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The Systematics and Biology of the Cave-Crickets of the North American Tribe Hadenoecini (Orthoptera Saltatoria: Ensifera: Rhaphidophoridae: Dolichopodinae)

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The Systematics and Biology of the Cave-Crickets of the North American Tribe Hadenoecini (Orthoptera Saltatoria: Ensifera: Rhaphidophoridae: Dolichopodinae) MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY. UNIVERSITY OF MICHIGAN NO. 156 The Systematics and Biology of the Cave-Crickets of the North American Tribe Hadenoecini (Orthoptera Saltatoria: Ensifera: Rhaphidophoridae: Dolichopodinae) I> y Theodore H. Hubbell and Russell M. Norton Osborn Memorial Laboratories, Department of Biology, Yale University Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MlCHIGAN August 1, 1978 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN FRANCIS C. EVANS, EDITOR The publications of the Museum of Zoology, University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 1913, serve as a medium for original studies bised principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contribu1.ions not within the scope of the Occasional Papers, are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, .4nn Arbor, Michigan 48109. MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 156 The Systematics and Biology of the Cave-Crickets of the North American Tribe Hadenoecini ' (Orthoptera Saltatoria: Ensifera: Rhaphidophoridae: Dolichopodinae) Theodore H. Hubbell and Russell M. Norton Osborn Memorial Laboratories, Department of Biology, Yale University Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN August 1, 1978 Hadenoecus subtemaneus, mating pair, Mammoth Cave, Kentucky, January, 1968 (photo by Norton) Hadenoecus copulates with the female above. The inset shows the position of the eversible elongate dorsolateral organs of the male enclosing the end of the female's abdomen. I everted glandular A4 1 organ 8 / j \ iii CONTENTS PAGE ABSTRACT ............................................... 1 INTRODUCTION ........................................... 1 ACKNOWLEDGMENTS ....................................... 5 PART I . A REVISION OF THE RHAPHIDOPHORID TRIBE HADENOECINI byT.II.Hubbel1 .......................................... 6 Material Studied ........................................ 6 DOLICHOPODINAE ....................................... 9 HADENOECINI .......................................... 12 Key to the Genera and Species ................................14 EUHADENOECUS new genus .................................. 18 E. puteanus (Scudder) ..................................... 19 E. adelphus new species .................................... 33 E. fragilis new species ..................................... 38 6 insolitus new species .................................... 44 HADENOECUS Scudder ..................................... 48 H. subterraneus (Scudder) ................................... 53 H. cumberlandicus new species ................................ 63 H . opilionoides new species .................................. 69 H . barri new species ...................................... 74 H. jonesi new species ...................................... 77 PART I1. THE PHYLETIC AND ZOOGEOGRAPHIC HISTORY OF THE RI-IAPHIDOPHORIDAE WITI-I SPECIAL REFERENCE TO THE HADENOECINI by T . W . Hubbell .......................................... 81 THE LAURASIAN SUBFAMILIES ...............................83 DOLICIlOPODINAE .......................................85 I-IADENOECINI .......................................... 86 Pleistocene Isolation and Speciation .............................91 The Sympatry of E. puteanus and E . adelphus ....................... 92 Zoogeography of the Cave Species .............................. 93 Parthenogenetic Populations ................................. 95 PART I11. THE BIOLOGY OF HADENOECUS SUBTERRANEUS AND I% CUMBERLANDZCUS. by R . M. Norton .......................... 100 Food and Feeding ........................................ 101 Daily Activity Rhythms ..................................... 103 Courtship. Copulation and Spermatophores ......................... 104 Oviposition. Eggs. Nymphs ................................... 107 Adult Behavior and Longevity ................................. 108 Predation and its Effects on Hadenoecus Morphology .................... 108 The Role of Hadenoecus in the Terrestrial Cave Community ................ 116 LITERATURECITED .......................................117 ILLUSTRATIONS FIGURES IN TEXT FIGURE PAGE 1 Diagram of phyletic relationships within the Rhaphidophoridae .............. 