recognized by morphological data. For instance, Wanless (1980, 1984, 1985) recognized the three distinct lyssomanine groups and the Spartaeinae. The Salticoida was strongly supported by many morphological characters (Maddison 1988, 1996, Maddison and Hedin 2003), except that the status of the hisponines was unclear. Wanless (1981) implicitly included the hisponines within the salticoids, while Maddison (1996) did not consider the hisponines in his listing of salticoid synapomorphies. Other groups whose previous formulation by morphology mostly or entirely matches their current boundaries by molecular data are the marpissines (Barnes 1958), euophryines (Prószyński 1976), amycines (Galiano 1968), heliophanines (Maddison 1987), dendryphantines (Maddison 1996), and plexippines (Maddison 1988). At the finer scale, morphological systematics gave us concepts for many genera that are concordant with more recent data. However, the first molecular data for salticid phylogeny as a whole (Maddison and Hedin 2003) uncovered several unanticipated groups, including the Amycoida, Plexippoida, and Marpissoida. Further data revealed the Astioida and Aelurilloida (Maddison et al. 2008), and later the Saltafresia (Bodner and Maddison 2012). These are major groups within the Salticoida, each uniting several subfamilies.

Deepest relationships

Our results help resolve or add strength to relationships at the deepest level of salticid phylogeny. Wanless (1980) recognized three major subdivisions of lyssomanines: (1) the New World genera and , (2) the Asian , and (3) the remaining Old World genera including . He suggested these three groups are so distinct that they may not belong together. The molecular data agree: the three groups’ divisions are so deep that their relationships have not yet been recovered, and it is possible, even likely, that they do not form a monophyletic group. Different analyses give different results of the relationships of these three, with some showing the New World genera as sister to the spartaeine-lapsiine-cocalodine clade (as recovered by Su et al. 2007), other results showing in that role, and others showing the three lyssomanine groups together. Spartaeines, lapsiines and cocalodines form a clade (node 1, Fig. 14). Although Rodrigo and Jackson (1992) concluded that spartaeines, PageBreak and the group form a clade (they were unaware of lapsiines), our analysis provides the first support for such an arrangement – their analysis included only a single taxon outside the group, and therefore it could not speak to the monophyly of the group. Our new result is intuitively appealing, as it groups together all of the extant medium-sized generalized non-salticoids/non-hisponines that are typically brown or gray. However, these presumably are or could be plesiomorphic traits; there had been no obvious reason to expect the spartaeines, lapsiines and cocalodines should have fallen together. There is no known morphological synapomorphy of this clade. Within this spartaeine-lapsiine-cocalodine clade, the subclade historically best known by morphology is Wanless’s (1984) narrow version of the Spartaeinae, delimited by the presence of a tegular furrow (Wanless 1984). The Spartaeinae sensu stricto is primarily Afro-Eurasian, with a few Australasian species. Outside of this clade, there are no clear morphological synapomorphies defining subclades, and yet there is a striking geographical pattern: all of the Neotropical species belong to a clade, thus forming the lapsiines, while all of the Australasian species belong to a clade, thus forming the cocalodines. It is unsatisfying that we lack morphological synapomorphies for the lapsiines or cocalodines. The data suggest that the lapsiines and cocalodines are sister groups, with spartaeines more distant (Fig. 14). Our results continue to support the relationship of hisponines with the Salticoida (node 2, Fig. 14; Figs 15–17; Maddison and Needham 2006, Bodner and Maddison 2012).
Figures 16–17.

Phylogeny from Non-salticoid dataset, All Genes analysis. Numbers beside branches show percentage of RAxML likelihood bootstrap replicates with clade. 16 Non-salticoid analysis with all taxa included (1500 bootstrap replicates used) 17 Non-salticoid analysis with , , and “” uplandicus excluded (500 bootstrap replicates used). Colors of branches are the same as those highlighting taxa in Fig. 14.

The placement of remains unclear. As noted under Results, the 28s and 18s genes of may be unreliable phylogenetically, although Maddison et al. 2007 found those genes to place among non-salticoid salticids. In our results likewise has no clear placement, except for being outside the clade of Salticoida + Hisponinae. This result appears in the Non-salticoid and Complete datasets, and with the separate analyses of wingless, CO1, and 16sND1.

