Orthoptera of the Nevada Test Site

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Brigham Young University Science Bulletin, Biological Series

Volume 4 Number 3 Article 1

9-1964

Andrew H. Barnum Dixie College, St. George, Utah

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ORTHOPTERA OF THE NEVADA TEST SITE

ANDREW H. BARNUM

BIOLOGICAL SERIES — VOLUME IV, NUMBER 3

SEPTEMBER, 1964

LIBRARY

Nuv 2 5 1964 HARVARD UNIVERSITY

Brigham Young University

Science Bulletin

ORTHOPTERA OF THE NEVADA TEST SITE

ANDREW H. BARNUM

BIOLOGICAL SERIES — VOLUME IV, NUMBER 3

SEPTEMBER, 1964 FOREWORD

This is another of a series of major pubhcations on desert ecology resulting from studies at the Nevada Test Site by the Brigham Young University Depart- ment of Zoology and Entomology in cooperation with the United States Atomic Energy Commission. Although some of the studies are the result of independent investigations by specialists who are not on our departmental staff, they are part of the major project initiated cooperatively by B.Y.U. and the A. E.G. to determine the effect of nuclear detonations on the native animals of the Nevada Test Site.

Dorald M. Allred and D Elden Beck Project Supervisors TABLE OF CONTENTS

INTRODUCTION 1

HISTORICAL REVIEW 1 ACKNOWLEDGMENTS 2

METHODS OF STUDY 2 Frequency and Abundance 3 Study of Individual Species 4

DESCRIPTION OF THE AREA 5 Location 5 Physiography 5 Vegetation 5 Regular Collecting Areas 6 Miscellaneous Collecting Areas 13

ENVIRONMENTAL RELATIONSHIPS OF THE ORTHOPTERA 15

CLASSIFICATION OF THE ORTHOPTERA 15 External Anatomy 17 Notes on Development 20

ANNOTATED LIST OF THE ORTHOPTERA AT THE NEVADA TEST SITE 20 Use of the Keys 20 Suborder Caelifera 22 SuperfamUy Acridc .dea 22 Family Tetri^ Jae, Subfamily Tetriginae 23 Family Eumastacidae, Subfamily Morseinae 24 Morsea californica piute Rehn and Grant 25

Family Tanaoceridae ' 27 Tanaocerus koebelei koehelei Bruner 28 Family Acrididae 29 Subfamily Romaleinae 31 Dracotettix plutonius Bruner 32 Tytthotyle maculata (Bruner) 33 Subfamily Cyrtacanthacridinae 35

Aeolopides tenuipennis ( Scudder) 37

Aeoloplides minor ( Bruner) 38 Hesperotettix viridis viridis (Thomas) 41 Hesperotettix viridis nevadensis Morse 42 Hesperotettix viridis terminus Hebard 42

Melanoplus aridus ( Scudder) 45 Melanoplus complanatipes canonicus Scudder 45 Poecilotettix sanguineus Scudder 47 Subfamily Acridinae 48

Eremiacris pallida ( Bruner) 51 Bootettix punctatus (Scudder) 52 Amphitornus coloradus ornatus McNeill 54 Cordillacris occipitalis cinera (Bruner) 55 Ageneotettix deoruni deorum (Scudder) 56 Fsoloessa delicatula delicatula (Scudder) 57 Ligurotettix coquilletti cantator Rehm 58 Arphia conspersa Scudder 60 Xanthippus corallipes. corallipes (Haldeman) 61 Leprus glaucipennis Scudder 63 Derotmema delicatulum Scudder 64 Page

Mexiohrcgmu impcxuvi Hchn 65 Trimerotropvi hitohalu Rfhn and Hebard 67 TrimewtropiH jontana Thomas 68 TriniL-rolropK alharcns McNeill 69 Trinienttrofm strcnua McNeill 70 TrimcrotTopli paUulipennui pallidipennis (Burmeister) 72 Trimcrotropis iiwon-spicua BniiiiT 74 Trimcrotropix ctjancipcimvs Bniner _ 75 Trimcrotropui sparm Thomas 76 Anconia integra SciicldcT 77

CiboLicris purviccps uridus ( Bruner) 78 Superfamily Tridactyloidca, Family Tridactylidae 81 SubordtT Ensifera, Supcrfiimily Tettigonioidea 81 Family Tettigoniidao 82 Subfamily Phaneropterinae 82 Insara elegans nuiculnta Bamum, new subspecies 83 Insara covilleae Rchn and Hebard 84

Arethaca brevicauila ( Scudder) 85 Subfamily 'lettigoiiiinae - 86

Cuptuihoti's fidiginosua ( Thomas ) 87

Capnohutcs uccidentalut ( Thomas ) 88

Anoplodusa urizonensii ( Rehn ) 89 Atcloplus luteus Caudell 90 Family Gryllacrididae 91 Subfamily Stcnopelmatinae 91 Stcnopelmattts fuscus Haldeman 91 Subfamily Rhapliidophorinae 94 Ccuthophilus nc'vadensis Bamum, new species 96 Ccuthophilus deserlicola Banuim, new species 100 Ceutliophitus hehardi Hubbell 103 Ccuthophilus fossor Hul)l)ell 105 Ccuthophilus Uinwllipes Relm 107 Pristoceuthophilus pacificus (Thomas) 109 Family Gnllidao 112 Subfamily Mogoplistinae 113 Cycloptilum comprchendens jortior Hebard 113 Subfamily Gryllinae 113 Acheta dssiniilis Fabricius _ 114 Subfamily OEcanthinae 115

OEcanthus californicus califamicus Saussure 115 OEccmtliu.s nigriioniis quadripunctatus Beutenmuller 116 Subfamily Myrmccophilinae 116

Mijrinccophilu tniniiti Schimmer 116

Stilx>r(ier Hhasmatoptera, Superfamily Phasmatoidea, Family Phasmatidae 118 Subfamily Pachymorphinae 119 Ptiwhdcillus hcsperus Hebard 119 Subfamily Heteronemiinae 119 Pseudoscrmijle stramineus (Scudder) 119 Sulx)rder DiL-tyoptera 121 Superf;miilv Mantodea, Family Manteidae 121 Subfamily Amelinae 121 IMuncutria minor (Scudder) 122 Subfamily Manteinae 122 StagmonuinILt californicus Rehn and Hebard 123 Superfamily Blattodea, Family Polyphagidae 124 Arenivaga crratica Rehn 125 Arenivaga apacha (Saussure) 126

EremobUitta subdiaphuna ( Scudder ) 126 SUMMARY AND CONCLUSIONS 127 LITERATURE CITED 128 APPENDICES

I. Depositories of Specimens Collected in This Study 130

II. Notes on Collecting and Preserving Orthoptera 130

III. Glossan^ 131

LIST OF ILLUSTRATIONS

Page

Plate I. Morphology of a typical acridid, Trimerutropis pallidipennis pallidipennis 19

Figure

1. Trimerotropis pallidipennis pallidipennis, female, caudal appendage 21

2. T. pallidipennis pallidipennis, female, pronotum, lateral view 21

3. T. pallidipennis pallidipennis, female, pro.\imal abdomen showing auditory appartus, lateral view 21

4. T. pallidipennis pallidipennis, female, caudal tarsus, lateral view 21

5. Capnohotes fuliginosus, female, distal femur and proximal tibia, showing auditory apparatus, lateral view 21

6. C. fuliginosus, female, caudal tarsus, lateral view 21

7. Trimerotropis pallidipennis pallidipennis, female, head, facial view showing insertion of antennae 22

8. Tridactylus apicalis, male, head, facial view, showing insertion of antennae 22

9. T. apicalis, male, cephalic appendage 22 10. Paratettix rnexicanus, female, distal tibia and tarsus of mesothoracic appendage, lateral view 23 11. P. rnexicanus, female, pronotum ;md tegmen, lateral view 23 12. Trimerotropis pallidipennis pallidipennis, female, distal segment of caudal tarsus showing claws and aroliiun 23 13. Morsea califomica piute, female, head and pronotum, lateral view 23 14. M. californica piute, female, head, facial view 23 15. Tanaocerus koebelei hoebelei, female, head, facial view 23 16. Morsea califomica piute, female, antenna, lateral view 25 17. M. califomica piute, male, cercus, lateral view 25 18. M. califomica piute, male, apex of abdomen, dorso-caudal view 25 19. Tytthottjle inaculatu, male, distal Ubia and pro.ximal tarsu.s of caudal appendage, lateral view 31 20. Dracotettix plutonius, female, distal tibia and proximal tarsus of caudal appendage, lateral view 31 21. Melanoplus complanatipes canonicus, female, prostemal spine, cephalic view 31 22. Dracotettix plutonius, female, head and pronotum, lateral view 32 23. Tytthotyle maculata, female, head and pronotum, lateral view 32 24. Hesperotettix viridis nevadensis, male, apex of abdomen, lateral view 36 25. Melanoplus complanatipes canonicus, male, apex of abdomen, lateral view 36 26. Aeoloplides tenuipennis, male, caudal femur, lateral view 36 27. A. minor, male, caudal femur, lateral view 36 28. Hesperotettix viridis termius, male, head, pronotum, and tegmina, dorso-lateral view 41 29. H. viridis viridis, male, head, pronotum, and tegmina, dorso-lateral view 41 30. //. viridis nevadensis, male, head, pronotum, and tegmina, dorso-lateral view 41 31. Melanoplus aridus, male, head, pronotum, and tegmina, dorso-lateral view 44 32. M. aridus, male, apex of abdomen, dorso-lateral view 44 33. M. complanatipes canonicus, male, apex of abdomen, dorso-lateral view 44 34. Amphitornus coloradus ornatus, male, head and pronotum, lateral view 49 35. A. coloradus ornatus, male, head and pronotum, dorsal view 49 36. Psoloessa delicatula delicatula, male, head and pronotum, lateral view 49 37. P. delicatula delicatula, male, head and pronotum, dorsal view 49 38. Eremiacris pallida, male, head and pronotum, dorsal view 49 Figurt- Page

39. Booti'tlix imnclatu.t. inali-. hi-al and proiiotum, lateral view 49 40. Cimlilliirris occipilalis ciiurea, male, liead and proiiotum, dorsal view 49 41. Ligiirotcltix coijuilletti cantator, male, dLstal tibia and tarsu.s of caudal appendage showing internal apical spines, lateral view 49 42. Ageneotettix deorum deorum. male, head and pronotum, dorso-lateral view 49 43. Ligurotcttix coquitletti cantiilor, male, head, pronotum, and tegmen, lateral view 49 44. CiiioUicris paniccps tiridus, male, head and pronotum, dorsal view 50 45. /Xnconiu integra, male, head and pronotum, dorsal view 50 46 Arphui compersa, male, mctaslenium and proximal abdominal stemites, ventral view 50 47. A. conspcTsa, male, pronotum, lateral view 50 48. Trimerotropis pallidipennis pallidipennis, male, metastemum and proximal abdominal stemites, ventral view 50 49. Xanthippws coraltipes corullipes, male, pronotum, lateral view 50 50. Leprux glaucipennii, male, pronotum, lateral view 50 51. Xanthijiini.s corullipes corullipes. male, pronotum. dorsal view 50 52. Lcprus gLiucipennis, male, pronotum, dorsal view 50 53. Mestohregmu inipexum, male, pronotum, lateral view 50 54. Derotmema delicutulum, male, pronotum, lateral view 50 55. Trimerotropis pullidipennit pullidipennis, female, pronotum, lateral view 50 56. T. strenua. male, pronotum, lateral view 50 57. Derotnwma delicutulum, male, pronotum, dorsal view 50 58. Trimerotropis bilohata, male, pronotum, lateral view 67 59. T. strenuu, male, pronotum, lateral view 67 60. Anoplodusa urizonensis, female, caudal tarsus, lateral view 82 61. Acheta aisimilis, female, caudal tarsus, lateral view 82

62. A. ussiniilis, female, head, pronotum, tegmina, dorso-lateral view 82 63. Anoplodusa arizonenxui, male, cephalic tibia showing auditory apparatus, lateral view 82 64. Captwbotes fuliginosu-i, female, caudal tarsus, lateral view 82 65. Itisara elegans nuicuUitu, female allotype, tegmen and wing 83 66. Arethuea brevicauda, male, tegmen and wing 83

67. A. brevicauda, male, modification of first abdominal tergite, cephalo-lateral view 83 68. Insara elegans maculata, female allotype, pronotum, dorso-lateral view 83

69. /. coiilleae, male, pronotum, lateral view 83

70. /. covilleue, male, pronotum and proximal tegmina showing striilulating mtxhanism, dorsal view 83

71. 7. eleguns maculata, female allotype, apex of abdomen and ovipositor, lateral view 84

72. /. elegans maculata, male holotype, pronotum and proximal tegmina showing stridulating mechanism, dorsal view 84

73. I. elegans maculata, male holotype, apex of abdomen, dorso-lateral view 84

74. /. covilleae, female, apex of abdomen and ovipositor, lateral view 85

75. /. covilleae, male, apex of abdomen, dorso-lateral view 85 76. Arethaea brevicauda, male, pronotum and proximal tegmina showing stridulating mechanism, dorsal view.. 86 77. Ateloplus luteus, male, pronotum and tegmina, dorsal view 87 78. Capnobotes fuligirwsus, male, prostenuim showing spines, cephalo-ventral view 87 79. C. fuliginosus, female, caudal femur, lateral view 87 80. C. fuliginosus, male, apex of abdomen, dorso-lateral view 87 81. Stenopelmatus fuscus, male, head, facial view 91

82. Ceuthophilus fossor, female, head, facial view 91

8.3. C. fossor, female, cephalic coxa showing spine 91 84. Pristoceutltophilus pucificus, male, apex of abdomen, dor.so-lateral view 94

8.5. /'. pacificus, female, distal valves of ovipositor, lateral view 94

8fi. P. pacificus. male, distal segment of caudal tarsus showing claws and sen.sory setae 94 87. Ceuthophilus lamellipes, female, distal valves of ovipositor, lateral view 94 88. C. tu'vadensis, male holotype, subgenital plate, caudal view 95

89. C fiissor, male, subgenital plate, caudal view 95 90. C. fossor, male, cephalic margin of cephalic femur, lateral view 95 91. C. heburdi. male, cephahc margin of cephalic femur, lateral view 95 92. C. lamellipes. male, cephalic margin of caudal femur. Literal view 95 Figure Page

93. C. lamellipcs, female, distal end of cephalic margin of caudal femur, lateral view 95 94. C. deserticola, male holotype, caudal tarsus, lateral view 95 95. C. deserticola, male holotype, distal abdominal tergites, dorsal view 95 96. C. deserticola, male holotype, subgenital plate, caudal view 95 97. C. hebardi, male, caudal tarsus, lateral view 95 98. C. hebardi, male, distal abdominal tergites, dorsal view 95 99. C. hebardi, male, subgenital plate, caudal view 95 100. Acheta assimilis, female, caudal tibia and tarsus, lateral view 112 101. Cycloptilum comprehendens fortior, male, caudal tibia and tarsus, lateral view 112 102. Myrmecophila manni, male, caudal appendage, lateral view 112

103. OEcanthus califoniicus califamicus , male, caudal appendage, lateral view 112

104. OE. c. californicus, male, detail of caudal tibia, lateral view 112

105. OE. c. californicus, male, proximal antennal segments, cephalic view 115 106. OE. tiigricornis quadripunctatus, male, proximal antennal segments, cephaUc view 115 107. Arenivaga erraticu, female, caudal femui showing distal spine, lateral view 124 108. A. erratica, male, concealed genital structures 125 109. A. apacha, male, concealed genital structures 125 110. Ceuthophilus nevadensis, male parat\pe, epiphallus 98 111. C. nevadensis, female allotvpe, distal valves of ovipositor, lateral view 98 112. C. nevadensis, female paratype, distal valves of ovipositor, lateral view 98 113. C. nevadensis, male holotype, apex of abdomen, lateral view 98 114. C. nevadensis, male holotvpe, subgenital plate, caudal view 98 115. C nevadensis, male holotype, distal epiproct 98 116. C. nevadensis, male holotype, abdominal tergites, dorsal view 98

1 17. C. nevadensis, male holotype, cephaUc margin of cephalic femur, lateral view 98 118. C. nevadensis, male paratype, cephaFic margin of cephalic femur, lateral view 98 119. C. nevadensis, male holotype, caudal femur, lateral view 98 120. C. nevadensis, male holotype, caudal tarsus, lateral view 98 121. C. deserticola, male paratype, epiphallus 102 122. C. deserticola, female allotype, distal valves of ovipositor, lateral view 102 123. C. deserticola, male holotype, apex of abdomen, lateral view 102 124. C. deserticola, male holotvpe, subgenital plate, caudal view .102 125. C deserticola, male holotype, epiproct 102 126. C. deserticola, male holotype, distal abdominal tergites, dorsal view 102 127. C. deserticola, male holotype, cephalic margin of cephalic femur, lateral view 102 128. C. deserticola, male holotype, caudal femur, lateral view 102 129. C. deserticola, male holotype, caudal tarsus, lateral view 102 130. C. hebardi, male, epiphallus 104 131. C. hebardi, female, distal valves of ovipositor, lateral view 104 132. C. hebardi, male, apex of abdomen, lateral view 104 133. C. hebardi. male, subgenital plate, caudal view 104 134. C. hebardi, male, distal abdominal tergites, dorsal view 104 135. C. hebardi, male, cephalic margin of cephalic femur, lateral view 104 136. C. hebardi, male, caudal femur, lateral view 104 137. C. hebardi, male, caudal tarsus, lateral view 104 138. C. fossor, male, epiphallus 106 139. C. fossor, female, distal valves of ovipositor, lateral view 106 140. C. fossor, female, distal valves of ovipositor, lateral view 106 141. C. fossor, male, apex of abdomen, lateral view 106 141. C fossor, male, subgenital plate, caudal view 106 143. C. fossor, male, epiproct 106 144. C. fossor, male, distal abdominal tergites, dorsal view 106 145. C. fossor, male, cephahc margin of cephalic femur, lateral view 106 146. C. fossor, male, caudal femur, lateral view 106 147. C. fossor, male, caudal tarsus, lateral view 106 148. C. lamellipes, male, epiphallus 108 149. C. lamellipcs, female, distal valves of ovipositor, lateral view 108 ) 1

Figure Page

150. C. lamfllipes, mail-, apex i)( alxlomrii. latiTal vww 108 151. C. knwUipes, male, suhgeiiital plate, caudal view 108 152. C. lamellipcs, male, cliMal alHlominal tergites, dorsal view 108 153. C. Uimcllipcs. male, cephalic margin of cephalic femur, lateral view 108 154. C. lamellipcs, male, caudal appi-ndage, lateral view 108 155. PrLstoccutliophilitx pucificus, male, epiphailu.s 11

156. P. pacifictis. female, di.stal valves of ovipo.sitor, lateral view Ill

157. P. iMcificus, male, apex of alKlomcii, lateral view Ill 158. P. pucificus, male, subgeiiital plate, caudal view ill

1.59. P. /JO(i/ic-ii,v, male, di.stal ahdominal tergites, dorsal view Ill

160. /'. pacifictis, male, caud.il Icuiiir. Literal view Ill

LIST OK DIAC:iUMS AM) lABLES Page

Diagram 1. Tvpical quadrate .showing position of can traps 7

Table

1. Sea.sonal distribution of the Orthoptera characteristics of the SaLmla hai)itat (Study IF) 7

2. Seasonal distrii)ution of the Orthoptera characteristic of the Grayia-Lijcium habitat (Studies IB, IG, 4A, 5E) 8

3. Seasonal distribution of the Orthoptera characteristic of the Larrea-Franseria habitat (Studies 5A, 5CQ) 9

4. Seasonal distribution of the Orthoptera characteristic of the Atriplcx-Kochia habitat (Study 6A) 10

5. Seasonal distribution of the Ortlioptera characteristic of the Culcogyne haliitat ( Stuches lOD, TA 11

6. Seasonal distribution of the Orthoptera characteristic of the piiiyon-juniper habitat (Studies 12A, 12E) .. 12

7. SeiLsonal distriI)ution of the Orthoptera characteristic of Cane Springs (Study CM) 13

8. Seasonal ihstribution of the Orthoptera 16

9. Size variation of Morsea californica piute 25

10. Size variation of Tanaoccnis koebelei koehelei 28 11. Size variation of Dracotetlix plutonius 32

12. Size variation of Tytthotyle maculata 34 13. Size variation of Aeoloplitlcs tenuipennis 37 14. Size variation of Aeoloplidcs minor 39 15. Size variation of Hesperotcttix liridis viridis 41 16. Size variation of Ilcsperotcttix viridis nevadensis 42

17. Size variation of Ilcsperotcttix liridis tcrmimis 44 18. Size variation of Mcltinoplus aridus 45 19. Size variation of Meltnwjtlus covipUnuitipcs canonicus 46 20. Size variation of Poecilotettix sanguineus 47 21. Size variation of Eremiacris pallida 51 22. Size variation of Bootettix punctatus 52 23. Size variation of Amphitornus coloradus orantus 54 24. Size variation of Cordillacris occipitais cinerca 55 25. Size variation of Ageneotcttix deorum deorum 56 26. Size variation of Psolocssa deliratuUi dcUccituhi 57 27. Size variation of Ligurotettix coijuilUlli innldtor 58 28. Size variation of Arphia conspcrsa 60 29. Size variation of Xanthippus coridlipes corallipes 62 30. Size variation of Lepras gluucipcnnis 63 31. Size variation of Derotmema delicatulum 64 32. Size variation of Mestohrcgnta impcxum 65 33. Size variation of Trimerotropus hilohata 67 34. Size variation of Trimerotropis fontnna 68 35. Size variation of Trimerotropis albescens 69 36. Size variation of Trimerotropis strenua 70

.37. Size variation of Triincrotro)ii\ pallidipcunis pallidipcnnis 72 Table Page

38. Seasonal distribution of Trimcrotropis pallidipcnnis pallidipennis 74 39. Size variation of Trimerotropis inconspiciia 74 40. Size variation of Trimerotropis cyaneipennis 75 41. Size variation of Trimerotropis sparsa 76 42. Size variation of Arwonia Integra 77 43. Size variation of Cibolacris parviceps aridus 78 44. Seasonal distribution of Cibolacris parviceps aridus 81 45. Size variation of Insara covilleae 83 46. Size variation of Arethaea brevicauda 86 47. Size variation of Capnobotes fuliginosus 87 48. Measurements of Capnobotes occidentalis 88 49. Size variation of Anoplodusa arizonensis 89 50. Size variation of Ateloplus luteus 90 51. Size variation of Stenopelmatus fuscus 93 52. Size variation of Ceuthophilus nevadensis 99 53. Size variation of Ceuthophilus deserticola 102 54. Seasonal distribution of Ceuthophilus deserticola 103 55. Size variation of Ceuthophilus hebardi 104 56. Size variation of Ceuthophilus fossor 105 57. Seasonal distribution of Ceuthophilus fossor 106 58. Size variation of Ceuthophilus lamellipes 107 59. Seasonal distribution of Ceuthophilus lamellipes 109 60. Size variation of Pristoceuthophius pacificus Ill 61. Size variation of Cycloptilum comprehendens fortior 113 62. Measurements of OEcanthus nigricornis quadripun 114 63. Measurements of OEcanthus nigricornis quadripunctatus 116 64. Size variation of Myrmecophila manni 117 65. Size variation of Pseudosermyle stramineus 121 66. Size variation of Litancutria minor 122 67. Measurement of Stagmomantis califomicus 123 68. Size variation of Arenivaga erratica 125 69. Measurements of Arenivaga apacha 126 70. Size variation of Eremoblatta suhdiapluina 127

LIST OF DISTRIBUTION MAPS

Map Page

1. Morsea califomica piute 26 2. Tanaocerus koebelei koebelei 26

3. Dracotettix plutonius and Tytthotyle maculata 26 4. Aeoloplides tenuipennis and A. minor 26

5. Hcspcrotcttix viridis 43 6. Melanoplus aridus and M. complanatipes canonicus 43 7. Poecilotettix sanguineus 43 8. Eremiacris pallida 43

9. Bootettix punctatus 53 10. Amphitornus coloradus ornatus 53 11. Cordillacris occipitalis cinerea 53 12. Ageneotettix deorum deorum and Psoloessa delicatula delicatula 53 13. Ligurotettix coquilletti cantator 59 14. Arphia conspersa, Xanthippus corallipes corallipes, and Leprus glaucipennis 59 15. Derotmema delicatidum and Mestobregma impexum 59 16. Trimerotropis bilobata, T. fontana, and T. albescens 59 17. T. strenua 71 18. T. pallidipennis pallidipennis 71 Map Page

19. T. mcuiuspictm, T. cyuucipciiiiis, and T. sparsa 71 20. AncoiiUi inft'gra VI 21. CiboUicris parviceps aridus 79

22. Inxara elegatts maculata and /. covillcac 79 23. Arcthaca hrevicaiida, Capnoholes ftiliginosus, and C. occidentalis 79 24. Anoplodusa arizuncnsis and Atcloplus luteus 79

15. Steiwpc'lmatiis fuscus 92 26. Ceuthophilus nevadensis, C. deserticola, and C. hebardi 92 27. C. fossor 92 28. C. Uimcllipcs 92 29. Prviloccuthophilus pacificus 110 30. Cijcloptilum comprcnhendens forlior and Acheta assimilis 110 31. OEcatilhus ailifornicus californicus and OE. nigricornis quadripunctatus 110 32. Mijrmccophiliu muiini 110 33. Pardhacillu.s Iwspcrux and Pseudosernujle strumineus 120 34. LitancutrUi minor and Stagjuonuintis californicus 120 35. Areniiuga erraticu and A. apacha 120 36. Eremobltttta xubdiuphana 120 ORTHOPTERA OF THE NEVADA TEST SITE by

Andrew H. Bamum" Reseiirch Associate

INTRODUCTION

This study is part of a larger ecological pro- the spring of 1961. PeriocUc trips extended ject to comparatively analyze the native animals througii March, April, and May. Extensive col- at the Nevada Test Site. The objectives of this lecting was done nearly every day tliroughout study were to (1) classify the species and pro- the months of June, July, and August, when vide taxonomic keys for their differentiation, these insects were most active. Periodic collect- (2) evaluate the populations, and (3) determine ing was again resorted to through September, the seasonal and geographical distributions of October, and November, until cold weather did native Orthoptera in areas disturbed by atomic not justify a return to the test site. Other collect- explosions as compared to those in undisturbed ing was done, as indicated, by field personnel areas, both contiguous and distant. instructed in the techni(|ues of collecting during The area encompassed by the Nevada Test all months of the years that the study was in Site and covered by this report lies principally progress. in the southeastern part of Nye County and Primary emphasis was directed toward a approximating both Clark and Lincoln counties. complete systematic and ecological study of The overall study was begun in 1959 and those ground-dwelling animals which may be continued into late 1963 with tlie periodic sam- selected as indicator animals because of their pling of Orthoptera from some areas of the test distribution and abundance in manv plant com- site. The use of special sunken can traps instru- munities throughout the test site. mented tlie collecting of ground-inhabiting species. These traps were establisiied in tran- Analysis of data was facilitated by an IBM sects or quadrates according to standardized punch card system. Field data were recorded techni(jues. In addition, thorough collecting on special fonns and were transferred to IBM was done at intervals bv field personnel. punch cards. The Brigham Young University The author began organized collecting at Computer Research Center analyzed tlie project the test site as soon as the weather pennitted in results with an IBM 650 Computer.

HISTORICAL REVIEW

The taxonomy and distribution of the Ameri- treatment of Orthoptera than any other indi- can Orthoptera are actually well known in com- vidual in the nineteenth century. He not only parison with other insect orders. The Orthop- named many new species, but revised many of tera of die Western United States, however, are the recognized groups into a unifonn order. still imperfectly known. The actual collecting of The first quarter of the present century was Nevada Orthoptera began in the early history dominated bv James A. C. Relin and Morgan of entomology when workers of the geological Hebard, both representing the Philadelphia and geographical surveys entered the territory Academy of Sciences. On a number of occasions and made limited collections of the more con- they entered the state and collected intensively, spicuous species. Of primary interest to tlicse particularly in the southern sections, as well as collections and the subsequent publication of collecting extensively throughout southwestern the infonnation were Cyrus Thomas and Lawr- United States. Not only did they build up a ence Bruner. Although he was never in the state, large collection of Orthoptera from die south- Samuel II. Scudder did more for the systematic west, which included a number of new species

°Di.xie College, St. George, Utah. BiiK.iiAM V<>i\<; L'mm.iimiv ScitNc t Bulletin

I'riinarilv on tin; basis (»l uliat liatl alrcads the Nevada Orthoptera because thev covered traiLs|)irccl. Dr. Ira La lii\('rs of liii' I'liiscrsitN- ol areas far distant Irorn the present study site, and NtAaiia t-nttTcd into a shidy of tlic Nevada very few species overlap into this area. The

Ortlioptcra, winch resulted in "A Synopsis of author is grateful to these individuals for pro-

Nevada Orllioplera" in uliieli lii' eonlribiited viding a basis upon which the present study is eonsiderahU; original information on tliu ecology made. of the Nevada Orthoptera hut little on the Other recent workers have attempted to

sysleniaties of the group. study till- ecology of some groups of Orthoptera (.'ontrary to the extensive work that has hi-en bv controlled laboratorv experiments, but tlie

done in systeniaties, there have been relatively ecological behavior of any s[x;cies differs vs ithin

few competent studies made of the ecolog)', the its own range, and is far different from anv so- life histories and habits of North American Or- called "controlled" laboratory situation. .Nothing thoptera. The earlier works in ecologv, such as of a laboratorv nature could be substituted for made bv \estal (1913), Ibibbeli (1922), Stro- ade(|nate field studies. '!"he present report con- heckcr ('l937), I.scly (1937, 1938), and L'rquhart tains field oliservations anil studies of all of the (1941), were important and served as a basis for species here recorded.

