Citation: Maddison WP, Li D, Bodner M, Zhang J, Xu X, Liu Q, Liu F (2014) The deep phylogeny of jumping spiders (Araneae, Salticidae). ZooKeys 440: 57–87. doi: 10.3897/zookeys.440.7891
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.
Jumping spiders, Salticidae, phylogeny, systematics
Salticid spiders, remarkable for their excellent vision (
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.
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 (
Specimens and sequences used in phylogenetic analyses, with GenBank numbers indicated. * marks previously published sequences. Specimen localities given in Suppl. material 1.
Reference | 28s | 18s | wingless | myosin HC | actin 5c | histone 3 | CO1 | 16sND1 | |
---|---|---|---|---|---|---|---|---|---|
Outgroups | |||||||||
Anyphaenidae: Hibana sp. | s318 | AY297295* | KM033091 | KM032961 | KM032929 | AY297422* | AY297295 / AY297358* | ||
Gnaphosidae: Cesonia sp. | s319 | AY297293; EF201663* | KM032996 | EU522700* | DQ665720* | AY297420* | AY296711 / AY297356* | ||
Miturgidae: Cheiracanthium sp. | s321 | AY297294; EF201664* | KM032997 | KM032928 | AY297421* | AY296712 / AY297357* | |||
Oxyopidae: Oxyopes birmanicus Thorell, 1887 | EF419032 / EF419065* | EF418998* | EF419126* | EF419097* | EF418969 / EF419150* | ||||
Philodromidae: Philodromus alascensis Keyserling, 1884 | GR011 | KM033130 | KM033092 | KM032998 | KM032962 | ||||
Thomisidae: Misumenops nepenthicola (Pocock, 1898) | EF419029 / EF419062* | EF418996* | EF419123* | EF419094* | EF418967 / EF419148* | ||||
Thomisidae: Xysticus sp. | s316 | AY297296; EF201665* | KM033093 | EU522701* | DQ665704* | AY297296* | AY296714 / AY297359* | ||
Lyssomanines | |||||||||
Asemonea sichuanensis Song & Chai, 1992 | SC-03-0055 | EF418986* | EF419082* | ||||||
Asemonea sichuanensis Song & Chai, 1992 | MRB084 | KM033131 | KM032931 | ||||||
Asemonea cf. stella Wanless, 1980 | MRB083 | JX145767* | KM033094 | KM032930 | JX145686* | ||||
Asemonea tenuipes (O. P.-Cambridge, 1869) | d186 | KM033132 | KM033095 | KM032999 | KM032963 | KM032932 | |||
Chinoscopus cf. flavus (Peckham, Peckham & Wheeler, 1889) | d273 | KM033133 | KM033096 | KM032888 | |||||
Goleba lyra Maddison & Zhang, 2006 | d051 | DQ665768* | KM033097 | KM033000 | EU522709* | DQ665707* | DQ665755* | ||
Lyssomanes amazonicus Peckham & Wheeler, 1889 | ECU11-6112 | KM033134 | KM032889 | ||||||
Lyssomanes antillanus Peckham & Wheeler, 1889 | d298 | KM033135 | KM033001 | ||||||
Lyssomanes cf. benderi Logunov, 2002 | ECU11-5402 | KM033136 | KM032890 | ||||||
Lyssomanes cf. jemineus Peckham & Wheeler, 1889 | ECU11-5682 | KM033137 | KM032891 | ||||||
Lyssomanes longipes (Taczanowski, 1871) | MRB086 | KM033138 | KM032933 | KM033208 | KM032892 | ||||
Lyssomanes pauper Mello-Leitão, 1945 | d297 | KM033139 | KM033002 | ||||||
Lyssomanes taczanowskii Galiano, 1980 | ECU11-4193 | KM033141 | KM032894 | ||||||
Lyssomanes tenuis Peckham & Wheeler, 1889 | ECU11-4869 | KM033142 | KM032895 | ||||||
Lyssomanes viridis (Walckenaer, 1837) | s160 | AY297231* | AY297360* | AY296652 / AY297297* | |||||
Lyssomanes viridis (Walckenaer, 1837) | d129 | KM033098 | KM033003 | EU522715* | DQ665715* | ||||