Spartaeinae

Our results strongly support the monophyly of the Spartaeinae sensu Su et al. (2007), placing and together with the Spartaeinae in the narrow sense. This is concordant with Wanless’s (1985) hypothesis that and formed a clade with the spartaeines to the exclusion of . The analyses of Su et al. (2007) did not sample , , or , but otherwise their results were largely concordant with ours, which are: (1) (with ) diverge deep, (2) is sister to , (3) , , and are monophyletic, (4) and are sisters, (5) is sister to and . There is strong support for through as a monophyletic group, and for their relationship with . By our data the exact placements of and are unclear. A few spartaeine taxa in our analyses were problematical in appearing unstable, having different placements by different analyses. One of these is PageBreak, for which we have only 16sND1 and CO1 data, both gene regions that appear to evolve too quickly for reliable phylogenetic placement at this level (Bodner and Maddison 2012, Zhang and Maddison 2013). The other is “” uplandicus, whose 28s sequence appears strongly divergent from others. This sequence is from Maddison and Hedin (2003, as “unidentified spartaeine”, vouchers 185 and 186), and it groups “” uplandicus with one species of , against the placements by morphology, CO1 and 16sND1. There is a chance that this gene was mis-sequenced in “” uplandicus. Because of the instability generated, we excluded and “” uplandicus from our analyses giving bootstrap results. Because of the concordance of our phylogenetic results with those of Su et al. (2007), our phylogeny continues to support their conclusions on the stepwise evolution of a complex predatory strategy in spartaeines.

Deep Salticoid relationships

The Salticoida’s basal divergence places the primarily-Neotropical Amycoida as sister group to an unnamed clade (node 3, Fig. 14) that contains most of salticid diversity. This surprising result, first discovered by Maddison and Hedin (2003), had very strong support in the analyses of Bodner and Maddison (2012). We here add support from two new genes, wingless and myosin HC, both of which independently resolve both the Amycoida and its sister group as monophyletic. There have been hints of a clade uniting the Marpissoida, Astioida and baviines (Bodner and Maddison 2012). In our analyses the clade does not receive bootstrap support above 50% in the Complete or Salticoida analyses. The maximum likelihood trees either show the three as monophyletic or not, depending on taxon inclusion and details of the analysis (e.g., Figs 15 and 18). At present we must conclude the relationship between these three and the Saltafresia is unresolved.

Astioida

The astioids as delimited by Maddison et al. (2008) continue to be resolved as a clade, with new support from myosin HC and wingless (Figs 18, 22, 23). Although the body form of resembles that of baviines and the marpissoid , our data clearly place it as an astioid.

Saltafresia

Bodner and Maddison (2012) proposed a clade, the Saltafresia, containing salticoids other than amycoids, astioids, baviines and marpissoids. They found this clade reasonably well supported – 0.78 likelihood bootstrap and 1.0 posterior probability – but no single gene supported it on its own. Our data here continue to support it when all genes are combined. Two genes support it separately, with the exception of single taxa: 28s (but is included) and wingless (but is excluded).

Hasarieae

Previous work had established and as close relatives of (Maddison et al. 2008). We here add several more genera to the group, all Asian. These are , and . The relationships among these genera are not clearly resolved except for a well-supported relationship between and (Figs 14, 19).

Salticinae

The relationship between and the group proposed by Maddison et al. (2008) receives additional support from wingless, along with previously-demonstrated support from 28s and actin. With high posterior probabilities (Bodner and Maddison 2008) and reasonable likelihood bootstrap values (Figs 15, 19), and supported by different genes independently (Figs 20, 22, 24), this relationship can now be considered sufficiently secure that we here formally place the genera of the group into a subfamily — the Salticinae. In addition to genera previously analyzed (, , , , , and ) the subfamily also includes , which is related to (Fig. 14).

Plexippoida + Aelurilloida + Leptorchesteae + Salticinae (Node 5)

A set of four major groups (plexippoids, aelurilloids, leptorchestines and the Salticinae) form a clade in our analyses (node 5, Fig. 14). This group is resolved in the All Genes analyses with high bootstrap values, and it appears, almost, in the independent analyses of each of three genes (18s, wingless, myosin HC). We say “almost” because three of the genes have one or two taxa missing from or added to the group (Fig. 14). While we believe the evidence is good that these form a clade, there is a possibility that the Euophryinae might also fall nested within it. For instance, in the analyses of Bodner and Maddison (2012) the euophryines were placed as sister to the plexippoids. In our analyses the Euophryinae is placed as sister to the Plexippoida + Aelurilloida + Leptorchesteae + Salticinae. This major clade is almost entirely Afro-Eurasian, with the plexippoid being the only exception with more than a handful of species (others are , , , , and , each with fewer than 15 described New World species).

