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New World Pholcid Spiders (Araneae: Pholcidae): a Revision at Generic Level

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New World Pholcid Spiders (Araneae: Pholcidae): a Revision at Generic Level NEW WORLD PHOLCID SPIDERS (ARANEAE: PHOLCIDAE): A REVISION AT GENERIC LEVEL BERNHARD A. HUBER Division of Invertebrate Zoology American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 254, 348 pages, 1357 figures, 9 maps, 4 appendices Issued June 30, 2000 Price: $29.30 a copy Copyright ᭧ American Museum of Natural History 2000 ISSN 0003-0090 CONTENTS Abstract ...................................................................... 04 Introduction ................................................................... 04 Materials and Methods ....................................................... 06 Acknowledgments ........................................................... 07 Phylogenetics ................................................................. 07 Terminal Taxa ............................................................... 07 Characters Scored ........................................................... 09 Characters Not Scored ....................................................... 32 Cladistic Analysis ........................................................... 34 Taxonomy—Pholcidae .......................................................... 41 Diagnosis ................................................................... 41 Description ................................................................. 46 Natural History .............................................................. 48 Composition ................................................................ 50 Key to New World Genera ................................................... 50 Metagonia Simon .............................................................. 53 Leptopholcus Simon ............................................................ 76 Ninetis Simon ................................................................. 80 Nerudia, New Genus ........................................................... 87 Kambiwa, New Genus .......................................................... 87 Gertschiola Brignoli ........................................................... 90 Ibotyporanga Mello-Leita˜o ...................................................... 94 Guaranita, New Genus ......................................................... 96 Galapa, New Genus .......................................................... 100 Enetea, New Genus ........................................................... 103 Aucana, New Genus .......................................................... 105 Pholcophora Banks ........................................................... 113 Tolteca, New Genus .......................................................... 117 Papiamenta, New Genus ...................................................... 121 Chisosa, New Genus .......................................................... 124 Priscula Simon ............................................................... 128 Tainonia, New Genus ......................................................... 145 Physocyclus Simon ............................................................ 148 Ixchela, New Genus ........................................................... 150 Aymaria, New Genus .......................................................... 153 Chibchea, New Genus ......................................................... 162 Mesabolivar Gonza´lez-Sponga .................................................. 189 Carapoia Gonza´lez-Sponga .................................................... 238 Coryssocnemis Simon ......................................................... 246 Mecoloesthus Simon .......................................................... 255 Kaliana, New Genus .......................................................... 271 Waunana, New Genus ......................................................... 274 Pisaboa, New Genus .......................................................... 281 Pomboa, New Genus .......................................................... 288 Litoporus Simon .............................................................. 292 Otavaloa, New Genus ......................................................... 305 Teuia, New Genus ............................................................ 313 Tupigea, New Genus .......................................................... 314 Canaima, New Genus ......................................................... 327 Blancoa, New Genus .......................................................... 330 Species Incertae Sedis ......................................................... 334 2 2000 HUBER: NEW WORLD PHOLCID SPIDERS 3 References ................................................................... 