Lyssomanes sp. [Esmeraldas] | d408 | KM033140 | KM032893 | ||||||
Onomastus nigrimaculatus Zhang & Li, 2005 | EF419031 / EF419064* | EF418997* | EF419125* | EF419096* | EF418968 / EF419149* | ||||
Onomastus sp. [Guangxi] | MRB085 | JX145768* | KM033099 | KM033004 | KM032964 | KM032934 | JX145687* | JX145910* | |
Pandisus cf. decorus Wanless, 1980 | d303 | KM033143 | KM033005 | ||||||
Cocalodines | |||||||||
Allococalodes madidus Maddison, 2009 | d236 | KM033144 | KM033006 | KM032896 | |||||
Cocalodes longicornis Wanless, 1982 | d291 | KM033145 | KM033007 | KM032935 | KM032897 | ||||
Cocalodes macellus (Thorell, 1878) | d230 | KM033146 | KM033100 | KM033008 | KM032936 | KM033209 | |||
Cucudeta gahavisuka Maddison, 2009 | d234 | KM033147 | KM033009 | KM032898 | |||||
Cucudeta zabkai Maddison, 2009 | d235 | KM033148 | KM033010 | KM032965 | KM032899 | ||||
Tabuina aff. baiteta Maddison, 2009 | d313 | KM033149 | KM033011 | ||||||
Tabuina rufa Maddison, 2009 | d232 | KM033151 | KM033013 | KM032900 | |||||
Tabuina aff. rufa Maddison, 2009 | d312 | KM033150 | KM033012 | ||||||
Tabuina varirata Maddison, 2009 | d233 | KM033152 | KM033014 | KM032901 | |||||
Yamangalea frewana Maddison, 2009 | d231 | KM033153 | KM033015 | KM032902 | |||||
Spartaeines | |||||||||
Brettus cf. adonis Simon, 1900 | SWK12-4323 | KM033154 | |||||||
Brettus sp. [Yunnan] | LiD-026-053-05 | KM033155S | KM033101S | KM033195S | |||||
cf. Phaeacius sp. [Sarawak] | SWK12-3728 | KM033156 | |||||||
Cocalus murinus Simon, 1899 | LiD-013-027-05 | EF419019 / EF419053* | EF418988* | EF419116* | EF419084* | EF418959 / EF419140* | |||
Cyrba algerina (Lucas, 1846) | EF419021 / EF419054* | EF418989* | EF419086* | EF418961 / EF419142* | |||||
Cyrba lineata Wanless, 1984 | MRB106 | JX145792* | KM033016 | KM032966 | KM032937 | JX145704* | |||
Cyrba ocellata (Kroneberg, 1875) | EF418990* | EF419087* | EF418962 / EF419143* | ||||||
Cyrba ocellata (Kroneberg, 1875) | MRB104 | KM033157 | |||||||
Cyrba sp. [Kenya] | EF419023 / EF419056* | EF418991* | EF419088* | ||||||
Gelotia cf. bimaculata Thorell, 1890 | d250 | KM033158 | KM033017 | KM032938 | |||||
Gelotia syringopalpis Wanless, 1984 | EF419024 / EF419057* | EF419118* | |||||||
Gelotia syringopalpis Wanless, 1984 | MRB105 | KM033019 | KM033212 | KM032903 | |||||
Gelotia sp. [Guangxi] | MRB199 | KM033018 | KM032939 | KM033210 | |||||
Gelotia sp. [Yunnan] | LiD002-053-05 | KM033102S | KM033196S | KM033211S | |||||
Holcolaetis vellerea Simon, 1910 | EF419025 / EF419058* | EF418992* | EF419119* | EF419090* | EF418963 / EF419144* | ||||
Holcolaetis cf. zuluensis Lawrence, 1937 | d036 | DQ665770* | KM033103 | EU522711* | DQ665721* | DQ665757* | |||
Meleon aff. kenti (Lessert, 1925) | d287 | KM033159 | KM032940 | ||||||
Mintonia mackiei Wanless, 1984 | SWK12-4202 | KM033161 | |||||||
Mintonia cf. melinauensis Wanless, 1984 | d441 | KM033160 | |||||||
Mintonia ramipalpis (Thorell, 1890) | SWK12-1442 | KM033162 | |||||||
Mintonia silvicola Wanless, 1987 | d104 | KM033020 | KM032904 | ||||||
Mintonia silvicola Wanless, 1987 | SWK12-1653 | KM033163 | |||||||
Mintonia silvicola Wanless, 1987 | EF418995* | EF419122* | EF419093* | ||||||