ACKNOWLEDGMENTS

Any scientific stiidv represents the combined thor in the field, and contributing to the effec- efforts of manv individuals and groups who have tiveness of collecting: C>live Jorgensen, field contributed to the success of the study, .\ppre- director; D. I'"lmer Johnson and .Merlin Killpack.

ciation is therefore extended to those individuals consultants antl specialists; (Carl Ingersoll, Mor- for the assistance renderi-d. ris (ioates. and Ci-rald Hichards. field biologists; The studv was made possible l)v a grant to and espcciailv lor the long hours of collecting Hrigham Young I'niversitv from the .Atomic witii and driving lor the author, field biologists I'jiergy (Commission on (-'ontract No. AT(ll-ll \\ illis .\. I'ackluini. Arthur .\nclerson. and .\mold

7S6. (iratelul acknovv letlgment is made to those OrtoiL individuals making this grant possible and par- The autlior is inrlJur indebted io Dr. .Ashlev (' ticularh to Dr. Dorald M. .-Mired, project of- H. (iurnev oi tlic . S. .National Museum for ficer, and Dr. D Klden Heck, associate investi- verifving and classilving certain six^cies of Or- gator in charge of invertebrate research, both of thoptera; and Dr. .Xrthur ('. (]ole, project con- the Department of Zoologv and iMitomologv ot sultant and specialist, lor tiie determination of Hrigham Young I'niversitv, without whom tliis several species of ants. research would not have been possible. -Appri'ciation is extended to those other indi- The following personnel contributed greatlv V iduals who contributed in anv wav to the com- bv collecting s[)ccimens, ai-companving tlic au- ])ictii)n of tiie |)rcsi'iit research.

Ml, 1 HODS OF SITDY

\\ lull the autlior began an Dii-silc iiivcsliga- The cans were' cmptictl regularly three times

tioii of the Orthoptera of the Nevada Test Site per wei-k in all ari>as, and the invertebrates col- in 1961. certain (|uailrate and transect studies lecled were placed in 70V alcohol. Major sam- had alreadv been established. A reconnaissance pling areas were run continuously over a onc-

of the test siti- was made to determine the most year period so that a total .seas(mal sampling ideal habitats lor Orthoptera and lo check addi- lould be achieved. .Ml the Orthoptera thus col- tional areas that might be sam|)led. Special I'cted have beiMi submitted to the author for sunken can traps ( Allred,

the Orthoptera collected were of minor signifi- tically at the tips of branches of shrubs, others cance to the overall study. near the groimd in the shade, and some on the Several of the field biologists carried collect- ground imderneath the vegetation. Very few ing nets and kept accurate information on the specimens can be foimd on die ground in full specimens captured. sun during the hot summer hours. To effect a systematic study, the author visit- Desert vegetation is topically that of scat- ed current study plots on every trip to the test tered plants, and it is possible to check an area site during die spring and autumn, and during in a siiort time by rapid walking between plants the summer visited study plots at least twice a to observe or capture die strong flyers, and by week, generally three times a week. Because of systematic visual or mechanical examination of the many miles between some study plots this the plants. systematic collecting occupied at least half the Night collecting was chiefly visual with time; the remainder of the time was spent in flashlights or lantenis and the use of aerial nets collecting from special areas, in between or to capture specimens. No systematic night light adjacent to the study plots, and in night obser- collecting was maintained, although some sam- vations and collecting. Because of the e.xtensive pling was done with black (ultra violet) light. area, some study plots were visited only once by Baits of rolled oats and/or molasses in can the author during the entire season. traps or scattered upon the ground were tried in The collecting method most generally em- some areas. No special advantage could be de- ployed, in addition to the special can traps, was termined, however, inasmuch as the cans fre- use of a sweeping net on shrubbery and other cjuently contained mice and other rodents or vegetation, and an aerial net to capture the other predaceous animals. As a matter of fact, great majority of specimens, as most of the as evidenced by parts of bodies, many ground- orthopteran inhabitants of the southwest deserts dwelling specimens captured in the traps were are strong fliers. A great deal of difficulty was consiuned b\- these animals, notably grasshopper encountered in sweeping desert plants because mice {Om/chomys) and less frequently by shrews

of their spinose nature. When these plants were ( S'orc.v ) .Wherever these rodents occurred in the sampled, an observation was first undertaken. cans, few or no arthropods were present. Some The entire shrub or plant was carefully examined lizards and predaceous arthropods, especially and notes taken on any orthopteran present. tenebrionid beetles and scorpions, were respon- Periodically an entire shrub was torn apart to sible for the destruction of large numbers of reveal the presence of specimens. Many insects s|iecimens. not visible because of their concealing colora- Notes were made, wliere\er possible, on the tion and patterns were thus captured. songs of the xarious species, both by day and During the hot summer months many of the night, though this is a minor contribution of desert shrubs lose their leaves. The most thor- the o\'erall study because of the seemingly in- ough, accurate, and speedy collection from active nature of so many of the desert species these slirubs was by trampling. Each shrub was and the absence from the test site of many trampled systematically, spirally from the out- stridulating nocturnal forms. side to the inside. It is believed that very few More than 8,000 specimens, both nymphs and orthopterans escaped when such methods were adults, were collected and preserved in the employed. An aerial net was used to captiu'e course of the investigation. As noted earlier, those specimens trying to escape. specimens collected from the can traps were Many of the data recorded are sight records. placed directly into separate vials of 10% alcohol. If all the observed specimens had been captured Some of the specimens of the most common there would have been insufficient time to ex- species were captured, examined, and later re- amine all the areas. leased in the same area. No special sweeping data were maintained with reference to length of stroke, distance from Frequency and Abundance

the ground, speed, etc. Most species of the No statistical frequency and abundance (i.e., desert are so different that sweeping methods numbers of specimens per sweep) was attempted must be adjusted to the habits of the various because of the general scarcity of orthopteran forms to achieve maximum effectiveness. forms at the test site. Some visual observations The height at which some species occur on on abimdance were made.

vegetation is variable according to atmospheric It must be emphasized that the present dis-

conditions. During the hottest hours of the sum- cussion is relative to the Orthoptera of the mer day many species are found characteris- Nevada Test Site (mlv during the years when the Bmicham Vol'nc Univebsity Science Bulletin study was in progress. Tlie same six;cies or com- a series and submit descriptions and drawings parative luiiiihers of s[x;ciinens may not be pres- on only one specimen without recognizing \ari- ent in any otlier year, before or after tlu- testing ation within the groiij). .Many new sjx-cies have program was begun. Cyclic appearance of cer- been describi'tl from unii|ue t\jx's. and in many tain species must he taken into consitleration, instances this has resulted in a long list of con- and tlie same specii-s tliat were numerous during fusing synonyms. the recorded period may actually be less numerous than some otlier sjK'cies at some otlier Study of Individual Species time. Each species represented by a series of speci- Nearly all grasshoppers fluctuate in numbers mens was studied for variability, and notes and from year to year. One year they may be very measurements in millimeters were made of rep- numerous, whereas the next year few will ap- resentative specimens of both sexes. Measure- pear. Such insects occur in small numbers for a ments were made with a standard micrometer in year or two, gradually increase, and when a a binocular microscope. The length of the body favorable season occurs appear in enormous and tegmen on large specimens was detennined numbers and may cause great damage, only to by metric callipers. disappear again for several years. The most accurate sjwcies analysis should be The reason for this fluctuation is apparent. made upon consideration of all measurements While grasshoppers are capable of increasing given, rather than reiving on a single measure- twenty to si,\ty times in one year, their enemies ment, such as total bod\' length, as has been used and diseases are capable of increasing several in the past. .\ccordingl\', the following measure- hundred and up to thousands of times in one ments were made on the series of specimens. season. While the grasshoppers are scarce, their Length of body. The measurement was parasites have a difficult time to find the hosts, made from \'ertex to tip of ovipositor of female and, as a result, the majority of the parasites or subgeiiital plate of male, but excluding teg- perish. Then, as the grasshoppers increase in miiia and wings that extend be\()nd the tip of numbers, the few parasites left have no trouble the abdomen. Although this is of the stand- in finding them and thev, too, increase enor- one ard measurements made on Orthoptera it is mously. The year tlie grasshoppers are most nu- variable and actually less valuable than some merous is often the year in which the parasites other measurements. The female that has been increase to such an e.xtent that jiractically no oN'ipositing or copulating often has the abdomen grasslio])pers or eggs are left to produce a bnxKl abnormalK- stretched; in cases the abdo- the following year. Hut they are not present in some men is abnormallv retracted. In the male, espe- sufficient numbers to cope with the swarms of cially in some groups such as Aeolaplulcs, the grasshoppers in the \ear in uhich they are most needed. abdomen is consistently upturned, and measurements are unreliable. In such cases the measure- The weather phu's an iiii[)(>rtant part in fluc- ments are gi\en to the most posterior part of the tuation of numbers. C-old, wet weather in tiie abdomen. spring will destro\' a large number of young grassh()])pi'rs. Hot, drv weather allows all eggs Greatest depth of bodv. This measurement

to hatcli .111(1 the Noting insects to thri\e. The was not used consistcntiv. The greatest liod\' same hot dr\ NM'ather burns up the vegetation depth in nearl\- all s]ieci('s was mcasm-ed from

so that there is less tor them to iced on. nrougiit the mesosteriHim to tlie iiicdian carina of the and grasshoppers often go together, especiall)' [jronotum.

if the drought extends through sc\eral years. Length of pronotum. I lie proiiotal length In some test site areas \isited regularlv a was taken in most cases, although it \aried be- large population of robber flies, bee flies, liz- causi' of caudal prolongation. In some specimens ards and otlu'r predaceous animals were present the pronotum was iiotit cabK' aberrt'iit, probably that might have accounted for the scarcit\ ot due to dr\cl(ipinrnlal iiijurN or malfomiation. specimens. In the author's experience of colltit- Cireatesl breatllli of pronotum. The greatest ing in desert environments, the specimens wru' ])n)iiotal hreadtli (iicms in most species on the far too few at the Nevada Test Site while tlic disk ol the metazona. sfiidv was in progress. Whenever a species was clisiDNcnd in aii\ Deptli of pronotum. The measurement is area, as large a series as possible was colk'cted of importaiKi" to some groups with a high me- to show variations. Too manv inor|)liologists dian pnmotal carina, and to otliers with modi- and taxonomists fail to realize the iinportaiice of fied lateral pronotal lobes. The measurement Biological Series, Vol. 4, No. 3, Septemiseh, 1961 was from the ventral edge of the lateral lobe to yond the tegmina. In nearly all species examined the highest dorsal part, usually the median the tegmina and wings are subequal in length. carina. Length of caudal femur. Measurements on

Length of tegmen. The tegminal length is the caudal femora have not been consistently considerably variable in some groups. The reported, but may be important to Orthoptera measurement was made of the wing in resting systematics. This structure shows less variability position from the angle of the radius, media, and than other body structures. The length was costal veins in the area of the pronotum to the measured from the anterior development to the tip of the tegmen. In some cases where the pro- greatest prolongation of the genicular lobe. notum is greatly prolonged the measurement is Greatest breadth of caudal femur. This given as projecting beyond the pronotum. This measurement, with the length, shows the salta- is individually stated in the account of the torial ability of the insect. species. No measurements were made on tlie Other miscellaneous measurements were total length of the wing, but in some species a made according to the species and are included measurement is given for wings projecting be- in the account of the individual species.

DESCRIFnON OF THE AREA

Location Playa, has a small empoundcd water area of approximately two hundred scjuare feet. The Nevada Test Site is located in Nye The water at Tippipah Spring, County, Nevada, contiguous to both Lincoln northwest of Yucca Playa, is restricted to the inside of tunnel, and Clark counties. It is appro.xiinately 65 a but provides miles northwest of Las Vegas, Clark County, water for some animals that venture into tile shaded interior. White Valley, Nevada, just off U. S. Highway 95. The test Rock north of Tippipah Spring, lias site encompasses some 1000 square miles, being a tiny amount of water from one spring. In addition an area appro.xiinately 40 miles from north to there are some few areas south by approximately 25 miles from east to to the northwest with minute amounts of permanent water, and a few wells west. The present study is limited by these have built for boundaries. Most of the collecting was restricted been industrial purposes. Such an to areas immediately surrounding the numerous environment is not conducive to some orthop- access roads within the area. terans, but is more typical of the habitat of the strong flying grasshoppers. Physiography VECETAriON The obvious features of the Nevada Test of Test is typical of Site are the two playa lakes, Frenclunan and Much the Nevada Site the Lower Sonoran Life Zone. southern part Yucca, and the very gradual sloping flats sur- The is typically Desert with its Larrea- rounding these areas. Scattered throughout and Mohave Fratiscria vegetation. More typical Upper Sono- actually isolating these areas is a series of moun- the section tains, especially prominent to the northwest. ran conditions are found in northern and around the bajadas adjacent to the northern The land is typically desert and very arid, hav- Desert. third faunal ing a total precipitation of approximately five limits of the Mohave The zone represented at the test site is the Transi- inches per year, tliis occurring largely in July valleys and December, with the most arid months being tional of higher elevations. Some higher are typical of the Great Basin Desert with its October and May. The soil is very poor and highly alkaline, especially around the playas associated Artemisia. where there is an associated, hazardous desert Immediately surrounding the completely bar- pavement, the small pebbles scattered over the ren Frenchman Playa of compacted silts and surface of the earth. Immediately below the clays is a fringe area of Lijcium pallidum, the Lij- surface is a very dusty, powdery soil. These dominant plant, with some Gratjia spinosa, areas extend to the bajadas and the mostly bar- cium andeisonii, Dalea pohjadcnia, Eurotia la- ren foothills and higher elevations, variously nata, and other plants. This fringe area of covered with pinyon and juniper. Lijcium. is bordered by a mucli larger, very ex- divaricata The only permanent water is restricted to tensive area of almost pure Larrca few areas. Cane Springs, west of Frenchman with its associated Frameria dumosa, Hijmeno- KitiniiAM VotNf: U.sivEHsi-n' Science Bulletin cU'a jusciculutti, Giuijui vpinosa. Lyciuin amler- inserted map, have been detailed by .Allred, sviUi. Ephedra iicuuiLiuiiA, and Dalca pultja- Beck, and Jorgensen (1%3). ilcniu. 'Iliu Luin-a-ir(iiij,cria vegetation is con- Hix;lii.-\r tiMiiuiis upon tiic bujuda;) to tlie wn.' steep and CoLLEC-n.NC Are.\s sharp lulls and ridges with their seattered The following collecting areas were visited grasses anil otlic-i vegetational types. regularly twice to three times per week during Sepaiat'.iig Frenchniaii and Yucca playas is a the months of June, July, and .August, and txvic-e seiies of hills and ridges \sith some growths of a month during .March, .April, .\Ia\, September, Coleugtjne lumoauisinui and Yucca brevifolia. October, and November, as outlined in the

Iminediatel) to tlie north of Yucca Pla>a is an "Methods of Study" above. The type of collect- association ul AtripU \ confcrtifolUi and Kochia ing was modified to suit each particular area anwricaiui, with some Eurotia laiuita and Arte- according to the vegetation present. iniiiti spincsccn^, designated as Atriplex-Kuchia. Area 1. (Yucca Flat, northwest of Yucca The next belt of vegetation, very extensive to Playa) Some of the most intensive collecting the north and east, less extensive to the west was done in Study IB, a radiating transect of and represented by a small fringe to the south eight lines running symmetricall\- from ground between the plava and the steep hills, is a belt zero, the point directly under the [X)int of detona- of Graijui-Lijcium. The two dominant species, tion. The lines were marked IBA, IBB, IBC, Grai/ui spinosa and Eijcitim audcrsonii, are as- etc., through IBH. Thirty stations were located sociated with some EuiutUi latuita. Atriph'x cane- along lines B, D, F, and H, each station being scens, Orijzopsis Iti/mcnoidcs. ArtemiA-ia spine- 264 feet apart. A total of 24 can traps were scens, Stipa speciosii. and other plants variously open continuously from March 9 to September scattered throughout the entire belt. Through 28. 1961; and from October 9, 1961, to Febniary the Gratjia-Lijciuni, at various ground zero loca- 15, 1962; and April 3 to May 18, 1962. Tliese tions where atomic detonations have occurred, same stations were open for three days in the are extensive areas of Sulsola kali, the first plant first and third week of each month. to appear in a new succession. All plants from ground zero to a radius of ap- To the nortliwest and northeast of the Graijia- pro.ximateh' one mile have been visibly affected Lijciitiii belt is a well-deNciopcd conimiinitx- of by the explosions, the damage being less severe Coleogijne, which is the dominant flora surround- progressing from ground zero to the ends of each ing Yucca Flat and extending to the various transect. In the immediate area where the plants mountain ranges. The flora of the canyon ap- were completely destroyed there has been an proaches to the higher mesas to the north and early plant succession of Russian tliistle, Salsola west is transitional. Oak, Qucrcus gamhclli, and kali. Near the maximum radius of total plant bitterbnish, Pitishui

These biolie eoinmtinities, shown b\ the lii.s ftisciis were collected in can traps. Biological Series, \'ol. 4, No. 3, Septemueh, 1964

Study IF was establislicd as a quadrate in a centrations of Haplofxipjnis cooperi, Graijw spi- Sahola habitat near ground zero. Collecting cans nosa. Eurotia lunutu, and nciu- the end of the were arranged 75 feet apart according to Dia- transect, Artcmiski tridcnUitu, Colcogijnc ramo- gram 1. This same plan was carried throughout siisinui, Hijmciwclca fascicuUita, and Aiicmma the major ijuadrate studies. .^yincscciis.

See Table I for a coinpk'te summary of speci- Study IC;, though primarily set up as a rep- mens collected in Study IF and the SiihoUi belt tile stud)- through Cratjia-Lijcitim, consisted of of Study IB. a quadrate of one hundred can traps marked from one through ten and from tlu-ough Most of the collecting in Study IB was done A J, around stations 19 through 23, a yariable belt each set at a distance of 35 feet. Although these of SalwUi, Ort/zopsl'i hijmcnoidcs. Hyrnciwclca regularly produced large numbers of ground- dwelling Orthoptera, jasciculata. Stipa speciosa, Chnjsothamnus tisci- the specimens were only occasionally diflorus. Ephedra ncvadcn.sis, and Lijcium un- preserved. Tlie area was regularly dersonii. Beyond station 23 were \arious con- swept for Orthoptera and produced a variety of species from time to time.

Area 4. (Immediately to the north of Area 1 described abo\e) Study 4A consisted of a -?® (]uadrate of twelve can traps open continuously from September 22, 19(i(), to September 23, 1961,

and from October 10-12, 1961. This is a typical Graijui-Ltjcium habitat similar to study IB or

IC;, but with larger shrubs. Desert pavement is common on the surface. A small sandy wash through most of the study is lined primaril) with Atriplcx canesccm. host to a variety of Orthop- tera during the hot mid-day hours. During the c(K)ler parts of the day tlie insects were common- s® ly found along the gravel in the bottom of the wash. Specimens collected in the Grayia-Lycium /o® habitat of study areas IB, IG, and 4 A are summarized in Table 2.

Area 5. (Frenchman Flat, southwest of /^ Frenchman Flaya) This area consisted of three very extensively collected studies, two quadrates and one line transect, each established with Dialer. im 1. Typical quadratL' .shouini; position traps(o). can traps for the capture of ground-dwelling HlllCMAM VolM. r.MXKMSIIY SciKNt K lUl.l.KI IN Biological Sehies, Vol. 4, No. 3, Septemueh, 1964 BiiKjiAM VouNf; Univuisitv Science Bulletin Biological Series, Vol. 4, No. 3, Septemueh, 1964 11 Iiiiii:iiAM ^nrsr. UvivKusi-n- Scientk Bii.LtTiN

Most of llif Ortlioptrra were colU'itcd fniiii series of hills and ridges was one of the best habi- tlic ciistiirlx-d ari'a, study 12A. This area also con- tats for Orthoptera at the test site. Clane Springs tained niort' sMfciilcnl annuals and liionnials was situated on a north slope with drainage from and smalliT pc-rennials, tlius proNiding a more the higher sl()[)es to Frenchman l'la\a below. suitable loot! supply for some species. The water originated from a man-made tunnel The n)()st numerous collections in tliese areas and was empounded in a small reservoir. were of camel crickets—two species of Cetitho- Natural vegetation of cat-tails, Tijphd dotnin- pliilus and I'rusloccutliopliihis—as well as Sleiw- f^ciisis, filled much of tlie reservoir, while dcxk, prlnuilux and the ant-lo\ing cricket, Myniwco- Rtimcx cri.spns. and wafer-parsnip. Bcruht rntia. philti. All specii's, excc-pt the Stcu(>i)clmtilus. were were in the depressions. Nearby was an associ- more numerous in the disturbed area, probably ation of grass, Elymii.s- cinereus, large shrubs, because of the loosening of the rocks and the Atriplex (aiicscnhs, and in the small valley below large fissures in the earth, providing places to a good growth of salt grass, DiMichlui stricta. hide during the day. One species of ('culhophihis The small reservoir, itself, was open at one end, was foinid in only two areas of the eiitir(> test but nomially contained \ariable growths of algae site —at the timnel at Tippipah Spring and in and aer, October, and in tlu- Cane Springs area. This table, how- November the studies were visited less fre- ever, does not include study CBA. quently. Transecting the upj)er edge of the Cane In addition to the can collecting, the sweep- Springs area was study CB.A, a line consisting of ing of vegetation, collecting and the of the a series of sixteen can traps in a mixed vegeta- strong fliers with aerial nets, man\ hours were tion association. The transect crossed most shrubs spent in overturning loosened rocks in the dis- found on the bajadas of the Nevada Test Site turbed stud}' as well as undisturbed rocks in and cxmtinued up the steep, sparsely vegetated both studies. Many fossorial Orthoptera were slope to a low ridge. obtained in this manner. The can traps on this line transect were open Because difficulty of the of access and the continuously from March 21, 1961, to March 24, distances involved, no night collecting was done 1962. They were checked three times per week, in these studies. and each of the different plants along the tran- Table ft summarizes the residts of collecting sect was checked thoroughly for Orthoptera. Be- in piiivoii-juniper the area of studies 12.\ and cause of the ideal situation of this transect, this 12E. study was checked most thoroughly for any pos- C:ane Springs. Immediately to the west of sible relationships between Orthoptera and the Frenchman Playa, but separated from it by a host plant. ?ioLociCAL Series, \'ol. 4, No. 3, Septemiseh. 1964 -

BiuuHAM VouNC Univkhsity Science Bulletin some Atriplcx amescetts. The soil types were an lected extensivelv on the few occasions they extreiiu' eontrast: from the very dark redcHsh- were visited. Due to the inaccessible nature of hrowii soils of the surroimding areas to tlie wliite the areas, however, they were only visited several ledges and the white gravels of tlu- stream beil. times during the course of the study.

Stiid\' 12(;F was established in the approaeh .\ ([uadrate vvas established in Midvalley, to Kowieh N'alley at tlie jimetion of tliat road sludv T.\, following the patterns already out- and the liainier Mesa road. Coneerted collect- lined, but the can traps were closed except for a ing was done on several occasions, resulting in few davs during each month. On several occas- the capture of Trimcrotropis foitlaiui and several ions while thev were opi-n, however, extensive other species. The area vvas one of sagebrush and collecting was done both with sweeping and bunch grass. aerial nets, and resulted in the discovery of an Sand Dunes-Target Rock .Areas. These sand apparent isolated community of Dracotettix dunes on the test site are stationary type dunes plutonius associated with Artemisia tridentata, in a very restricted area approaching south which is dominant in the valley. The results of Rainier Mesa. The area vvas visited as frecjuently collecting are summarized in Table 5 with as possible during August through November. Coleogyne, as the sagebrush here is considered No early season collecting was done there, al- a subclimax to Coleogijne.

though it may have resulted in some species Study TCB, originally set up as a transect peculiar to that environment. The study known through the bottom of one of the small driiiii- as EC.\ was originally a transect through the age canyons from the higher elevations to Yucca dunes. Bitterbrush, Purshia gUindulosa, is pri- Flat, was another of the inaccessible areas visit- marilv associated with the dunes themselves. ed only a few times. The vegetation in the bot-

.\lso in the area is Juniperus ostcosjwrma, a tree tom of the small rock-covered canyon was pre- cholla, Opiintia sp., Atriplex canesccns. Erio- dominently Purshia glandulosa, vvitli some Atri- gonum fusciaikittiin, and along the flats many plex canesccns, Eriogonttm fasciculatum, and species of small plants. (^ucrciis gambclli. The ortliopteran Morsea lali- Five can traps were established in this study fornica was common on Piushiii and Lepras and maintained from .August 16 to September glattcipennis was found along the steeper slopes. 23, 1961. They were placed to take advantage A new subspecies of Insara elegans was also of tlie best possible movement of fossorial Ortho- collected on Purshia. ptera. Study TE, known as Tippipah Spring, con- Tlie dunes were of very light-colored sand, sisted of a man-made tunnel dug in the liillside. and the light-colored Trimcrotropus aJhcsvcns A perennial water supply was found in the pro- vvas abundant. Also collected in tliis area were tected confines of the tunnel, making it cool Me.vtohregma impcxum (the only collection for and humid. Several species of birds, rabbits, the test site), Mchinophts complatuitipcs (found and one snake were found in the tunnel. This in direct opposition to the habitat of very heavy was the type locality of a new species of Ceittho- vegetation of Cane Springs), and Morsea calif philus. The onlv other place this species was ornica (associated with Pttrshui). found, as discussed previously, was on Rainier Study ECB, originally a line transect, was -Mesa associated with the cracks and fissures of also known as the Target Rock area. It consisted the disturbed area. No otliiT ortliopteran was of the same type of white sand as at the sand found in the tunnel proper, but some acridids dunes and vvas located in a long narrow valley were found in the immediate vicinity. The approach to the south end of Rainier Mesa. It vegetation was primarily sagebrush. was covered by a rather dense growth of sage- Numerous other studies were established at brush, Artemisia tridcntata, and Atriplex canc- the test site as (luadrates or transects. A con- scens on the shoulders of the graded road. Some siderable amount ol collecting was done between junipers were in the area, especially along the the major established studies. Can traps were marginal hills. Tlie area vvas not extensively col- open in inanv areas for ten-dav pericxis during lected, but some time was spent on sevejal oc- late 1961 and early 1962. These were visited casions in overturning rocks in an attempt to in the course of the study by the author or by captiu-e secretive that might anv orthopterans field personnel associated with the ecxilogy pro- otherwise captured. can traps were not be No jects, but are not outlined here because of the established in this ;irea. limitetl collivting reported. Further tximments Tippipah Spring-Midvalley Areas. These on these areas will be made with the report of areas, situated at higher elevations, were col- the individual species. Biological Series, \'ol. 4, No. 3, Septemueh, 1964 15 ENVIRONMENTAL RELATIONSHIPS OF THE ORTHOPTERA

The environmental factors which determine gardless of their comparative abundance to other the distribution of the Orthoptera are not clearly species. It is assumed that each species is as understood, particularly as they pertain to a abundant during any season as the conditions restricted area such as the Nevada Test Site. of its environment will permit, and the broadest

This is complicated by the fact that the Ortho- part of each line therefore represents the maxi- ptera as a group are not restricted to a particular mum seasonal abundance and is equal to 100 plant. Isely (1937) based his studies in Te.xas per cent of the possible population density under on the correlation of the Acrididae and the dis- the existing conditions. tribution of soil types. Cantrall (1943), on the The immediate environment of the fossorial other hand, found that in Micliigan it was easier Orthoptera is incident to that of the major plant to correlate the distribution of the Orthoptera communitv. The presence of any species is with vegetation. At the Nevada Test Site some necessarily dependent upon such conditions as of the species can be correlated with vegetation, subterranean runways and nests of rodents, fis- which is necessarily determined by soil types and sures and caves, rocks, and ground debris, or, in other physiographic features. Because of the the case of the symbiotic Mijrmecophila, the severe environment it is more meaningful to presence of an ant colony. Tliese Orthoptera, correlate the distribution of the Orthoptera, for nevertheless, are included with the major plant the most part, with the vegetation. communities if they were collected in that parti-

Tables 1 through 7 have been prepared to cular plant habitat. show the seasonal relationships of tlie various Where there is insufficient evidence to cor- specit'S of Orthoptera in the major plant habitats. relate the relative distribution or abundance of a

The earliest and latest observed records for each species, each record for that species is plotted species are shown. The broadest part of the as ".x" on the table. line in each case represents the maximum abund- A complete summary of the species of Ortho- ance. It is of the same width for all species, re- ptera collected at the test site is given in Table 8.

