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United States National Museum Bulletin259 SMITHSONIAN INSTITUTION MUSEUM O F NATURAL HISTORY UNITED STATES NATIONAL MUSEUM BULLETIN259 Systematics, Speciation, and Distribution of the Subterranean Amphipod Genus Stygonectes (Gammaridae) JOHN R. HOLSINGER SMITHSONIAN INSTITUTION PRESS WASHINGTON, B.C. 1967 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States National Museum Bulletin. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the fields of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. This work forms number 259 of the Bulletin series. Frank A. Taylor Director, United States National Museum U.S. GOVERNMENT PRINTING OFFICE WASHINGTON : 1967 For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C., 20102 Price 75 cents (paper cover) Contents Page Introduction 1 Objectives 2 Historical review 3 Procedure 6 Terminology 9 Taxonomy 12 Acknowledgments 14 Systematics 15 Family Gammaridae 15 Crangonyx section 16 Genus Stygonecies 17 Evolution 124 Distribution and origin 124 Speciation and dispersal 132 Conclusions 168 Summary 170 Literature cited 172 Figures Page 1. Preserved specimen of Stygonectes reddelli Holsinger from Whit eface Cave, San Saba Co., Texas facing page 16 2. Stygonectes emarginatus (Hubricht) 24 3. Stygonectes viundus, new species 27 -i. Stygonectes morrisoni, new species 29 5. Stygonectes cooperi, new species 32 G. Stygonectes stellmacki, new species 34 7. Stygonectes gracilipes, new species 37 8. Stygonectes conradi, new species 41 9. Stygonectes spinatiis, new species 44 10. Stygonectes pizzinii (Shoemaker) 48 11. Stygonectes indentatiis, new species 54 12. Stygonectes tenuis tenxtis (Smith) 59 13. Stygonectes tenuis potomactis, new subspecies 63 14. Relation of antennal length to standard length (body length excluding appendages) in Stygonectes tenuis potomacus, new subspecies, from seeps in Burleith Woods, Washington, D.C 64 15. Stygonectes allegheniensis, new species 67 16. Relation of antennal length to standard length (body length excluding appendages) in Stygonectes allegheniensis, new species, from seeps, springs, and wells in Herkimer and Onondaga Counties, New York 70 17. Stygonectes hayi (Huliricht and Mackin) 72 18. Stygonectes alabamensis alabamensis (Stout) 76 19. Stygonectes montanus, new species 83 20. Stygonectes elatus, new species 85 V VI FIGURES Page 21. Stygonectes barri, new species 88 22. Stygonectes bifurcatus, new species 93 23. Stygonectes russelH, new species 96 24. Stygonectes russeUi, new species 98 25. Stygonectes clantoni (Greaser) 103 26. Stygonectes ozarkensis, new species 108 27. Stygonectes bowmani, new species 112 28. Stygonectes pecki, new species 118 29. Stygonectes dejectus, new species 120 30. Distribution of Stygonectes in North America 126 31. Distribution of species of (S^y^onecies in the central Appalachians . 133 32. Suggested phylogeny of emarginatus and flagellatus groups of Sty- gonectes 134 33. Distribution of species of Stygonectes in central Texas 140 34. Distribution of species of Stygonectes in eastern United States .... 145 35. Distribution of tenuis group of Stygonectes in south-centi-al United States 151 36. Suggested phylogeny of tenuis group of Stygonectes 152 Tables Page 1. Representative North American genera of the Crangonyx section of Gammaridae 16 2. Comparison of six representative North American genera of the Cran- gonyx section 19 3. Appendage to body length ratios in females of Stygonectes gracilipes . 39 4. Comparison of mean size of sexually mature individuals from different populations of Stygonectes pizzinii (Shoemaker) 50 5. Variation in the ratio of length of 1st antenna to length of body in different populations of Stygonectes russelli, new species 100 6. Variation in the ratio of length of 1st antenna to length of body in two populations of Stygonectes clantoni 105 Systematics, Speciation, and Distribution of the Subterranean Amphipod Genus Stygonectes (Gammaridae) ^ Introduction Amphipod crustaceans are among the largest and least studied groups of North American fresh-water invertebrates. Five famihes, including Coropliiidae, Haustoriidae, HyalelHdae, Pontogeneiidae, and Gammaridae, are represented in the fresh waters