Mintonia tauricornis Wanless, 1984 | d249 | KM033164 | KM033021 | KM032941 | KM032905 | ||||
Neobrettus tibialis (Prószyński, 1978) | LiD-001-055-05 | EF419030 / EF419063* | EF419124* | EF419095* | |||||
Neobrettus sp. [Sarawak] | SWK12-1040 | KM033165 | |||||||
Paracyrba wanlessi Zabka & Kovac, 1996 | EF419033 / EF419066* | EF418999* | EF419098* | ||||||
Phaeacius lancearius (Thorell, 1895) | d111 | DQ665775* | KM033022 | DQ665759* | |||||
Phaeacius malayensis Wanless, 1981 | EF419034 / EF419067* | EF419000* | EF419099* | EF418970 / EF419151* | |||||
Phaeacius sp. [Guangxi] | LQ-24-06 | KM033166S | KM033104S | KM033213S | KM032906S | ||||
Phaeacius sp. [Hainan] | EF419035 / EF419068* | EF419001* | EF418971 / EF419152* | ||||||
Phaeacius sp. [Sarawak] | SWK12-4541 | KM033167 | |||||||
Portia africana (Simon, 1886) | EF419037 / EF419069* | EF419003* | EF419128* | EF419101* | |||||
Portia crassipalpis (Peckham & Peckham, 1907) | SWK12-2354 | KM033168 | |||||||
Portia fimbriata (Doleschall, 1859) | LiD-001-04 | EF419038 / EF419070* | EF419004* | EF419129* | EF419102* | EF418973 / EF419154* | |||
Portia heteroidea Xie & Yin, 1991 | EF419039 / EF419071* | EF419005* | EF419130* | EF419103* | EF418974 / EF419155* | ||||
Portia jianfeng Song & Zhu, 1998 | EF419040 / EF419072* | EF419006* | EF419104* | EF418975 / EF419156* | |||||
Portia labiata (Thorell, 1887) | S206 | AY297232* | AY297361* | AY296653 / AY297298* | |||||
Portia cf. schultzi Karsch, 1878 | d131 | DQ665776* | KM033105 | KM033023 | KM032967 | EU522718* | DQ665708* | ||
Portia quei Zabka, 1985 | EF419042 / EF419074* | EF419008* | EF419132* | EF419106* | EF418977 / EF419158* | ||||
Portia taiwanica Zhang & Li, 2005 | MRB103 | KM033169 | KM032942 | KM033214 | KM032907 | ||||
Portia sp. [Sichuan] | SC-03-0011 | EF419043 / EF419075* | EF419009* | EF419133* | EF418978 / EF419159* | ||||
Sonoita lightfooti Peckham & Peckham, 1903 | d226 | KM033170 | KM033215 | ||||||
Sonoita aff. lightfooti Peckham & Peckham, 1903 | MRB200 | JX145791* | JX145705* | JX145927* | |||||
Sparbambus gombakensis Zhang, Woon & Li, 2006 | d251 | KM033171 | KM033024 | KM032943 | |||||
Spartaeus jianfengensis Song & Chai, 1991 | EF419045 / EF419076* | EF419011* | EF419109* | EF418980 / EF419161* | |||||
Spartaeus platnicki Song, Chen & Gong, 1991 | SC-03-069 | EF419046 / EF419077* | EF419012* | EF419135* | EF419110* | EF418981 / EF419162* | |||
Spartaeus spinimanus (Thorell, 1878) | S199 | KM033216 | KM032908 | ||||||
Spartaeus thailandicus Wanless, 1984 | BV-004 | EF419047 / EF419078* | EF419013* | EF419136* | EF419111* | EF418982 / EF419163* | |||
Spartaeus uplandicus Barrion & Litsinger, 1995 | S185/S186 | AY297233* | AY297363* | AY296655* | |||||
Spartaeus wildtrackii Wanless, 1987 | EF419048 / EF419079* | EF419014* | EF419137* | EF419112* | EF418983 / EF419164* | ||||
Taraxella sp. [Johor] | d246 | KM033172 | KM032944 | KM032909 | |||||
Taraxella sp. [Pahang] | d248 | KM033173 | KM032945 | KM033197 | |||||
Taraxella sp. [Pahang] | LiD-001-003-06 | KM033106S | KM033217S | KM032910S | |||||
Yaginumanis wanlessi Zhang & Li, 2005 | EF419050 / EF419081* | EF419016* | EF419139* | EF419114* | EF418985 / EF419166* | ||||
Lapsiines | |||||||||