CLASSIFICATION OF THE ORTHOPTERA

The classification of the Orthoptera has It is exceedingly difficult to present a clear shown an evolution that parallels other insect analysis of any major group in a limited area. groups. Interpretation of morphological char- Certainly a complete revision of a group is need- acteristics by different individuals has resulted, ed to point out special relationships. This present in the past, in an emphasis of different entities study, however, is not an attempt at revision of above the rank of genus, for which there is no any group. From an evaluation of data derived established priority. Many students in the Ortho- from similar studies others can more properly ptera have elevated lesser groupings to the rank bring about the major revisions. As an aid to of order, thus creating two or more orders out such a study, this tiiesis will have been worth- of what should actually be one diversified order. while. Numerous recent studies on detailed internal The classification as used herein is modified morphology, particularly the phallic complex from Rehn and Grant ( 1961 ) to include those of the male, the spermatheca and accessory or- insects definitely found or likely to be found at gans of the female, as well as a re-evaluation of the Nevada Test Site. the external morphological characteristics has re- Order Orthoptera sulted in a more complex system of classification that actuallv simplifies an understanding to the Suborder Caelifera Orthoptera. Some of the problems that have Superfamily AcRrooiDEA existed with reference to the relationships of cer- Family Tetrigidae' tain groups are cleared up. As these studies Family Eumastacidae continue this classification will undoubtedly Family TANAOCERroAE change, as it has in the past. Family Acrididae

'Families included only as hypothetical owitij^ to th lack of extensive environmental areas at the Nevada Test Site in whieli members of the family would be foil nd. UiiK.iiAM ^'ouNG Univkhsity Sciencf Bulletin Biological Series, Vol. 4, No. .3, Septembeh. 1964 HiiiciiAM \orN(: UsivmsiTY Science Bulletin ally onlv two are prfst'nt in tlu- Ortlioptcra, and not at all transparent. In the resting position usually overlapping flii'v arr rart'ly not i-vidi-nt or absent), are typi- they repose and are and cally loiatitl, one at the niidcllc- of tlif frontal closed over the dorsum of the abdomen and the thus protecting costa, a little- bi-low the base of the antennae, more delicate hind wings, them. and t\yo paired lateral ocelli close to the upper Till- hint! wings are usiialK more niembranous. front inari^ins of tin- coniponnd eyes. transparent (sometimes brightly colored), and The appendages of the head are the antennae furnished with radiating or divergent veins. ant! the inoiithparts. The antennae are ninltiarti- When not in use they are folded like a fan un- culate. In some gronps they are nnich longer than der the tegmina. The hind wings are the active the body and consist of a \ery large ninnher of appendages where flight is possible, the tegmina small segments; in other groups a determinate being used more for balance. On the wings and number of segments, general!)' a relati\ely small on the tegmina is a complicated system of number (not more than 28) of fairly large seg- longitudinal and transverse veins, a study of ments, the length of which seldom exceeds that which venation is often important in the classifi- of the entire hotU-. Tin- shape of the antennae is cation of a species. usually filiform and e(|ually broad throughout, The limbs of the Orthoptera arc simple in but in some species they are yariously modified, structure, or variously modified, and consist of ensiform or clavate. Tlie mouth is mandibulate. coxa, trochanter, femur, tibia, and tarsus. The The i7iandibles are usually of a grinding type. coxa is often provided with spines or projections In addition to the pair of mandibles, a pair of meaningful to taxonomy. Tlie tibia, and fre- ma.\illae with fi\e-jointed palpi, a labium with (juently the femur, is armed with spines (spurs,

three-jointed palpi, and a labrum, which struc- calcars, tul)erclcs, spinules, etc. ) on the sides

ture is continuous with the clypeus, but distinct and at the apex. The tarsus is one- to five-jointed,

from it by definite sutures, are present. terminating in two claws, between wliich, in The thorax consists of three segments, the some groups, is an arolium. The proximal or first prothorax, mesothorax, and metathorax, each of joint may also bear a plantula. The front limbs which bears a pair of legs. The prothorax is are ambulatorial, or occasionally enlarged, moveable to a degree, frequently developed into pincer-iike and developed for grasping prey. The a dorsal structure which covers most or all of the middle limbs are always ambulatorial, and the mesonotum and sometimes tlie metanotum, and liind limbs are adapted for running, jumping, or in one family even most or all of the abdomen. walking. The shape and details of the pronotum are of The abdomen consists of several segments, great importance in the systematics of the Ortho- connected by feebly sclerotized elastic mem- ptera, especially its raised ridges or carinae and branes. Each segment is divided into upper and transverse furrows. The mesothorax and meta- lower halves, called tergites and sternites, re-

thorax are usually not freely moveable. The lower spectively, .'^t the lower margin of each tergite surface of the prothorax, or the prosternum, be- a small opening, the spiracle, is visible, while tween the bases of the front legs, is more deeply the first tergite bears a large structure, the tym- sunk than that of the mesosternum and meta- panal organ. This organ, if present, is located on sternum. In some groups of acridids this pro- the protibia of the Tettigoniidae and Gryllidae. sternum is provided with a raised tubercle or The number of \isible segments is different in spine of different sliapes. It is used as a criterion the two se.xes, while, even in the same sex, their for separating sufjfamilies and for classification arrangement is different on the dorsal and ven- of some species within that subfamily. Its pre- tral side.

sence, however, is not restricted to one sub- In tlie male there are eleven tergites, the 9th family. and 10th being partly or wholly fused. The 11th

The tvvo posterior segments of the thorax, tergite is represented by the epiproct. Laterally the mesothorax and the metathorax, bear, in to the epiproct. and arising independently from addition to the ventral or latero-ventral legs, the membranous areas behind the posterior mar- the dorsal tegmina and wings, except in the ap- gin of the 10th tergite, are the cerci, developed terous forms. In some species opposite sc.xes are as a single unit or multiariculate, and often im- alate and apterous, the female usually being the portant in taxonomy. apterous sex. The true, or hind wings, attached On the ventral surface nine sternites are to the metanotum, are sometimes very greatly re- visible, the 1st being fused with the metasternum, duced. The front wings, or tegmina, are fixed to but still distinguislied from it. Tlie distal part the mesonotum. They are relatively coriaceous of the 9th sternite forms the subgenital plate, an (liard or leathery), not folding and scarcely or inflated structure in the male under which the Biological Series, \'ol. 4, No. 3, Septemher, 1964 19

Plate I. Morphology of Trimerofropis paRidipennis pallidipennis. Fig. 1, female, lateral view; Fig. 2, female, head, facial view; Fig. 3, female, head and pronotum, dorsal view; Fig. 4. female, tegmen and wing; Fig. 5, female, ape.\ of abdomen, lateral view; Fig. 6, male, apex of abdomen, lateral view.

, FRONTAL SULCUS

RONTAL CARINA

FIG. 2

FIG. 3

^EPIPROCT ^PARAPROCT

FIG. 5

,tPIPROCT 'PARAPROCT

CERCUS

FIG. 6 FIG. 4 20 Ukiuiiam Vijung Univejisitv Science Buu.in'iN copiilatory organs are concealed. The copulatory The genital structures of both sexes are imjK)rt organs of the male are usually syniinetricai, very ant criteria to the classification of the Ortho- nirelv asyiniiictricai. The lOtli sternito of tlie ptera. male is not visible externally in the adult and the lltli sternite consists of the latcro-veiitral Notes on Development plates or paraprocts associated with the epiproct, The first nymphal instars are recognizable represented by the 11th tergite. The abdomen of as Orthoptera and differ from the parents in size, the male thus terminates in a more or less conical, in the total lack of wings, and external genitalia, or obtuse, genital plate. and in the proportionately large head. Their The se.xes of the Orthoptera are easily dis- metamorphosis is gradual paurometabolous of tinguished. In the female there are also eleven ( some authors), the growth to adults being dorsal tergites, the 11th segment forming the accomplished by a series of stages or instars, epiproct, as in the male. On the ventral surface, during which they feed ravenously. These stages however, there are onlv eight sternitcs, the 8th are separated bv periodic molts being usually considerably longer than the others (ecdyses) of the chitinous exoskeleton. After ecdysis the in- and called the subgenital plate. The 9th and sect increases rapidly in size before the body wall 10th sternitcs are not visible externally and the becomes rigid. 11th forms the paraprocts as in the male. The tip of the female abdomen is formed by the two The wings, if present in the adult, appear pairs of \alves ol the ovipositor which resemble in the third instar as sliglit backward outgrowths strong chitinous hooks or appendages. The shape of the second and third thoracic nota. These of the ovipositer varies in different species in wing pads increase in size with each subsequent accordance with the conditions under which ecdysis. They may extend over several segments the eggs are laid, since its function is to prepare of the abdomen in the last immature instar. The the hole in which the eggs are deposited. In the instar of the nymph can generally be determined species that insert their eggs into the ground, the by the comparati\'e size of the wing pads. valves of the ovipositor are shaped like strong The external genitaha are present in the ulti- hooks, while in those species that lay their eggs mate or penultimate nymphal instar. After the in the stems of plants, the valves are built on the series of usually five ecdyses the insec-t attains principle of a saw. Thus, different modifications the adult form and does not again shed its of the ovipositors exist in the different groups. exoskeleton.

ANNOTATED LIST OF THE ORTHOPTERA AT THE NEVADA TEST SITE

sary to check a complete description given by a Use of the Keys recent author. Descriptions used in this paper Keys based on only the more salient char- are, for the most part, incomplete and may be acters are imperfect instruments. They are only (juite useless in some cases of mistaken identity partialK' descriptive, and are for convenience or establishment of new records. only, as the\' are intended as a short-cut in identi- No taxonomic work is so complete as to be fication. In cases involving any doubt of identi- absolute. The keys presented here are c-onfined fication, comparisons should be made with accur- to the species definitely known from the Nevada ately determined specimens. In some instances a Ti'st Site and some definitely known from siu- full description of the species in question mav be rounding areas. Others mav e\entually be found checked. In the matter of descriptions the worker within the confines of the test site, necessitat- may run into difficulty. Early entomologists ing a re\ision of the keys. published descriptions to species that are com- Most of the structures made use of in the pletel\' inadecjuate, if not entirely useless. These keys are discussed in "External Anatomy" and are descriptions perhaps identified the species further identified by separate figures. These and known at the time of publication, but the con- ailditioual structures, about which there may be stant addition of new species to the literature some uncertainty, are illustrated in other draw- has limited the use of the original description to ings inserted near the keys as characters for easy present-day useage. It may, therefore, be neces- reference. Biological Series, \'ol. 4, No. 3, Septembek, 1964 21

Key to the Suborders of the Orthoptera

{ MdclitiLil from Utlin and Grant, 1961. Any rtfcrfiicc to groups not found thf prest'nt study area

h IS Ixcn omitted. )

Caudal limbs saltatorial in type, the femur great))- enlarged (Fig. 1); pronotum generally developed as a large saddle-shaped stnicture co\ering most of thora.v laterally and dorsally (Fig. 2); evident auditory organs generally present on pro.ximal abdominal segment or on cephalic tibia; tarsi always less than five-jointed 2

Caudal limbs ambulatorial or cursorial in type; pronotum compressed, flattened above and below or not markedly different from mesonotum and metanotum, never as in alternative; evident auditory organs not present; tarsi five-jointed 3

Antennae short, usually no longer than the head and pronotum combined (the chief exception being the family Tanaoceridae in which the antennae are slightly less than the total body length); antennal segments less tlian thirtv' in number, filiform or sometimes flattened; auditory apparatus, when evident, placed on each side of the pro.ximal abdominal tergite (Fig. 3); female ovipositor composed of four short valves; caudal tarsi with three segments (Fig. 4) Suborder Caelifera, page 22

Antennae usually long, setaceous and with many minute segments, almost always exceeding the body in length; auditory apparatus present on cephalic tibiae (verv rarelv absent) (Fig. 5); female ovipositor usually long and and well developed, often spear- like or sword-

shaped, rarely reduced in length, composed of four or si.\ valves; caudal tarsi usually with four segments (Fig. 6) Suborder Ensifera, page 81

Body very slender and elongate, of cylinderical, stick- or twig-like form; pronotum shorter than mesonotum or metanotum; tegmina and wings completely absent; cerci a single segment; no styles present on the subgenital plate of the male; ovipositor of female formed of

si.x short valves, not surpassing the subgenital plate; limbs ambulatorial, long, similar in type; coxae never elongate, well separated Sul^order Phasmatoptera, page 118

Body ranging from slender and elongate (in .\Ianteidae) to stock)' and robust, somewhat

flattened (in Polyphagidae ) ; pronotum ovate or elongate, larger and more conspicuous

FIG. 1 FIG. 3 FIG. 2

Figs. 1-6. 1. Trimcrotroph palUdipciiim palUdijwnuifi. female, caudal appendage. 2, T. p. piiUkUpennis, female, showing auditory apparatu.s, lateral pmnotum, lattral \ic\\ . 3, T. p. pallidipcuuin, female, proximal abdomen view. 4. T. p. pallidipennii, female, caudal tarsus, lateral view. 5. Capnohotcf^ fuliginosus, female, distal femur and proximal tibia, showing auditory apparatus, lateral view. 6, C. fuliginosus, female, caudal tarsus, lateral view. Hhk.ium Vounc UsiNKiiM-n- ScitNcE Bullktin

than thf mesonotum or metanotnm; tegmina and wings generally present, but often rcxluced or absent; eerei innltisegmented; styles present on tlie subgenital plate of adult male; ovipositor of female little de\elo[x;d; limbs all cursorial, or the cephalic pair strongly modified and developed as efficient raptorial organs, in the latter ease with the covae greatly lengthened, median and caudal coxae closely placed, often in contact Suborder Dictyoptera, page 121

Suborder C aixifkra

Key to the Superfarrulies of the Caelifera

Antennae inserted mesad of the eyes (Fig. 7), and almost aiwa\s with more than twelve segments; cephalic limbs of ambulatory type; cerci c-omposed of a single segment; ovipositor of female formed by two pairs of opposed valves; caudal tarsi with tliree segments Superfamily Acridoidea, page 22

Antennae inserted below the eyes (Fig. 8), and with from six to twelve segments; cephalic

tibia of s|i(ci.ili/.cd tspc. aclaptcci lor tossorial iiabit ( Fis^. 9); eerei composed of one or two segments; ovipositor of female composed of four divergent valves; caudal tarsi with a single segment. Total length less than 10 mm. Superfamilv Tridactvloidea, page 81

FIG. 8

FIG 7

Figs. 7-9. 7, Trimerotropis pallidipennis paUidipennis, female, head, facial view showing insertion of antennae. 8. TrUlactylus iipicalix, male, head, facial view showing insertion of antennae. 8, T. apicalis, male, head facial view, sliowiiig insertion of antennae. 9, T. apicalis; male, cephalic appendage.

,. , ., , range from the 8 mm. eumastacid male, the

SuiXMtaillllv .ACKIIKJIDE.V ii . . .1 xi i t- . ' • smallest s|H'cies oeeurrmg at the Nevada Test

The general form of tiiese insects is largely Site, to the 50 mm. body, 100 mm. wing spread of the so-called "grasshopper" type, although the of the acridids. Some of the tropical acridids, not body ma\' be \ariousl\- de\elopecl from extremely of this area, may aehie\e a wing-spread of over elongate to siiort and flatti'ued. In size they -00 mm.

Key to the Families of the Acsudoidea

/ Mo.lilird hoiM Urlin and Cirant. UJOP

1. Cephalic and median tarsi two-segmented (Fig. 10), caudal tarsi three-segmentetl, no aroh- um present between tlie tarsal claws; pronotmn always extended caudad. covering all of mesonotum, metanotum, and generally, most or all of abdomen (Fig. 11); prosternum developed into a broad apron-like sternomentum which encircles a portion of the moutli parts; frontal costa ventrad of median ocellus always a single carinate ridge

l'"ainil\' Tetrigidae, page 2,'^

All tarsi three-segmented, arolium present between tarsal claws (Fig. 12); pronotum rarely extended caudad sufficiently to cover the remainder of the thoracic nota and much of the abdomen; prostermnn not developed into a definite sternomentum 2

2. Abdominal spiracles situated in the terga; pro.xiinai abdoininal segment generally with evident lateral tympanic membrane Famil\ Aerididae, page 29

Abdominal spiracles situated in the latero-dorsal membrane between the terga and sterna; proximal .ibdoniinal Icr^ilcs without rxidciit lateral t)'mpanic membrane, wingless 3 Biological Series, Vol. 4, \o. 3, September, 1964

(Fig. frontal 3. Antennae short and stubby, shorter than the caudal femora in both sexes 13); costa bicarinate ventrad of the median ocellus (Fig. 14); no specialized stridulatory organ present on tlie sides of tlie abdomen in tlie male se.\ Family Eumastacidae, page 24

Antennae very long in tlie male exceeding the body length, in the female at least equalling the the caudal femoral length; frontal costa, especialh in the male, unicarinate ventrad of median ocellus (Fig. 15); specialized stridulatory organ present in the male sex laterad on the third abdominal tergite Family Tanaoceridae, page 27

Cte=^

FIG 10

FIG II

FIG. 12

FIG

Figs. 10-15. 10, Parutcttix mexicanus, female, di.stal tibia and tarsus of mesothoraeic appendage, lateral view. 11, P. mexicanus, female, pronotum and tegmen, lateral view. 12, Trimcrotropls pcillidipcium pallidipennis,

female, distal segment of eaudal tarsus showing elaws and arolium. 1.3, Mor.sea cutifornicu piutc, female, head and pronotum, Literal view. 14, .\/. c. piute, female, head, facial view. 15, Taiiaocerus koebclei koebelei, female, head, facial view.

Family TETRicmAE The female may be recognized by the serrulate Subfamily Tetriginae ovipositor with sharp diverging extremities.

(Figures 10, 11) The Tetrigidae are found in practically all liabitable areas of the earth, even to relatively Members of this family are among the small- higli elevations. They occupy a variety of terres- est of the Acridoidea and are commonly referred trial habitats, but are nearly always associated to as the pygmy or grouse locusts. They may be with damp situations or water and are generally readily recognized by the prolonged pronotum, common along streams and other bodies of frequently with a very high median carina, water. They feed upon algae, lichens, mosses, which covers most or ail of the abdomen, often sprouting seeds, sedges, and other tender plants including the terminal abdominal appendages. and debris. This specialization pr<)\'ides protection for the delicate wings and replaces the tegmina which Their coloration is protective, resembling the have been reduced to small oval lobes or scales. soil background in pattern, and, altliough often The wings are usually present and well develop- varied, is never such as to make the insect con- ed, the length varying with the length of the spicuous in its habitat. Individual and local pronotum. Both long and short winged individ- variations are obvious and overlap geographic- uals, accordingly those with a long or short pron- ally, adding to the complexity of the group. otum ( macropronotal and brachypronotal of Representatives of this family have not been Rehn) are found in the same species. actually collected from the Nevada Test Site, but

The prosternum projects forward as a chin the area is within the range of two species. These piece, the sternomentum, encircling the caudal insects could survive only in the areas within section of the mouthparts. The arolium between the confines of the test site where there is a the tarsal claws is absent, which may be correlat- perennial water source, such as at Cane Springs ed with the habit of resting on the ground in- or Whiterock, questionably on Rainier Mesa, and stead of living on plants. The front and middle perliaps could become established around some tarsi are two-segmented, the caudal tarsi three- of the water tanks in the areas. Numerous segmented. The subgenital plate of the male is attempts were made to secure these insects in conical or triangular. The cerci are very small. the possible habitats at the test site. )

24 Hlllc.IIAM ^

Pitnitcttix incxiranu.i (Saiissiirc) has a very Family Kl'.mastacidae widi- tlistrihiitioii ovt-r imicli of tlii- western Subfamily Mohskinae United States, and is definitely known from The name by which these insects are com- Heattv and Asli Meadow in Xve Countv, and moiiK' known is eumastacids, although some en- Las Vegas, CiKiik Co., N'e\ada ( Relni and Grant, tomologists have applied the name "monkey 1957). grasshoppers" because of the agility by wliicli Paratcttix azlccus (Sanssure) has been col- they move along the branches and twigs of lected in the Panamint Mountains and Pana- brush and chaparral. mint Valley, as well as other localities in Inyo They are of relatively small size (the small- Co., California (Rehn and Grant, 1957), and est acridoid insect collected at the Nevada Test conceivabK' could be found in the confines of (lie Site was a male of this family measuring 8.2 test site. mm. in total body lengtli) and completely wing- The genus Faiatcttix can be recogni/etl by less. The famiK' as now ri'cogni/ed is predo- the fastigium, which is hardly, if at all, produced minantly a tropical one, is well distributetl cephalad of the eyes. As seen from the dorsum throughout the .southwestern United States, but the fastigium is narrower than one of the eyes. infrc([uently encountered because of the natural iuibits of the insects. The two species which may be found in the head of the eiunastacids is remarkabK area are distinct according to the condition of the The enlarged, projecting dorsad beyond the cephalic median femora. In mcxicaniis, the median femora 12- have two or three pronounced lobes on the margin of the pronotum. The or 13-segment- \entro-external margin and are iobulate or un- ed antennae are shorter than the combined length of the head pronotum. dulate (or even simply Ciirinate) on the dorsal and The male siibgenital plate is simple, the proximal abdo- margin. In aztecus, the ventro-e.xtemal margin minal tergite without evident lateral tympanic is at most undulate or loculate, never truly membrane. Abdominal spiracles are situated in lobate. The preferred habitat of mexicanus is on the latero-dorsal membrane. Females have one or near mucky ground, in the vicinity of stand- or more abdominal tergites preceding the epi- ing or running water, but it has been collected proct with a marked medio-longitiidinal fold or on bare sand in temponuy dry creek beds. "In fissure. Tlie mesosternum and metasternum are its desert environments, while the surroundings strongly united into a more ob\iously single may otherwise be exceedingly dry and arid, it structure than in the Acrididae. The caudal is met with only near water, if only a small seep femora are without the dorso-proximal over- or a tiny rill, or in a spot where limited ground hanging lobe found in most acridids. Tlie caudal moisture is present." (Rehn and Grant, 1961) tibiae have two pair of distal spurs ( calciiria On the other hand, aztecus "is more definitelv and a tarsal arolium is well developed.

saxicolous . . ., its general preference being for Only one species has definitely been collected gravelly stony or situations near water. These at the Nevada Test Site, but because of the may be the stony margins of rivers or streams, range of distribution of other species approximat- along rills on rock slopes, or about 'tinajas', or ing the test site boundaries, tliey, too, are in- rock water pockets." (Rehn and Grant, 1961) cluded in the keys and are discussed.

Key to the Genera of the Eumastacidae

(Modifi.-d from Relin and Crant. 19fil)

Antennae with thirteen segments, node on \i'ntral surface of lOth segment (Fig. 16); male cerci subcompressed, falcate distad (Fig. 17); subgenital plate of male with a recur\'ed, median dorso-proximal sclerotized linguifonn process (Fig. LS). Form as a whole more slender; frontal costa narrower proportionately than in (lie same sex of the alternate category Morsea Scudder

Antennae witli twelve segments, node on ventral surl;Hc ol 9th seguienl; male cerci tapering, not falcate; subgenital plate of male without a recur\(>d median dorso-proximal sclerotizetl linguiform process. Form as a whole more robust: frontal costa broadi-r proportionately than in the same sex oi (lie a!l<'niatc category Pstjrlionuisldx Rehn .nid Ilebard )

Biological Series, Vol. 4, No. 3, September, 1964 25

Test Site represents nymphs from 2.9 mm. long to adults. Size variation of adults selected at random are gi\en in Table 9. As discussed previously, the total length of

the insect is probably the least accurate measurement because of the abnormal stretching of the FIG. FIG. FIG. 16 17 18 abdomen of the female and the abnormal curvature of the abdomen of the male. Tlie above Figs. 16-18. Morsea californica piute. 16, female, antenna, lateral view. 17, male, cercus, lateral measurements of total body length of females view. 18, m.ile, apex of abdomen, dorso-caudal are considerably larger than previously publish- view. ed measurements. There is a possibility, of course, that the Nevada Test Site specimens re- Genus Morsea Scudder present atypical forms, intermediate between

M . californica piute and M. californica califor- 1898. Morsea Scudder, Psyche, VIII, p. 179. nica. a more western form which averages larger. The proper placement cannot be made without Morsea californica piute comparison to typical specimens of that sub- Rehn and Grant species.

( Figures 13, 14, 16-18; Table 9; Map 1 Coloration. This subsj^ecies as represented

1958. Morsea californica piute Rehn and Grant, by the series from the test site is more vari- Trans. Amei. Entom. Soc, LXXXIV, pp. 239- able in color phases than in morphological 250, figs. 18-22. characteristics. This same variation is noticeable in a series of specimens from Utah (3 miles Distinctive Features. Fonn slender and southwest of Shivwits Indian Reservation on U.S. elongate, the males especially with deep, narrow Highway 91, Washington Co., I'tah, September head and very large and prominent eyes. Anten- 9, 1959, A. H. Bamum, on desert-almond, Prunus nae witli tliirteen segments, witli a xentral spini- fasciculata, and cliff-rose, Cowania mexicana, form tootii on the 10th. Pronotum long in relation and from Nevada ( Lee Canyon, Clark Co., Nev- to depth, about twice as long as deep in males; ada, August 13, 1961, A. H. Barnum, from bitter- ventro-caudal angle of lateral lobes rounded, brush, PursJiia glandulosa. In coloration the especially evident in females. Male cerci falcate, iivmphs are light brown or tan, occasionally gray- the subgenital plate with a median, dorsal, brown, rarelv dark; the adult coloration is anteriorly projecting linguiform process. Caudal brownish-black, with an occasional gray-brown femora with mcdio-dorsal and dorso-lateral min- or tan individual. No correlation could be made ute, spiniform processes; genicular lobes similarly with reference to habitat, se.x, or color phase, as armed. Tarsal claws asymmetrical. individuals collected even from the same shrub, Size Variation. The series from the Nevada or from the same group of shrubs in one area

Table 9. Size variation of Morsea californica piute. 26 BiticiiAM Voi'Nc Univehsity Science Bulletin

I TrrTHOrvLC HACUIATA Biological Series, \'ol. 4, \o. 3, Septemuer, 1964 27 exhibited degrees of variation. The majority' of Genus Psychomasfax specimens have the caudal margins of the lateral Rehn and Hebard lobes of the pronotum contrastingly light colored. 1918. Psychomastax Rehn and Hebard, Trans. This light coloration extends (especially in Amer. Entom. Soc, XLIV, pp. 225, 242. males ) along the entire ventral border of the Specimens belonging to this genus have not lateral lobes. The males usually have darker been collected at the Nevada Test Site, but the postocular bars, this condition obviously some- genus is restricted in its distribution to south- what reduced in darker phases. western Nevada and southern California, and Distribution. In their recent revision of the may eventually be found on the test site. genus, Rehn and Grant listed the range of this This genus is more robust but less attenuate subspecies as extensive in the Great Basin. It in form than Morsea, but it resembles that genus

extends from California, in the Mono Lake region in that they are both apterous. The head is not to Beaver County, Utah, and from northwestern as deep in relation to its breadth. The antennae Arizona to White Pine County, Nevada. It inter- have twelve segments with the ventral spiniform grades with the other four subspecies in the peri- tooth present on the 9th segment. The pronotum pheral margins of its distribution. is deeper in relation to its length, the ventro-

caudal angle of the lateral lobes is angulate rat- Habitats. The preferred host plant of this her than rounded. Male cerci are styliform, flat- insect is bitter-brush (Purshki g^latutulosa), and tened on the mesal surface;' the male subgenital the insect was captured wherever this shrub was plate without a dorsal linguiform process. The present on the Nevada Test Site. It was also caudal femora have their apices (including the collected from Artemisia tridentaia in one area genicular lobes ) armed as in Morsea, but the where Purshi

of the inaccessibility of the areas where it is Nevada Test Site is Psychomastax rohusta He- found. The earliest collection of nymphs was bard, the type locality being Charleston Peak, June 22 and adults appeared as early as July 13. Nevada. It has also been collected at Lee Canyon The last recorded appearance is September 30. at 6,000 feet and in eastern California. It is It almost assuredly occurs in adult form into apparently "restricted to the mountainous areas October and possibly November. Its greatest of southwestern Nevada and adjacent south-

abundance is during the month of August. eastern California" (Rehn and Grant, 1961). It may possibly be found on some of the higher Localities Represented. Specimens exam- mesas and mountains of the Nevada Test Site.

ined ( nymphs and adults ) : 65. Study TCB, near Tippipah Springs, 37 speci- Family TANAOCERroAE mens, from June 22 to July 16, on Piirshia ghirul- Members of this family, which was recently ttlosa. removed from the Acrididae (Dirsh, 1955), ap- Study sand dunes, 25 specimens, ECA, on pear very early and are seldom encountered by from August 11 to September 30, on Purshia collectors. They are remarkably agile, especially glamlulosa. the males. Study 12A, Rainier Mesa (disturbed area), They are of medium size, the male moder- 2 specimens, August 21, on Purshia glamlulosa. ately elongate, the female relatively robust. Study 12E, Rainier Mesa (undisturbed area), They differ from the Acrididae by the frontal 1 specimen, August 24, on Artemisia tridentata. costa which consists of a single carina ventrad BiiiGiiAM YouNc U.sivEnsiTY Science Bulletin of flic iiH-cliaii oci'lliis, siiggestivf of tlie Tt'trigi- profile; lateral carinae continuous with those of diic in tliat rt'spect. The antennai- arc .slii;litlv frontal costa which are rather widely separated longer than the body (males), or slightly shorti-r and only weakly sulcate between. Head, in (females). Wings are completcK' ahsiTit. The frontal aspect, with moderately marked supple- stridtilatorv ridge is locati-d on the sitle of the mentary carinae. Median carina of pronotum third ahdoininal segment and on the lower inter- moderately evident, entire, usuall)' finely sulcate; nal hasal margin of the eandal femur. The cau- cephalic margin subtrnncate, caudal m;irgins dal tibiae lia\'i- the external apical spine present, truncate or very lowly arcuate with shallow as in the Homalcinae of the Acrididae. Because crenulations; lateral cariruie absent; lateral lobes of the presence of this e.vternal apical spine the with caudal margin weakly sinuate. Medio-dorsal group \sas considered to belong to the Romalci- carina of abdomen tectate, finely, but evidently, nae. Recent research on the male and female sulcate in proximal segments; lateral carinae internal genitalia, however, by many entomo- absi'nt; males with stridulatory ridge on side logists, has straightened out the complexity of of third abdominal tergite; epiproc-t apically many of these aberrant species. The subgenital acute, especially in females. Subgenital plate plate of the male is composed of two or three truncate in females. sclerotic plates connected bv a membrane. Nymphs can be recognized by their long

.\ccording to Relin and Grant ( 1961 ) "the antennae. This is the only acridid found at the tanaocerids are an endemic North American Nevada Test Site with extremely long antennae, family restricted to southwestern United States the antennae of instars of all other spec-ies being and extreme northern liaja California, Mexico. very short. All instars are represented in the Tile family is known from the states of Utah, collection from the Nevada Test Site, the small- Nevada, Arizona and California. It is found only est, representing the first instar, being 4.6 mm in in the Upper and Lower Sonoran Life Zones." total bod\' length and with antennae 3.5 mm One representative of this family is known long. from the Nevada Test Site. Morphological Variation. Considerable vari-

ation is noted in the degree of surface tubercula- Genus Taiuioccrus Bruner tions among specimens. Some specimens are 1906. Tanuoccnts Bruner, Biol. Centr.-Amer., much more rugose than others. Orth., II, 191. p. Size Variation. The series from the Nevada Test Site represents nymphs from 4.6 mm long, Tarmoccrus koehcici koehelci Bruner as previously noted, to adults. Sizes of adults

(Figure 15; r.iiilc 10; Map 2 I selected at random are gi\'en in Table 10. 1906. Tanaocerus kocbelei Bnmer, Biol. Centr.- Coloration. The series of sjx'cimens from Amer., Orth., II, p. 192. the Nevada Test Site shows considerable varia- Distinctive Features. Surface nnxlerately tion in coloration, the base color being dark rugose, pronotum with numerous short rugae mottled with buff or grayish-white. A inmiber and rounded tubercles. of specimens are particularly maculate, the de- Fastigium, in profile, sloping ventro-cephalad gree of maculation cxirresponding to the degree continuous with frontal costa which is arcuate in of rugae and tubercles. ( Tliis maculate tendency

10. .Size variation of Tuiiuuccru.s hu'lu'lci koehcici.