of North America. The first four families are represented by only four genera and five species; the last, however, is represented by nine genera and numerous species. For further, comprehensive reviews of the taxonomy and distribution of North American, fresh-water am- phipods, the reader is referred to excellent treatments by Pennak (1953) and Hubricht (1959). For a detailed account of the sys- tematics of fresh-water amphipods of glaciated North America, the thorough work of Bousfield (1958) is indispensable. Of the nine genera of Gammaridae with representatives in North America, seven have invaded subterranean waters of which five are known only from subterranean and/or interstitial habitats. Exclud- ing Anisogammarus and Gammarus, which also occur in marine waters, the remaining seven North American genera of Gammaridae are exclusively fresh-water inhabitants and have been assigned to the Crangonyx section of this family (see Schellenberg, 1936; Shoe- maker, 1942a; and elsewhere in this paper). A hst of six of these genera (excluding Allocrangonyx, see p. 16) and their breakdown into numbers of species is given in table 1 (see p. 16). Although pre- liminary studies indicate a wealth of undescribed species in the Crangonyx section (especially subterranean forms), the present investigation totals 29 described species in Stygonectes alone, making this the largest genus of North American, fresh-water amphipods known to date. 1 A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the University of Kentucky. Present address of the author: Department of Biology, East Tennessee State University, Johnson City, Tennessee 37601. 1 2 U.S. NATIONAL MUSEUM BULLETIN 2 59 Objectives The objective of this study is threefold: (1) to revise the systematics of the subterranean amphipod genus Stygonectes based on all available collections and all pertinent hterature; (2) to define, as clearly as possible, the patterns of species distribution and to correlate these patterns with extrinsic barriers and theoretical dispersal limits; and (3) to utilize the data derived from systematics and geographic distribution to construct a tenable model for explaining speciation in this genus. Previous to this investigation there had been no attempt to com- pletely revise the taxonomy of any of the North American, fresh- water ampliipod genera. The earher paper of Weckel (1907) and the more recent papers of Hubricht and Macldn (1940), Shoemaker (1942a), and Hubricht (1943), while all monographic in scope, did not deal specifically with a given genus but covered a wide range of species in a number of different genera. A more recent monograph on fresh-water amphipods of this continent is that by Bousfield (1958); although this paper treated comprehensively 30 species and subspecies, it was geographically limited to forms known primarily from glaciated North America. With the exception of short papers by Greaser (1934), Mackin (1935), and Shoemaker (1938, 1942a, 1942b, 1945), few papers in the 20th century have dealt specifically with North American, subterranean amphipods. JVIore than one-half of all American, fresh-water amphipod species occur exclusively in subterranean waters. The taxonomy of this large biota is still incompletely known, and its ecology and physiology is virtually unknown. Currently, not a single Hfe history is com- pletely known for an obligatory subterranean amphipod species in this country. Moreover, comprehensive studies on the evolutionary biology of North American, subterranean amphipods have been practically nonexistent until now. The present study of Stygonectes is the first in a series of several designed to ultimately revise, according to modern standards of systematics, the six North American genera presently assigned to the Crangonyx section of Gammaridae. These studies, along with ad- ditional observations on several "crangonycid genera" that occiu" out- side of North America, are intended to increase oiu- knowledge of the systematics, zoogeography, and speciation of this group of fresh- water amphipods. It is fiu-ther suggested, but perhaps a bit pre- matiu-ely, that when the proposed series of revisions is completed, sufficient
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