Galianora bryicola Maddison, 2006 | d124 | DQ665771* | DQ665741* | KM033025 | EU522706* | DQ665717* | DQ665758* | DQ665727* | |
Galianora sacha Maddison, 2006 | d116 | DQ665766* | DQ665734* | KM033026 | KM032968 | EU522707* | DQ665716* | DQ665754* | |
Lapsias canandea Maddison, 2012 | d442 | KM033174 | |||||||
Lapsias guamani Maddison, 2012 | UBC-SEM AR00191 | KM033175 | KM033027 | ||||||
Lapsias lorax Maddison, 2012 | UBC-SEM AR00194 | KM033176 | KM033028 | ||||||
Soesiladeepakius lyra Ruiz & Maddison, 2012 | GR130 | JQ312077 | KM033029 | JQ312074* | JQ312079* | ||||
Thrandina bellavista Maddison, 2012 | d396 | KM033177 | KM033030 | ||||||
Thrandina cosanga Maddison, 2012 | d395 | KM033178 | |||||||
Thrandina parocula Maddison, 2006 | d123 | DQ665779* | KM033107 | EU522720* | DQ665718* | DQ665761* | DQ665726* | ||
Thrandina parocula Maddison, 2006 | d394 | KM033031 | KM032969 | ||||||
Eupoa | |||||||||
Eupoa nezha Maddison & Zhang, 2007 | d220/MRB102 | EF201648* | EF201666* | KM033032 | EF201668* | EF201667* | |||
Hisponines | |||||||||
cf. Tomocyrba sp. [Madagascar] | d305 | KM032881* | |||||||
Hispo macfarlanei Wanless, 1981 | d404 | KM032882* | KM032970 | ||||||
Hispo sp. [Madagascar] | d309 | KM032883* | |||||||
Jerzego cf. alboguttatus Simon, 1903 | SWK12-4787 | KM032884* | |||||||
Jerzego corticicola Maddison, 2014 | SWK12-2900 | KM032885* | KM032887* | ||||||
Massagris contortuplicata Wesolowska & Haddad, 2013 | d082 | DQ665772* | KM033108 | KM033033 | DQ665705* | DQ665722* | |||
Massagris schisma Maddison & Zhang, 2006 | d081 | DQ665762* | KM033109 | KM033034 | DQ665728* | ||||
Tomobella andasibe (Maddison & Zhang, 2006) | d127 | DQ665780* | DQ665752* | KM033035 | KM033198 | DQ665725* | |||
Tomocyrba sp. [Madagascar] | d306 | KM032886* | |||||||
Tomomingi sp. [Gabon] | MRB243 | JX145764* | KM033110 | KM033036 | KM032971 | JX145850* | JX145684* | ||
Salticoida | |||||||||
Agoriines | |||||||||
Agorius constrictus Simon, 1901 | d172 | KM032953 | |||||||
Agorius constrictus Simon, 1901 | d213 | KM033119 | KM033072 | KM032921 | |||||
Agorius sp. [Selangor] | d299 | KM033189 | KM033073 | ||||||
Synagelides cf. lushanensis Xie & Yin, 1990 | d214 | KM033074 | |||||||
Synagelides cf. palpalis Zabka, 1985 | MRB050 | KM032922 | |||||||
Synagelides cf. palpalis Zabka, 1985 | d225 | KM033190 | KM033226 | ||||||
Amycoids | |||||||||
Cotinusa sp. [Ecuador] | MRB024 | JX145746* | KM033120 | KM033075 | KM032987 | JX145832* | JX145671* | JX145896* | |
Hurius vulpinus Simon, 1901 | S213 | AY297239* | AY297368* | AY296662 / AY297306* | |||||
Hurius cf. vulpinus Simon, 1901 | d156 | KM033076 | EU522712* | KM033203 | |||||
Hypaeus aff. miles Simon, 1900 [Ecuador] | d130 | EU815499* | KM033121 | KM033077 | KM032988 | EU522702* | KM032923 | ||
Sarinda cutleri (Richman, 1965) | MRB193 | JX145744* | KM033078 | KM032954 | JX145669* | JX145895* | |||
Sitticus floricola palustris (Peckham & Peckham, 1883) | d030 | DQ665778* | KM033122 | KM033079 | KM032989 | KM033204 | DQ665760* | DQ665729* | |
Astioids | |||||||||
Arasia mollicoma (L. Koch, 1880) | d046 | EU815483* | EU815532* | KM032990 | JX145834* | KM033205 | EU815598* | EU815550* | |