1 ) .

Biological Series, Vol. 4, No. 3, Septemuer, 19fi4

is also seen in a series in the author's collection Area 12, 1 specimen, March 16, vegetation from Washington County, Utah [in the vicinity unidentified. of Terry s Ranch, Beaver Dam Wash, April 17, Area 15, 1 specimen, November 28, vegeta- 1952, and April 2, 1960, all adult]. The environ- tion unidentified. ment is verv similar to that at the Nevada Test Study JA, Jackass approach of mi.xed vegeta- Site. tion, 17 specimens, January 9 to April 8 (adults), Distribution. The distribution of this form and August 13 to December 28, also January, occurs "from e.xtreme southvi'estern Utah, west February and March (nymphs). to Nye County, Nevada, and Inyo County, Cali- Study CM, Cane Springs, 7 specimens, Feb- fornia. From its northern limit, the subspecies ruary 26 to March 14 and (one nymph) Nov- is found southward in eastern Kern, western San ember 27. Bernardino and eastern Los Angeles counties, Study NCC, 1 nymph, November 7, vegeta- Cahfornia. It intergrades with T. k. albatus in tion unidentified. southeastern Los Angeles County, southwestern San Bernardino County and northwestern River- side County" (Rehn and Grant, 1961). Family Acrididae It has been collected, but not found common, This family comprises the exceedingly num- throughout most of the range of the Nevada erous and common "short-horned" grasshoppers Test Site. It is probaby more common than is (as distinct from the "long-horned" grasshoppers indicated by the records because of its early belonging to the superfamily Tettigonioidea ) appearance and difficulty of collecting. They are found throughout the entire year but are common and abundant from early spring to Habitats. According to observed records this late autumn. The family includes all the econ- species inhabits both the Upper and Lower omically important migratory locusts or grass- Sonoran life zones over a broad range of desert hoppers, so well-known throughout recorded environments. Nymphs are generally associated history. Over one thousand genera and well over with vegetation and can best be collected by ten thousand species of world-wide distribution sweeping with nets. The adults apparently prefer are known in the family. All members of this the ground covered with small pebbles or coarse family feed on plant material and often are im- gravel, in which environment they are adequately portant insect pests. Most of the damage is done concealed and seen only when disturbed. by a small number of species, but many others Characteristic areas are vegetated by Yucca may do some damage at times. brevifolia and early spring ephemerals. They are characterized by relatively short Seasonal Occurrence. Adults have been col- antennae, usually shorter than the body, filiform, lected from January 9 to May 1. One first instar ensiform, or clavate, with the segments distinct. was collected in July and immature specimens There are three ocelli, two laterad and one mesad have been collected in all months to the middle in the frontal costa. The frontal costa is well of March. The series of nymphs shows a gradual marked, never replaced ventrad of the median increase in size. They overwinter and appear ocellus by a single carina as in the Tanaoceridae. as adults when optimum conditions are reached The lateral temporal foveolae are usually present, throughout late winter and early spring. frequently well marked and sharply outlined, varying in position, and important the classi- Localities Represented. Specimens examined to fication of the group. The eyes are lateral, us- (nymphs and adults): 66. ually relatively large. Study TA, Midvalley, 26 specimens, from The pronotum is proportionately large, vari- March 15 to August 18 (July and August speci- ously produced and ornamented. The dorsal sur- mens all nymphs), in an area of Artemimi face, or disk, usually has a distinct medio-longi- triclentata. Very few specimens actually collected tudinal carina which may be elevated into an en- off vegetation. tire or interrupted crest. The lateral carinae Study lOD, 1 specimen, March 13, in Coleo- defining the lateral margins of the disk may or gijme ramosussima area. may not be present. A single transverse sulcus Study 5A, 8 specimens, January 9 to August is almost invariably present, this being the caudal 31, in Larrea divaricata area. sulcus; anterior sulci may also be indicated. Area 4, 4 specimens, March 13 and August That portion of the pronotum anterior to the

26, vegetation unidentified but probably in principal sulcus is termed the prozona, that

Graijia-Ltjciiim. posterior is the metazona. Bmiciiam Voi'sr. Univehsity Science Bulletin

The tegmina are fully developed and iisnaily The phallic structures for many species of narrowly elongate, occasionall)' hraciiypteroiis, Acrididae ha\e been described by several auth- or even apterous, sonietiines pre.sent in tt)i> male ors, and the relationships of the subfamiUes and absent in tlie female. When present tliey are and families of the .•\cridoitlea have been reveal- alwavs at least as long as flie wings, usually ed. As a matter of fact, the present scheme of duli-eolored and thickened or coriaceous. In Orthoptera phylogeny is a result of these studies. numerous cases certain veins of the marginal The .'\cridida<' contains species which are or discoidal fields mav he specialized for stridu- highly variable. The large body size and environ- lating purposes. Hind wings are developed ac- mental differences to which species are subjected cording to the tegmina, from fully developed to are undoubtedly responsible for this \ariation very abbreviate, or absent, but never exceeding for the most part. This is especially true of the tegmina in length. In some c;ises the radiate species from western North .America which may veins of the wings are thickened and rod-like, be subjected to extremes of altitudes, tempera- and their surfaces are specialized, along with tures and general habitat, and also for species certain veins of the anterior field and the anal throughout the western hemisphere. complex, for stridulating purposes. The mem- Environmental conditions pla\- important branous hind wings may be briglitly colored and an in variation external morphological char- very attractive, especially noticeable in the part of acters, but internal characters such as the phallus species found in the southwestern deserts. should be less variable, except as may be in- The auditory organs are with few exceptions cidental to external changes. These highly vari- ( in some apterous s[X'cies ) on the side of the able characters can not often be relied upon as a proximal tergite. The external genitalia of both basis for classification. .Apparently the more sexes are basically as in the other families of the bizarre a structure may be, such as a cristate Acridoidea. The abdominal spiracles are placed median pronotal carina, the more variable it ventrad in the sides of the dorsal sclerites, not is. In a number of recent studies based on the in the membrane as in the Tetrigidae and Eum- internal genital structures of the male, on the astacidae. The prosternum is with or without a other hand, it was found that the so-called median spine or tubercle. When present this is phallic complex and especially the epiphallus of usually well marked, and its form varies in dif- the male, and the shapes and relationships of the ferent groups. valves of the cingulum and the apical valves of Cephalic and median limbs are short, relati\'e- the penis are not .subject to variation to any ly slender, usually sul)e(jual in size and general great degree, and when it does occur it is found form. The caudal femora arc much larger, sait- in a complex species group which occurs over atorial. Stridulating organs are often located an extended area. on the inner side of the caudal femora. All There appears to be little variation in tlie species in the family show the outer surface genital structures of any species. Such variation of the caudal femora with the same ts'pe of may result from the different technitjues of dis- impressed striae. The caudal tibiae have their secting or treatment of the dissected structures. exten.sor margins armed with spaced spines The degree of sclerotization of parts is obvious for the greater part of their length, and two and especially apparent when teneral specimens distal spurs ( calcaria ) are found on each side. are examined. The sclerotization of internal The tarsi of all limbs are composed of three seg- structures apparently proceeds more slowly than ments, the proximal one (the metatarsus) with externally, and specimens which are apparently three pads or pulvilli. The tarsal claws ha\e a completely sclerotized externally show internal distinct pad or arolium between them. teneral conditions. The determination of whether In recent years application of the male ami or not a specimen is fully mature is problem- female genital structures, both external and inter- atical, but important, and can best be determined nal, have been regularK- ap]5lied to many orders b\' the internal structures. of insects. These structures have been especially Anyone who is going into a serious study of useful to show phvlogenv both within a group the Acrididae would benefit by a detailed study and the relationships of different groups. Tlie of the internal genitalia, especially of the male. male organs of intromission have been especially As previously statcnJ, these structures are less useful in re\isions of several acridoid genera, variable than the external characters. Only the and overall studies of the epiphallus of the external characters are used as criteria in the superfamilv .\cridoidea have been made to es- following key, so it is limited in use to the im- tablish certain families and subfamilit's. mediate area. )

Biological Series, \'ol. 4, No. 3, September, 1964

Key to the Subfamilies of AcRiDroAE

1. External distal spine of caudal tibia present, with the appearance of three apical external spurs' (Figs. 19 and 20); median carina of pronotum ranging from virtually obsolete to strongly elevated; wings reduced or fully developed in both sexes Romaleinae, page 31

External distal spine of caudal tibia absent (in species at the Nevada Test Site) 2

2. Prosternum armed with some type of process, usually a spiniform tubercle (Fig. 21); wings reduced, sometimes with marked se.xual difference, or fully developed Cyrtacanthacridinae, page 35

Prosternum rarely armed with any type of process, generally relatively flat (if a flat transverse prosternal process is apparently present, then the species is elongate, the form being very slender and linear); wings generally well developed, rarely reduced, never apterous, often brightly colored; pronotum often with a marked well elevated median carina Acridinae, page 48

FIG. 20 FIG. 2 I

FIG. 19

Fig.s. 155-21. 19, Tytthotyle maculata, male, distal tibia and pro.\imaI tarsus of caudal apix-ndage, lateral view. 20, Dracotcttix plutonius, female, distal tibia and pro.\imal tarsus of caudal appendage, lateral view. 21, Mchinoplus complanatipes canonicus, female, prosternal spine, cephalic view.

are completely wingless or have these organs Subfamily Romaleinae greatly reduced. Tlie family is principally tropical The common name of "lubber grasshoppers" in distribution. has been inappropriately applied to the members The subfamily exhibits a great diversity in of this subfamily, inasmuch as many of them are external appearance, and the two species found winged ( often with brightly colored hind wings at the Nevada Test Site show little e.xternal and very graceful in appearance, though large. similarity. The internal genitalia must be studied Others, suggestive of the true "lubber" condition to show this true relationship.

Key to the Genera of Romaleinae (Modified from Rehn and Grant, 1959b)

Median carina of pronotum very high, lateral earinae well developed (Fig. 22); tegmina and wings shorter than the abdomen in both sexes; fastigium with a marked rostral development; pronotum with surface rugose, caudal margin of disk with distinct spaced nodules Dracotcttix Bruner

The world-wide subfamily Cyrtacanthacridinae shows the general presence of the external distal spine of the caudal tibia. The spine is absent, however, in two genera, one of which is North American. The genus Sjxiuiacris is apparently limited in distribution to extreme southern Cahfornia in the Coachella and Imperial valleys. There is no obvious "third spine" in the genera presently under consideration, and this key character is reliable only for this immediate area.

^The subfamily Oedipodinae is now considered a.s a part of the Acridinae. The variability of intermediate forms with respect to such external characters as the median carina of the pronotum, the lateral profile of the face, the presence or absence of stridulatory mechanism on the caudal femora, the presence or absence of an intercalary vein of the tegmina, can not be relied upon for the separation of these groups. Recent studies on the internal genitalia load to a synonymizing of this old subfamily Oedipodinae and an inclusion of these genera in the Acridinae. (See Rehn and Grant, 1960, for discussion of this problem.) 32 BiiiciivM ^l)l^(; I'nivkiisitv Sciiknc k Bfi.LhrnN

Mcdi.iii liiriiia ol pronotimi ohsdli-tr to siibohsolc'c. no l.itci'.il caiiiiac |>ri'si'iit on disk ( Fig. 23); iiilK winncil in liolh scvcs; raslit^iiini latlifr hroadk rounding inio tin- liontal costa; pron-

olnni rclaliM'K' snioolli, tlic pro/ona ol tlii' cliA idinidcil transvcisi'h' Ti/lllioliilc Scndder

FIG. 22 FIG. 23

Figs. 22-2.3. Drill nil Ilix y.liilimiiis. Icni.i Iliad and pruiiotiiiii. lad il \ II u 2}, ri/llluitiilc timiiilula, female, lic.id and piipiKiliini. I.ilrr.d \iiw.

(iciiiis l)rmnliilix Hnincr (il ihe tegmina ;iiid the candal femor.i. (Coloration. In color the speiies pri'si'nls an 1SS9. Diiunlvllix i5iunci-. i'roc. U.S. ,\at. .\Ius., interesting contrast of darks and lights, the base .Ml. p. 50, pi. I, Fig. 1. olor of whitish gray oNcrlaid with a luscous to ilull black p;ittern. This pattt'rii consists of dark longitudinal tlonds on tin- head and proiiotnm Dmcdiiilix philoniiis Unini'r and ilark distal trans\t'rse clouds on all the abdo- (Fii;iires 20, 22; Tal.l.' II; M.ip 3) minal tergites with sullnsions ol darkness on the lateral abdominal tergites. The caudal Icmora 1893. Diacotctlix pliitoniits Hrnncr, \oitIi .Anicr- ha\e two, or sometimes a third, transverse dark itan Fanna, V'll, p. 267. cloudings which are obsolete on the external

Di.stincti\e Features. Tlicrc is nn (itlur lace. The tegniina are less contrasted, being

spi'C'ii's in tlic area witli wliitli tliis could he mn- gia\ -brow n. w itli (he scnation somewhat darker. fnsod. The prodiici'd lostium (hfst scon in later- Distribution. Tliis species was originally al ontlint'), the distincti\(.' pronotal crest, the described Iroin the I'ananiint N'allev and .\rgus broad lingnilorni proslcnial sjiinc, aTul (lie ali- \I(iiiii(,(iiis ill lu\() C"oimt\', California. The dis- bre\iafed tegniina and wings will dislingnisli trihndon gi\(ii 1)\ Kehn and CIrant (1959b) is tlie species. ,i liinltcd ;irc.i ol lu\(i (]ount\', California, be-

Size Variation. Tliiri' is a rein.irkable sex- tween (lie Muiiiii(.iiiis (o the east of Owen's Lake

ual diniorpliisni in this species, the females be- (which is now ;i s.dine sink) and the western ing considerabU- larger. No coinpari.sons were side ol the l'a!iarnint Range, which borders and made with other specimens, bnt apparently the o\erhangs Death \'allev on the west", from series varies only in size from the pre\ionslv "ele\ations of Ironi 45()() to possiblv as high as known material. The most noticeable differenci's, C^!)()() Icel". The li\c sjiecimens from the Nevada both in the male and the female, are in the length T(s( .Site arc (lie onl\ specimens collected out

Tal.l .Si/r N.iri.ilion of Drill nlftlix pUilotiiiis.

§ „ E

cT, TA, April 8, 1961 cif (^ , MeaMiremrnIs

Rehn and Cr.int ( I'Kil )

'ICC. \l.it. Il 21. I'Jfil 9 , April Hi. MKil 9 , lA. I!), 9 , 'IW. .Septenilxr 1961 of 9 , M( .iMwiimnts

Relin and Cr.int ( H)01 ) Biological Series, Vol. 4, N*o. 3, Septemuek, 1964 of tlie Invo County area and the State of Cali- areas hv the senior author later in the year dur- fornia. Tliis locality at the Nevada Test Site now ing the period of greatest heat ( August and Sep- represents tlie most eastern (and northern) dis- tember), and in three different vears, failed to tributional limits known for the species. produce the species." One adult female collected on September 19, Habitats. According to Rehn and Grant 1961, extends the seasonal distribution to at least (19.59b), the species occurs in the "upper part late summer. It is probably present, though of the Lower Sonoran Life Zone, and probably never common, from early spring (the earliest enters the Upper Sonoran, although we ha\e no spring record being March 24, 1961 ) througli definite evidence except known elevations to late summer, or early autumn. support the assumption". The authors furtiicr Localities Represented. Specimens examin- state that "little information is available on the (adults): 5. e.xact conditions where phitonius has been taken. ed

Apparently ... it occurs above the Larrea belt Studies TA and TCC, Midvalley, 2 males, and below that of juniper and pinyon. No clear- 3 females, from March 24 to April 16 and Sep- ly associated vegetational notes are available, tember 19, all on Artemisia tridcntata. No im- but the localities where the species has been mature specimens were collected. taken by others have been visited by the senior

author, but unfortunately not at the time when Genus Tt/ttlioti/Ie Scudder Dracotcttix has been taken". 1897. Ttjttliotijle Scudder, Entom., Several trips were made into the area where Canad. .WIX, 74. the original discovery was made in March, and p. extensive collecting by the author produced only Tijttliotijlc macidata (Bruncr) one specimen, the female collected on April 16, 1961. The preferred vegetation of the insect at (Figures 19, 23: T.ililc r2; Map 3)

the Nevada Test Site is definitelv sagebrush 1889. Thrinchtis {?)maculatus Bruner, Proc. U.S. (Artemisia tridcntata), of what might be con- Nat. Mus., XII, p. 79. sidered the Upper Sonoran Zone. Attempts at Distinctive Features. This large, fully w ing- securing this species by sweeping the brush were ed (in both sexes) species is so distinct it should futile. It was also useless to try to spot the insect not be confused with any other acridid in the visually. The one specimen obtained by the southwest. Tile body surface is rather smooth, author was from a shrub that had been inspectctl not rugose, although a few minute tubercles are visually and then kicked forcefully at the base. generally present on the metazona of the prono- It was found that a strong kick at the base of tum. The median carina of the pronotvim is the shrub would be violent enough to remove faintly evident, the lateral carinae al)sent. Three the other Orthoptera, so this techni(|ue was used, transverse sulci are evident on the pronotum, the finally, in securing the specimen. After being most anterior sulcus ending on the disk and not kicked out of the sagebrush it tried to recover continuous on the lobes as are the other two. to the bush immediately in a series of cpiiek The metazona is distinctly longer than the pro- hops, the wings being useless to the large body. zona. The tegmina and wings are long, well The insects apparently rest on the stouter vertical surpassing the apices of the caudal femora. brandies where they are almost perfectly concealed bv the patches of light and dark bark Morphological Variation. There is consider- of the shrub. The feeding habits of the insect are able morphological variation in this species, as is unknown, but from an examination of the area found in many of the large-bodied acridids. The the insect undoubtedly feeds on Aitemisia tridcn- relative length of the pronotum is especially tata. variable, due to the angulation of the caudal The other four specimens in the series were margin of the disk. The surface of the pronotum, "accidental" discoveries, the insects hopping from as well as the carinations and sulcations of the the bush when disturbed. Many hours of concen- head, is variable.

trated efforts by several collectors by visually Size Variation. As noted in Table 12, vari-

spotting or sweeping have not produced more ation in size of body and structure is apparent. than the five specimens. Coloration. "The general pattern seen in

Seasonal Occurrence. Rehn and Grant this species is a multi-maculate overlay of tones (1959b) comment as follows: "clearly a Spring of brown of varying density and shade on a form, all known material of phitonius was se- base which may range from chalky white cured in April and May. Field work in the same through pale vellowish to a light buff, or rarely ,

Biiif.iiAM Vi)i N<; (.'mvkiisity Scieste Bulletin

TaliK- 12. .Size- varialion of TijIIIidIijU- macuUita.

cf, CB, Aug. 28, 1959

, 1901 cf MCC, ,\iig. 12. (^, j.\, j.iK 18. iwn ^ Mc.isiirimciils ol Hill (;i..i.t I nmi i .5M, Jiilv 24, 1961 9 , .5DA, Jiil) 1.5, 1961 9 , .Sept. 26, 1961 9 , 5M, 9, Mfiisiircminls of Uilm ;iikI Cniiif (1961) ,

JiOLOGiCAL Series, \'ol. 4, Xo. 3, September, 1964 at the Nevada Test Site the species was gener- some are decidedlv colorful, thev are generally alh' found on \egetation other than Larrea, e\ en drab, less attractiv ely colored, the coloration be- in areas where Larrea was present. In all areas, ing generally protective. This coloration is main- however, it was characteristic of the so-called ly a combination of olivaceous, yellow and brown "desert pavement", the numerous small rocks of varying shades with ornamental touches of scattered about on the surface of the ground. In reds or blues. the recorded captures it was noted that the in- As with the other groups of the Acrididae, in sect was found at the tips of the shrubs during order to present a comprehensive picture of the the hottest part of the day and could best be subfamily, a study of the internal genital struc- obtained by capti^ire with the fingers, inasmuch tures would be required. Apart from these as many of the shrubs upon which it was found structures, the most reliable e.xternal features were spiny, and sweeping would end in failure are the form of the cercus and other terminal of capture. abdominal appendages of the male. In Mekino- pliis, one of the largest of the North Seasonal Occurrence. Adidts have been re- American genera of grasshoppers, a knowledge of these corded as earlv as April 7 and as late as October structures is absolutely essential. The females 8, with immatures found late in March and into of this genus are very difficult to classify owing April. At the Nevada Test Site the earliest adult to the variability of external characters. occurrence was June 10, with September 26 representing the latest date. The greatest activitv The important economic species which are of the species was in August. No nvmphs were found in cultivated areas and on the ranges are collected during the course of the studv. found in this group. Most of the economic species belong to the genus Melariaphis. Many Localities Represented. Specimens examin- are omnivorous, others are selective in food ed (adults): 9. habits, feeding primarily on dicotyledons, especially the perennial members of the plant family Studv 5.\, I specimen, Jime 10, on groimd, .\steraceae. The migrations and plagues of grass- in area of Larrea divaricata. hoppers throughout history were primarily due Study 5DA, 1 specimen, Julv 15, in area of to species belonging to this subfamilv. Larrea divarieata. Because of their seeming preference for cul-

Area 5, 2 specimens, Julv 24 and September tivated areas or meadows, the subfamily is poor- 26, vegetation unidentified. ly represented at the Nevada Test Site. This area is well Area 6, 1 specimen, .\ugust 15, on Coleo'^i/iie within the distribution of a number of ramosissinia. species that may never be found at the test site because of this preference, unless, of course, Studv CB, 1 specimen, .\ugust 28, C. on they are found in migration. This ecological raniosi'isima. factor undoubtedly explains the absence of such Study .\1CC, 1 specimen, August 12, vegeta- common species as Sehistocerea shoshone tion unidentified. ( Thomas ) , Melanoplui inexieaints ( Saussure ) Studv JA, 2 specimens, July 18 and August .\/. femur-rubrum (DeCeer), A/, paekurdi Scud- 19, on Ltjciiun andersonii. der. A/, differentialis nigrieam Cockerell, M. bivittatus (Say), and A/, ijarroui (Thomas).

It is possible that Ocdaleonottts borekii orien- SubfamiK Cvrt.\( anth.\cridinae tis Hebard may be found at higher elevations on

The species of this subfamilv, commonly the Nevada Test Site, but is not included in the known as the "spine-breasted" locusts, mav be key. It would fit the first couplet (with Mela- easilv recognized by the presence of a conical nophis), but is distinct from that genus by the or cvlindrical elevation, termed the prosternal presence of distinct lateral keels on the pro- spine, projecting from the prosterum. The face notum. The species was described from speci- is usuallv vertical, the head decidedly roimded. mens taken at Lee Canyon, a few miles from The tarsal pulvilli are exceptionallv large, a the Nevada Test Site. feature correlated with the characteristicallv The key presented is a very artificial one, but plant-loving habitats, and a well-developed will be (juite adequate for the few species found arolium is present between the tarsal claws. in this area. Anyone attempting to utilize the key These insects are customarily found on should be cautioned by the variability, both as vegetation, in sharp contrast to the soil-frecjuent- to color and as to the development of the wings ing habits of most of the other acridids. Although of individual species. )

36 Bkiciiam Vmsr. L'nivkhmtv Science Bulletin

Key to the Genera of Cybtacanthacridinae

1. Siil)^t'iiital pl.itf ol malf uitli a distinct siibapical tulxTcIc (Fig. -i): body color green (or grcenisli) or h.iff, usualls with contrasting colors on pronotum or tegmina 2

Siihgeiiital plate of male without a subapical tubercle (Fig. 25); body color dark. no\er as abo\e; tegmina and wings variously developed from completely alate to reduced to small nonrniictional pads Melanuplus Stal

2. Bod\- \ariouslv marked witli coutrasting red and yellow; tegmina and wings projecting beyond the caudal femora by at least the breadtii of the tegmina Poecilolctlix Scudder Body not marked with contrasting red and yellow; tegmina and wings variously developed from short nonfunctional pads to completely alate 3

3. Body bright green (occasionally lighter) with dorsal white stripes on pronotum and white lateral patches on thorax; tegmina bluish-green, with very narrow white stripes Hesperotettix Scudder

Body uniformly colored, buff or tan, without light stripes, or if present never bright green. (There mav be a dark median dorsal stripe and postocular stripes on head and pro:iotum, but not as in alternate. Aeoloplkles Caudcll

FIG. 24 FIG. 25

Figs. 24-2.5. 24, Hesperotettix viridis nevadensis, mile, ape.\ of abdomen, later.il \icw. 2.5, Melanoplus complanatipes canonicus, male, ape.t of abdomen, lateral view.

Genus Acoloplidcs Caudell

Genus Acoloplidcs Caudell" 1915. Acoloplidcs Caudell. Proc. U.S. Nat. Mus.. 49, p. 2S.

Ke\- to the Species of Aeoloplides

(Modified from Wallace, 19.5,5.)

Ventral basal wedge on hind femur of male projecting ventrad for a distance appro.\imateIy

etjual to its width at base when viewed laterad (Fig. 26); tegmina e(|ualling or surpassing the abdomen and/or hind femora A. Iciuiipcnnis (Scudder)

Ventral basal wedge on hind femur of male projecting ventrad for a distance less than its

width at ba>.e when viewed laterad ( Fig. 27); tegmina not reaching the tip of abdomen and/ or hind femora, sometimes not much longer than pronotum A. minor (Bruner)

FIG. 27 FIG. 26

Figs. 26-27. 26, .Aco/o/j/k/c.v Icmiipcwiis. m.ile. e.iiid.il femur, l.itcr.il view. 27. A. niiitor. ni.ilc r.iiul.il Iriiiiir, lateral view.

"The name Acoloplidcs wa.s proposed by C.uidell to replace the name Acoloplus of Scudder, now considered lo be a synonoym of ^^clalU)plus. See Wallace, 1955, for the historiciJ development involving the name change. Biological Series, Vol. 4, No. .3, September, 1964 37

Aeoloplides fenuipennis (Scudder) the ventral basal wedge 0.31 mm; breadth of the (Figure 26; Table 13; Map 4) \entral basal wedge 0.48 mm. The a\erages of seven female specimens are as follows: length Proc. U.S. 1897. Aeolophts tenuipeiinis Scudder, of the \entral basal wedge 1.78 mm; breadi of Nat. Mus., XX, p. 68. the ventral basal wedge 0.42 mm.

Established Synonomy. Aeoloplus arizone- A comparison of the Nevada Test Site speci- nsis Scudder; Aeoloplus ociihitus Scudder. mens was made with typical tenuipennis determined by Wallace and used in his revision of the Distinctive Features. This species, supposed- genus. The specimens compare favorably except ly, has tlie ventral basal wedge on the hind in the most important external character, the femora extended more than in minor, and the ventral basal wedge of the caudal femur. tegmina of both se.xes are "almost always" sur- Coloration. The predominant color of this passing the tip of the abdomen. Wallace (1955) insect is light greenish- states that the "xentral basal wedge of hind gray with a yellowish or femur well developed in both sexes, in male yellow tinge, older individuals being darker and usually projecting ventrad for a distance about brownish. A median dorsal brown stripe extends from the occiput of the head over the pronotum; twice its width at base in lateral view, in female light usually projecting about two-thirds of width at a sti-eak, the same as the ground color, is usually present in the center, starting as fine base in lateral view". This statement is incorrect, a line at the anterior margin of or inasmuch as, even in typical forms, the ventral the pronotvim, occasionallv beginning at the basal wedge projects onl\' approximately equal transverse incision of pronotum. to the basal width of the wedge. In his illustra- the A postocular stripe is sometimes faintly indicated on the head and sides of tions Wallace shows tliis to be about equal. the pronotum. Tlie tegmina are usually light Size Variation. The length and breadth of the olive-gray, often with small, indefinite dark ventral basal wedge of the caudal femur is diffi- inaculations. The hind femora contain three cult to measure. Comparative measurements broad indefinite bands on their external surface; were made, however, using a standard grid the hind tibiae, in specimens from the Nevada micrometer in a stereoscope. The averages of Test Site, are light mauve, this color becoming five male specimens are as follows: length of less intense dorsally and distally.

Table 13. Size variation of Aeolopides tenuipennis. Ri(ii;iiAM Vdlng L'mvkhsitv Science Bulletin

The nvmphs, represented only by later in- minor from N'almes'. Ne\ada. have the tegmina stars from tlie Nevada Test Site, are often diffi- slightly surpassing the tip of the abdomen in the c lilt to tell from llcspcrotcltix t iridis nymplis, in- male and nearly that long in the female. In asiniieli as tliey are often briglit green, with or minor the length of the tegmina is not clearly without a pronotal stripe, or more freipiently tan, corri'lated with north-south distribution as it whieh color phase is also loiind in nymphs of is in tenuipennis. Specimens of minor from

tirspnolctlix. It is <|nite iinpraetieal to try to Lovelock. .\i'\;ida. have the wings sufficiently ciistinguish the n\niplis of tctniipcimis and minor. deselopcd for Might, uhile those from Wads- Additional eomiiients, I'speeially as to com- uorth. 5.S miles to the southwest, are brachy- parative fi-atnres and iiiori)li()logical \ariation, ptlTOUS. distrihntion, habitats, anti seasonal occurrence, "The ilifference in the yentr;il l)asal wedge are given under the discussion ol Acoloplidcs of the hind femur, particularly in the male, ap- minor. pears to be dependable (except in the few available specimens of tenuipennis from Coconino Localities Represented. .Specimens examin- County, .\rizona, some of which have a femoral ed (nvmphs and adults): 3fi. \\edge similar to that of minor), but it is a small Studv CM, Cane Springs, 10 nymphs, June difference in degree of development. The shape 22 to August 22; 9 adults, June 17 ti) August 22, of the prosternal spine will separate most speci- all on Alriplcx canc.sccii\. mens of these two form.s ( including the Coconino

Stiidv IF, I nv-mi-)]!, |unc 19, on SaLsola CountX' specimens ) but it is somew hat \ ariable kali. and cannot be used as an absolute criterion.

