THE SPIDER SUBFAMILY AMAUROBIOIDINAE (ARANEAE, ANYPHAENIDAE): A PHYLOGENETIC REVISION AT THE GENERIC LEVEL MARTIÂN J. RAMIÂREZ Fessenden Research Fellow, Division of Invertebrate Zoology American Museum of Natural History Research Scientist, Museo Argentino de Ciencias Naturales, Consejo Nacional de Investigaciones Cientõ®cas y TeÂcnicas Av. Angel Gallardo 470 C1405DJR Buenos Aires Argentina BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 277, 262 pp., 139 ®gures, 29 tables Issued August 13, 2003 Copyright q American Museum of Natural History 2003 ISSN 0003-0090 2 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 277 CONTENTS Abstract ....................................................................... 3 Introduction .................................................................... 4 Materials and Methods .......................................................... 6 Format of Descriptions ........................................................ 6 Cladistic Analysis ............................................................... 7 Representatives ............................................................... 7 Outgroups ................................................................... 8 Characters ................................................................... 8 Coding ...................................................................... 8 Character Description and Optimizations ........................................ 8 Characters Not Included ...................................................... 23 Evaluation of Cladistic Hypothesis ............................................. 24 Tree Searches ............................................................... 24 Character Weighting ......................................................... 24 Lists of Synapomorphies ..................................................... 28 Indices of Support ........................................................... 28 Weighting Functions and Support .............................................. 31 Discussion .................................................................... 31 Taxonomy .................................................................... 38 Anyphaenidae Bertkau ....................................................... 38 Malenellinae RamõÂrez ........................................................ 38 Anyphaeninae Bertkau ....................................................... 39 Amaurobioidinae Hickman .................................................... 41 Morphological Remarks .................................................... 44 Tribe Amaurobioidini Hickman ................................................ 51 Amaurobioides O.P.-Cambridge ............................................. 52 Axyracrus Simon .......................................................... 57 Aysenia Tullgren .......................................................... 60 Aysenoides, new genus ..................................................... 67 Acanthoceto Mello-LeitaÄo .................................................. 74 Ferrieria Tullgren ......................................................... 78 Coptoprepes Simon ........................................................ 79 Gamakia, new genus ....................................................... 91 Negayan, new genus ....................................................... 96 Selknamia, new genus ..................................................... 105 Josa Keyserling .......................................................... 108 Tribe Gayennini, New Rank ................................................. 120 Gayenna Nicolet ......................................................... 120 Gayennoides, new genus .................................................. 125 Arachosia O.P.-Cambridge ................................................. 130 Sanogasta Mello-LeitaÄo ................................................... 142 Philisca Simon ........................................................... 176 Tomopisthes Simon ....................................................... 192 Araiya, new genus ........................................................ 210 Oxysoma Nicolet ......................................................... 214 Tasata Simon ............................................................ 229 Phidyle Simon ........................................................... 242 Monapia Simon .......................................................... 247 Acknowledgments ............................................................ 256 References ................................................................... 256 Index of Speci®c and Generic Names ........................................... 261 2003 RAMIÂREZ: SPIDER SUBFAMILY AMAUROBIOIDINAE 3 ABSTRACT A cladistic phylogenetic analysis at generic level of the subfamily Amaurobioidinae is pre- sented. 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 identi®ed by jackkni®ng and Bremer indices are in general those that vary in resolution with different character-weighting schemes. Seven outgroup representatives were included (Malenella nana RamõÂrez, from Malenellinae, and six Anyphaeninae species). In this analysis Anyphaeninae, previously identi®ed as sister group of Amaurobioidinae, is paraphyletic, but forcing its monophyly does not alter the group- ings within Amaurobioidinae. The monophyly of the genera is in general well supported, but some particularly con¯icting 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 classi®ed 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 ®rst 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-LeitaÄo, Clubiona paduana Karsch, Gayenna excepta Tullgren, Gayenna exigua Mello-LeitaÄo, and Tomopisthes lebruni Simon). The previously revised genera Acanthoceto Mello-LeitaÄo 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 de®ned 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-LeitaÄo, Gayenna duplovittata Mello-LeitaÄo, Gayenna bonneti Mello-LeitaÄo, Oxysoma dubium Berland, Oxysoma bifasciatum Mello-LeitaÄo, Oxysoma cubana Banks, Oxysoma polytrichium Mello- LeitaÄo, 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-LeitaÄo 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-LeitaÄo, Gayenna alticola Simon, Gayenna bonariensis Mello-LeitaÄo, Gayenna ru®thorax Tullgren, Gayenna x-signata Keyser- ling, Gayenna approximata Tullgren, Samuza minuta Keyserling, and Tomopisthes backhau- seni 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 de- scribed. The males of Monapia carolina RamõÂrez and Monapia angusta (Mello-LeitaÄo) 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 rede®ned to include Liparotoma Simon. Clu- biona tripunctata Nicolet and Clubiona gayi Nicolet are also transferred to Philisca. The