Helpis minitabunda (L. Koch, 1880) | d265 | KM033123 | KM033080 | KM032991 | KM032955 | KM033227 | |||
Ligurra latidens (Doleschall, 1859) | d175 | JX145749* | KM033081 | JX145835* | JX145898* | ||||
Ligurra latidens (Doleschall, 1859) | LiD-001-027-05 | EF418993* | EF419120* | EF419091* | |||||
Mopsus mormon Karsch, 1878 | d018 | EU815470* | EU815529* | KM033082 | JX145836* | KM033206 | EU815586* | ||
Myrmarachne sp. [Pahang] | d162 | EU815507* | KM033124 | KM033083 | KM032992 | JX145837* | EU815616* | EU815565* | |
Neon reticulatus (Blackwall, 1853) | d283 | KM033191 | KM033125 | KM033084 | KM032993 | KM032956 | |||
Nungia epigynalis Zabka, 1985 | d221 | KM033192 | KM032924 | ||||||
Simaetha sp. | d027 | EU815477* | KM033126 | KM033085 | JX145839* | EU815592* | EU815546* | ||
Trite pennata Simon, 1885 | d035 | EU815478* | KM033086 | KM032957 | KM033207 | EU815593* | EU815547* | ||
Baviines | |||||||||
Bavia aff. aericeps Simon, 1877 [Sabah] | d079 | EU815490* | KM033127 | KM032958 | EU815603* | KM032925 | |||
Stagetilus sp. [Selangor] | MRB079 | KM033193 | KM033087 | KM032959 | KM032926 | ||||
Marpissoids | |||||||||
Afromarengo sp. [Gabon] | MRB262 | JX145758* | KM033128 | KM033088 | KM032994 | JX145842* | JX145682* | JX145905* | |
Dendryphantes hastatus (Clerck, 1757) | d043 | EF201646* | KM033129 | KM033089 | KM033228 | KM032927 | |||
Platycryptus californicus (Peckham & Peckham, 1888) | d316 | KM033194 | KM033090 | KM032995 | KM032960 | KM033229 | |||
Rhene sp. [Pahang] | LiD-001-021-05 | EF419044* | EF419010* | EF419134* | EF419108* | EF418979 / EF419160* | |||
Tisaniba mulu Zhang & Maddison, 2014 | SWK12-1244 | KM032876* | KM032880* | ||||||
Saltafresians | |||||||||
Aelurillus cf. ater (Kroneberg, 1875) | d140 | EU815504* | EU815536* | KM033037 | KM032972 | JX145831* | KM033199 | EU815615* | EU815564* |
Amphidraus complexus Zhang & Maddison, 2012 | JXZ035 | KC615380* | KM033038 | KC616069* | KC615640* | KC615806* | |||
Athamas cf. whitmeei O. P.-Cambridge, 1877 | JXZ345 | KC616286* | KC615649* | KC615822* | |||||
Bacelarella pavida Szüts & Jocqué, 2001 | d195 | EU815511* | EU815538* | KM033039 | KM032973 | KM032946 | EU815618* | EU815569* | |
Bathippus macrognathus (Thorell, 1881) | JXZ372 | KC615407* | KM033040 | KC616305* | KC615835* | ||||
Bianor maculatus (Keyserling, 1883) | d017 | EU815469* | KM033041 | KM033200 | EU815585* | EU815542* | |||
Bristowia afra Szüts, 2004 | JXZ363 | KC615409* | KC616301* | ||||||
Bristowia afra Szüts, 2004 | MRB230 | KM033042 | KM033218 | ||||||
Cheliceroides longipalpis Zabka, 1985 | d222 | KM033111 | KM033043 | JX145830* | KM033219 | EU815579* | |||
Cheliceroides cf. longipalpis Zabka, 1985 | d415 | KM033179 | |||||||
Chinattus parvulus (Banks, 1895) | d009 | EU815464* | EU815525* | KM033044 | JX145848* | KM033201 | EU815581* | ||
Chinophrys pengi Zhang & Maddison, 2012 | JXZ145 | KC615416* | KM033045 | KC616146* | KC615843* | ||||
Corythalia locuples (Simon, 1888) | JXZ315 | KC615390* | KM033046 | KC616260* | KC615645* | KC615816* | |||
Cosmophasis umbratica Simon, 1903 | EF419020* | EF419117* | EF419085* | EF418960 / EF419141* | |||||
Cytaea nimbata (Thorell, 1881) | JXZ229 | KC615474* | KM033047 | KC616197* | KC615693* | KC615899* | |||