Area 5, 1 nvmph, July 1; 8 adults, June 20 to The width of the intermetasternal space of the September 26, vegetation unidciitilicd, hut pro- female presents a small difference which is not bably on Atriplex coiifciiifolid. too dependable. The differences in the phallic structures of the two are slight Studv 5K, 2 n\mphs. May 27 to June 15; 4 forms and not dependable, though small but constant differences adults, June 17 to Jul) 14, on Atriplex conferti- e.xist between all of the other species of the folia. genus. Further collecting in the areas of contact Area 3, 1 nxinph. .\ugust 15, on Atriplex of overlap of the ranges coiifrrtifolid. of these two forms may demonstr;ite the regular occurrenc-e of specimens internu'diate in all of these characters. The few Aeoloplicles minor (Bruner) specimens a\;iihd)le from these areas indicate ( Figure 27; Table 14; Map 4 ) that the txvo forms are distinct. Specimens of 1904, Aeoloplm- minor Ikuner. Colorado Agri- tenuipennis from Cima in California. Beatty in cidtural K\p. Sta. Bull. 94, pp. 60-61. Nexada and Zion National Park in Utah are typical of the species xxliile sfx>cimens from the ad- Established Svnononiv. Aeoloplu.i eremiO' jacent localities Cuchenberry Ranch in Calif- philo llebiU'd. ornia. Spring Mountain in Nex'ada and Pipe Ciomparative Features. Of this s[K'cies. Wal- Springs in .\rizona are typical of minor. The

lace (1955) states that it resembles tenuipennis Coconino County popidation of tenuipennis is

])ut is usuallv much smaller and the tegmina separated from minor by the Crand Ciuixon so

seldom reach the tip of the abdomen ( usually interbreeding cannot occur. In any case, the two much shorter) and ;ire e\'enlv narrowed to a forms are certainK' distinct enough to be re-

rounded apex. The following st;itement is made tiiined ;is subsjx'cies, exen if thex' should be

in its entirety from Wallace's revision of the determined to be conspecific.

genus: "Tinkham (19.38. p. ,347) referred to 'Aeolo- "The difficultv encountered in constructing plus tenuipennis tenuipennis Scudder' indicating that portion of the kev which separates tcniii- that he recognizi'd at least two subspecies of penni.s from minor emphasized the possibilit\' tcmiijxnnis. llelxud, in notes in the possession

that minor is a subspecies of trniiipcnni.s. Length ol r. II. Ilubbell. listed under Aeoloplus the of tegmin.i. the Tuost obxioiis tlistinguishing form 'tenuipennis minor Bruner 1904' with the char;tcter. is not \('rv s;itisfactorv for chagnosis uncompleti'd comment: 'Reduced to race by in this genus. In tentii- ' most . . of the specimens of . . .\ se;irch of the literature has failed to pcnni-t from Williams in C^oconino Co.. .Arizon;i. reveal th;it minor has been reduced to sub- iuid northward the tegmin;i rciuh \'er\- slightK' s|ietific rank in a public;ition. Until more mater- short of the tip of the ;d)domen. though longer i.d from the areas of cont;ict or oxerlap of these

(egmina are tvi)ic;>l of the s[)ecii's. Specimens of two tornis is studied, it seems Ix'st to retain Biological Series, \'ol. 4, No. 3, Septembeh, 1464

minor as a distinct species, which, in truth, it Co., .\rizona, near Pipe Springs, and Delta Co., may be." Colorado, at Delta, the type locality! In collecting at the Nevada Test Site, and It is interesting to note that the tvpe locality in a study of the entire collection from that is so far distant from the Great Basin confines. area, I find it difficult to justify conclusive Specimens before me, not pre\ iously record- evidence of typical tcnuipennis. Specimens have ed, that seem typical of minor are from VVashing- been found with long tegmina, surpassing the ton County, Utah (Shivwits Indian Reserva- abdomen bv more than their own width, a sup- tion, September 9, 1959, Andrew H. Barnum and posed characteristic of tenuipennis. The abdo- one mile north of St. George, June 10, 1959, men as a comparative structure in size, especially Andrew H, Barnum); Sanpete Countv, Utah in Aeoluplides males cannot be relied upon as a (Palisade Park, August 6. 1961, Andrew H. criterion because of the usual nature of the up- Barnum); Grand County, Utah (Westwater, turned abdomen, and the pwssibilitv of stretch- August 9, 1953, Andrew H. Barnum, and Ruby ing the abdomen in the female. Canyon, on the Colorado River near Westsvater, August 1.3, 19.50, Andrew H. Barnum); and Mesa Size Variation. The averages of the ventral Countv, Colorado (Grand Junction, Julv 26, basal of caudal as follows: wedge the femur are 19.5,3, and August 12, 19.56, Andrew H. Barnum). three male specimens length 0.2 mm, breadth The distribution of tenuipennis, by contrast, 0.4 mm; three female specimens, length 0.15 is given bv Wallace as being south and east of mm, breadth 0.47 mm. minor, forming a semi-circle in distribution around that form. Coloration. The color is ver\' similar in all parts of the insect to tenuipennis except that the Habitats. At the Nevada Test Site, Aeoh- dark postocular stripes on the head and pronotum plidcs is associated with the various species of are sometimes well defined and of the same Atripk'x. No absolute correlation could be made color as the median dorsal stripe. The dark with reference to the two species. Generallv, spots on the sides of the first two or three abdo- A. minor frequents Atriplex confertifolia. but is minal segments are present in about three- found with regularity on Kochia iiinericutui in fourths of the specimens (according to Wallace). the Kochia-Atriplex association. On the other Darker specimens of minor are more common hand, A. tcnuipennis is most frequently encoun- than lighter specimens. tered on Atriplex canescens.

Distribution. The distribution of minor is The nymphs are easily obtained bv sweeping given by Wallace as being well restricted to the the vegetation, but the agility of the adults makes Great Basin to include e.xtreme southwestern them more difficult to collect. The adults were Idaho, southeastern Oregon, northeastern Cali- never common and could not be obtained in fornia, and southeastern California across the larger series. state line from Las Vegas, Nevada, all of which The adults are fre([uentlv encountered near are within the confines of the Great Basin. Ac- the ends of the stems of the shrubs, but when cording to Wallace's records the only specimens disturbed jump into the center of the bush. The from outside the Great Basin are from Coconino best method of collecting, where the insects are

Table 14. Size variation of Aeolopides minor.

rf. Bmk.iiam Voisc Univehsity Science BtuLETi.v

so uncommon, is to tramplt- the hush. The speci- No types have been studied to confirm these minis art- satisfactoriK' dislodgi'd, but are so opinions, however, or to test the hypothesis of wary tliat tlu'v rccovi-r to anollicr slinib nctv minor bi'iug an absolute synonym.

(jiiicklv. Tlii'N' cliaracti-risticallv jump, hut will not ht'sitati- to IK' short tlistain-cs. CJenus llcspcrolcltix Scudder

Seasonal Occurrence. Aeoloplides is typi- 1875. Hcspcrolcttix Scudder, Bull. U.S. Ceol. cally a mid-summer form and no dctinite cor- Surv. Ti'rr., II, p. 262. relation could he made between tlie occurrence of minor and Icniiipcniiis. Adult iiiiiior was Hcspcrolcttix viridus complex found only in July and August, wheri'as tcnui- 1872. Cdloptcniis tiridi-s Thomas, U.S. Geol. pcituwi occurred earlier and later. Smpn. Hep. Mont. .Adj. Terr., p. 4.50. Localities Repesented. Specimens i\aniiii- Kstablished S\nonomy. llcspcrotcttix fcstiv- eti (nxinplis and adults): 22. iis Scudder. Stud\ 6A, If n\inpiis, June Hi to August Hi; Distinctive Features. This group, as present .3 adults, JuK 12 to .Vugiist 16, on both Atriplrx confertifolid and Kocliiu americana. at the Nevada Test Site, consists of a single variable species which has been sul3di\ided into a Study IG, 2 nymphs. May 28 and JnJx 19; 2 number ol different subspecies. The adults are adults, July 10 and 12, on Alii})lr.\ conjcrti- distincti\c and should not be confused with folia. an\- other form from the area. Nymphs, however, Study ECH, 1 adult. August 11, on Atriplrx may be confused with Aeoloplides, as certain cancscens. color phases of Aeoloplides are suggestive of Additional Remarks. In a comparison of the Hesperotettix. serii's before little correlation total me there is Most of the specitni'us collected at the in wing length, a very unreliable character in the Nevada Test Site show an intermediate condition Orthoptera, in areas. is little especially arid There between two t\pical subspecies. Others are typi- correlation in the extension of the yentral basal cal of one subspecies. The Nevada Test Site is wedge of the caudal femur, and no correlation apparently an ;u-ea of intergradation where all in the width of the intermetasternal space of the three forms arc found. Tlie group could, con- is difference date female. Tliere only a slight in ceivably, be discussed as a single, variable of habitat. appearance and species. Subspecifie differences can not be dis- assignments herein on tlie txisis The made tinguished in any immature stage, and such im- of faulty. all of the aboxe cliaractcrs ma\' be The mature specimens are assigned only on the basis differences noted in morpholog\- ma)' l)e due to of adult collections. environmental differences. Study 6A, most The species can be recognized by the follow- characteristic of minor, is a very arid area of low ing morphological features and color pattern: Afriplcx confertifolid and Kochiii umcriaina. In profile, the face is noticeabU' slantinJ, These environmental situations are suggestive of es])eciall\' in males. The vertex is very narrow size sliortness sniallness of and of wings. betuei'u the eves, but expanded immediately in According to the complete series from the front of the eves. The median and lateral carinae Ti'st liave to Nevada Site which been assigned of the pronotum are absent, or nearly so, but the two species, little reliance can be given to a marked bv contrasting colors. The tegmina and single character. All of the specimens were at wings are variously developed according to the first considered to be minor, but on the basis characteristics given in the key to the subspecies. of Wallace's re\isionary study, after comparison The hind femora are elongate and slender. of this si'ries with specimens in the author's col- Morphological N'ariation. In its typical lection from I'tali and (lolorado, after compari- eoiulitiou the species is highly v .triable in size, son to original material designated b\' Wallace in brilliancy and intensity of marking, which is his stud\-, and after considerable hesitation, tlie largely if not entirely in keeping with the luxur- present assignments are inatle. ious and green through light vellowish-brown to A comparison of a\ailable specimens with brow n color of tlii' plants on which it lives. those from the ti-st site indicates that subspecia-

tion is presi-nt in the tcntiifHiinis-minor complex, C^oloration. .\ most striking color pattern

in which case Tinkham and llebard were cor- exists. It is bright green to greenish brown (in

rect in their subspecifie assiginnents, inasmucli as older specimens ) marked with thin whitish or

tfnnipcnnis has prioritN'. The N't'vada Test Site is vi-Ilowish longitudinal streaks. The median car-

an area of inti-rgratlation. ina is striped vv ith a whitish line which arises on )

Biological Series, Vol. 4, No. 3, September, 1964 41

Key to the Subspecies of Hesperotettix viridis

1. Distal margin of tegmen truncate, subequal to the length of the pronotuni (Fig. 28) H. viridLs tertnius Hebard Distal margin of tegmen acutely produced, never truncate, variable in length from short to

long' - - 2

2. Tegmen extending to end of caudal femiu-, or slightly beyond, its apex broadly rounded (Fig. 29) H. viridis viridis (Thomas) Tegmen generally short, subequal to the length of pronotum, often considerably longer,

its apex pointed (Fig. 30) H. viridis nevadensis Morse

FIG. 28 FIG. 29 FIG. 30

Figs. 28-30. 28, Hesperotettix viridis termius, male, head, pronotum, and tegmina, dorso-lateral view. 29, H. V. viridis, male, head, pronotum, and tegmina, dor^o-hileral view. 30, H. v. nevadensis, male, head, pronotum, and tegmina, dorso-lateral view.

the occiput, bordered on each side with a thinner the Nevada Test Site concurrently with nvmphs blackish line, viuiously developed and often ab- from June to September, with the greatest occiu- sent, and extending to the posterior margin of the rence being in late July and early August. pronotum. A whitish line, sometimes indistinct, arises on the margin of the occiput against the Hesperotettix viridis viridis (Thomas) comjxjund eye and continues along the humeral ( Atypical angle of the pronotum, and continues down the (Figure 29: Table 15; Map 5) humeral angle of the tegmen. A third white line Morphological Variation. This species is not generally extends from the compound eyes across found in the typical condition at the Nevada the lateral lobes of the pronotiim. The typical Test Site, but is intermediate between viritlis and specimen is marked with black or dark brown on nevadensis. Because some specimens are more the lateral lobes of the pronotvim between the representative of viridis. the group is included two light lines. The light coloration is variously herein. developed on the thorax and caudal femora. The intermediate viridis-nevaderisis forms The nymphs of the species are generally collected are slightlv smaller than typical viridis marked with the single dorsal white line. and lack the pink coloration of the hind femora,

Distribution. The species is characteristic- which color character is tvpical of the viridis ally a western form, common from the Great c-ondition. The black markings on the lateral Plains to the Pacific Coast. It is recognized as lobes of the pronotum are variously developed. subspecies or geographic varieties throughout its Distribution. Typical viridis is absent from range. most of the desert portions of the southwest, but

Seasonal Occurrence. Adults are found at is widespread, abundant, and generally distri-

Table 15. Size variation of Hesperotettix viridis viridis.

t ^r= § 2 J a. ^6

^, 3CH, July 22, 1961 (small) )

42 liKII.IIAM VoL'NC UnIVKIISITV SCIENCE BULLKTIN

billed over the Croat Plains and al liiglior flo\'a- i\i'ly more decided, and is in turn supplanted fions (if the southwest. by the much more local and less numerous

Thesi' iiitei'ine(h'ate hiiiiis were colleeted in liiidi.s Icniiiw^ in the extensivi- desert areas of onl\' tme ari'a at the Nevada Test .Site, and all large Wfsteru |iortions of the state". nymphs loileetcd hen- were assigned to (Ills II both races are indeed found at the Nevada sanii- iiileniiediate group. It is impossiliie to Test Site the same comments with reference to distingnish any difterenees between (lie iivniplis numbers and distribution apply. colleeted over tlie test site, and all other F.(Halilies IU|)resented. Specimens exiunin- nymphs have been assigned to either tcmiius ed (n)-mplis ami adults): 19. or nectulcnaifi on the basis of adult collections. Study 6CR. 2ailults, September 19, on Cairo-

Loculities Represented. Specimens examin- fltjni' rtiniosissiina (not a natural host, but prob- ed (nymphs and adnlts): 25. abh' an accidental occurrence).

Study .3CJII, July 22, on ('linisotliniiniiis li.sci- Study T.\. 13 nymphs and adnlts. June 23 (Hflorus and Tctnuhjmia sp. to August 31, on ('liii/sotliatimus liscidiflonis.

StiuK- 3("C;, 1 adult. June 27, \egetation not clctenniMrd. Hrsj)ci(}liilix riiidi.s uctdclcitsis ,\h)rsc Stu(l\ I'.M, 2 adults. July 2-4, vegetation not (iMgurcs 21, 28; Table 16; Map 5) (li'Icrmined, probabK Cliiysathammtfi viscidi- 190.3. Ucs'pcrotellix nevailcnsis Morse, Ps\che, flonts. X, p. 115, Study C;B.\. 1 iidult. July IS. on Clinjso- fhainniis liscidiflonis. Ivstahlished Syiumoiny. Ilehard in 1931 eslahlislicd //. <^illctlri Hruner as a synonym. //.

Sciidder is also a cuilipcnnui Henderson nee llcspciotctlix viridis nevadensis (Atypical) synonym. Two males (measurements given below with Morphological Variation. Specimens from ncvddouis) are considered intermediate be- the Ne\ada Test Site which have been assigned tween nccadcmis and fennitis, in each case there as typical iwvadcnsis show a reduced condition being an "in between" condition of the pointed of the legmina and wings. t\pical tein;de One and truncate apex of the tegmina. In one speci- was collected in copula with an interinetliate men a variability was noticed in each tegmen twvddciisi.s-tcntiiiis male. The j)resent series is on tlie same specimen, one tegmen being t\ pi- indistinguishable from those marked intermedi- ca! ol e.icli subsjM'cies. ate viridus-ncvadensis except for the short tegmina and wings which are typically pointed at the apex. Ilcspciotrtli.x viridis fenniiis Hcbartl

(I'igurc .30; Table 17; Map .5) Distribution, .\ccording to llebard (1920a) the present race, in the state of Utah, "supplants 1918. Jlcsperotcllix ncvademis tennius llebard. virUlLs- ciridis wiiere aridity has become progress- Trans. .Amer. I'aitom. Soc., XI.IV, p. 16.3.

Table 16. Size variation of llcspcrotettix viridis nevud mui.

cf , 6CR, September 19, 1961 (typical) 12.5

Tiiblc 17. Sizf variation of llcsperotctiix ciritlis termius.

a Biological Series, Vol. 4, No. 3, September, 1964

Melanophts aridiis (Scudder) Localities Represented. Specimens examin-

(Figures 31, 32; Tabic- 18; Map 6) ed { nvmphs and adults ) : 10.

1879. Pezotettix aridus Scudder, Proc. Bost. Soc. Studv TA, Midvalley, 2 nymph females, 1 Nat. Hist., XX, pp. 84-&5. adult male, August 17, on ArtemiMa tridentata.

Distinctive Features. Brachypterous, the Study TCB, 1 adult female, October 14, tegmina and wings shorter than the pronotum, vegetation not recorded, probably on A. tri- the tegmina reduced to short oval pads. An- dentata. tennae of the male conspicuously long. Posterior Studv CNL Cane Springs, 1 adult female, margin of the pronotum subtruncate. Cerci of September 30, vegetation not recorded. male very slender, elongate. Furcula small, Stvidy JA, Jackass approach, 1 adult male, bluntly tipped, appro.ximately one-fourth the 1 adult female, in copula. October 15, on length of the epiproct. Coleogyne ramosissima.

Coloration. Brownish flavous, marked with Studies 5C and 5M, Frenchman Flat environs, black. with a narrow mesial black stripe Head I adult male, September 3, 2 adult females, broader postocular band, continuous, but and a September 26 and October 3, vegetation not re- generally interrupted on the lateral lobes of the corded. pronotum. Hind tibiae glaucous, changing to bluish gray or brownish in dead specimens. Additional Remarks. This species is only tentatively referred to M. aridus. One male and Distribution. This species is widely distri- one female were sent to Dr. Gurnev at the U.S. buted throughout the southwest from California National Museum for confirmation, who extract- to Te.xas from lower elevations in the Lower ed the genital complex of the male but found Sonoran deserts to high altitudes (the Canadian that the tips of the aedcagal valves had been zone to at least 9400 feet, according to Ball et al. broken off. Therefore, the correct placement 1942). At the Nevada Test Site it found was could not be made. widely distributed, though uncommon, in Mid- This may or may not represent a different valley, at Cane Springs, Jackass approach, and species of the aritlus group or a new species. A in the Frenchman Flat area. further study of the group will have to be made Habitats. At the test site the species was to determine the correct placement. found on Arteinma tridentata and Coleogijne ramosissima, the only recorded vegetation. In other areas of distribution it could have been Melanoplus complanatipes canonicus Scudder associated with Larrea divaricata and vegetation (Figuri-s 21, 25, 33; Table 19; Map 6) associated with Gratjia-Lijcium. 1897. Melanoplus canonicus Scudder, Proc. Seasonal Occurrence. Adults were collect- Amer. Phil. Soc, 36: 26, 34. ed from August 17 to October 15. Only two nymphs were collected, both late instar females, Distinctive Features. Tegmina and wings on August 17. moderately slender and gently taj>ering, extend-

Table 18. Size variation of Melonaplus aridus.

n Bhicham Vounc Univebsity Science Bulletin ing In-yond tlie apices of the hind femora. is frequently associated uith the vegetation on Caudal margin of pronotiim obtiise-angulat<', and near sand dunes. MH'diaii carina distinct on the metazona, more Habitat. This group was known earlv in (il)solfte on the pro/.ona. (,'erci of male slender, the literature as a sagebrush inhabitant. In this narrouing on basal third, the middle third nar- i'n\ironment the gray and rusty colors harmon- rower, then expanding to a nearly equal slight- ize with its surroundings making it extremely ly spatnlate tip. Fnrcula of male very broad on difficult to detect when at rest. basal third, tapering to parallel appendages, two- It was found in only two areas at the Nevada thirds as long as the epiproct. Test Site, both ha\ing a different etnirotunent. Morphological Variation. There is consider- It was common, though not abundant, in the able variation in the production of the tcgmina sand dune area, where it was collected only bevond the abdomen, especially the females, in after persistence because of its remarkable ability the series examined from the Ne\ada Test Site. of flight and escape. The other habitat was in It should be re-emphasized that the use of the the very heavy vegetation (Rumex) at Cane abdomen as an organ for comparison is ques- Springs, where it was cjuite common but difficult tionable because of the stretching or contracting to collect because of its escape into the tall t-dt- of that organ. A comparison of two extremes tails when disturbed. tegmina projecting 1.1 showed the 18.3 mm long LaRivers (1948) gave some details of the 21.0 mm beyond the abdomen and the mm habits and habitats that compare favorably with long projecting 4.6 be\()nd the tegmina mm obser\ations from the test site: "In addition to abdomen. sagebrush I have taken it on Chrysothamnus, C^oloration. Brownish fuscous, sometimes Oryzofysis hytnemndes, Dalea pohjadenui and several with a ferruginous tinge, more or less feebly other utudentitied succulent sand dune flecked with obscure maculations. I'ostncular plants. While it has a considerable distribution ill sagebrush areas, I it black streak extends to pronotal lateral lobes ha\e found in swarm- ing conditions only \icinit\' between transverse incisions. Mind femora in- in the of sanded regions, particularly acti\e distinctly marked by two black bands, strong dunes. W'hen disturbed, dorsallv, with some red on inner face of caudal coinpkinotipcs flies to on adjacent bush or plant, it non-stop if distance is femora. Hind tibiae very pale glaucous. making the only a matter of a few feet; if the distance is The nymphs are characteristicalK' colored greater, the insect usually alights on the ground and can be recognized easily b\' the conspicu- and almost immediately flies off again to sanctu- ously striped pronotum. The median carina is ary long before the collector can get within outlined with a light stripe, the lateral carinae disturbing range again. At several sand dune are outlined with dark stripes which extend onto localities, large series could be obtained by the head. merely sweeping a net rapidly in front of the Distribution. This species, described from collector while walking through the insects." It the (Jrand C^anvon of Arizona, is common in the might be noted that nowhere at the Nevada Test sagebrush areas of the Great Basin Desert. It Site was the race that numerous.

Tubii- 19. Size variation of Melanoplus complanatipes canonictis.

SQ .

Biological Series, Vol. 4, No. 3, September, 1964

Seasonal Occurrence. The earliest occur- Coloration. This beautiful, long-winged rence of the insect was May 27 (nymphs). species will not be confused with anything else Adults were found from June through October on the Nevada Test Site. The general body color 14. Nyinphal instars were found as late as Aug- is yellow to olivaceous, the long tegmina pale ust. The insect is most abundant during August greenish, and the posterior tibiae dark bluish and September. green. The body is conspicuously marked with contrasting red, one median line on the head and Localities Represented. Specimens examin- along the median carina of the pronotum; the ed (nymphs and adults): 71. posterior margins of the lateral lobes of the Study 12CF, Kowich Valley approach, 3 pronotum are outlined in red. Liirge red spots specimens, August 12 and August 21, on Ar- are present at the humeral angles of the prono- temisia tridentata. tum, variously arranged on the head and on the Study EGA, sand dunes, 35 specimens from femora. The metazona of the pronotum is con- August 11 to October 14, on the various plants spicuously punctate with black spots, especially growing on the sand dunes. noticeable in the females.

Study CM, Cane Springs, 33 specimens, from Distribution. This species is an inhabitant May 27 (nymph) to August 22 (late instar of the southwest, being found in the Lower and nymphs and adults), on Rtiinex sp. and Ttjpha Upper Sonoran life zones of California, Nevada, domingetisis. Utah, and Arizona. The type locality is Brad- shaw Mountain, Arizona. Genus Poecilotettix Scudder At the Ne\ada Test Site it was found widely distributed at lower ele\ations was collect- 1897. Poecilotettix Scudder, Proc. U. S. Nat. and ed from four areas. Mus., XX, pp. 385-386. Habitats. In keeping with the colors of the

Poecilotet^x sanguineus Scudder insect, green \egetation is the characteristic (Table 20; Map 7) habitat of the species. It was recorded from 1897. Poecilottettix sanguineus Scudder, Proc. Hijmenoclea salsola, Coleogyne ramosissima, Chnjsothainnus U. S. Nat. Mus., XX, pp. 387-389. viscidiflorus, and Ephedra viridis. C. ramosissima is the only non-green shrub Established Synonomy. This species is also listed. No preferred vegetation could be deter- P. longipennis recognized as (Townsend) mined because of the uncommon occurrence of ( Dactylotum longipennis ) = Townsend the species at the test site. Distinctive Features. The species shows a Seasonal Occurrence. The earliest collect- general relationship to Mekinoplus, but can be ing date for adults was June 19; the latest Sep- distinguished not only by the bright colors but tember 8. No nymphs were collected. morphologically by the tuberculate abdomen. In this respect it resembles Hcsperolcltix, which Localities Represented. Specimens examin- genus has a subapical tubercle. In Poecilotettix ed (adults): 10. the tubercle is apical. The prosternal spine is Study IG, 5 adults, June 19 and July 10, on very slender. C. viscidiflorus and E. viridis.

Table 20. Size variation of Poecilotettix sanguineus.

•^ Bhh:iiam VoLNf; University Science Bulletin

Study 4A, 1 adult, August 11, on H. salsola. the species from the Nevada Test Site. These are the conspicuous grasshoppers of every-day Study JA, Jackass approacli, 3 adults, July occurrence. They are well represented in a IS and SeptcrnlKT 8, on C. mntosissimti. desert environment and are common c\ery- Studv CM, Cane Springs, 1 adult, August 3, where. //. sdlsold. on .\lthough general and specific collecting has l>een rather extensive at the test site only two- Siibfamilv Acridinae lliirds of the expected species have been found. This subfamily, as now recogni/od, IikIikIcs In addition to those species reported here as definite records and included in the keys, the Acridinae (or Tryxalinac ), the so-called "slant- faced locusts," and the Oedipodinae, the "hand- following species mav be fovmd that are not uingi'd locusts." These two groups were until iMciiidcd: subfamilies, dis- recently considered as distinct Orplnilclla coinpta Scudder, a form of the the basis, primarily, of the cliar- tinguished on Lower Sonoran life zone, although found com- face in lateral view slant- acters of the head, the monlv in moist areas, should be presi'ut along sub-perpendicular, a characteristic that ing or .some of the foothills, and higher grassv areas; has alwavs lead to confusion in trying to separate Aiilocara elliotti ( Thomas ) common in short the genera; the characters of tlie pronotum, the grass o\er much of its range; Dussosteira sfnircata presence or absence of distinct median and later- Saussure; Trepkluhts rosaceits (Scudder), pre- reliable for most genera, fail al carinae, while \iouslv collected in the \icinitv of Las Wgas, completelv as a criterion in soiue; the intercalary Nevada; Conozoa sulcifroris stilcifrans { Scud- \ein of the tegmen, used as a characteristic, leads der) and Conozoa wallula (Scudder); Circo- coloration of the wings, and to difficulties. The tcttix crofcdttni Rehn. described from Lee Can- been useless in the all other characters have von, Nevada, perhaps present at higher eleva- absolute separation of some genera. tions at the test site; Cratypedes negJcrttis The recent emphasis of the internal genital (Thomas) should be found at some of the high- structures has proved the close relationships and er elevations; Xcnicri.s inininiiis (Scudder). one has led to the grouping of these two subfamilies of the smallest acridids; Coniana snowi Caudell, under one, the Acridinae. found onlv on sand dunes or in the vicinitv of As now recognized, the sublamilv includes a dunes; Ccininulu pclluckia (Scudder), while not great varietv of species showing a diverse ex- a desert species, mav be encountered at some ternal morphology, and includes the majority of of the higher elewitions.

