The spider subfamily Amaurobioidinae (Araneae, Anyphaenidae): a phylogenetic revision at the generic level

Publication Type:Journal Article
Year of Publication:2003
Authors:Ramírez, MJ
Journal:Bulletin of the American Museum of Natural History
Volume:277
Pagination:1-262
Abstract:

A cladistic phylogenetic analysis at generic level of the subfamily Amaurobioidinae is presented. The analysis is based on a dataset of 93 representative species scored for one behavioral and 199 morphological characters. Tree searches were made under equal and implied weights according to homoplasy, and the results were compared in terms of sensitivity to jackknife resampling. Mildest weighting functions produced trees more robust to resampling, and those results were selected as the working phylogenetic hypotheses. Groups of weak support as identified by jackknifing and Bremer indices are in general those that vary in resolution with different character-weighting schemes.
Seven outgroup representatives were included (Malenella nana Raḿırez, from Malenellinae, and six Anyphaeninae species). In this analysis Anyphaeninae, previously identified as sister group of Amaurobioidinae, is paraphyletic, but forcing its monophyly does not alter the groupings within Amaurobioidinae. The monophyly of the genera is in general well supported, but some particularly conflicting groups are discussed. In contrast, the relationships among genera are in general problematic.
Amaurobioidinae is diagnosed by a pronounced indentation at the base of male palpal tegulum, and by a particular male copulatory bulb conformation, with a paramedian apophysis. The subfamily is classified in two tribes (Gayennini and Amaurobioidini); the genus Josa Keyserling, probably sister group to Gayennini, is not assigned to either tribe.
The tribe Amaurobioidini is mainly diagnosed by an apical loop of the sperm duct in the male copulatory bulb. It includes 10 genera: Amaurobioides O.P.-Cambridge is restricted to seashores of southern continents. Clubiona chilensis Nicolet, transferred to Amaurobioides, is the first true record of the genus for South America. The male of Axyracrus elegans Simon, three species of Aysenia Tullgren, and three of Coptoprepes Simon are newly described. Four new genera are proposed in Amaurobioidini: Gamakia, Selknamia (described for one new species each), Aysenoides (for three new species), and Negayan (type species Gayenna tri- dentata Simon, including also Axyracrus coccineus Mello-Leita ̃o, Clubiona paduana Karsch, Gayenna excepta Tullgren, Gayenna exigua Mello-Leitão, and Tomopisthes lebruni Simon). The previously revised genera Acanthoceto Mello-Leitao and Ferrieria Tullgren are also in- cluded in the tribe. The basal branch and most intergeneric branches of the tribe have low support values. Amaurobioides and Negayan, however, are relatively well supported.
The tribe Gayennini is well defined by a homogeneous conformation of male and female genitalia, with a distinctive secondary conductor and spherical spermathecae. It includes 11 genera: Gayenna Nicolet includes only G. americana Nicolet from Chile and adjacent Argen- tina. Arachosia O.P.-Cambridge comprises many species previously assigned to Oxysoma. Abuzaida striata Keyserling, Anyphaena oblonga Keyserling, Gayenna proseni Mello-Leita ̃o, Gayenna duplovittata Mello-Leita ̃o, Gayenna bonneti Mello-Leita ̃o, Oxysoma dubium Berland, Oxysoma bifasciatum Mello-Leita ̃o, Oxysoma cubana Banks, Oxysoma polytrichium Mello- Leita ̃o, Phidyle bergi Simon, and Samuza praesignis Keyserling are transferred to Arachosia. The males of Arachosia bergi (Simon), A. honesta Keyserling, and Arachosia praesignis (Key- serling) are newly described. Arachosia is easily recognized by the thick setae on the anterior lateral spinnerets, and it has good support values. A very diverse group of species here as- signed to the genus Sanogasta Mello-Leita ̃o is paraphyletic in terms of Arachosia. It includes many of the species formerly placed in Gayenna Nicolet. Anyphaena maculatipes Keyserling, Clubiona maculosa Nicolet, Gayenna paucilineata Mello-Leitão, Gayenna alticola Simon, Gayenna bonariensis Mello-Leitao, Gayenna rufithorax Tullgren, Gayenna x-signata Keyser- ling, Gayenna approximata Tullgren, Samuza minuta Keyserling, and Tomopisthes backhauseni Simon are transferred to Sanogasta. The female of Sanogasta alticola (Simon), the males of S. x-signata (Keyserling) and S. approximata (Tullgren), and four species are newly described. The males of Monapia carolina Ram ́ırez and Monapia angusta (Mello-Leita ̃o) are newly described. A new species of Oxysoma Nicolet from southern Brazil is described, and Gayenna saccata Tullgren is transferred to Oxysoma. Phidyle Simon is removed from the synonymy of Oxysoma Nicolet; the male of its only species Phidyle punctipes (Nicolet) is newly described. The genus Philisca Simon is redefined to include Liparotoma Simon. Clubiona tripunctata Nicolet and Clubiona gayi Nicolet are also transferred to Philisca. The male of Philisca hahni Simon and two species are newly described. The genus is reasonably supported, except for one basal species of questionable placement. Anyphaena punctata Keyserling, Gayenna fuscotaeniata Keyserling, Gayenna tripunctata Mello-Leitao, Gayenna reticulata Mello-Leitão, Gayenna taperae Mello-Leitao, Oxysoma quinquenotatum Simon, Oxysoma unipunctatum Simon, Oxysoma novum Mello-Leitao, Oxysoma lineatum Tullgren, and Tomopisthes frenatus Mello-Leitao are transferred to Tasata. The males of Tasata parcepunctata Simon, T. variolosa Mello-Leitao, and three species are newly described. Tasata albofasciata Mello-Leitão is transferred to Tupirinna Bonaldo, in the Corinnidae. Tomopisthes Simon includes only three species from Chile and adjacent Argentina. Clubiona horrenda Nicolet and Clubiona pusilla Nicolet are transferred to Tomopisthes. The male of Tomopisthes pusillus (Nicolet) is newly described. Two new genera are proposed in Gayennini: Araiya (Gayenna pallida Tullgren, type species and Gayenna coccinea Simon) and Gayennoides (for two new Chilean species).