Euophryinae

The 14 euophryine taxa in the analyses are resolved strongly as a monophyletic group. This is a stronger test of monophyly than that of Zhang and Maddison (2013), because PageBreakit includes additional genes and more non-euophryine taxa. The All Genes analyses, along with wingless and myosin HC individually, suggest that the euophryines are the sister group to node 5 (Fig. 14).

Agoriines

Morphologically, the antlike agoriines and are puzzling, with strangely contorted legs and unusual genitalia (Szüts 2003, Logunov and Hereward 2006, Prószyński 2009). While they appear to be salticoids, morphology has given little guidance as to their placement. As noted already, their 28s and 18s genes appear anomalous, and give no clear indication as to their relationships. In the All Genes analysis their placement is ambiguous, though they appear to be salticoids. In an attempt to determine their placement, an additional analysis was done, using a dataset that included and a chimera of cf. lushanensis and cf. palpalis (to have a single taxon with three genes). The aberrant nuclear ribosomal genes of agoriines were excluded from the analysis. The other taxa included were the 70 taxa having at least 4 genes other than CO1 and histone 3. A RAxML likelihood analyses placed and within the sister group of the Amycoida (node 3, Figure 14) with high support (bootstrap percentage 88), but exactly where was highly unstable. Among the 100 likelihood non-bootstrap search replicates were 7 different placements: sister to leptorchestines, sister to baviines, sister to node 5 in Figure 14, sister to the Saltafresia, sister to astioids + marpissoids + baviines, sister to node 3, or sister to node 3 without the baviines. While a relationship with the leptorchestines is appealing, as it would allow their antlike body forms to be homologous, the best we can say at present is that agoriines likely belong within the sister group of amycoids (node 3).

Generic limits

Most of the genera for which we have multiple species – e.g., , , , , – are inferred to be monophyletic in our analyses, corroborating existing concepts based on morphology. The clearest exception is , in which and the similar aff. rufa fall apart from the type species , which had been anticipated as a possibility by Maddison (2009). , , and are reconstructed as paraphyletic, but in each case the bootstrap values are low. The placement of cf. [Sarawak] as sister to , with strong molecular divergence from the other species, would justify establishing a new genus for it.

Behaviour of individual genes

Previous work (Maddison and Hedin 2003, Bodner and Maddison 2012) has suggested that 28s and actin 5C are phylogenetically informative to a reasonable degree for PageBreakdeeper salticid phylogeny, insofar as their results are concordant with summed genes analyses, morphological resemblances, and biogeographical patterns. 16sND1 is useful at the shallower levels (Hedin and Maddison 2001) but has difficulties recovering deeper relationships, while CO1 struggles through both shallow and deep levels (Maddison and Hedin 2003, Bodner and Maddison 2012). One surprise in our analyses was the informative behaviour of CO1 in deeper relationships among the non-salticoid salticids. Although CO1 is almost nonsensical in its inferred relationships within the Salticoida, it succeeds in recovering the Spartaeinae, the Spartaeineae sensu Wanless, the lapsiines, and the Salticoida as monophyletic. Two new genes added, wingless, myosin HC, both show clear concordance with the 28s and previous all genes analyses. supports many of the previously recognized clades, including the Salticoida, Amycoida, the sister clade to Amycoida, Plexippoida, Marpissoida (in part), Astioida (in part), Spartaeinae sensu Wanless, and lapsiines. We find it encouraging that a haphazardly chosen protein-coding gene, independent from 28s, supports previous molecular results in Salticidae. There are still, however, many aspects of salticid relationships yet to be resolved, such as the deepest relationships in the family, including the relationships among the three subgroups of lyssomanines, the placement of and the agoriines, and the relationships among astioids, marpissoids, baviines and the Saltafresia. With the coming era of genomic data, we expect large quantities of new data will be available for exploring these relationships.
  15 in total

1.  A combined molecular approach to phylogeny of the jumping spider subfamily dendryphantinae (araneae: salticidae).

Authors:  M C Hedin; W P Maddison
Journal:  Mol Phylogenet Evol       Date:  2001-03       Impact factor: 4.286

2.  MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform.

Authors:  Kazutaka Katoh; Kazuharu Misawa; Kei-ichi Kuma; Takashi Miyata
Journal:  Nucleic Acids Res       Date:  2002-07-15       Impact factor: 16.971

3.  RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models.

Authors:  Alexandros Stamatakis
Journal:  Bioinformatics       Date:  2006-08-23       Impact factor: 6.937

4.  Molecular phylogeny, divergence times and biogeography of spiders of the subfamily Euophryinae (Araneae: Salticidae).

