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Novitaltesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK NovitaltesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK. N.Y. 10024 U.S.A. NUMBER 2705 OCTOBER 28, 1980 WILLIAM A. SHEAR A Review of the Cyphophthalmi of the United States and Mexico, with a Proposed Reclassification of the Suborder (Arachnida, Opiliones) AMERICANt MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2705, pp. 1-34, figs. 1-33, tables 1-4, 1 map October 28, 1980 A Review of the Cyphophthalmi of the United States and Mexico, with a Proposed Reclassification of the Suborder (Arachnida, Opiliones) WILLIAM A. SHEAR' ABSTRACT The species of cyphophthalmid opilionids The new family Pettalidae is proposed for Pettal- known from the United States and Mexico are us, Purcellia, Parapurcellia, Neopurcellia, Spe- surveyed. The genus Neosiro is considered a syn- leosiro, Rakaia, and Chileogovea. the subfamily onym of Siro; Siro sonoma is described as new. Stylocellinae Hansen and Sorensen is raised to The male genitalia of S. exilis, S. kamiakensis, family rank and redefined to include only the ge- and S. acaroides are illustrated for the first time, nus Stylocellus. For the genera Ogovea and Hui- and the male of Neogovea mexasca is newly de- taca, the new family Ogoveidae is proposed, and scribed. A new scheme of family-level classifi- for the genera Neogovea, Paragovia, and Meta- cation is proposed for the suborder worldwide. govea, the new family Neogoveidae. The new ar- The family Sironidae Simon is redefined to in- rangement is based upon a cladistic analysis. Ar- clude the genera Siro, Parasiro, Tranteeva, guments against Savory's 1977 proposal to Odonotosiro, Metasiro, Paramiopsalis, and Su- consider the Cyphophthalmi an order of Arach- zukielus. Troglosiro is placed here provisionally. nida separate from Opiliones are presented. INTRODUCTION Opilionids of the suborder Cyphophthalmi "harvestmen." Because of their restrictive are mitelike creatures inhabiting caves, for- habits, these animals are not likely to dis- est litter, and soil crevices. Their short legs tribute themselves with ease and therefore and armored, egg-shaped bodies are far from could be of considerable importance in bio- the usual concept of "daddy-long-legs" or geography (Juberthie and Massoud, 1976). 1 Research Associate, Department of Entomology, American Museum of Natural History; Department of Biology, Hampden-Sydney College, Hampden-Sydney, VA 23943. Copyright © American Museum of Natural History 1980 ISSN 0003-0082 / Price $2.35 2 AMERICAN MUSEUM NOVITATES NO. 2705 At first considered among the rarest of arach- phophthalmids (Shear, 1977, 1979a, 1979b) nids, the cyphophthalmids have been the it has become obvious that the present subject, in the last two or three decades, of scheme of classification, especially at the a virtual explosion of knowledge and inter- level of the family, no longer reflects all that est, which, paradoxically, has not been fully we have learned about the group. In partic- incorporated into their classification. Fur- ular, as Hoffman (1963) accurately predicted, ther, no comprehensive review of the group the grouping of all forms into one family Si- in America has been carried out. ronidae, with two subfamilies Sironinae and The first cyphophthalmid opilionid known Stylocellinae, is quite inadequate to repre- from the United States, Siro acaroides (Ew- sent the knowledge that has accumulated ing) of Oregon, was described in 1923. Dur- concerning the probable evolutionary rela- ing the next 20 years, only two new species tionships of the described genera and were described: Siro americanus Davis from species. Therefore, the purpose of the sec- Florida (Davis, 1933) and Neosiro kamia- ond part of this paper is to propose a new kensis Newell from Washington (Newell, family-level classification for the suborder 1943). Only the work of Newell, including a Cyphophthalmi. In addition, the recent pro- redescription of acaroides published in 1947, posal of Savory (1977) that the suborder be can be said to meet modern standards for raised to full ordinal status will be critically taxonomic work. By far the most valuable examined. contribution on the suborder in America was that of Hoffman (1963), who described Siro ACKNOWLEDGMENTS exilis from the central Appalachian region, critically reviewed previous work, and set up I am grateful to Dr. Norman Platnick of a new genus, Floridogovea, for S. ameri- the American Museum of Natural History canus. He also drew attention to the useful- for the loan of specimens and for his com- ness in classification of the sternal region of ments on the manuscript. Specimens were the cephalothorax, particularly the relation- also lent by Dr. H. W. Levi, Museum of ships of the coxal elements to the gonos- Comparative Zoology, Mr. Henry Dybas, tome. Hoffman did not examine the male Field Museum of Natural History, Dr. Paul genitalia of any of the American species, al- Arnaud, California Academy of Sciences, though it had been known for some time that