Wheeler Ward C, Coddington Jonathan A, Crowley Louise M, Dimitrov Dimitar, Goloboff Pablo A, Griswold Charles E, Hormiga Gustavo, Prendini Lorenzo, Ramírez Martín J, Sierwald Petra, Almeida-Silva Lina, Alvarez-Padilla Fernando, Arnedo Miquel A, Benavides Silva Ligia R, Benjamin Suresh P, Bond Jason E, Grismado Cristian J, Hasan Emile, Hedin Marshal, Izquierdo Matías A, Labarque Facundo M, Ledford Joel, Lopardo Lara, Maddison Wayne P, Miller Jeremy A, Piacentini Luis N, Platnick Norman I, Polotow Daniele, Silva-Dávila Diana, Scharff Nikolaj, Szűts Tamás, Ubick Darrell, Vink Cor J, Wood Hannah M, Zhang Junxia
Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th St., New York, NY, 10024, USA.
Smithsonian Institution, National Museum of Natural History, 10th and Constitution, NW Washington, DC, 20560-0105, USA.
Cladistics. 2017 Dec;33(6):574-616. doi: 10.1111/cla.12182. Epub 2016 Dec 12.
We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher-level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb-weavers, compatible with their non-monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, with Amphinecta, Mamoea, Maniho, Paramamoea and Rangitata (transferred from Amphinectidae); Ischaleinae, with Bakala and Manjala (transferred from Amaurobiidae) and Ischalea (transferred from Stiphidiidae); Metaltellinae, with Austmusia, Buyina, Calacadia, Cunnawarra, Jalkaraburra, Keera, Magua, Metaltella, Penaoola and Quemusia; Porteriinae (new rank), with Baiami, Cambridgea, Corasoides and Nanocambridgea (transferred from Stiphidiidae); and Desinae, with Desis, and provisionally Poaka (transferred from Amaurobiidae) and Barahna (transferred from Stiphidiidae). Argyroneta is transferred from Cybaeidae to Dictynidae. Cicurina is transferred from Dictynidae to Hahniidae. The genera Neoramia (from Agelenidae) and Aorangia, Marplesia and Neolana (from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, with Gasparia, Gohia, Hulua, Neomyro, Myro, Ommatauxesis and Otagoa (transferred from Desidae); and Toxopinae, with Midgee and Jamara, formerly Midgeeinae, new syn. (transferred from Amaurobiidae) and Hapona, Laestrygones, Lamina, Toxops and Toxopsoides (transferred from Desidae). We obtain a monophyletic Oval Calamistrum clade and Dionycha; Sparassidae, however, are not dionychans, but probably the sister group of those two clades. The composition of the Oval Calamistrum clade is confirmed (including Zoropsidae, Udubidae, Ctenidae, Oxyopidae, Senoculidae, Pisauridae, Trechaleidae, Lycosidae, Psechridae and Thomisidae), affirming previous findings on the uncertain relationships of the "ctenids" Ancylometes and Cupiennius, although a core group of Ctenidae are well supported. Our data were ambiguous as to the monophyly of Oxyopidae. In Dionycha, we found a first split of core Prodidomidae, excluding the Australian Molycriinae, which fall distantly from core prodidomids, among gnaphosoids. The rest of the dionychans form two main groups, Dionycha part A and part B. The former includes much of the Oblique Median Tapetum clade (Trochanteriidae, Gnaphosidae, Gallieniellidae, Phrurolithidae, Trachelidae, Gnaphosidae, Ammoxenidae, Lamponidae and the Molycriinae), and also Anyphaenidae and Clubionidae. Orthobula is transferred from Phrurolithidae to Trachelidae. Our data did not allow for complete resolution for the gnaphosoid families. Dionycha part B includes the families Salticidae, Eutichuridae, Miturgidae, Philodromidae, Viridasiidae, Selenopidae, Corinnidae and Xenoctenidae (new fam., including Xenoctenus, Paravulsor and Odo, transferred from Miturgidae, as well as Incasoctenus from Ctenidae). We confirm the inclusion of Zora (formerly Zoridae) within Miturgidae.