Authors:  Jun-Xia Zhang; Wayne P Maddison
Journal:  Mol Phylogenet Evol       Date:  2013-03-28       Impact factor: 4.286

5.  The biogeography and age of salticid spider radiations (Araneae: Salticidae).

Authors:  Melissa R Bodner; Wayne P Maddison
Journal:  Mol Phylogenet Evol       Date:  2012-06-23       Impact factor: 4.286

6.  Tisaniba, a new genus of marpissoid jumping spiders from Borneo (Araneae: Salticidae).

Authors:  Jun-Xia Zhang; Wayne P Maddison
Journal:  Zootaxa       Date:  2014-08-14       Impact factor: 1.091

7.  Ribosomal RNA trees misleading?

Authors:  M Hasegawa; T Hashimoto
Journal:  Nature       Date:  1993-01-07       Impact factor: 49.962

8.  Jerzego, a new hisponine jumping spider from Borneo (Araneae: Salticidae).

Authors:  Wayne P Maddison; Edyta K Piascik
Journal:  Zootaxa       Date:  2014-08-19       Impact factor: 1.091

9.  MAFFT version 5: improvement in accuracy of multiple sequence alignment.

Authors:  Kazutaka Katoh; Kei-ichi Kuma; Hiroyuki Toh; Takashi Miyata
Journal:  Nucleic Acids Res       Date:  2005-01-20       Impact factor: 16.971

10.  Structure of the retinae of the principal eyes of jumping spiders (Salticidae: dendryphantinae) in relation to visual optics.

Authors:  M F Land
Journal:  J Exp Biol       Date:  1969-11       Impact factor: 3.312

View more
  10 in total

1.  A phylogenetic and taxonomic review of baviine jumping spiders (Araneae, Salticidae, Baviini).

Authors:  Wayne P Maddison; Imara Beattie; Kiran Marathe; Paul Y C Ng; Nilani Kanesharatnam; Suresh P Benjamin; Krushnamegh Kunte
Journal:  Zookeys       Date:  2020-12-16       Impact factor: 1.546

2.  The execution of planned detours by spider-eating predators.

Authors:  Fiona R Cross; Robert R Jackson
Journal:  J Exp Anal Behav       Date:  2016-01       Impact factor: 2.468

3.  A genome-wide phylogeny of jumping spiders (Araneae, Salticidae), using anchored hybrid enrichment.

Authors:  Wayne P Maddison; Samuel C Evans; Chris A Hamilton; Jason E Bond; Alan R Lemmon; Emily Moriarty Lemmon
Journal:  Zookeys       Date:  2017-09-04       Impact factor: 1.546

4.  Multilocus genetic and morphological phylogenetic analysis reveals a radiation of shiny South Asian jumping spiders (Araneae, Salticidae).

Authors:  Nilani Kanesharatnam; Suresh P Benjamin
Journal:  Zookeys       Date:  2019-01-16       Impact factor: 1.546

5.  A new lapsiine jumping spider from North America, with a review of Simon's Lapsias species (Araneae, Salticidae, Spartaeinae).

Authors:  Wayne P Maddison
Journal:  Zookeys       Date:  2019-11-21       Impact factor: 1.546

6.  Sitticine jumping spiders: phylogeny, classification, and chromosomes (Araneae, Salticidae, Sitticini).

Authors:  Wayne P Maddison; David R Maddison; Shahan Derkarabetian; Marshal Hedin
Journal:  Zookeys       Date:  2020-04-08       Impact factor: 1.546

7.  First description of the female of the jumping spider Balmaceda nigrosecta Mello-Leitão (Salticidae, Dendryphantini, Marpissina).

Authors:  Gonzalo D Rubio; Julián E Baigorria; G B Edwards
Journal:  Zookeys       Date:  2016-02-15       Impact factor: 1.546

8.  Phylogenetic placement of the unusual jumping spider Depreissia Lessert, and a new synapomorphy uniting Hisponinae and Salticinae (Araneae, Salticidae).

Authors:  Wayne P Maddison; David R Maddison; Junxia Zhang; Tamás Szűts
Journal:  Zookeys       Date:  2016-01-05       Impact factor: 1.546

9.  Miocene spider Maevia eureka nov. sp. (Araneae: Salticidae).

Authors:  Francisco Riquelme; Miguel Menéndez-Acuña
Journal:  PeerJ       Date:  2017-07-25       Impact factor: 2.984

10.  Sumakuru, a deeply-diverging new genus of lyssomanine jumping spiders from Ecuador (Araneae: Salticidae).

Authors:  Wayne P Maddison
Journal:  Zookeys       Date:  2016-09-01       Impact factor: 1.546

  10 in total

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