Key to the Genera of .\c.-ridi.\.\e

1. Face stronglv receding so that the \ertex is angled sharply with the front of the face (Fig. 34), to the extent that the lateral foveolae of the vertex are iinisible dorsad (Fig. 35) 2

Face more or less vertical, the vertex broadly rounded into the face (Fig. 36), the lateral foveolae of the vertex, if present, visible dorsad (Fig. 37) 5

2. Antennae ensiform; lateral carinae of pronotum parallel or subp;irallel (Fig. 38) Eremiacris Heb-

ard .Antennae simple, if slightlv flattened the lateral carin.ic of pronotum curved 3

3. Head distinctly elevated above pronotum which is strongly scUate (Fig. .39) Bootettix Bruner Head not distinctly elevated above pronotum wliiili is tl.it or nearly so 4

4. Lateral carinae of pronotum parallel or subparallel; antennae simjile; fastigium of vertex with surface largely convex, lacking a conspicuous infra-marginal impression (Fig. 35)

^ Ampliitonuis McNeill

Lateral carinae of pronotum well indicated in ailor, divergent in middle of pronotum; antennae subensiform; fastigium of vertex with surface largely concave, with a conspicuous infra-marginal impression (Fig. 40) CordiUacris Rehn

5. Hind wings not brightlv colored, never m;uked with a conspicuous black band; median carina of pronotum nt)t stronglv elevated 6 black Hind wings brightlv colored, red, vellow oi liliu , ,uid/or marked with a conspicuous baud; median c;irina of pronotum variously cK\atcd, low to strongly carinate 10 Biological Series, Vol. 4, No. 3, Septemder. 1964 49

6. Internal apical spines of caudal tibiae unequal in length (Fig. 41); lateral foveolae of the vertex well marked on all sides (Fig. 42) 7

Internal apical spines of caudal tibiae subequal in length; lateral foveolae of the vertex not well indicated 9

7. Costal field of the tegmen broadly expanded by one series (in males) or by two series (in females) of enlarged hyaline cells (Fig. 43) Ligurotettix McNeill

Costal field of the tegmen not broadly expanded as in alternate 8

8. Lateral carinae of pronotum continuous and sharply constricted in middle; prozona shorter than metazona (Figs. 36, 37) Psoloessa Scxxdder

Lateral carinae obsolete on prozona (Fig. 42); prozona longer than metazona At'eneotettix McNeill

FIG. 35 FIG. 36 FIG. 37

FIG. 38 FIG. 39 FIG. 40

FIG. 41

Figs. 34-4.3. 34, Amphitornus colowdus ornutiis, male, head and pronotiim, lateral view. 35, A. c. ornatus, male, head and pronotum. dorsal view. 36, Psolofstd (Iclicatuhi dclicatuhi, male, head and pronotum, lateral view. 37, P. d. dcticatula, male, head and pronotum, dorsal view. 38, Eremiacris fudUdo, male, head and pronotum, dorsal view. 39, Bootcttix inmctcitus, male, head and pronotum. lateral view. 40, Corddlacris occipitalis cincrcii, male, head and pronotimi, dorsal view. 41, Ligurotettix coipnlletti cuntutor, male, distal tibia and tarsus of caudal appendage showing internal apical spines, lateral \iew. 42, Agencotettix deorum deorutn, male, head and pronotum, dorso-lateral view. 43, L. c. cuntutor, male, head, pronotum, and tegmen, lateral view.

9. Occiput with a series of transverse carinae medio-caudad of compound eyes, the fastigial impression elongate in both sexes; metazona of pronotum not greatly expanded (Fig. 44) Cibolacris Hebard

Occiput with a series of transverse carinae medio-caudal of compound eyes, the fastigial impression broadly rounded in both sexes; metazona of pronotum conspicuously enlarged, much wider than prozona (Fig. 45) Anconia Scudder

10. Interspace of metasternum linear, or distinctly longer than broad in male, narrower than interspace between the mesosternal lobes in female (Fig. 46); median carina of pronotum distinct, intersected by one transverse incision (Fig. 47) Arphia Stal

Interspace of metasternum broad, quadrate in male, transverse in female (Fig. 48) 11 so Bmr.HAM VoiNG UNr\EB.siTY Science Bulletin

11. Median carina of pronotiim intersected by two sulci, the anterior one of which is shallow, lateral carinae long and intersected by the posterior sulcus (figs. 49, 50, 51, 52); form robust ' 12

Median carina of pronotum intersected by two nearly equal sulci; lateral carinae of pronotiirn distinct or not intersected by the posterior sulcus (Fig. 53); form slender 13

12. Median carina of pronotum conspicuous and well elevated (figs. 49, 51); wing disk red Xanthipptis Saussure Median carina of pronotum slight (figs. 50, 52); wing disk blue Lepnis Saussure

13. Lateral lobes of pronotum acutely produced (Fig. S3) Mestobregma Scudder Lateral lobes of pronotum rounded (figs. 54, 55), a small tooth may be present on lower lateral lobes (Fig. 56) ' 14

14. Lateral prominences present near median tariua of [ironotum, more pronounced in male (figs. 54, 57) Denrtmcina Scudder

Disk of pronotum without high lateral prominences near median c-arina (figs. 55, 56) Trimerotropis Stal

FIG. 52 FIG. 53 FIG 56

Figs. 44-57. 44, Ciholacris parciceps aridus, male, head and pronotum, dorsal view. 45, Anconia intcgra, male head and pronotum, dorsal view. 46, Arphia conspersa, male, metastemum and proximal abdominal stem ites, sentral view. 47, A. conspersa, male, pronotum, lateral view. 48, Trimcrotropis pallidipcnnis pctlli clipennis, male, metastemum and proximal alKloininal steniites, ventral view. 49, Xaitthippus coriillipes, male, pronotum, lateral view. 50. Lcpriis ^kucipcniiis, mali'. pronotum. Literal view, 51, .V. c. corcillipcs, male, pronotum, dorsal view. 52, L. gUnuipciiiiis, male, pronotum, dorsal \iew. .53. Mcstohrcgma impcxum male, pronotum, lateral view. 54, Dcrotmenia dclicatttlum, male, pronotum, lateral view. 55, T. p. /xj//i- dipennis, female, pronotum, lateral view. .56, T. strcmiii. male, pronotum, lateral view. 57, D. delicatulum male, pronotum, dorsal view. Biological Series, \'ol. 4, No. 3, September, 1964

Genus Eremiacris Hebard of color in the areas collected, the specimens do vary according to habitats. In the sagebrush area 1929. Ercmkicris Hebard, Proc. Acad. Nat. Sci., the insects tend to be more grav or brown; those Phila., 81:303-425. insects collected in and among grasses were rather consistently light. Some of the intermedi- Eremiacris pallida (Bruner) ate areas, between the two extremes showed both phases. (Figure 38; Table 21; Map 8)

Distinctive Features. A species of slender Distribution. This species is widely distri- form, small to medium in size, with the head buted throughout the southwest. Its known strongly slanted. The wings are variously de- range extends from north central New Mexico veloped, generally reaching to the end of the west to California. At the Nevada Test Site it abdomen. A poorly developed prosternal spine is one of the more common species and has a wide distribution. is present, which character led to the placement of the genus in the subfamily C\rtacan- Habitats. The species is most often associat- thacridinae until recent years when a study of ed with Oryzopsis hijmcnoidcs. It is \ery active the genital characters showed a true relationship and a strong jumper, especially in the nymphal of the group. stage, and is most difficult to collect, not only Coloration. The degree of color variation because of its agility but because of its color- in the species leads to the assumption that per- ation. It readily escapes in \egetation where it haps two or three species are actually found in is well hidden. Or if it happens to alight on the area. The color of the insect ranges from light the ground it is well concealed because of its tan or decidedly yellowish and yellowish-green resemblance to the small bleached desert sticks through darker. These different color phases to and t\\igs. some extent may be correlated with vegetation In the Cane Springs area the insect was most upon which the insect lives. Most of the s{>eci- common on Ehjmus ciiicretis and Distichlis mens show a marking of lateral bars on the head stiictua, the common grasses. and pronotum, which markings are suppressed in Seasonal Occurrence. Nymphs of Eremiacris the yellowish and greenish specimens. An oc- were collected from May 13 (where they were casional unifomi yellow or uniform light green is common in study area IBF) to mid-.\ugust (as encoimtered, and yellowish tegmina on a uni- sub-adults). Adults were collected in early June form light green body is found. In some of the and were present to October 4. They were most light brownish specimens there is a suffusion of common and numerous during and are a white on the head, on the pronotum and on the July, summer insect at the test site. Their numbers caudal appendages. All of the nymphs collected are maintained throughout the hottest months. showed a definite pallid coloration. The darker colors are apparently found only in the adult Localities Represented. S[)ecimens e.xamin- specimens. ed (nymphs and adults): 112.

Nearly all of the specimens collected at the Study TA, Midvalley, 22 nymphs and adults, Nevada Test Site were of the light, yellowish- June 22 to September 19 (most common in Aug- green phase; about twentv percent were of the ust at this higher station), on Oryzopsis lujmen- darker phase. Although there was no consistency cndes.

Table 21. Size variation of Ereiyiiacris pallida.

t )

52 Bkkmiam VoiNf: Univebsity Science Bl'lletin

Study 3CD, 4 ii\ niplis, June 27, on O. hijmni- pleura and limbs. The tegmina ha\e small black oules. dots. An occasional brownish specimen is found. Study fiCL, 5 adults, July 13 and It, on Bni- These two extremes (green and brown) corre- mils tectorum. late to till- Lcirrca divaricata, upon wliicli it is always found. This shrub varies from a deep green Studv 5A, 1 adult, August 31, vegetation un- to brown. ( See additional remarks under "Habi- rt'corded. tats".) Studies C;M and CB, at Cane Springs, 38 nvinplis and adults. May 27 to August 10, on Distribution. This insect is found wherever E. citwrcus and D. strictus. Larrea grows in the Lower Sonoran life zone of Miscellaneous Graijui-Lycium studies (IB, the southwest. M the Nevada Test Site the species collected wherever the shrub was IG, etc.), 10 nvmplis and adults. May 13 to Aug- was present, and was collected, even, in some areas ust 17, on O. Iii/mciu>i([e.s. where the shrubs are very scattered, not at all Miscellaneous mixed \egetation studies (JA, common. EGA, ECU, ACC, TCB), 32 nymphs and adults, It is com[)letelv restricted in habitat to the June 6 to October 4, vegetation, wliere recorded, creosote bush and the distribution is about ecjual O. hijmenoides. to that of the host plant within the borders of the United States, Genus Bootettix Bruner Habitats. Recognition is difficult because of 1889. Bootettix Bnmer, Proc. U. S. Nat. Mus., the rich olive-grL-en base color which blends so Xll, p. 57. completeh' with the foliage of the creosote bush. Tlie markings produce the effect of the silvery sheen of the seed capsules of the shrub. This is Bootettix punctatus (Scudder) one of the few species of insects strictly limited (Figure 39; Table 22; Map 9 to one shrub, and shares with Insara covilleac as

1890. Gtjttinrs punctatus Scudder, Psyche, V, p. being one of the two orthopterans found only on 440. Larrea. Larrea achieves a \ery deep green and dense DistiiK'tive Features. This insect is one growth under optimum conditions, especially which shows a strong slant to the face. A along the margins of roads where it recei\es distinct angle is formed with the vertex, and the more moisture than its neighbors away from the lateral foveolae form a right or acute angle with road. M the Nevada Test Site, as with other the plane of the fastigium. The head is distinct- areas of the arid west, its growth is very stunted ly elevated above the saddle-shaped pronotum. and it takes on a brownish foliage. Bootettix most The antennae are short and simple (filiform). commonly fretjuents the dense growth, but is The coloration is the most distinctive feature. also found on stunted, brown shrubs in few

Coloration. This is one of the most interest- numbers. Collecting indicated that the majority ing insects found in the southwestern United of specimens taken in the dense, deep green States. It has an unusual rich green coloration shnibs were predominanth' of the gi"een pluise; with silvery white or mother-of-pearl markings in the areas where the shrub was more brown in addition to brown and black on tlie pronotum. the incidence of the brown phase in the insects

Table 22. Size variation of Bootettix jmnctatus.

! Biological Series, \'ol. 4, No. 3, Septembeh, 1964 53 Bhiciiam Vounc Univeiisitv Science Bulletin

was very high, pointing out the vahie of protec- Anipliilornus coUnadus ornatus tive coloration as an advantage to the insect. McNeill The species is most commonly collected by (Figures 34, 35; Table 23; Map 10) sweeping the outer branches with a net. By 1897. Ainphilornus ornatus McNeill, Proc. actual comparison, ver)' few specimens were Davenport Acad. Nat. Sci., VI, 224-225. visually spotted in collecting before sweeping pp was resorted to, in some instances several min- Established Synonomy. Acentetus unicolor utes of close observation were retjuired to spot McNeill; Acciitctii.'i cariiuitus Scudder; Steno- a group of specimens. They characteristically rest hothrus biculor Thomas. at the tips of the branches on the upper side and Distinctive Features. The vertex of the head will frequently not even move when touched is a little declivent, advanced in front of the with the finger. Tlie insect is a very excellent eyes, the antennae are slightly flattened. The jumper and a good flyer and if removed from pronotal disk is well rounded and the lateral the shrub to the ground will very quickly regain carinae are extremely faint and not interfering the protecti\e confines of the shrub, either by with the rounded outline of the humeral angles. flight or a series of f|uick jumps. The median carina is distinct and accompanied The nyniphs are easy to recognize as of that by more or less distinct supplementary carinae species, inasmuch as they are colored like the on the disk. All of these carinae are intersected adults, but are also difficult to spot and collect by the posterior principal sulcus only a little or except by sweeping the branches. considerably behind the middle. Tlie posterior The stridulation of the males is distinct. angle of the disk is moderately rounded. The Seasonal Occurrence. The insect is a sum- tegmina are well developed. mer species. The earliest occurrence was June Coloration. The insect is dull brown with 15 (nymphs) and the latest date of collection of fine yellow bars on the sides of the pronotum. A adults was October 22. Adults occurred early in dorsal light stripe may be present on the pro- July, while n)niphs were present into August. notum and, if present, generally extends onto the The greatest activity of the insect was during head. Two black bars are present on the outer July and August. face of the hind femora. The posterior tibiae Localities Represented. Specimens examin- are bluish. ed (nymphs and adults): 320. Distribution. This race is widely distribut- Area 5 (5A, 5CQ, 5HP), Frenchman Flat, 280 ed throughout western North .America, in the nymphs and adults, 15 October 22. June to Lower Sonoran, Upper Sonoran and Transition Study 3CD, 16 adults, June 27 to August 15. life zones. At the Nevada Test Site its distribu- Studies JA (Jackass Approach) and CB tion was limited bv the grasses with which it is (Cane Springs), 23 nymphs and adults, June 24 associated and it was uncommon in all collect- to October 15. ing areas.

Study ACC, 1 adult, October 2. Habitats. Amphitomus was found in some of the same situations as Eremiacris, with Genus Amphitomus McNeill wliich it should not be confused because of 1897. Amphitornus McNeill, Proc. Davenport its different appearance and different habits. Acad. Nat. Sci., VI, p. 223. Amphitornus seeks the denser grass and tries to

Table 23. Size variation of Amphitomus coloradus ornatus.

cf , 6CL, Jiilv 13, 1961 cT, CM, Auguvt 22, 1961

(if, TA, September 19, 1961

9 , 12CF, August 12, 1961

9 , 12CK, August 21, 1961

9 , CM, August 22, 1961 )

Biological Series, \'ol. 4, No. 3, Septemuer, 1964 55 escape by concealment in the bunches instead cated in color, but obsolete or subobsolete in of tn'ing to escape by movement. The insect contour. A distinctive feature of the group is the loses its caudal legs very readily when picked up conspicuous infra-marginal impression (concave and care must be exercised in collecting good surface) of the fastigium of the verte.x. Tlie specimens. The insect was most common on tegmina and wings reach to the end of the Distichlis strictus (at Cane Springs), Eltjmiis abdomen. cincreus (at the approach to Kowach Valley) Coloration. This insect is buff colored with and Onjzopsis hymenoides (in Midvalley). brown markings. A dark brown stripe extends Seasonal Occurrence. No accurate seasonal from the posterior margin of the eve, widening occurrence can be given for this group because on the sides of the genae to the anterior edge of of the few numbers collected. The earliest adults the pronotum, then continuing across the pro- were collected, however, on July 13; the latest notum to encompass the upper lateral lobes. Im- on September 19. No nymphs were collected. mediately below on the head and lateral lobes of the pronotum is a cinereous area, the cinere- Localities Represented. Specimens examin- ous repeating on the lower half of the caudal feed (adults): 16. mur. These markings are bent abrupdv inward Study 6CL, 1 adult, record of the July 13, no on the disk of the pronotum. The tegmina ha\e vegetation with which it was associated. The dark brov\Ti and cinereous spots, giving the in- area is very rocky. sect a grizzled appearance. The posterior tibiae Study 12CF, the approach to Kowich Valley, are testaceous. 4 adults, August 12, on Elijmus cinereus. Nymphs are easily recognized by the pro- Study TA, Midvalley, 4 adults, September 19. notal markings, the ensiform antennae and the on Onjzopsis hijmeiundes. projection of the \ertex. In these markings it Study CM, Cane Springs, 7 adults, August 22, could be confused only with Psoloessa delicatida on Distichlis strictus. dclicatula (Scudder), the only other species of the Nevada Test Site with similar markings, but the groups can be recognized by the projection Genus Cordillacris Rehn of the head.

1901. Cordilktcrk Rehn, Canad. Ent., .XXXIII, One male and t^vo females ( from studies TA, p. 271. IBF and 12E) have a less maculate appearance, but with a dark brown stripe extending down the Cordillacris occipitalis cinerca tegmen half its length. Except for the color pat- (Bruner) terns the entire series is quite consistent. The caudal tibiae are testaceous, except in about half ( Figure 40; Table 24; Map 1 1 the males which have a pinkish cast. 1889. Ochrilidia {?)cinerea Bmner, Proc. U. S. Distribution. This race is widely distribut- Nat. Mus., XII, p. 52. ed throughout the Great Basin and east to the Established Synonomy. Cordillacris affinis Colorado Rockies and Arizona. It was common Morse. in most of the studies maintained at the test site.

Distinctive Features. A slender insect with Habitats. The species is found t)nlv in areas slanting head and subensiform antennae. The of short grass, especially Onjzapsis lujmenoides, lateral carinae of the pronotum are well indi- where it is well concealed. It is a fairly strong

Table 24. Size variation of Cordillacris occipitalis cinerea.

•^ )

56 Bkk.iiam VoiNG Umvebsity Science Bvlletin fixer and acti\c jumper and wlien disturbed will female. The face is nearly \ertical and rounded leave tlie spot to remain concealed in its new at the vertex. The distinct median carina of the location. It is difficult to collect because of its pronotum is continuous, though low, and inter- habits and markings. sected bv one sulcus. The lateral carinae are obsolete on the prozona, which is longer than the .-Kilidts been col- Seasonal Occurrence. have metazona. lected as earl)- as Ma) 13 and as late as August Coloration. The general color is dull brown 31. The greatest activity is in June. above, vellowish-white below, the tegmina Localities Represented. S[X'cimens examin- brown or gravish-brown, usualK' with numerous ed (nvmphs and ailults): 5S. small darker brown cpiadrate spots, these some- Studies IG, IF and IB, 28 specimens. May times contined to a median row. The sides of the 13 to July 12, on O. hijmcnoicles. head and pronotum ha\e black bars or spots. The caudal tibiae are bright coral red, with a Stud\- 3C"C, 1 specimen, June 27, on Bramtis tcctomni. white or lightened proximal ring. The antennae are conspicuousK white or light colored. Area 3, miscellaneous c-ollecting, 1 specimen, Julv 15, vegetation not recorded. Distribution. The insect has a wide distri-

Studv lOD, 1 specimen, June 14, on O. bution throughout tlie entire west, from the hi/inciioides. Cireat Plains and north central states to the west, and from Ganada to Texas. .\t the Ne\ada Test Study 12E, Rainier Mesa, 1 specimen, June Site in 26, vegetation not rec-orded. it was found onlv the intermediate vallevs of the Upper Sonoran life zone and into the low- Studv EGA, near sand dunes, 5 specimens, er Transition life zone. June 27 to .August 12, on O. hijmcnaides. Study T.\, Mid\alley, 21 specimens, June 22 Habitats. This is one of tlie most important to August 31, on O. htjmetioides. range and grassland grasshoppers in the west,

where it appears abinidantK' in some areas. It

feeds on grasses and other low plants. It is prob- Genus Ageneotcttix McNeill ablv more abundant at the Ne\ada Test Site

1S97. A':,ciwolcttix McNeill, Psyche. X'lll. p. 71. than the records indicate. It is difficult to c-ol-

lect imless it appears in large numbers, because

of its rapid movement, small size, and concealing Agciicotettix deorum dcorum pattern bv which it blends with the desert vege- ( Scudder tation. It is found on the ground where it is ex- (Figtire 42; Table 25; Map 12) ceedingly difficidt to detect. It is only associated 1876. Chnjsochmon deontm Scudder, Bull. Geol. with Ori/zi>ims hi/mcnoidcs. Geogr. Surv. Terr., II, p. 262. Seasonal Occurrence. Ageneatettix has a Established Synonomy. Aidocaru scntddcri summer appearance. It was first collected on Bruner; At(itix occidcnUdis Bruner; A

Distinctive Features. Tiiis species is short, Localities Represented. Specimens examin-

rather stout, the male distinctlv smaller than the etl ( adults ) : 8.

Table 25. Size variation of Ageneatettix deorum deorum.

tl to a)i

cf , TA, August 31, 1961 13.3 cC, TCB, Julv 16, 1961 cf, TCB, July 16. 1961 .31, 1961 9 , TA, Aupist

9 , ECA, Stpfcinlier 4. 1961 ;5(», 1961 9 . KCA, StptrmlH-r

9 , TA, Augu-st 17, 1961 Biological Series, \'ol. 4, \o. 3, September, 1964

Study EGA, sand dunes, 2 adults, September length to show the size and color variants of 4 to September 30. this forni.

Studv TA, Midvalley, 3 adults, August 17 to Distribution. Psoloessa shiires nearK' the August 31. same distribution as Ageneotettix, from the Great Study TCB, near Midvalley, 3 adults, July 16. Plains westward. It has a wide distribution over the Nevada Test Site.

Habitats. This species is found on the Genus Psoloessa Scudder ground associated with Ortjzopsis, Haplopappus,

1875. Psoloessa Scudder, Proc. Boston Soc. Nat. and probablv other perennial plants. It is a very Hist., XVII, p. 512. active jumper and is well concealed on the ground among the desert debris. When disturb-

ed it generally flies into or near the shrub or Psoloessa delicatula deUcatula grass. (Scudder) Seasonal Occurrence. The species appears ( Figures 36, 37; Table 26; Map 12 ) early in the spring and remains throughout the 1876. ScijUina delicatula Scudder, Bull. U. S. summer. The first specimen was collected on Geol. Surv., II, p. 263. April 15, the last on September 30. Its most abundant occurrence was in Mav. No specimens Established Synonomy. Psoloessa colaradcn- were collected during the month of Julv, and no sis Thomas; Stiraplcura decussata Scudder; nvmphs were collected during the course of the Stirapleiira teinticarina Scudder; Psoloessa (?) studw so there is no indication of whether or not eurotiae Bruner. tlie August and September specimens were from Distinctive Features. This insect closely re- a second brood. Agetwotettix, but distinguished sembles can be Localities Represented. Specimens examin- its larger size and the color of the caudal by ed (adults): 24. tibiae. In Psoloessa the tibiae are pink with no sharp demarcation of white on the pro.ximal end. Study IB, 15 adults, April 15 to June 21, It further differs from that insect by the con- associated with On/zopsis hipueiioidcs and Hap- tinuous lateral carinae of the pronotum which lopappus sp. are sharply constricted in the middle, making the Studv 3CD, 1 adult, June 27, vegetation not prozona shorter than the metazona. recorded.

Area 12, Rainier Mesa, 1 adult, June 26, Coloration. In coloration and pattern this vegetation not recorded. insect most closely resembles Cordillacris. It Study EGA, sand dunes, 2 adults, Septem- should not be confused with that insect, how- ber 30, vegetation not recorded, probably O. ever, because of the nearly rounded vertex. hijmenoides. Cordillacris is typically slant-faced. The dorsal 1 adult, abdomen under the wings is bright, colored the Study JA, Jackass Approach, August same as the caudal legs. The distinct markings 31, vegetation not recorded. and maculations are very contrasting. One fe- Study TA, Midvalley, 4 adults, June 22 to male, perhaps a teneral specimen, had very August 31, apparently associated with O. hijmen- bright markings. Rehn (1942) has published at oides.

Table 26. Size variation of Psoloessa dclicutulit dcUcatuh.

I§ 58 Biiir^iiAM ^()^^o Univeiisity Science Bulletin

Genus LiiJ^iiroteltix McNeill scription, this subspecies is found t\pic;dl)- in the L'pper .Sonoran life zone. .\t the Ne\ada 1897. Li'^motvttix McNeill Proc. Da\cii()()it Test Site, again, it was more numerous in the Acad. .Sci., VI, pp. 198, 2.57. Lower Sonoran life zone.

Habitats. The most obvious character about

l.tf^iiroli'llix coijuillriti ((inlatur Helm this insect is its ability to produce sound. It is, (Figures 41, 43; Table 27; Map 13) in fact, yer\' noisy but difficult to see in the shrubs. In order to study the habits of the in- 192.3. IJfiurolrttix coquilletli cantator Rehn, sect some were \'isually spotted and captured by Trans. .Amer. Entom. Soc, XLIX, p. 64. hand. The insect is not characteristically strong

and any disturli;ince results in its jumping into Distinctive Features. Lateral hni-olae of tlie the center of the sliruh. It generally rests the fastigiuin dccpK' impressed, trapezoidal in shape; on outer limbs. antennae short; e\es prominent, occiput higher .\hmy of the desert shrubs drop their lea\es than disk of pronotum; medi;m carina slight but iluring the hot summer, and Ligurotettix distinct, intersected near the middle by the was especially upon these shnibs. It principal sulcus; lateral carinae obsolete; pro- common was Larrea other leaved shrubs, sternum furnished with a large pyr;unid;il spine; foimd on and but not to any great extent, comparatively. Sweep- sciipular ;irea of tegmina greatly expanded, one- ing for these insects is not successful of third the widtii of the tegmina, hyaline, with because the spiny nature of the shrubs. strong, curved oblitjue veins, forming an efficient To collect a series it was found that the best method was to organ for the production of sound. completely trample the shrub, working system-

Coloration. The body of the insect is brown, iiticiilh' ;ir()und the periphery first beeeuise the the tegmina grav suffused with brown, the cand- insects ne;uly alw;i\s hop into the center branch- id femorii with two black bands and dark genicu- es. B\' the time the shrub is completeK- broken lar lobes. The caudal tibiae iirc gray. The in- down most of tin- insects ha\e tried to escape to sect is well concealed in its habitat, on the leaf- another shrub, where they can be captured if less branches of the .xerophytic shrubs. spotted. The notes of Rehn (192.3) on the biolog\- of of the Distribution. This insect is typical the group are interesting. "The scattered gray higher western Ney;ida areas, extending into the green bush of the Nevada Basin, a land of broad regions of parts of Death Valley and the Inyo desert plains and \allevs and long mountain California. The type loc;dity is Mason, Lyon Co., ranges, is the favored habitat of this subspecies. very during Nevada. The insect was common In the Great Basin greasewood {Scircobatus), in Test the summer in all lower areas at the Nevada Atriplcx. and other species of the wiry ;md spiny .Site, ;ind could be heard during the hottest hours shrubs which sparsely clothe the valle\s and of the d;i\-. present series ha\e not been The lower mountain slopes of this region, it will bi' with t\pical material, be compared so may found at home. In but two phices (Daylight ;itvpic;d toward kitnzci. The drawings and de- Spring and Hole-in-the-Rock Spring) w;is it presented Rehn ) show the scriptions by ( 1923 found on creosote bush (Coiilica) [now Lar- insect is mori- typic;il of rcintator. rea] which is so much favoretl by the other sub-

According lo the notes with the origiiKil ile- species, which is reatliK understood when it is

Table 27. Size variation of Ligurotettix coquilletti aintator.

t Biological Series, \'ol. 4, \o. 3, September. 1964 59 . ) )

Bhicha.m Vocng U.nivebsity Science Bclletin

realized that tliis plant is almost absent from tlie 13 to September 3, on C. ramosi

Seasonal Occurrence. Tiie i-arjiest idIIci lion der; Arphia jri'^ida Scudder; Arphia infcnudis was made on June 20, the latest on September Saussure; Arphia Icporata Scutlder; Arphia can- 30. Because of the numbers present at that time ora Rehn.

it undoubtedh- is a part of the fauna well into Distinctive Features. This is the first of the October, at least. It was about ecjually numer- so-called "band-winged" grasshop[>ers found in ous during August and September. No July, the area and so common throughout the south- n\inphs were collected. west. The best characteristic to differentiate this Localities Represented. Specimens examin- species from the other orthopterans with bright- ed (adults): SS. (This number represents only ly-colored wings is the condition of the ineta- is distinctly the collected specimens. Actually the insect is sternal interspace. In Arphia this

very common and present in large numbers, ac- linear ( longer than broad ) in the male and nar- cording to the stridulations. An actual estimate rower tlum the interspace between the meta- sternal IoIh's in the female. pronotum is of numbers is difficult to make. The carinate, with one incision. Area 1 (studies IB and IC), 14 adults, June 24 to September 4, on Gratjia spiuosa, Lijnttm Morphological Variation. This is a very vari-

pallidum and L. andcrsonil. Other \egetati()n able species as is indicated b\- the abo\e svnon- not recorded. omv. Hebard (1937) published at length on the Study 3CD, 2 adults, .\ugust 15, on Lurrca phases of conspersa. and established fi\e phases, (livaricata. e;ich with a geographic distribution. Each phase is subject to decided individual \ariation, and Study 5.-\, 11 adults, June 20 to August 31, on shows such a remarkable response to ctinditions L. divdricatii. of the immediate en\ironment "that its extremes Study 5K, 12 adults, Jiuie 24 to September 2, are often i|uite as widely different as any of the on Lijciiim p(dlidum and Dalca pohjadcnia. typical representatives of the other phases. lOD, 2 adults, August 14 and 16, on Study Moreover, in some cases, one phase fades into Colco<^tjnc osis^m ram a ;uiother gradually over a wide extent of territory.

Study CM, Cane Springs, 17 adults, Septem- For these reasons, though it long setMned possible

ber 30, on G. i^pitiosa. C. ramosissima and /-. that some \alid races existed, I do not feel justi- pallidum, principally. fied in recognizing any geographic races what-

.\p[M()aeh, '>() ailults, I'ser in the case of cons^pcrsa." Study J.\, Jackass July

Table 28. Size variation of Arphia conspersa.