Mr. Rod Crawford, University of Washing- the form of the penis is of great importance ton (Seattle) Museum, Dr. S. Tuxen, Uni- in taxonomy. He was also evidently unaware versitets Museum, K0benhavn, and Mr. F. of a paper by Juberthie (1960) redescribing Wanless, British Museum (Natural History). S. americanus in detail, and giving it the new Dr. William B. Muchmore lent material from generic name Metasiro. Juberthie provided his personal collection, as did Dr. Richard the first illustrations of the male genitalia of L. Hoffman. an American species. Finally, Shear (1977) Knowledge of the distribution of Siro aca- described, from female specimens, the first roides comes largely from a sampling pro- troglobitic cyphophthalmid from the New gram carried out by Dr. Ellen Benedict, Port- World, Neogovea mexasca, collected in a land State University, Portland, Oregon. She cave in Oaxaca, Mexico. generously allowed me to sort opilionids The purpose of the first part of this paper from her material. is to review briefly our knowledge of a bio- Drs. Jochen Martens, Ray Foster, and Jur- geographically important group of soil ani- gen Gruber read the second section of the mals in North America, providing compre- manuscript and made valuable comments, hensive data on distribution and illustrating but the ideas expressed there are the full re- taxonomic features not heretofore described. sponsibility of the author. I am also most In addition, a remarkable new species from grateful to Mr. Theodore Savory for his cor- California has been discovered. During the respondence. course of this investigation and others on cy- Finally, I thank the Faculty Research 1980 SHEAR: CYPHOPHTHALMI 3 Committee of Hampden-Sydney College for generous financial support. ABBREVIATIONS AMNH, American Museum of Natural History, New York MCZ, Museum of Comparative Zoology, Harvard University, Cambridge FMNH, Field Museum of Natural History, Chi- cago CAS, California Academy of Sciences, San Fran- cisco UWM, University of Washington Museum, Se- attle WBM, personal collection of William B. Much- MAP 1. Distribution of cyphophthalmid opili- more, Rochester onids in the United States. Open circle enclosed in WAS, personal collection of William A. Shear, solid line, Siro exilis. Filled circles, Metasiro Hampden-Sydney americanus. Open triangles enclosed in solid line, Siro acaroides. Filled triangles, Siro kamiakensis. THE NORTH AMERICAN Filled square, Siro sonoma. Open square, single female specimen of undescribed probable new CYPHOPHTHALMID FAUNA species. Hoffman (1963), Juberthie (1970) and Shear (1977) have discussed in some detail the usefulness of various taxonomic charac- ters in the study of the Cyphophthalmi. To Cyphophthalmus Joseph, 1868, p. 249 (type summarize, the characters most useful at the species, C. duricorius, Joseph by original des- level of species seem to be the form and lo- ignation). Rosas Costa, 1950, p. 145 (complete references). cation of the adenostyle of the male, the Holosiro Ewing, 1923, p. 388 (type species, H. shape and spination of the penis, the rela- acaroides Ewing, by original designation). tionships of the elements of the ventral tho- Roewer, 1927, p. 266. Hinton, 1938, p. 332. racic complex to the gonostome, and the pro- Neosiro Newell, 1943, p. 416 (type species, N. portions of the bodies of both sexes. These kamiakensis Newell, by original designation). characters, if concepts are appropriately Rosas Costa, 1950, p. 142. Hoffman, 1963, p. broadened, remain useful for distinguishing 133. Juberthie 1969, p. 1383. New Subjective related groups of species as genera, but to Synonymy. these we must add the degree of fusion of DIAGNOSIS: Metasiro is the only other si- the coxae to one another and to the dorsum, ronid genus found in North America; the the presence or absence of teeth on the males of its single species have a fimbriate claws, modifications of the male anal region, adenostyle rim giving the appearance of a the position of the ozophores, and the den- tuft of setae; in Siro species the adenostyle tition of the cheliceral fingers. is lamelliform. DESCRIPTION: See Juberthie (1967, 1969). FAMILY SIRONIDAE SIMON The former reference is to a detailed rede- SIRO LATRIELLE scription of S. rubens, the type species. DISTRIBUTION: Europe; Appalachians in Siro Latrielle, 1804, p. 329 (type species, S. ru- North the Pacific coast bens Latrielle, by original designation). Hansen eastern America, and S6rensen, 1904, p. 88, p. 107. Roewer, from Washington to northern California; 1923, p. 52; 1927, p. 266. Hinton, 1938, p. 331. western Washington and adjoining regions of Rosas Costa, 1950, p. 145 (complete references northern Idaho. to 1948). Hoffman, 1963, p. 132. Juberthie, NOTES: Species of Siro are united by the 1969, p. 1383. Martens, 1978, p. 60. form of the adenostyle, ventral complex, and 4 AMERICAN MUSEUM NOVITATES NO. 2705 penis, as well as some additional characters plate has a median ridge (figs. 6, 11), and in discussed by Juberthie (1969). Neosiro was S.
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