Diolenius varicus Gardzińska & Zabka, 2006 | JXZ349 | KC615480* | KM033048 | KC616290* | KC615695* | KC615905* | |||
Diplocanthopoda marina Abraham, 1925 | d209 | KM033180 | KM032947 | KM033220 | KM032911 | ||||
Eburneana sp. [Gabon] | MRB231 | KM033181 | KM033049 | JX145858* | KM033221 | KM032912 | |||
Echeclus sp. [Selangor] | MRB089 | KM033182 | KM032948 | KM033222 | KM032913 | ||||
Euophrys frontalis (Walckenaer, 1802) | JXZ137 | KC615536* | KM033050 | KC616139* | KC615960* | ||||
Evarcha proszynskii Marusik & Logunov, 1998 | d096 | DQ665765* | KM033112 | EU522704* | DQ665723* | ||||
Evarcha proszynskii Marusik & Logunov, 1998 | d323 | KM033051 | KM032974 | ||||||
Freya decorata (C. L. Koch, 1846) | d211 | EU815521* | EU815539* | KM032975 | EU522705* | JX145908* | |||
Gedea cf. tibialis Zabka, 1985 | MRB090 | KM033183 | KM032949 | KM033223 | KM032914 | ||||
Habrocestum cf. albimanum Simon, 1901 | d132 | EU815500* | EU815611* | EU815562* | |||||
Habronattus borealis (Banks, 1895) | d207 | KM033184 | KM033052 | KM032976 | KM032950 | KM033224 | KM032915 | ||
Hasarius adansoni (Audouin, 1826) | d295 | KM033113 | KM033053 | KM032977 | |||||
Hasarius adansoni (Audouin, 1826) | S130/S131/S324 | AY297281* | AY297409* | ||||||
Heliophanus cupreus (Walckenaer, 1802) | d044 | DQ665769* | KM033114 | EU522710* | DQ665710* | DQ665756* | KM032916 | ||
Idastrandia cf. orientalis (Szombathy, 1915) | d108 | EU815535; EU815496* | EU815535* | JX145852* | EU815608* | EU815560* | |||
Langerra aff. longicymbium Song & Chai, 1991 | d182 | KM033185 | KM033054 | KM032917 | |||||
Leptorchestes berolinensis (C. L. Koch, 1846) | d086 | EU815491* | EU815534* | KM033055 | EU815604* | EU815556* | |||
Longarenus brachycephalus Simon, 1903 | MRB258 | JX145798* | KM033056 | KM032978 | KM032951 | JX145707* | KM032918 | ||
Nannenus sp. [Pahang] | d105 | EU815493* | KM033057 | KM032979 | JX145853* | EU815558* | |||
Naphrys pulex (Hentz, 1846) | JXZ081 | JX145760* | KM033115 | KM032980 | JX145844* | KC615749* | JX145907* | ||
Omoedus orbiculatus (Keyserling, 1881) | d008 | KC615792* | |||||||
Omoedus orbiculatus (Keyserling, 1881) | JXZ136 | JX145762* | KM033116 | KM033058 | JX145846* | KM033202 | |||
Omoedus papuanus Zhang & Maddison, 2012 | JXZ286 | KC615619* | KM033059 | KC616234* | KC615790* | KC616042* | |||
Pellenes peninsularis Emerton, 1925 | d057 | DQ665774* | KM033117 | KM033060 | JX145864* | DQ665712* | |||
Pellenes peninsularis Emerton, 1925 | d400 | KM032981 | |||||||
Phaulostylus grammicus Simon, 1902 | d304 | KM033186 | KM033061 | ||||||
Philaeus chrysops (Poda, 1761) | d025 | EU815475* | EU815530* | KM033062 | JX145855* | EU815590* | EU815545* | ||
Phintella sp. [Gabon] | d402 | KM033187 | KM033063 | KM032982 | |||||
Plexippus paykulli (Audouin, 1826) | LiD-001-029-05 | EF419002* | EF419127* | ||||||
Plexippus paykulli (Audouin, 1826) | MRB016 | JX145784* | KM033064 | EU522713* | |||||
Plexippus paykulli (Audouin, 1826) | S73 | AY297384* | AY296674 / AY297317* | ||||||
Pochyta cf. pannosa Simon, 1903 | MRB257 | JX145806* | KM033065 | KM032983 | KM032952 | JX145715* | KM032919 | ||