& )

Biological Series, \ol. 4, No. 3, Septemuer, 1964

Coloration. From the Nevada Test Site the show a variable condition of the tibiae, ranging insect is dark gra\ish brown, mottled with dark- (in dr\' specimens) from very light blue to \el- er brown and black. Most of the specimens e.\- lowish. The condition of the tibiae of the Ne- hibit a pale dorsal stripe on the tegmina (this \ada Test Site specimens is suggestive of Arphia condition is variable in the series ) . The hind ramomi Rehn from California, but that insect, wings are red" with a black outer band, the spur represented in the author's collection by speci- of the black band extending well inward to the mens from Riverside County, California, and one base of the wing. Between the spur and the ape.x female from Baja California, is quite different is an area of dark veins and lighter cells, with the morphologically from the present series. ape.x infuniate, slightly lighter than the dark These insects suggest a close relationship of band. The ventral sulcus of the interior femur eonspcrsa and aberrans and the present series basal is bluish, the hind tibiae bluish with a pale is assigned to conspersa after much consider- annulus. ation.

Distribution. This species has a very wide Arphia hchrensi Saussure has been reported distribution, ranging from Alaska to Mexico and from Nevada by Baker, Essig and LaRivers (in from Texas to Nevada. It was found in only one Ormsby County). I have not seen this sup- area at the Nevada Test Site. posedly yellow-winged species, so have made

no comparison of it witli the series from the Habitats. ArphUi was collected at or near test site. the top of one of the mountains between French- man and Yucca flats. It was found in an area of sparse grasses and small shrubs, typical of its Genus Xanthippus Saussure fairly strong flyer habits and habitats. It is a I8S4. Xanthippus Saussure, Mem. Soc. Geneve, and is deceptive, trying to hide after flight. XXVIII, pp. 46, 88. Three biologists were unable to collect any females after extensive collecting. The females are, however, more sluggish and remain hidden. Xanthippus corallipcs corallipcs

It is quite possible that the females appear at ( Haldeman a different time than the males. No nymphs (Figures 49, 51; Table 29; Map 14) were found. 18.53. OEdipoda coraUipes Haldeman, Appendi.x Seasonal Occurrence. The only collection C in Stansbury, E.xploration of Great Salt Lake, dates of the insect were July 13 and July 14. The p. .371, PI. X, Fig. 2. insect has been collected by the author in similar Established Synonomy. OEdipoda paradoxa habitats from May to however. July, Thomas; Hippiscus {Xanthippus) conspicuus Localities Represented. Specimens examin- Scudder; Hippiscus (Xanthippus) macuhtus Scudder; Hippiscus {Xanthippus)crcniitus Scud- ed ( adults ) : 6. der. Area 5, south end of French Petik, 6 adult males, July 13 and 14, not found on vegetation Distinctive Features. This large, ponderous and the only observation of vegetation from the grasshopper is easily recognized by its very large area was Bromiis teetotum. size (especially the females) and markings. It is the largest acridid found at the Nevada Test Additional Remarks. This insect has not Site and is very robust in appearance. The pro- been reported from Nevada previously and is notum is enlarged, extending over the occiput assigned here pending complete revision of the approaching the eyes. The head and pronotum genus. It is identical to one specimen in the are very rugose, the median carina of the pro- author's collection from Las Vegas, Clark Co., notum partially obliterated. The tegmina and Nevada, and is slightly atypical to a series of wings extend beyond the abdomen. specimens from several localities in Washington County, Utah. The bluish hind tibiae with the Morphological Variation. This species is pale basal annuli agree with tliose specimens very variable over its entire range, and has been from Utah, and are very much like a series of grouped into eight or nine subspecies through- specimens from southern Arizona, supposedly out its distribution. The specimens from the Ne- Arphia abeirans Bruner. A series of eonspcrsa vada Test Site are very typical of those from from north and east in Utah and from Colorado the Salt Lake Valley, the type locality.

book, is "Authors have iist-il Ridgwiiy (1912) a.s a means of [itif\ini; color of specimens. The however, difficult to obtain for c-omparative purposes. 62 Bhiciiam Vounc University Science Bulletin

Table 2y. Size variation of Xaiitliippus coruUipes coriillipes.

! Biological Series, Vol. 4, No. 3, September. 1964 63

Genus Lepras Saussure darkened with two other bands on some speci-

mens. The posterior tibia is dull with a light 1861. Lepnis Saussure, Revue et Magasin de subpro.ximal area. The dorsal tergites of the Zoologie, 2e Ser., XIII, p. 398. abdomen (especiallv in the males) are washed with blue. The inner face of the caudal femur Leprtis glaucipennis Scudder is blue with one large basal and one preapical (Figures 50, 52; Table 30; Map 14) black band.

1900. Lepnis glaucipennis Scudder, Psyche, IX, The hind wings are light blue to deep blue pp. 75-76. with a very broad black band and a dark spot at the apex of the wing. Distinctive Features. Morphologically this

species most nearly resembles XantJiippus but is Distribution. The species is distributed from distinct because of tlie slight median carina of southern California, Nevada, and Arizona south

the pronotum, not elevated as in Xantliippiis. into Mexico. At the Nevada Test Site it was The species is generally not as robust as tliat found only at intermediate elevations in two

species. The pronotum is very rugose. general areas.

Coloration. The species is easy to recognize Habitats. Leprus is imcommon on the test in flight because of the bright blue wings. Col- site, but the somewhat robust size and sluggish lected specimens can further be recognized by movements make them eas)- to capture once they the prominent dark spots on the tegmina. fly. Until they do fly they blend well with the

There is considerable variation in the series background. The typical habitat upon which from the Nevada Test Site. Tlie head of some they are found is one of small to medium sized specimens shows a definite ash-gray color, other rocks among sparse \egetation in the can\ons specimens are darker. The pronotum is variable of footliiUs. None was found directly associated from light to very dark (one female) and even with vegetation. a reddish suffusion in some specimens. Tire Seasonal Occurrence. The species was col- basal ground color of the tegmina is of the same lected from June 22 to September 19. It was general color as the pronotum (except the one most common during the month of August. Two dark female) to the first dark tegminal bar, subadults were collected in July. which extends across the entire tegmen; the next light space is lighter. The second dark bar is Localities Represented. Sjiecimens examin-

present only anterior to the median vein. The ed ( subadults and adults ) : 22. next distinct area, exceedingly light, is followed Study CM, Cane Springs, 1 adult, Septem- series light background, by a of dark spots on a ber 2. the dark decreasing in intensity to the tip of the Studies TA and TCB, Midvalley area, 21 tegmen. Some specimens show less obvious subadults and adults, June 22 to September 19. banding on the tegmina, the dark areas prac-

tically continuous with each other. There is a Additional Remarks. This series from the Ne- light yellowish Hne extending along the humeral vada Test Site is very variable in size and color, angle of the tegmen. The caudal femur has one less variable in morpholog)', and is assigned to conspicuous preapical dark band, variously glaucipennis pending a complete revision of the

Table 30. Size variation of Leprus glaucipennis.

£ s

{^, TA, August 31, 1961

cT , CM, September 2, 1961 ^, TA, AugiLst 17, 1961 J-, TCB, July 16, 1961

9 , TA, July 19, 1961

9 , TA, August 17, 1961

9 , TCB, August 7, 1961 Bhiciiam Vounc University Science Bulletin genus. Typical specimens have greenisli-blue Distribution. The general range of the in- hind wings (hence tlie name), and tlie speci- sect, as listed by Kehn and Hebard (1908) is mens from tliis area would have to be atypical the ".Mohave and Yuma deserts, ranging from of that species on the basis of wing color. the western edge of the Mohave at Mohave and Lancaster, California, to at least Sentinel, Mari- The species was at first considered to be L. copa County, Arizona." It was uncommon at interior Bruner, to which it is closely related, if lower elevations in both not synonymous. many of the stations at Frenchman and Yucca flats and adjac-ent areas.

Habitats. The most commoa habitat for the Genus Derotinema Scudder species at the Nevada Test Site was the Atriplcx- 1876. Derotmema Scudder, Appendi.x H9 of .\p- Kochia vegetation on the margins of Yucca Ann. Rep. Chief Eng. U. S. Geogr. pendi.x Jj of Playa. It has not been associated with \egeta- Sur\'. W. lOOth Merid., p. 513. tion, inasmuch as the insect is found on bare

ground and it attempts to escape by flight Derotmema cleliaitulum (Scudder) rather than to escape into the vegetation. This is of the few orthopterans collected around (Figures 54, 57; Table 31; Map 15 i one lights at night, in the Mercury Area. 19(X). Derotmema delicatulum Scudder, Proc. of Amer. Acad. Arts & Sci., XXXV, p. 390. Seasonal Occurrence. Tlie earliest record Distinctive F'eatures. This small sized, very n\mphs of Derotmema is June 21, with adults oc- active insect can be recognized by the enlarged curring July 3. The last series of adults were cx)l- head and prominent eyes. The pronotal disk lected on September 4. Nymphs were present has prominent rugae and lateral prominences only into July. It was equally abundant in July near the median carina. It is less rugose than and August.

is typical in the genus, the females being less Localities Represented. Specimens examin- rugose than the males. The posterior angle of ed ( nymphs and adults ) : .36. the metazona is broadly rounded or slightly angulate in some specimens. The tegmina and Study IB, 4 adults, September 4; the domin- wings are variously produced, always reaching ant vegetation in tlie area was Graijia-Lycium. beyond the abdomen, excessively so in some Study 3CD, 3 adults, August 15, no record specimens of both sexes. of vegetation around which the insects were The nymphs can be recognized by the very found. large eyes and rugose pronotum. Study 5A, Frenchman Flat. 5 adults, July 3 to August 10. Tlie vegetation was practically all Coloration. The insect is pallid testaceous, Larrea divaricata. but, of course, the insects flecked more or less with fuscous and with no were not determined to be associated uith the distinct banding on the tegmina. A series of dark vegetation. spots on the proximal tcgmen adjacent to the Study 5E, 2 adults, July 13 and August 17, no metazona gives tlie pronotum an elongated ap- vegetation associated with the speci- pearance. Tlic hind wings have a light yellow record of mens. disk and a variable black band. The posterior tibiae are light, generally grayish. The antennae Study fiA, on the margins of Yucca Playa, arc pallid, interrupted with fuscous. 14 nymphs and adults, June 21 to August 16.

Table 31. Size variation of Derotmema delicatulum.

•5-

t t c = JO

8.6 1.9 cf,6A, July 10, 1961 12.3 2.6 14.8 12.9 2.4 16.3 9.2 1.8 J', ,5E, July 3. 1961 2.1 cf,6A. July 12, 1961 15.1 2.8 15.1 8.7 lily 1961 19.4 3.2 20.3 10.4 2.2 9 , MD, J 15, 1961 20.1 3.8 21.0 10.5 2.4 9 , 5E, July 13, 21.4 4.1 22.2 10.8 2.8 9 , 5M, July 20, 1961 )

Biological Series, Vol. 4, No. 3, September, 1964 65

Study JA, Jackass Approach, 7 adults, July sexes), with black fuscous band and several 15 to August 22, no record of vegetation. maculate areas in the clear wing tip. Mercury, 1 adult, July 15, attracted to lights The caudal femur has one black subapical at night. band and a second incomplete band. The caudal tibiae are bluish gray mottled with Genus Mestobregma Scudder brown. Distribution. 1876. Mestobregma Scudder, Hayden's BuU. U.S. Mestobregma impexum is found from northern Arizona, southern Nevada Geol. & Geogr. Surv., II, p. 264. and southern California, tlirough Utah and into Idaho. The species was described from speci- Mestobregma impexum Rehn mens taken at Milford, Beaver Co., Utah (type ( Figure 53; Table 32; Map 15 locality) and Cima and Bird Spring Mountains, 1919. Mestobregma impexum Rehn, Trans. Amer. California, from August 11 to September 5. "The Entom. Soc., XLV, pp. 239-242, Plate XXVI, species was scarce at Milford, occurring on sage figs. 9 and 10; Plate XXVIII, figs. 13 and 14. covered ridges at 5000 feet and on relatively bare slopes, with scattered sage and yellow- Distinctive Features. Size medium, form flowered bushes, at 4900 to 5000 feet elevation." slender. The median carina of the pronotum is (Rehn 1919) At the Nevada Test Site it was intersected by two nearly equal sulci, the carina found only in the sand dune area. is elevated and bilobate on the prozona, reduced on the metazona. There are accessory projec- Habitats. This has been reported as "a rare tions on either side of the median carina in the species found in sandy or dry soil with scatter- middle of tlie pronotal disk. The pronotum ed clumps of short grass in the sagebrush des- is moderately rugose on the dorsal surface. The ert." (Ball, et al., 1942). As indicated previously, tegmina and wings surpass the apex of the abdo- both specimens were collected from the sand men. The lateral lobes of the pronotum are dune area. The collecting was made from an acutely produced, which character will dis- area of scattered shrubs {Eriogonum sp.) and tinguish it from both Derotmema and Trimer- small annuals and perennials with scattered otropis. grasses, rather than on the dunes themselves, although the areas are adjacent to each other. Coloration. The color pattern is light, suf- Extensive collecting was done during August fused with dark maculations. Tlie posterior and into November throughout this same area, margins of the pronotum are outlined in light, but no other specimens were found. without maculations, the sides of the pronotum The species was very active, flying some marked with black in the male (absent in the distance when disturbed. The red wings of the female). The lower margins of the lateral lobes male made this a conspicuous insect in flight, are light in both sexes. The tegmina have two but the yellow wings of the female could dark bands extending from the edge, indistinct scarcely be discerned from the yellow-winged and broken into suffusions of maculations be- Trimerotropis albescens, so common in the area. yond the humeral angle, the area between the The habits of the two were quite different, two bands without maculations, the distal two- however, the Trimerotropis being a more slug- fifths suffused with dark. gish insect, moving only short distances when Only two specimens were collected at the disturbed. Nevada Test Site, one male and one female. The hind wings in the male are red, in the fe- Seasonal Occurrence. The two specimens male yellow (either color can appear in both were collected on July 30 and August 12.

Table 32. Size variation of Mestobregma impexum.

s JO% i Bitic.iiAM VoL'Nc Univeiisity Science Billetin

Localities Represented. Specimens examin- flat, smooth or granulate to minutely tuberculate

L-d (adults): 2. on metazona, which is strongly broadened. .Median carina of pronotiun \'ariable, cristate, Study EGA, sand dune area, 2 adults, July even bilobate, or low on prozona, alwavs less 30 and August 12, no association was made with elevated on metazona, and with two transverse vegetation. incisions. .Metazona much longer than prozona, from one and one-half to more than twice as Genus Trimerotropis Stal long. Lateral carinae indistinct or absent except occasionally front 1873. Trimerotropis Stal Recensio Orthopteror- on of metazona and prozona. Posterior margin of pronotum distinctK' angu- um, 1, p. 118. late; lateral lobes of pronotum parallel, posterior .More species belong to the genus Trimero- angle well rounded. A tooth projected down- tropis than anv other genus found at the Nevada ward from the posterior angle of the lateral lobe Test Site. One of tiiese (T. pallidipcnnis palli- is characteristic of two species. The tegmina dipcnnis) is flie most widespread acridid found. and wings surpass the apex of the abdomen, oc- .Members of the genus be recognized by the can casionally are plain or commonlv maculate over following combination of characters: the entire surface of the tegmen, or much more Head of moderate size, the lateral foveolae frecjuentlv ananged in three well-marked bands, distinct and triangular, the antennae filiform, of the two proximal bands generally much better variable length. Disk of the pronotiun nearly defined than the distal band.

Key to the Species of Trimerotropis

1. Median carina of prozona cristate, the anterior and median lobes distinctly bilobate (Fig. 58); tegmina without solid, distinct fasciations or bands T. bilobata Rehn and Ilebard

Median carina of prozona neither bilobate nor highly cristate, the lobes indistinct and more or less fused (Fig. 59) 2

2. Disk of wing yellow, always with a well defined black band 3

Disk of wing blue or bluish, with or without a well defined black band 7

3. Gaudal tibiae yellow or yellowish, occasionally light brownish in some drit>d sj^cimens, but never blue, green or red'- T. palUdipcniiis pallidipcnnis (Burmeister)

Caudal tibiae not yellowish 4

4. Gaudal tibiae red; posterior angle of the lateral lobes of pronotum with a small tooth (Fig. 59) T. strentta McNeill

Gaudal tibiae never red; posterior angle of lateral lobes of pronotum without a tooth 5

5. Caudal tibiae greenish (occasionally drying to tan) T. inconspicua Bnmer

C;uidal tibiae blue or bluish 6

6. Ground color white; tegmina white or very pale with thrt-e narrow dark bands; caudal tibiae light blue, sometimes drying to pale gray T. albescens MciNeill

Ground color brown, never white; caudal tibiae dark blue, sometimes dr)'ing to almost black T. fontanel Thomas

7. Wing deep blue, with a well defined dark band T. cyaneipennis Bnmer

Wing light blue, without a dark band T. s}>arsa Thomas

'•An (icfasioiiai tlriod .-ipfcimcii of 7'. iuctm.spiciui will show a \ellowish tibia; liut tho color is actually grceii-

i.sh. They can In- ilistini;iiis}u-ci from piilliilipcnnis i)y size, liowcNcr. Tiu- ptillUliiwnnis males arc larger than 20 inconspicua the mm, tile fc'males larger than 29 mm; iiicompicua i.s alw.iys smiillcr than tliesc measurements. In first proximal fasciation of tlic tegmen is not conspicuous be.ausc of the dark coloration of the tegmen from this fasciation to the base of llu' tcgmrn. in palUdipcwiis the first fasciation is distinct. Biological Series, Vol. 4, No. 3, September, 1964 67

Figs. 58-59. 58, Trimcrotropis bitobata, male, pronotum, lateral view. 59, T. strenua, male, pronotum, lateral

Trimerotropis bilobata Rehn & Hebard Distribution. The type locality of the species

(Figure 58; Table 33; Map 16) is Antlers, Mesa Co., Colorado, flebard (1929) commented: "We believe that it is very widely 1906. Trimerotropis bilobata Rehn & Hebard, distributed over the Great Basin, in desert en- Proc. Acad. Nat. Sci. Phila., LVIII, 382-385. pp. vironment at lower elevations." This species has Distinctive Features. The great elevation of now been collected throughout the western the prozonal section of the median carina of the states, the eastern limits being Arizona, Colo- pronotum definitely characterizes this from any rado, Wyoming and Idaho. other species of the genus found at the Nevada The species was common in Study 6.\, the Test Site. only locality at the Nevada Test Site where it was found. This study was located on the mar- Morphological Variation. This species sup- gin of Yucca Playa. posedly has no distinct projecting process on the ventro-caudal angle of the pronotum, but Habitats. In its original description, the the specimens from the Nevada Test Site are authors commented on the habitat and habits variable in this character. In a series of eight of the species. "This species was found in the time, a specimens collected at one two show arid valley of the Crand river ( the Colorado show only definite tooth, while the remainder River ) near Antlers station, where the only vege- a slight projection. Three specimens taken one tation was a heavy growth of low cactus inter- week later are without any trace of a tooth. spersed with occasional sage. Specimens were bv no means uncommon and could have been Coloration. Body color light brown, with easily taken in numbers had the cactus not inter- dark markings and lighter color on the head and fered so much with collecting, as when alarmed pronotum. Tegmina with two complete trans- the individuals would invariably seek refuge in verse bars, the distal third with small irregular the dense beds of cactus." maculations chiefly along the veins. Wing disk yellow with a dark band. The caudal tibiae, in M the Nevada Test Site it was found only the specimens from the test site, are grayish- in the Atriplcx-Kuchia vegetation {A. confcr- blue with a definite pro.ximal ring, the colors tifolki and K. amcricana), although its habits fading in dried specimens. were not tied in with either shrub. It is a

Table 33 Size variation of Trimcrotroi)is bilobata.

cf , 6A, July 10, 1961 cf , 6A, July 10, 1961 cf, 6A, July 19, 1961 J-, 6A, July 19, 1961 ^, 6A, July 19, 1961

9 , 6A, July 12, 1961

9 , 6A, July 19, 1961

9 , 6A, July 19, 1961 Bkicmam Vol'nc UNivtiisiTY Science Bulletin inoderaff stridulator during fliglit and on oc- merotropis calignona .McNeill; Trimerotropis casion a good flier. It was mucli easier to cap- caeruk'ipes Scudder. ture than Trimeralropis sparsa, witli wliicli it Distinctive Features. This species, as well was associated. It was always found on bare as many of the of is ground and would invariably fly to another members the genus, best characterized by bare area. color and pattern rather than morphology. Seaiional Occurrence. The adults were collected from June Hi to .August 16. They were Coloration. The ground color of the species most numerous during the month of July. is ash-brown, the head and pronotum the darkest and without distinct markings. The tegmina Localities Represented. Specimens examin- show the t\pical trifasciation of the genus, the ed ( adults ) : 22. No nymphs were collected. proximal fuscous band being the darkest and Study 6A, 22 adults, June 16 to August 16. extending from the costal margin half way across

the tegmen; the middle band, which is about Additional Remarks. In describing this in the middle of the tegmen, extends nearly or species, the authors remarked: "This species is (juite across the wing; the distal band is indis- one of a number which might with almost equal tinct and situated about one-third the length propriety be placed in either Conozoa or Tri- from the apex of the wing; mcrotiopis. but which we have placed here the apical portion of the tegmen is transparent, marked with a few chiefly because McNeill has considered its allied pale fuscous spots. The wings are pale trans- species as a member of Trimerotropis rather than parent \cil()w at the base, with a rather nar- Conozoa." In his catalogue to the world Ortho- row, fuscous band ;uk1 transparent apex with ptera, Kirby placed it in Conozoa. The state- dark veins. The posterior femora arc black ment indicates that many of these related forms (possibly bluish-black vary and might be confused with one another when living) internally at the base, with a black band toward the apex; to some extent. the apex is black internally and fusa)us extemal- Over its complete range it is \ery variable in 1\'. The antennae appear to be marked in some many external characters as well as the phallic specimens with indistinct pale annulations. complex of the male. The species should ob- The caudal tibiae are deep blue in living specimens, viously be subdivided into geographic races. changing to a dark color in dried specimens.

Trimerotropis fontanel Thomas Distribution. The type IcK-alit)' of the species (Table 34; Map 16) is Spring Lake, Utah Co., Utah. Tlie present distribution includes all states of the Rock\' Moun- 1876. Trinierotropis fontana Thomas, Proc. tain area west to the Pacific Coast and north into Davenport Acad. Nat. Sci., I, 255-256. pp. British Columbia and Vancouver Island.

Established Synonomy. Trimerotropis juluina At the Nevada Test Site it was found only in Scudder; Trinierotropis- fcrruginea McNeill; Tri- one area (Kowitli \',i!lc\ Junction), near Rainier

Table 34. Size variation of Trimerotropis fontana.

a g 69 Biological Series, Vol. 4, No. 3, September, 1964

Mesa, but it is probably present in most of the Coloration. The ground color is whitish, habitats suggestive of that study area. sparsely punctate with fuscous on the pronotum and conspicuouslv banded with fuscous on the Habitats. The grasshopper is a loud stridu- tegmina and posterior femora. The tegmina are lator during fhght, the topical fhght of the indi- whitish like the bodv, with the basal band nar- vidual being ten to fifteen feet. Tlie species is row and nearl)' solid, the median and third bands verv warv and flies often. Some specimens re- narrow and obviouslv made up of maculations, quired nine or ten attempts before capture. One but very conspicuous; beyond the third fuscous attempt to collect the species during cloudy, band a few groups of fuscous annuli are present. rain-tiucatening weather, indicated that the in- All of the light areas are very broad and im- sect would trv to escape by hiding in the shrubs punctate, except the basal, with a few dusky or grass, rather than fly. The insects were found points, and an oblique fuscous dash just beyond on very dark soil in an area of large clumps of the edge of the pronotimi, best seen when the Ehjmiis cinereiis and Artemisia tridentata. tegmina are at rest over the abdomen. The Seasonal Occurrence. Collections in this area wing disk is light yellow, nearly transparent, were made only on August 12 and August 21. bordered by a few fuscous clouds representing Males and females were found on both dates. the fuscous band; the ape.x is hyaline. The pos- No nymphs were collected. terior femur has the lower sulcus light except for a narrow stripe on the basal h;df and a band Locah'ties Represented. Specimens examin- preceding the preapical light spot. The e.xterior ed (adults): 12. face is whitish, except for a \ery distinct fuscous Study 12CF, 12 adults, August 12 and 21. band preceding the preapical light band and a Additional Remarks. The established synon- few faint clouds representing the other bands. posterior tibia is blue with the base black, omy indicates that variation is found within The followed distinct light annulus. the group. The species is fre(jucntlv referred by a N\niplis in to in the literature as jiiliatui. Specimens collect- resemble adults body color and ed near the type locality bv the author show a iiie alwa\s found on the ground where they tendency towards typical cincta in the black blend in with the environment. The specimens at the Nevada Test Site banding across the face. This character is shown in the minority of specimens in the series, how- are somewhat \ariable according to the color of ever. sand upon which they are found. They are always very pale, however, and found on light Triinerotropis albescens McNeill sandy soils. Specimens found at lower eleva-

(Table 35; Map 16) tions, where the sand is very white, show an ex- cei'tlingly light color; those at higher elevations, 1901. Triinerotropis albescens McNeill, Proc. U. where the sand is more yellowish, show a yel- S. Nat. Mus., XXIII, pp. 418-419. lowish suffusion of the ground color.

Distinctive Features. This species is small in The caudal tibiae are alwaj's light blue, size for the genus, but the best distinguishing lre(|uently with blue under the femora, the characters are found in the color and markings. ventral band may be solid, broken, or nearly in-

Table 35. Size variation of Trimewtropis albescens.

-5 C )

70 Bhigiiam Young Univeksity Science Bulletin distinct. Specimens collected early in the season Trimerot ropis strenua McNeill

( these specimens were from lower elevations (KiguRs .56, 59; Tahli- 36; M.ip 17) averaged considerably paler than later specimens 1901. Trimciolropis strenua McNeill, Proc. U. S. at higher elevations. Nat. .\Ius., XXIII, pp. 432-433. Distribution. Tlie species is apparently limit- Established Synonomy. Trimcrotrapis mon- ed in distrii)iition to California and adjacent taiui .McNeill. areas in Nevada. At the Ne\ada Test Site the species is found only in very light colored sand. Distinctive Features. The proz(jnal carina of

the pronotum is slightly elevated. The lateral Habitats. LaRivers (1948) commented that lobe of the pronotum is armed with a distinct the insect is an "effortless and wary flier." In tooth (in the Nevada Test Site specimens) c-oUccting albescens from the test site it was which character will distinguish it from tlie other determined that the insect is ver\' active, though members of the genus. The caudal tibiae are wary, with a distinct undulating flight and a coral red. loud stridulation during the flight. Generally, if the insect was not captured on the first attempt Coloration. This species closely resembles after the first flight, it would escape because of pallidipcnnis in size, coloration and markings, its remarkable resemblance to the environment but can be distinguished by the following mark- and because of the scattered desert shrubs un- ings: The tegmen has a \erv narrow basal band, der which it would eventually hide. distinct and well defined, the median and apical As with other members of the genus, albc- bands are less distinct, composed of dark macu- scens is found on sand, never on vegetation until lations, the intervening light areas entirely un- the extreme heat of the day forces them onto the spotted. The wing disk is yellow, the fuscous branch tips of shrubs. band rather broad. The disk of the posterior femur has a black inner face, with two yellow Seasonal Occurrence. Adults were collected bands on the apical half and one subapical black from June 22 to October 14. Three subadults band. The lower sulcus is yellow. The outer face were found during the months of and Aug- July is plain, with a single fuscous transverse sub- ust. during Adults were most common July and apical band. The most distinct difference be- August. tween pallidipcnnis and strenua is the c-olor of Localities Represented. Specimens examin- the caudal tibiae. In strenua they arc coral red; ed (subadults and adults): 40. in pallidipennis, buffish-yellow.

Study EGA, sand dunes, 18 specimens, July Distribution. The distribution of this species 22 to October 14. ranges from Oregon and Idaho south and east Study EGB, target rock area, 9 specimens, to western Colorado, New Mexico and western June 22 to August 11. Texas and into northern Chihuahua, Mexico. The Area E, miscellaneous collecting near Area species lias been described by Rehn and Hebard 12 garbage dump, 8 specimens, July 23 and 24. (1909) as a "Great Basin and interior desert form." Study 3GH, 1 specimen, June 27. More speci- At the Ne\'ada Test Site it was widely distri- mens were seen in this area, but were exceeding- buted, but limited seasonally. ly difficult to collect.

Area 3, miscellaneous, near Study 3GH, 4 Habitats. At the test site the species was specimens, July 15. found onl\' in sandy areas where they were cap-

Table 36. Size variation of Trimcrotropis strenua.

i Biological Series, Vol. 4, No. 3, September, 1964 71 72 Biti(;iiAM VouNc Univeiisitv Science Bulletin

tiirc'd only with ditficultv because of their con- Triiitcnilropis pallidipcnnis pallidipcnni.s triistiiii; markings and strong fliglit. The onlv (Burmeister) feeding records are Sahulu kali, upon upon I; (I'l.ite Kigiires 48, 55, Tables .j7, ,3«; Map 18 i wliich tlu'V were found in stud\- II'". 1838. OEdipoda piUlidipennis Burmeister, Seasonal Occurrence. Adults were collected liandb. Ent., II. p. 641. from August 9 to September 4. All but two of Fstablished Synonomy. Trirnerotropis vin- the specimens were collected during the month ctddld Scudder; Trirnerotropis similis Scudder. of August. At the Nevada Test Site it apparently has a \er\- short adult life. No ininphs xvcre Distinctive Features. At the .Nevada Test collected. Site this insect is the most widely distributed and most common acridid appearing through- Locah'ties Represented. Specimens examin- out the entire year. It can be recognized among ed (subadults and adults): 1.5. the nuTiibers of the genus Trirnerotropis h\ its large size, sharing the large size in common with Stud\' IM, 1 specimen, .\ugust 16. T. strcnua, and b\' bc-iiig tlie onlv member (jf Study IF, 3 specimens, August 9 and li. tlic genus witli \elic)\\ wings and yellow caudal feeding on Sal.iola kali. tibiae.