8 2 Structural details of Euhadenoecus species .......................... 21 3 Ovipositors of Euhadenoecus species ............................. 23 4 Euhadenoecus insolitus. and tibia I11 of Euhadenoecus species ............... 24 5 Male terminalia of Hadcnoecini ................................. 27 6 Phallus of Hadenoecus opilionoides. fully extruded ..................... 28 7 Map showing distribution of Euhadenoecus puteanus .................... 29 8 Comparison of femur 111 lengths in Euhadenoecus puteanus and E. adelphus ...... 35 9 Map showing distribution of Euhadenoecus adelphus and E. puteanus in the southern Appalachians ................................37 10 Map showing distribution of the cavernicolous species of Euhadenoecus and Hadenoecus .......................................41 11 Map showing distribution of Euhadenoecus fragilis ..................... 42 12 Ifadenoecus o. opilionoides. and tibia 111 of Hadenoecus spccies .............. 49 13 Female subgenital platcs of Hadenoecus species ....................... 51 14 Ovipositors of Hadenoecus species ............................... 52 15 Map showing distribution of IIadenoecus subterraneus and of its predator Neaphaenops tellhampfii .................................. 60 16 Map showing distribution of Hadenoecus cumberlandicus and of its predator Darlingtonea hentuckensis .................................66 17 Map showing distribution of Hadenoecus opilionoides. H. barri. and (in part) of H. jonesi and Euhadenoecus insolitus ..........................72 18 Map showing distribution of Hadenoecus jonesi and (in part) of Euhadenoecus insolitus ...................................80 19 Hypothetical phylogeny of the Hadenoecini ......................... 87 20 Scasonal distribution of adult Hadenoecus subterraneus in Marion Avenue. deep within Mammoth Cave. Ky .............................102 21 Seasonal distribution of copulating pairs of Hadenoecus subterraneus seen in Marion Avenue. deep within Mammoth Cave. Ky .................. 106 22 Seasonal distribution of first-instar nymphs of Hadenoecus subterraneus in defined areas of Mammoth Cave. Ky ...........................106 23 Hind tibia and ovipositor lengths of adult individuals of Hadenoecus subterraneus from predated and nonpredated populations ......................113 24 Mean ovipositor lcngth versus mean hind tibia length in adults of predated and nonpredated populations of Hadenoecus subterraneus ................ 114 25 Mean ovipositor lcngth versus mean hind tibia length in adults of predated and nonprcdatcd populations of Hadenoecus cumbalandicus ............... 115 PLATES (Plates 11-V follow p . 124) I Copulating pair of Ifadenoecus subterraneus. Mammoth Cave. Ky I1 Male terminalia of Hadenoecini in distodorsal aspect 111 Male subgcnital plates of Hadenoecini in distovcntral aspect IV Scanning electron microgrzaphs of structures of Hadenoecini V Electrophoretic starch gel stained for Tetrazolium Oxidase. from samples of Hadenoecus cumberlandicus and H . subterraneus ABSTRACT The rhaphidophorid tribe Hadenoecini, restricted to the eastern United States, comprises two genera-Euhadenoecus gen. nov., with four species of which three are new, and Hadenoecus Scudder, with five species of which four are new. Two of the species of Euhadenoe- cus are forest-dwellers; all the other members of the tribe are obligatory cavernicoles that reproduce only in caves but emerge from them at night to feed. In parts I and I1 of this study, by T. H. Hubbell, the tribe, genera and species are described, and their distribution, phylo- geny and evolutionary history are discussed. Part 111, by R. hl. Norton, deals with the life history, behavior and ecology of the two best-known species, H. subterraneus and H. cumberlandicus. INTRODUCTION Among the larger, more conspicuous and more often observed of the insects that inhabit caves in the eastern United States are the pale, spidery "cave-crickets"' of the rhaphidophorid subfamily Dolicho- podinae, tribe Hadenoecini. Commonly seen clinging singly or in groups to the cavern walls and ceilings, they are wingless, with compact bodies and very long, slender legs and antennae, and have small but functional dark eyes. The guides in commercial caves often refer to them as "white cave-crickets" to distinguish them from the more robust, shorter-legged, brown-maculate "camel-crickets" of the genus Ceuthophilus that frequent cave entrances and the twilight zones. Actually they are not white; adults are pale yellowish or yellowish-
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