Saitis barbipes (Simon, 1868) | JXZ147 | KC615589* | KM033066 | KC616147* | KC615767* | KC616011* | |||
Salticus scenicus (Clerck, 1757) | d003 | DQ665777* | KM033118 | KM033067 | KM032984 | EU522719* | DQ665713* | JX145663* | AY296707 / AY297352* |
Thiania bhamoensis Thorell, 1887 | LiD-001-028-05 | EF419049 / EF419080* | EF419015* | EF419138* | EF419113* | EF418984 / EF419165* | |||
Trydarssus cf. nobilitatus (Nicolet, 1849) | MRB270 | KM033188 | KM033068 | KM032985 | JX145847* | KM033225 | KM032920 | ||
Tusitala lyrata (Simon, 1903) | MRB226 | JX145771* | KM033069 | JX145856* | JX145689* | JX145912* | |||
Yllenus arenarius Menge, 1868 | d013 | EU815527* | EU815583* | EU815541* | |||||
Yllenus arenarius Menge, 1868 | JXZ173 | JX145766* | KM033070 | KM032986 | JX145851* | ||||
Zabkattus furcatus Zhang & Maddison, 2012 | JXZ218 | KC615503* | KM033071 | KC616190* | KC615928* |
Our sample targeted especially the non-salticoid salticids, those that lie outside the major clade of familiar salticids (
Some previously-published data from non-salticoid salticids was either excluded or represented under a different species name here. Excluded are sequences of Hispo cf. frenata, because its limited data made it unstable in the analyses (see
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. Phaeacius [Sarawak]” (Figs 1, 2) is known from a single female and juvenile from Lambir Hills, Sarawak. It resembles Phaeacius but the legs are shorter, and the epigynum is distinctively different. Phaeacius sp. [Sarawak] (Figs 3, 4) is a fairly typical Phaeacius whose epigynum resembles that of Phaeacius leytensis Wijesinghe, 1991, but with the atria elongated posteriorly. Onomastus sp. [Guangxi] is shown in Fig. 5. Sonoita aff. lightfooti (Fig. 6) has longer grooves for the openings of the epigynum than Sonoita lightfooti, and is distinctive in gene sequences as well. Gelotia sp. [Guangxi] (Fig. 7) has a palp resembling Gelotia syringopalpis, but the tibial apophyses are much shorter. Echeclus sp. [Selangor] (Figs 8, 9) was identified as an Echeclus 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 (
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. Phaeacius [Sarawak], voucher SWK12–3728 3–4 Female Phaeacius sp. [Sarawak], voucher SWK12–4541 5 female Onomastus sp. [Guangxi], voucher MRB085 6 Female Sonoita aff. lightfooti, voucher MRB200. 7 male Gelotia sp. [Guangxi], voucher MRB199. The drawing is reversed so as to appear to be the left palpus 8–9 Male Echeclus sp. [Selangor], voucher MRB089 10–11 Male Taraxella 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 Taraxella 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.
Eight genes were used for this analysis. Two are nuclear ribosomal genes, 28s and 18s (
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
For most wingless sequences, the forward and reverse primers used were respectively Spwgf1 and Spwgr1 (
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 Cyrtophora citricola (Genbank accession AAM97635.1;
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 (Gelotia sp. [Guangxi]) and MRB231 (Eburneana sp. [Gabon]). Given that CO1 had little resolution, these are unlikely to have affected the results.