我们利用一个包含932种蜘蛛的数据集进行了蜘蛛的系统发育分析，这些蜘蛛代表了115个科（仅Synaphridae科未被代表）、700个已知属以及26个未鉴定或未描述属的其他代表。盲蛛目、触肢蛛目、裂盾蛛目和尾鞭蝎目的11个属被纳入作为外类群。该数据集包括来自线粒体（12S、16S、COI）和核基因组（组蛋白H3、18S、28S）的六个标记，并通过多种方法进行分析，包括使用来自转录组数据的高度支持的主干树进行约束分析。我们以良好的支持度恢复了蜘蛛树的大部分高级结构，包括中纺亚目、后纺亚目、原蛛亚目和新蛛亚目。我们的一些分析恢复了Hypochilidae科和Filistatidae科为姐妹群，正如先前转录组分析所表明的那样。Synspermiata得到了有力支持，Trogloraptoridae科和Caponiidae科被发现是Dysderoidea的姐妹群。我们的结果支持失落气管类群，包括幽灵蛛科、四斑蛛科、迪盖蒂蛛科、盘腹蛛科和Pacullidae科（恢复地位），它们与四斑蛛科分开。Scytodoidea包括奥氏蛛科以及刺客蛛科、绞蛛科、Drymusidae科和Periegopidae科；我们的结果对于最后这两个科的分离尚无定论。我们没有恢复单系的Austrochiloidea和Leptonetidae科，但我们的数据表明这两个类群与圆柱形腺管类群的关系比与Synspermiata更密切。我们的分析没有恢复Palpimanoidea，但也没有强烈反驳。我们发现了对Entelegynae和Oecobioidea（Oecobiidae科加Hersiliidae科）的支持，以及筛器圆网蛛的位置不明确，这与它们的非单系性相符。Nicodamoidea（Nicodamidae科加Megadictynidae科）以及园蛛总科（Araneoidea）的组成和关系与最近的分析一致。我们没有得到泰坦蛛类（Titanoecidae科和Phyxelididae科）的分辨率，但后侧胫节突类群得到了很好的支持。Penestomidae科，可能还有Homalonychidae科，是Zodarioidea的一部分，尽管后一个科最近被转录组分析分开。我们的数据支持一个我们称为马龙类群的大群体（包括园蛛科、Desidae科、皿蛛科、微蛛科、Stiphidiidae科、漏斗蛛科和毒蛛科）。这里重新定义了大多数马龙类科的范围。园蛛科包括园蛛亚科和临时的Macrobuninae亚科。我们将Malenellinae亚科（来自Anyphaenidae科的Malenella属）、Chummidae科（Chumma属）（新同义名）和Tasmarubriinae亚科（来自Amphinectidae科的Tasmarubrius属、Tasmabrochus属和Teeatta属）转移到Macrobuninae亚科。Cybaeidae科被重新定义为包括Calymmaria属、Cryphoeca属、Ethobuella属和Willisius属（从微蛛科转移过来），以及Blabomma属和Yorima属（从皿蛛科转移过来）。Cycloctenidae科被重新定义为包括Orepukia属（从漏斗蛛科转移过来）以及Pakeha属和Paravoca属（从园蛛科转移过来）。Desidae科被重新定义为包括五个亚科：Amphinectinae亚科，包括Amphinecta属、Mamoea属、Maniho属、Paramamoea属和Rangitata属（从Amphinectidae科转移过来）；Ischaleinae亚科，包括Bakala属和Manjala属（从园蛛科转移过来）以及Ischalea属（从Stiphidiidae科转移过来）；Metaltellinae亚科，包括Austmusia属、Buyina属、Calacadia属、Cunnawarra属、Jalkaraburra属、Keera属、Magua属、Metaltella属、Penaoola属和Quemusia属；Porteriinae亚科（新等级），包括Baiami属、Cambridgea属、Corasoides属和Nanocambridgea属（从Stiphidiidae科转移过来）；以及Desinae亚科，包括Desis属，以及临时的Poaka属（从园蛛科转移过来）和Barahna属（从Stiphidiidae科转移过来）。水蛛属从Cybaeidae科转移到皿蛛科。Cicurina属从皿蛛科转移到微蛛科。Neoramia属（来自漏斗蛛科）以及Aorangia属、Marplesia属和Neolana属（来自Amphinectidae科）被转移到Stiphidiidae科。毒蛛科（恢复地位）包括两个亚科：Myroinae亚科，包括Gasparia属、Gohia属、Hulua属、Neomyro属、Myro属、Ommatauxesis属和Otagoa属（从Desidae科转移过来）；以及Toxopinae亚科，包括Midgee属和Jamara属，以前的Midgeeinae亚科，新同义名（从园蛛科转移过来）以及Hapona属、Laestrygones属、Lamina属、Toxops属和Toxopsoides属（从Desidae科转移过来）。我们得到了一个单系的椭圆形纺管类群和双爪类；然而，狼蛛科不是双爪类，而是可能是这两个类群的姐妹群。椭圆形纺管类群的组成得到了确认（包括管巢蛛科、Udubidae科、栉足蛛科、猫蛛科、Senoculidae科、水蛛科、Trechaleidae科、狼蛛科、Psechridae科和蟹蛛科），肯定了先前关于“栉足蛛”Ancylometes属和Cupiennius属不确定关系的发现，尽管栉足蛛科的一个核心类群得到了很好的支持。我们的数据对于猫蛛科的单系性不明确。在双爪类中，我们发现核心的原遁蛛科首先分裂，不包括澳大利亚的莫氏遁蛛亚科，它们与核心原遁蛛科距离较远，在平腹蛛类中。其余的双爪类形成两个主要类群，双爪类A部分和B部分。前者包括大部分斜向中隔反光层类群（Trochanteriidae科、平腹蛛科、Gallieniellidae科、Phrurolithidae科、Trachelidae科、平腹蛛科、Ammoxenidae科、Lamponidae科和莫氏遁蛛亚科），以及Anyphaenidae科和管巢蛛科。Orthobula属从Phrurolithidae科转移到Trachelidae科。我们的数据无法完全解决平腹蛛类科的问题。双爪类B部分包括跳蛛科、Eutichuridae科、Miturgidae科、盗蛛科、Viridasiidae科、妖面蛛科、管巢蛛科和Xenoctenidae科（新科，包括从Miturgidae科转移过来的Xenoctenus属、Paravulsor属和Odo属，以及从栉足蛛科转移过来的Incasoctenus属）。我们确认将Zora属（以前的Zoridae科）纳入Miturgidae科。
