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Spiders

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4 BULLETIN AMERICAN

4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 178 the Araneoidea (the Textricellidae, Micropholcommatidae, and Mimetidae) are palpimanoids as well. ACKNOWLEDGMENTS Specimens were obtained from the collections of the American Museum of Natural History, New York (AMNH), the Field Museum of Natural History, Chicago (FMNH, Dr. L. Watrous), the Museum of Comparative Zoology, Harvard University (MCZ, Dr. H. W. Levi), the National Museum of Victoria, Melbourne (NMV, Dr. A. Neboiss), the Otago Museum, Dunedin (OMD), the Queensland Museum, Brisbane (QMB, Drs. V. E. Davies and R. J. Raven), and the University of California, Berkeley (UCB, Dr. C. Griswold). We thank each of these curators and also Drs. A. F. Newton, Jr., and M. Thayer of Harvard University for getting the spiders from their 1982 Chilean expedition to us in time for them to be included in this study. Drs. Bruno Lamoral of the Natal Museum and Roland Legendre of the Universite des Sciences, Montpellier, generously provided material for comparative studies. We are indebted to the Eppley Foundation for Research for its generous funding of the second author's 1981 fieldwork in Chile, which was instrumental in our recognition of the surprising diversity of the Chilean mecysmaucheniid fauna, and to the Scientific Distribution Committee of the Golden Kiwi Lottery Fund for its continuing support of the first author's work. Technical assistance was provided by Dr. Mohammad Shadab, Ms. Joan Whelan, Mr. Paul McGuffin, Ms. Eileen Polk, and Ms. Rachel Reiss at the American Museum of Natural History and by Ms. Denise Hesson and Ms. Ruth Newlove at the Otago Museum. Scanning scope facilities were made available by the University of Otago Medical School. We especially thank Dr. Jonathan Coddington of Harvard University for making available a copy of his study of orb web monophyly prior to its publication. Reviews ofvarious drafts ofthe manuscript were kindly provided by Dr. Coddington and by Drs. Lyn Forster of the Otago Museum, Willis Gertsch of the American Museum, Robert Raven of the Queensland Museum, and William Shear of Hampden-Sydney College. The family Archaeidae was established by Koch and Berendt, in a posthumously published paper (1854), for three new species from Baltic amber which they placed in the newly erected genus Archaea. In a footnote in the same paper, Menge (who edited their original manuscript for publication) added three new species from the same source. These six fossil species remained the only known representatives of the family until 0. P.-Cambridge (1881) described (as a theridiid) the first living species, Eriauchenius workmani, from Madagascar. Simon soon placed two additional species in the family, for each ofwhich he established a new genus. He first (1883) described Landana petiti from the Congo, suggesting that both Landana and Eriauchenius are close to Archaea, and then (1884) described Mecysmauchenius segmentatus from Chile. Later (1895), having had the opportunity to ob- TAXONOMIC HISTORY serve Landana in the field and to examine more material, he concluded that he was mistaken in his original views on the relationships of Landana and transferred that genus to his group Meteae in the Tetragnathinae. The two remaining genera, Archaea (which he considered a senior synonym of Eriauchenius) and Mecysmauchenius, he placed in separate groups-the Archaeae and Mecysmauchenieae. Subsequently, Petrunkevitch (1928) listed these two taxa as subfamilies and in a later paper (Petrunkevitch, 1939) argued strongly for the reinstatement of Landana within the Archaeidae. Wilton (1946) established a new genus, Zearchaea, for a minute spider from New Zealand which he placed in the Archaeidae, stating that it possessed characters intermediate between Archaea and Mecysmauchenius. In that paper he apparently accepted Petrunkevitch's views on the inclu-

1 984 FORSTER AND PLATNICK: ARCHAEID SPIDERS 5 sion of Landana within the Archaeidae but pointed out that Landana differs more from the other three genera (Archaea, Mecysmauchenius, and Zearchaea) than they do from each other, and he proposed that the subfamily Mecysmaucheniinae be dropped. Shortly after this, Forster (1949) described three new "archaeids" from New Zealand, two of which were placed in Zearchaea and the third in Archaea. After additional material, including specimens from Australia, became available, he (1955) transferred one of these species from Zearchaea to the newly established genus Pararchaea (along with two newly described species, one from Australia and the other from New Zealand). In that same paper, the new genus Holarchaea was established for the New Zealand species originally described in Archaea. Lehtinen (1967) rejected Petrunkevitch's suggestion that Landana be returned to the Archaeidae and also stated that the Recent species of Archaea could not be considered congeneric with the fossil species, which he believed probably represented at least two genera. He gave no evidence in support of these views, however, either with regard to the generic allocation of the fossil species or their relationship with the Recent taxa. He suggested that Archaea and Eriauchenius be retained in the family Archaeidae, which he felt might also include Holarchaea, but that Mecysmauchenius, Pararchaea, and Zearchaea should be placed in a separate family Mecysmaucheniidae. Lehtinen also maintained that these two families are not related and should be placed in entirely different branches within the classification he proposed. The Archaeidae he placed in the Araneoidea, within his branch Araneides, whereas the Mecysmaucheniidae were located in his branch Zodariides. His placement of the Mecysmaucheniidae in the superfamily Zodarioidea was apparently influenced by the reduction of the posterior and median spinnerets (which actually occurs in only two of the three genera he grouped together), but again no direct justification for this action was given. Lehtinen did specify (as usual, without supporting characters) that his group Mecysmaucheniidae "seems to be related to Palpimanidae, and possibly also to the genus Plectophanes" (1967, p. 290). Legendre (1970a, 1977) concurred with Lehtinen on the misplacement of Landana in the Archaeidae but rejected his dismemberment of the family. Although Lehtinen (1975) nonetheless maintained his diphyly argument, he seems subsequently to have abandoned it, for Lehtinen (1980, p. 495) includes the self-contradictory statement that "Mecysmaucheniidae and Archaeidae are not closely related to each other, but they seem to belong to the same main group of spider evolution, in which there are no other recent families." Brignoli (1980a) noted that although Mecysmauchenius has haplogyne genitalia, those ofPararchaea and Holarchaea are much more complex; elsewhere (1 980b) he commented on the cheliceral and vulval similarities between Mecysmauchenius and Palpimanus. However, he concluded only (1 980a, p. 31) that "Il y a donc la possibilite que le taxon 'Archaeidae' soit polyphyletique." Assessments as mutually contradictory as those of Lehtinen have recently been published by Levi, who (1981) supported the conclusions of Lehtinen and Legendre regarding the placement of Landana, although he synonymized the genus with Dolichognatha (on the phenetic grounds that "there are intermediates") and placed it in the Tetragnathinae. In that paper, he argued that the Archaeidae must be excluded from the Araneoidea, citing (p. 280) as "the most important difference" the fact that "the median spinnerets are minute, and the posterior spinnerets are much smaller than the anterior ones" and also mentioning that, unlike araneoids, the single metatarsal trichobothrium is distally situated. In this (1981) paper, Levi specifically included in the Archaeidae the genera Archaea and Zearchaea, and, judging by his references to Forster's (1955) illustrations, intended to include at least Pararchaea as well. Despite these comments on excluding archaeids from the Araneoidea, Levi (1982) published a new classification of spiders in which the genera Archaea and Holarchaea are placed in the Archaeidae and retained in the Araneoidea (Archaea was erroneously characterized as having no posterior spiracles; the New Zealand Holarchaea was also erroneously cited as Australian). The three genera Mecysmauchenius, Zearchaea, and

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