Automatic multiple sequence alignment was performed by MAFFT (
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 first 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 analyses using maximum likelihood were run using RAxML version 7.2.8alpha (
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 (
The data underpinning the analyses reported in this paper are deposited in the Dryad Data Repository at doi: 10.5061/dryad.v53h1.
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-salticoid and Salticoida datasets. Figures 20–27 show the results for individual genes analyzed separately.
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 Eupoa, agoriines, Spartaeus spinimanus and “Spartaeus” 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 Eupoa 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 Eupoa 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 Eupoa, Spartaeus spinimanus, and “Spartaeus” 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 Agorius and Synagelides 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. Eupoa was not only difficult to sequence (
Many of the salticid clades now recognized by molecular data had been previously recognized by morphological data. For instance,
However, the first molecular data for salticid phylogeny as a whole (
Our results help resolve or add strength to relationships at the deepest level of salticid phylogeny.
Spartaeines, lapsiines and cocalodines form a clade (node 1, Fig. 14). Although
Within this spartaeine-lapsiine-cocalodine clade, the subclade historically best known by morphology is
Our results continue to support the relationship of hisponines with the Salticoida (node 2, Fig. 14; Figs 15–17;
The placement of Eupoa remains unclear. As noted under Results, the 28s and 18s genes of Eupoa may be unreliable phylogenetically, although
Our results strongly support the monophyly of the Spartaeinae sensu
A few spartaeine taxa in our analyses were problematical in appearing unstable, having different placements by different analyses. One of these is Spartaeus spinimanus, 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 (
Because of the concordance of our phylogenetic results with those of
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
There have been hints of a clade uniting the Marpissoida, Astioida and baviines (
The astioids as delimited by
Previous work had established Habrocestum and Chinattus as close relatives of Hasarius (
The relationship between Salticus and the Philaeus group proposed by
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
This major clade is almost entirely Afro-Eurasian, with the plexippoid Habronattus being the only exception with more than a handful of species (others are Pellenes, Sibianor, Evarcha, Phlegra, Paramarpissa and Salticus, each with fewer than 15 described New World species).
The 14 euophryine taxa in the analyses are resolved strongly as a monophyletic group. This is a stronger test of monophyly than that of
Morphologically, the antlike agoriines Agorius and Synagelides are puzzling, with strangely contorted legs and unusual genitalia (
Most of the genera for which we have multiple species – e.g., Asemonea, Portia, Mintonia, Phaeacius, Cyrba – are inferred to be monophyletic in our analyses, corroborating existing concepts based on morphology. The clearest exception is Tabuina, in which Tabuina rufa and the similar Tabuina aff. rufa fall apart from the type species Tabuina varirata, which had been anticipated as a possibility by
The placement of cf. Phaeacius [Sarawak] as sister to Phaeacius, with strong molecular divergence from the other species, would justify establishing a new genus for it.
Previous work (
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. Wingless 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 Eupoa 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.
We thank David Maddison for obtaining the sequences of wingless and 28s in Lapsias bellavista and Lapsias guamani. Geneviéve Leduc-Robert and Teresa Maddison assisted with molecular laboratory work. We thank Todd Blackledge and Cheryl Hayashi for their unpublished protocols for myosin HC. Gustavo Ruiz, Martín Ramírez, and G.B. Edwards gave helpful comments on the manuscript. For assistance on the 2008 expedition to Papua New Guinea, we thank Stephen Richards, Bruce Beehler, William Thomas, Luc Fimo Tuki, Aislan Tama Wanakipa Indiaf, Pingisa Saiké, Yainé Ribson, Agustus Kore, Muse Opiango, Banak Gamui, Robert Sine, Conservation International, Porgera Joint Venture, and the PNG Department of Environment and Conservation (for more details, see
Specimens used in phylogenetic analyses, with localities and GenBank numbers of sequences indicated.
Authors: Wayne Maddison, Daiqin Li, Melissa Bodner, Junxia Zhang, Xu Xin, Qinqing Liu, Fengxiang Liu
Data type: Occurence; geographic locality; sex.
Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.