337 Appendix 1. Matrix for Phylogenetic Analysis ................................... 342 Appendix 2. Cladogram ....................................................... 343 Appendix 3. Species of Wrong or Doubtful Generic Position ....................... 344 Appendix 4. Complete List of Pholcid Genera Worldwide ......................... 346 Index of Generic and Specific Names ........................................... 347 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 254 ABSTRACT New World pholcids are revised at the generic level, with an emphasis on South America. A total of 47 extant genera occur in the New World, 22 of which are newly described. A preliminary key to the genera of the New World is presented. Redescriptions are given for 11 genera and for 59 species, and 106 species are newly described. Three generic names and 14 specific names are newly synonymized (see below). A numerical cladistic analysis is per- formed using a matrix of 61 taxa (48 of them New World pholcids) and 61 morphological characters. The main results of the analysis are as follows: (1) Pholcids are strongly supported as a monophyletic group. (2) Pholcids are separated into the following clades, which are tentatively named to emphasize their character as a working hypothesis: ‘‘ninetines,’’ ‘‘phol- cines’’ (Metagonia Simon and the Pholcus group sensu Huber), ‘‘holocnemines’’ (Holocnemus group sensu Timm, Artema Walckenaer, Physocyclus Simon, and Priscula Simon), and the ‘‘New World clade.’’ Their interrelationships are not definitively resolved. (3) New World pholcids are an assemblage of representatives of all major clades within the family, but most genera are part of the New World clade, which is the only clade restricted to the New World. (4) A hypothesis concerning the evolutionary transformation of characters is given for 41 of the 54 traits scored that vary among pholcids. The following names are newly synonymized: Anomalaia Gonza´lez-Sponga, 1998, with Metagonia Simon, 1893; Blechroscelis Simon, 1893, with Priscula Simon, 1893; Myrmido- nella Berland, 1919, with Ninetis Simon, 1890; Blechroscelis coeruleus (Keyserling, 1891), with Coryssocnemis [now Mesabolivar] togatus (Keyserling, 1891); Blechroscelis irroratus Mello-Leita˜o, 1947, Psilochorus browningi Roewer, 1951, and Blechroscelis virescens Mello- Leita˜o, 1947, with Blechroscelis [now Mesabolivar] aurantiacus (Mello-Leita˜o, 1930); Ble- chroscelis viridis Mello-Leita˜o, 1918, with Litoporus [now Mesabolivar] brasiliensis (Moenk- haus, 1898); Hypsorinus conwayi Mello-Leita˜o, 1947, with Priscula binghamae (Chamberlin, 1916); Priscula ranchograndensis Gonza´lez-Sponga, 1996, with Priscula venezuelana Simon, 1893; Litoporus abrahami Mello-Leita˜o, 1947, with Coryssocnemis [now Litoporus] uncatus (Simon, 1893); Litoporus coccineus Simon, 1893, Litoporus imbecillus (Keyserling, 1891), and Litoporus fulvus Moenkhaus, 1898, with Litoporus [now Mesabolivar] luteus (Keyserling, 1891); Micromerys occidentalis (Mello-Leita˜o, 1929), with Micropholcus fauroti (Simon, 1887); Pholcus dubiomaculatus Mello-Leita˜o, 1918, with Pholcus phalangioides (Fuesslin, 1775); Physocyclus dubius Mello-Leita˜o, 1922, with Physocyclus globosus (Taczanowski, 1874). INTRODUCTION One of the major incentives for this study survey in a single state (Departamento del was the gross imbalance between the appar- Valle) yielded a total of 52 morphospecies, ent diversity and species richness of pholcid only one of which could be identified (Flor- spiders on one hand, and the seemingly ez, 1996). In another faunistic study in north- hopeless taxonomic situation within the fam- ern Peru, Silva (1996) reported over two doz- ily on the other hand, especially regarding en morphospecies from a restricted rainforest the Neotropics. The first point has been made area, while only four species were previously clear in a number of recent, scattered studies recorded from all of Peru. Species numbers that have shown that throughout the tropics, have increased tremendously in the few cases pholcids are not only vastly more diverse where more or less comprehensive surveys than suggested by the number of nominal were made (e.g., Gertsch, 1982, who de- species, but that in some areas they are even scribed 44 new species of Anopsicus Cham- among the dominant web-building spider berlin and Ivie in a region where only 19 families. For example, while only five phol- were previously known; Huber, 1998c, who cid species were previously recorded from described 10 new species of Modisimus Si- Colombia (two of them were actually col- mon in Costa Rica, where only one was pre- lected in Hamburg from bananas shipped viously recorded). There are hints that the from Colombia: Schmidt, 1956), a spider situation in the Old World is not much dif- 2000 HUBER: NEW WORLD PHOLCID SPIDERS 5 ferent: Deeleman-Reinhold (1986a) reported not surprising that subsequent
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