The genus Josa Keyserling, distinguished by a femoral apophysis on the male palp, is extremely diverse in Andean cloud forests and tropical America. It is one of the better supported groups of the analysis. Anyphaena keyserlingi L. Koch, Gayenna andesiana Berland, Gayenna simoni Berland, Gayennella riveti Berland, Haptisus nigrifrons Simon, Haptisus analis Simon, Haptisus maurus Simon, Olbophthalmus lojensis Berland, Olbus personatus Simon, Olbus gounellei Simon, Tetromma luteum Keyserling, and Tomopisthes chazaliae Simon are transferred to Josa. The male of Josa riveti (Berland) and one species are newly described. The following names are newly synonymized: Cluilius Simon, with Amaurobioides O.P.- Cambridge; Schiapellia Mello-Leitao, with Axyracrus Simon; Schiapellia gerschmanni Mello Leitão and Amaurobioides boydi Forster, with Axyracrus elegans Simon; Tomopisthes magel- lanicus Simon and Gayenna strigosa Tullgren, with Clubiona (now Negayan) paduana Karsch; Tetromma Keyserling (preoccupied), Haptisus Simon, Olbophthalmus Simon, and Gayennella Berland, with Josa Keyserling; Anyphaena pilosa Keyserling and Gayenna riveti Berland, with Tetromma (now Josa) luteum Keyserling; Pelayo insignis Banks, with Haptisus (now Josa) nigrifrons Simon; Samuza Keyserling, Abuzaida Keyserling, and Gayennina Gertsch, with Arachosia O.P.-Cambridge; Tomopisthes tripunctatus Mello-Leitão, with Samuza (now Arachosia) praesignis Keyserling; Oxysoma ramboi Mello-Leitao, with Arachosia honesta Keyserling; Sanogasta intermedia Mello-Leitao, with Anyphaena (now Sanogasta) maculatipes Keyserling; Gayenna monticola Chamberlin, with Gayenna alticola Simon; Clubiona sternalis Nicolet, Anyphaena ignota Keyserling, Gayenna affinis Tullgren, Gayenna dubia Tullgren, Tomopisthes conspersus Simon, Tomopisthes modestus Simon, Tomopisthes taeniatus Simon, Gayenna skottsbergi Berland, and Tomopisthes injucundus Simon, with Clubiona (now Sanogasta) maculosa Nicolet; Tomopisthes kraepelini Simon, with Gayenna approximata Tullgren; Liparotoma Simon, with Philisca Simon; Philisca navarinensis Tullgren, with Philisca hahni Simon; Heteromma Karsch (preoccupied), with Tomopisthes Simon; Tomopisthes im- manis Simon, Heteromma fuegiana Karsch, Philisca sica Strand, and Nonianus argentinus Mello-Leitao, with Clubiona (now Tomopisthes) horrenda Nicolet; Gayenna chilensis Tullgren, with Clubiona (now Tomopisthes) pusilla Nicolet; Gayenna stellata Simon, with Gayenna (now Araiya) coccinea Simon; Oxysoma punctipes Nicolet, Oxysoma aurata Nicolet, Oxysoma longipes Nicolet, Oxysoma lineata Nicolet, and Aporatea valdiviensis Simon, with Oxysoma punctatum Nicolet.
The following names, previously listed in Anyphaenidae, are considered nomina dubia: Anyphaena pampa Holmberg, Clubiona albiventris Nicolet, Clubiona citrina Nicolet, Clubiona gemella Nicolet, Clubiona gibbosa Nicolet, Clubiona lepida Nicolet, Clubiona limbata Nicolet, Clubiona lineata Nicolet, Clubiona nigricans Nicolet, Clubiona nubes Nicolet, Clubiona pulchella Nicolet, Clubiona puella Nicolet, Clubiona versicolor Nicolet, Oxysoma auratum Nicolet, Oxysoma delfini Simon, and Tomopisthes aethiops Simon.

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