Study 3CD, 1 specimen, -August 15. Coloration. This is the large, flashy, yellow- Study .5A, 1 specimen, August 31. winged species of the desert. Specimens from

Area 6, miscellaneous collecting, 2 sjjeci- the -Nevada Test Site ha\e a defim'te tendency mcns, August 15. for "X" markings on the pronotum because of the contrasting colors. The insect has the usual dark Stud)' EGA, sand dunes, 7 specimens, Aug- band on the hind wings, typical of the entire ust 12 to September 4. group of band-winged grasshoppers. As stated previously, the caudal tibiae are yellow, often .Additional Remarks. The original description drying to a yellowish-tan. of strcnua indicates its closeness to T. californica The species has a ver\- definite color as a re- Brunei. TIicn ' inay be but varieties of an c.\- sponse to soil, being light in light-colored soil, ceedingK- variable species." Rehn and Hcbard dark in dark-colored soil, more gra\'ish in gray also commented on the closeness of these species. soil, and even (not observed at the Nevada Test In an examination of the phallic structures of the Site) reddish in red soil. male the author found tlie two species practically inseparable, certainly no more than subspe- Distribution. Tiiis grasshopper has, without cies, but the original designation of species is a doubt, as great a distribution as any other maintained until complete revision is made of grasshopper in the world. Relm (1940) sum- this very difficult genus. marized its distribution as follows:

Table 37. Size variation of Trirnerotropis pallidipennis pallicHjK'nnis.

t Biological Series, Vol. 4, No. 3, September, 1964 73

"This wide-ranging and quite variable sub- mens, but the female flew away. Again I fol- species is one of that interesting group of forms lowed her and the course took her in pro.ximity which have marked discontinuous distribution. of a second male, who carried out the same Almost universally distributed over the western manners as the first, but without actually mount- United States west of the eastern edge of the ing the female. As he hopped slowly away from Great Plains e.xcept in boreal areas, and occur- the female I was able to capture them both. ring from depressions below sea-level to con- In the late season and as the temperatures siderable elevations, where the form is often the decrease with the approach of winter, the species sole geophilous acridid, it extends southward in is found less frequently, but even during wann- arid or semi-arid Mexico at least as far as the er days of winter it can be found in the bright state of Oaxaca .... From tliis point southward sunshine. The colder temperatures bring about a it is absent until sub-Andean conditions in change in body color, and after the first cold southern Ecuador and Peru are reached ... a night the specimens exhibit a darkened sternum third area, in which the race is apparently as and abdomen, very suggestive of Dissosteira abundant locally as in the western United States, Carolina. Bolivia Argentina border extends from at the The species was commonly obscr\ed about ." ...... eastward. . . southward and the lights at night. It is found distributed over the entire Nevada Seasonal Occurrence. Adults were first col- Test Site except at highest elevations such as lected on March 11; the last collecting date was Rainier Mesa and other similar areas, where it November 28. Nymphs were first collected on should be present, but was not collected. December 8 and present into July. No speci-

Habitats. This acridid is at times a very loud mens, nymphs or adults, were collected during stridulator during flight, is a very strong flier, the months of January and February, probably and difficult to approach during optimum tem- because collecting activities had slowed down peratures. Tlie species is invariably found in considerably, but the one nymph present in De- a clearing, except during extreme temperatures cember, and again in March, would indicate of the summer when it may be found on vege- that they would have been present on warm tation, usually the highest tips of the shrubs. days during those months, seeking protection

When disturbed it always lands in a clearing during the cold days, and that they overwinter and when approached increases its flight dis- in the nymphal stage. They were nearly equal tance with each stop as it is pursued. in abundance during the months of June through October. In November their numbers sharply In the course of chasing and capturing one decreased. ( See Table 38 for summary of speci- female three attempts had been made and the mens of pallkUpennis throughout the test site.) next flight took her in the area of a male of the same species. The male was not observed until Localities Represented. Specimens examin- he gave a quick short jump. He was observed ed ( nymphs and adults ) : 299. approaching the female from a distance of about Sahol^ (studies IF and 5P), 19 specimens, six feet in a series of quick, short, jerky move- 19 3. ments (about one body length at a time), often June to November flexing his hind legs. In the meantime the fe- Grayia-Ltjcium (studies IB, IG and 4A), 78 male had flexed her hind legs in the same man- specimens, April 5 to December 8 (last adult ner. At a distance of about twelve inches he collected in November; December 8 record was stopped and the only movement of either was one nymph). the characteristic waving of the antennae. He Lijcitim (Study 5E), 18 specimens, March then quickly approached her and jumped on IS to July 27. her back from the side in a position of copula- Larrea-Franseria (studies 5A, 5CQ, 3CD), off tion. After about two seconds he jumped 27 specimens, April 8 to October 1. almost immediately flew about 25 feet. An and Atriplex-Kochia (studies 6A and miscellane- attempt was then made to capture the female, ous), 17 specimens, April 27 to August 14. but she was very elusive and after six or eight Colcogijne (studies lOD and Area 6 mis' attempts and covering at least 75 feet in a circle cellaneous), 24 specimens, June 14 to August 28. she again lit in the vicinity of the same male. Artemisia (studies ECB, TA, TCB and mis- He went through the same movements as be- cellaneous), 14 specimens, June 22 to August 18. fore, except that he stayed on the back of the female for about ten seconds. Almost immediate- Cane Springs (CBA and CM), 24 specimens. ly, upon parting, I tried to collect both speci- May 27 to October 14. 74 Bitrr.iiAM VouNc UNrv'EnsiTY Science Bulletin Biological Series, Vol. 4, No. 3, September, 1964 75 posteriorly, while the apical portion is nearly Study IG, 2 specimens, July 10. destitute of markings e.xcept for the infuscation Study 3CD, 2 specimens, August 15. here and there of a few veinlets. The wing has Study 3CF, 1 specimen, June 27. a very pale greenish-yellow disk, crossed about Study 5A, 1 specimen, July 18. the middle by a narrow fuliginous band, with a Area collection, transparent apical portion beyond the dark band. 5, miscellaneous 4 specimens, September 26. The lower sulcus of the caudal femur is yellow or at least with two pale bands. Tlie hind tibiae Area 6, miscellaneous collections, 1 specimen, are pale greenish or slightly yellowish, except August 11. on the extreme base where they are dark brown Study lOD, 4 specimens, June 14 to Aug- and they are somewhat infuscated beyond the ust 16. subbasal pale annulus and apically. The front Study CM, Cane Springs, 1 specimen, ' Aug- and middle legs and antennae are well marked ust 19. with dusky annulations. Area E, miscellaneous collecting near Area Distribution. The type locality of this 12 garbage dump, 1 specimen, July 24. species is Palisade, Mesa Co., Colorado. At the Studies JA and CBA, 23 specimens, June 12 time of description it had very limited known to October 15. (These two studies are grouped distribution. The species has been collected in because of a mixed vegetation.) Arizona, and Henderson's description of viridi- Study TA, Midvalley, 9 specimens, June 22 tibialii' was from Central Utah. At the Nevada to August 17. Test Site it is widely distributed, though not Additional Remarks. This species shows con- abundant, in many of the areas. siderable variation in both morphology and color Habitats. The species is not an active flier, pattern throughout its distribution at the Nevada moving only five or six feet (males) or not at- Test Site. Specimens were sent to the U. S. Na- tempting to move before capture (females). No tional Museum for confirmation. attempt was made to correlate it with any vegetation types because Trimerotropis, generally, is found on bare ground and upon alighting re- Trimerotropis cijaneipcnnis Bruner turns to bare ground. (Table 40; Map 19)

Seasonal Occurrence. The species was col- 1889. Trimerotropis cyaneipennisn [Sic] Bru- 12 October 15. Adults were lected from June to ner, Proc. U. S. Nat. Mus., XII, pp. 68-69. equally common through the months of June to Established Synonomy. September. One nymph was assigned to the Trimerotropis ajanea Scudder. species, and only one subadult was found. It is likely that the nymphs of incontpicua could be Distinctive Features. Coloration is the main confused with those of pallidipcnnis, and may distinguishing character of the species. The only be told only by the general pronotal characters, other dark blue winged species, Leprus glauci- the smaller size, and the banding on the femora. peiinis Scudder, is so distinct from the genus Tri-

Localities Represented. Specimens examin- merotropis morphologically that it should not be

ed ( nymphs and adults ) : 49. confused with the present species.

Table 40. Size variation of Trimerotropis cyaneipennis.

1 76 BiiiciiAM VouNC Univehsity Science Bulletin

C^oloration. Tlir gcnrral IkkIv color is dark There is no correlation between vegetation gr.iy, ottci) with a Icrnigiiious tinge, piofiisi-ly and the species. inottlc'l and markfil witli tiiscoiis. Tlic lifaci is Seasonal Occurrence. .Adults were collected mottlcil will) gray and hrouii. Tlic proiioliim is Irom June 2ti to October 4. Most of the speci- marked with tlie saine contrasting colors; tlie mens were collected during the month of July. tegmina arc rnottied with rather large quadrate No specimens were collected during the month brownish spots, which in most specimens are of September, as no collecting trips were made grouped info tliree bands or patches, the first onto Hainier Mesa, and only one specimen was occupying the iiasal third, the second the center captured in October. No nvmphs were collected. of the middle, and the third the outer third of the tegmen, not forming definite bands as in Localities Represented. Specimens examin- most of the other members of the genus. ed (adults): 76.

Wings \er\' dark blue on their basal half, Study SMQ, 4 specimens, August 12 and 13. crossed bevond hv a rather \\ ide fuliginous band The occiu-rence of this species at this low ele-

that docs not continue annmd towards the anal vation is surprising. It was an area of Sahola angle; the apical third hyaline \\ ith tlie veins kali. black. Tlie caudal femora are crossed externally Studies 12A and 12E, Rainier Mesa, 58 speci- by three moderately broad oblique brown bands, mens, June 26 to August 21. At least 90 percent internally with the basal half and a single black of the specimens were collected in Study 12A, band in advance of the light yellow preapical an- the disturbed area. nulation. Caudal tibiae deep cocrulean blue Study ACC, 1 specimen, October 4. with a light basal annulation, the spines black- Area E, miscellaneous collecting near the tipped. Abdomen deep blue above in some Area 12 garbage dump, 10 specimens, July 23 specimens, inclining to greenish along the sides, and 24. suffused ^\^th gray below. Study TCB, 3 specimens, .\ugust 12 and 13. Distribution. This sj:>ecies was described from specimens collected in the Salt Lake Val- ley, Utah, near the mouth of Ogden Canyon. It Trimerotropi-s sparsa Thomas ranges from western Texas west to California (Table 41; Map 19) Idalio. At and north to southern the Nevada 1S75. OEiUpoda sparsa Thomas, Wheeler's Re- Site it (juite on Rainier Mesa, Test was common port, Ckx)g. Geol. Expl. Sur^'. West 100th Merid- less common in the other arejis where it was ian, pp. 883-884. found. Established Synonomy. Trimerotropis azure- Habitats. The species is a strong flier and scens Bruner; Trimerotropis perplexa Bruner. loud stridulator. One specimen was observed flying for more than 1()0 feet before alighting. Distinctive Features. The species is distinct TTie flash of blue as the insect flies is very as being the only member of the genus without obvious. a black band. In flight it may not be recognized The specimens from Study 5HQ average because of the transparent light blue wings. smaller and are considerably lighter in color, suggesting a response to soil environment, inas- C^oioration. The body of the insect is dull much as Area 12 has very dark soil; Area 5, much gra\ish brown throughout, sprinkled with fus- lighter soil. cous dots. The tegmina are somewhat darker on

Table 41. Size varLition of Trimerotrofm sparsa.

£ — U to Biological Sehies, Vol. 4, No. 3, September, 1964 77 the basal third, the other two thirds being lighter Anconia integra Scudder and sparsely sprinkled with the fuscous dots. The (Figure 45; Table 42; Map 20) wings are a transparent light blue with promin- 1876. Anconia integra Scudder, Appendix H9 ent and strong veins. The caudal tibiae are of Appendix of Ann. yellow. JJ Rep. Chief Eng. U. S. Geogr, Surv. W, 100th Meridian, p, 515, Throughout its range, especially to the northeast there is considerable variation in color and Distinctive Features. There is a remarkable intensity of the wings, and the development of size difference in the sexes of this species. The the wing band. The specimens from the Nevada females are large, the males small. Morpho- Test Site, however, are quite consistent in pat- logically they are distinct from other species tern and coloration. found at the Nevada Test Site. The head and most of the anterior lobe of the pronotum are Distribution. The species is widely distri- smooth, the posterior lobe of the pronotum pro- buted in all of the western states from North fuseh' punctulate. The head and pronotum are Dakota, Montana, and Alberta, Canada, south to both small in comparison to the expanded meso- western Te.xas Mexico (the local- and New type sternum and metasternum. The tegmina and ity) and west into the Great Basin. At the Ne- wings are very long, the caudal femora very long vada Test Site it was found in only one sti^idy, and narrow. The pronotum is broadly rounded in the Atriplex-Kodiia association immediately posteriorly. adjacent to Yucca Playa. Coloration. This species is variable in color Habitats. A very loud stridulator and strong from ash gray to yellow with minute fuscous flier, this species flies short distances of 20 to 25 maculations on the pronotum and head, and feet, loudly stridulating with each flight. It is with larger maculations on the tegmina. The a very wary insect on the ground and, at times, lateral carinae of the pronotum are noticeably it has been impossible to approach any nearer marked with a cream-color to produce an "X" than ten feet. At least a dozen attempts were on the pronotum. Occasionally, especially the made on one specimen, which finally escaped by females, the entire body, including the tegmina a longer flight and apparently hid in an Athplex. and caudal femora, is green with yellow markings and brown maculations. The wings are Seasonal Occurrence. Adults were collected transparent, clear, or slightly smoked in some from June 28 to August 16. They were most specimens, with the larger veins fuscous. The numerous in July. No nymphs were collected. caudal femora have two indistinct black bands Localities Represented. Specimens examin- both externally and internally, the outer central ed (adults): 10. area of the caudal femora with ash-gray wash. the Study 6A, 10 specimens, June 28 to Aug- The caudal tibiae are the same color as ust 16. body, bifasciate proximally. Klost of the body markings fade somewhat in drying. Genus Anconia Scudder At the Nevada Test Site the brown specimens 1876. Anconki Scudder, Appendix H9 of Ap- predominate, the green phase was only occasionally found, and specimens exhibiting the true pendix JJ of Ann. Rep. Ghief Eng. U. S. Geogr. Surv. W. 100th Meridian, pp. 514-515. yellowish phase were not collected.

Table 42. Size variation of Aticonia integra.

1 t J (C

21.0 4.1 cf , 6A, July 10, 1961 21.9 4.6 J', 5E, August 13, 1961 22.0 4.7 J' , 5E, May 27, 1961 April 34.5 7.0 § , 5E, 29, 1961 1961 36.7 7.9 9 , 5E, September 9, 38.0 7.6 ? , SCI, June 29, 1961 78 Bkkmiam Vounc University Science Bulletin

Distribution. TIh- range of the sjx'cios is now a late spring and very early summer form. known to extend from Las Vegas, Nevada, and Localities Represented. Specimens examin- Death \alley, Cahfornia, south to Indio, Calif- ed (adults): 37. ornia and Y'uma, Arizona, and from the western Study IB, 1 adult, .\ugust 27, vegetation not portion of the Molui\e Desert to at least tlie recorded. vieinitv of Tueson, Arizona. Tlie Nevada Test

.3CI, 1 Site distribution is limited to lower elevations. Study adult, June 29, vegetation not recorded, probably on A. confertifolia. Habitats. The eolleeting results at the Ne- Study 5A, 1 adult, .August 31, vegetation not \ada Test .Site somewhat contradict some of the rec-orded, although the area is predominantly pMhlished comments on this species. Caudell Larrea divdricata and Franseria dumosa. (1908) reported that "these grasshoppers are Study 5E, 28 adults, March 14 to September wild and hard to catch, especially as they often 9, always on A. cimfcrtifolia or on alkaline fly in thorny shrubs, where they are \ery diffi- ground. cult to get. They are protectively colored when on the ground and when flushed fly long dis- Study 6A, 6 adults. June 14 to July 10, on Atriplex tances, especially the females which fly much confertifolia. Oni- specimen was found farther than the males." Rehn and Ilebard on Kochia anicricitna. (1908) reported: "At Tucson this species was taken among high weeds both in damp and dry Genus Cibolacris Hebard locations. They were very wary and alert and 1937. Cibolacrui Hebard, Trans. Amer. Entom. when missed flew for some considerable dis- Soc, LXIII, pp. 368-369. tance. A preference to alighting on the ground when pursued rather tlian on weeds and bushes Cibolacris fmrviceps aridus (Bruner) was observed, tliough invariably first discovered among vegetation." (Figure 44; Table 43; Map 21)

.At the Nevada Test Site this grasshopper 1889. Thrincus (?) avidus (= ariilus) Brunei, was found only on alkaline outwashes. Several Proc. U. S. Nat. Mus., XII, p. 78, pi. 1, figs. 2 specimens were found directly on Atriplex con- and 3. fertifolia. More often, especially the males, they Distinctive Features. Form moderately ro- were found on the groimd, where they were bust, the general contour of the head suggesting well concealed. It was observed that these speci- Anconia, the vertex with fine lateral Ciirinae sud- mens were weak fliers, never flying high nor far. denly and strongly convergent distad and briefly Manv of the specimens were collected during the separated at the apex of the fastigium. The an- middle of the day when they should be most tennae are very short, the eyes more prominent. active. The pronotum is weakly sellate, definitely cx)n- No nymphs were collected, but the species stricted in the cephalic portion (but not as de- did show a preference to A. confertifolia. in cidedly so as in Ariconia), the cephalic margin which they were well concealed, and upon of the disk with two, usually definite, small adja- which they fed. c-ent median con\exities. The disk of the pro-

Seasonal Occurrence. Adults were collected notum is broadly rounded posteriorly. The from March 14 to September 9. Tliey were most caudal femora are short and robust. There is no numerous during April, May and June, and are trace of the band on the hind wing.

Table 43. Size variation of Cibolacris parviceps aridus.

I t ^1 t a J £ 2 ^c3

cf, 5CQ, July 10, 1961 16.4 3.2 15.5 8.7 2.5 cf , CM, July 6, 1961 17.3 3.0 17.4 9.1 2.8 cT, JA, July 6, 1961 18.1 3.2 17.1 9.8 2.8 4.2 24.0 13.9 3.6 9 , 5B, July 3, 1961 27.8 .5B, 23.7 13.5 3.8 9 , July 3, 1961 28.2 4.9 26.4 13.6 3.8 9 , 5E, July 13, 1961 31.6 5.7 Biological Series, Vol. 4, No. 3, September. 1964 )

80 Bricham Younc UNi\'EHsrTY Science Buluetin

Coloration. The color variation in fin's siib- watching and pursuing actually hundreds of all specit-s is interesting. The general gronnd color specimens, male, female and nymphs, in varies from \er\ dark gray, brown, and even red- stages of development. brown, to verv light gra\-, buff, and almost while. Hebard ( 19.37) commented on the habitat of There are usually rather coarse and scattered this group: "From the series before me it is evi- dark dots in \ar\ing degrees of contrast, some dent that this insect prefers pebbly or coarse specimens presenting a decided speckled ap- gravelly areas in washes particularly near or at pearance. The insect has a strong tendency to the bases of the desert hills and mountains of duplicate the soil coloration. This is more notice- the southwestern United States, but is able to able over its entire range. From the White Sands reach considerable elevations (as high as 6950 area of New Me.xico it is almost white; from feet) in such environment. It is ven,' often en- southern Arizona, (juite yellow; and from the countered, but is seldom numerous and almost red sands of southern Utah it takes on a red disappears in the dr)' and hot valleys. Adults are appearance. From the Nevada Test Site it has present almost throughout the year as well as a tendency to intermediate colors, the yellows, small immatures. I am inclined to believe, how- grays, and light brownish-reds. The mauve- ever, that the largest number of adults are pre- colored macuiations blend in rather well with sent from May to early July over most of its the mau\e-colored rocks of the desert pavement range." upon which it may be found. The specimens Contrary to what Hebard remarked about show a definite tendency to the light "X" mark- this insect not being found commonly in the val-

ings of the pronotum, so common with many leys, at the Nevada Test Site it is most common of the desert acridids. This marking definitel)' in those areas of low shrubs immediately sur- blends in with the background. rounding the playa lakes, wherever small peb-

Tlie wings are pale bluish-green; the Ciiudal bles iire scattered on the ground. It is never

tibiae delicate blue with a white basal annulus. found in the alkali outwashes, but is always as- Nymphs mav be recognized by the contrast- sociatcnJ with the desert pavement. It is also ing colors and the definite tendency of the "X" found at higher elevations (but not on Rainier on the pronotum. Mesa nor at the highest elevations on the test site) on rocky terrain. Distribution. This species is a true faunal in- It is definitely attracted to lights at night. dicator of the Ixnver Sonoran life zone, its distribution extending through the southwestern des- Seasonal Occurrence. Nvmphs were collect- erts from southwestern Texas and northern Chi- ed from March 11 to the middle of June. Adults huahua, Me.xico, to southern California and were found from .\pril 1 to October 14. The

north to southern Nevada and Utah. group is most numerous from .\pril through July. Site it is as widely dis- At the Nevada Test ( See Tal)le 44 for comparative distribution. tributed as Trimcrotropus jxiUidipcnnis pallidi- pcnnls, but has not been collected throughout Localities Represented. Sjiecimens examin-

the vear as has that species. This may be a case ed ( nymphs and adults ) : 287. of concealment in the habitat (jxillidipcnnis be- Salsola studies (IF), 13 specimens June 19 actually being a showv insect) rather than not to August 16. ing absent from the environment. Graijia-Lijcium studies (IB, IG, 4A, 5E), 101

Habitats. This species is exceedingly difficult specimens, April 8 to August 16. to collect, though often numerous in the environ- Larrea-Franseria studies (3CD, 5A, 58, 5CQ, ment. Their ability to blend in with their en- 5M), 149 specimens, March 11 to October 2. vironment is most remarkable of any other Atriplcx-KcH-hia studies, no specimens col-

geopliilous acridid at the test site. lected. This is one of the two major associations Thev are always found on desert pavement, where the species was not found. high temperatures when except during extremely Coleogijne studies (lOD), 5 specimens, June be found resting on the vegetation off they can 16 to August 14. the ground. Tliey are rarely seen until they Artemisia studies (TA), 1 specimen, June 16. move. When pursued they always alight in the Pinvon-Juniper studies, no specimens col- open, flving onlv a short distance, but always are lected. The group was not found at this high well c-oncealed upon alighting. Tliey apparently elevation on the test site. have the abilitv to detect the spot in the area where they will be remarkablv well camou- Mixed vegetation studies (ECA and JA), 12 flaged. Tliese statements can be made after specimens, April 28 to October 14. Biological Series, Vol. 4. No. 3, September, 1964 81 82 BiiiciiAM YouNC University Science Bulletin

*^ * FIG 60 FIG. 61 r FIG 63

Figs. 60-63. 60, Anoplodusa arizonensis, female, caudal taiMLs, lateral view. 61, Acheta assimilis, female, caudal tarsus, lateral view. 62, A. asHmilis, female, head, pronotum, tegmina, dorso-lateral view. 63, A. arizon- eiuiis, male, cephalic tibia showing auditor^' apparatus, lateral view. but in .some groiip.s arc clcnsf and opa(jue. gans of hearing situated on the front tibiae, and

Stridulation is accoinplislicd by the sound- tegmina, if fully developed, with the larger part producers by the modified anal or dorsal field of their surfaces sloping at the sides of the body. of the tegmina. One tegmen is rubbed over the The tegmina of the males are modified to form other. Auditory organs, if present, are on the a sounding-board for the stridulating apparatus. cephalic tibiae. This is located near the base of the tegmina and The female ovipositor is usually long and consists of a trans\erse ridge bearing a series of well de\el()ped, sometimes spear-like or sword- teeth which act upon a stiffened edge on the shaped, often sharply upturned and greatly outer tegmen, causing both to vibrate and pro- curved, and composed of four or six valves. duce a scraping sound. Winter is usually passed in the egg stage and hatching takes place in the spring. Growth Family Tetiigoniidae is rapid and maturity reached in midsummer. Many different and distinct forms can be The oviposition of the females in early autumn found among the long-horned grasshoppers. ends the cycle. They can be distinguished, however, by the These insects are most attractive in appear- characters given in the key: the extremely long, ance and many of them have a distinctive song. finelv tapering antennae, the four-segmented Most species can be identified by their songs tarsi, without arolia between the claws, a com- and may often be caught by following the sound pressed, blade-like ovipositor in the female, or- at night. They are commonly noc-turnal insects.

Key to Subfamilies of TFrrricoMiDAE

First two tarsal segments lacking a lateral groove; posterior margin of hind tibiae with its two series of spines continued to tibial apex. Long, slender-winged species Subfamily Phaneropterinae, page 82

series continued tibial First two tarsal segments with lateral groove ( figs, 60, 64 ) ; spine not to apex; wings long or greatly reduced. If wings long, the insects are large and hea\y-bodied.

Subfamily Tettigoniinae ( = Decticinae), page 86

FIG 64

Fig. 64. Capnohotcs fuligino.siis, female, caudal tarsus, lateral view.

Subfamily Phanemopterinae

Key to the Genera of Ph.\neropterinae

Comparatively robust species; tegmina broad, barred with white or maculate; hind wings not over 7 mm. longer than tegmina (Fig. 65). Usually found in trees and on higher shrubs Insara Walker

Extremely slender, long-legged species; tegmina, if present, uniformly colored, hind wings more than 7 mm. longer than tegmina (Fig. (i6); male with first abdominal tergite specialized as in Figure 67. Usually found in grass and on low plants Arcthaca Stal .

Biological Series, Vol. 4, No. .3, September, 1964 83

FIG. 66

Figs. 65-67. 65, Insara elegans maculata, female allotype, tegmen and wing. 66, Arethaea brevicauda, male, tegmen and wing. 67, A. brevicauda, male, modification of first abdominal tergite, cephalo-lateral view.

Genus Insara Walker

1869. Insara Walker, Cat. Derm. Salt. British Museum, II, p. 267.

Key to the Species of Insara

Dorsum of pronotum deplanate (Fig. 68); tcgmina marked with a slight herringbone pattern of light green, conspicuously maculate with dark dots. (Fig. 65)

I. elegans maculata Barnum, new subspecies

Dorsum of pronotum extremely sellate (figs. 69, 70); tegmina conspicuously marked with a series of large white or pale greenish spots /. covilleae Rehn and Hebard

FIG. 69 FIG. 68 FIG. 70

Figs. 68-70. 68, Itisara elegans maculata, female allotype, pronotum, dorso-lateral view. 69, /. covilleae, male, pronotum, lateral view. 70, / covilleae, male, pronotum and proximal tegmina showing stridulating mechan.- ism, dorsal view.

Insara elegans maculata Barnimi, elegans. The ovipositor is bent more decidedly new subspecies" upward than in elegans. Compared to /. elegans (Figures 65, 68, 71-73; Map 22) consuetipes the tegmina are very maculate.

Holotype Male, NEVADA, Nye Co., Nevada Description. General body markings as in Test Site, one-half mile south of Tippipah elegans, the head greenish, more pronounced Springs (Study TCB), July 16, 1961 (A. H. than the rest of the body except the tegmina. Barnum, collector). Tegmina with a distinct herringbone pattern of Ught and dark green, about as in elegans, but Allotype Female. Same locality as Holoty[>e conspicuously punctate with dark purplish dots, male, October 14, 1961 (A. H. Barnum, col- these diu-k areas often completely filling cells, lector ) generally less extensive. Pronotum and stridu- Both Holotype and Allotype are deposited lating mechanism of male as shown in Figure in the U. S. National Museum. 72. Ovipositor of female (Fig. 71) with distal half Comparative Features. Tliis insect shows few half of ventral valve brilliant green, basal green, all valves structural differences when compared to I. ele- and dorsal valves dull yellowish tips. male cercus and gans elegans. It is somewhat smaller. The distal terminated by dark The Fig- portion of the tegmina and wings are narrow, terminal abdominal appendages as shown in the marginal field of the tegmina narrowing ure 73. Tibiae, distal palpi and basal antennae abruptly distad from the proximal third as in brilliant green, the distal half of antennae an-

' 'Named after the maculate appearance of the tegmina. IliiK^iiAM VouNc Univeiisity Science Bulletin

Male Biological Series, Vol. 4, No. 3, September, 1964 85

In addition to the above distribution, the species is now definitely known from Washing- ton Count)', Utah, the northeastern distribu- tional limit.

Habitats. The insect can best be collected by sweeping the tenninal clusters of leaflets of Larrea. Differing from Bootettix, the other com- FIG. 74 pletely creosote bush restricted orthopteran, 7n- FIG. 75 sara coviUcac will frequentlv fly when disturb- Figs. 74-75. Insara coviUeae. 74, female, apex of abdo- ed. Although their alar ability is men and ovipositor, lateral view. 75, male, apex remarkable of abdomen, dorso-lateral view. they generally fly only onto an adjacent Larrea, flying seldom more than forty or fifty feet, but and white markings, abdomen otherwise spotted on occasion farther. They can be spotted visual- with minute purplish dots. Tibiae presenting an ly, but are difficult to collect without a net. annulate appearance with a subdistal hght spot, Seasonal Occurrence. Nymphs were collect- slightly narrower than the proximal dark spK)t; ed from June 15 to mid-.'\ugust; adults were caudal femora with subdistal light area around found from early July to October 1. Most of the entire appendage. adults were collected in July. The species is marked with the same contrasting colors as Bootettix pitnctatus (Scudder), Localities Represented. Specimens examin- but is more easily seen in its habitat. ed ( nymphs and adults ) : 47. Study lOD, 3 specimens, July 19. Distribution. The species is absolutely and Studies completely limited to the creosote bush, Larrea 5A and 5CQ, 41 specimens, June 15 divaricata, although, according to the original to October 1. description the distribution of Larrea is greater Study JA, 3 specimens, June 24 to August 21. than that of Insara coviUeae. This species, described from Hill, Tumamoc Genus Arethaea Stal Tucson Mountains, Pima Co., Arizona, extends from "Lordsburg, New Mexico, westward 1876. Arethaea Stal, Bihang Svenska Akad., IV through the desert portions of Southern Arizona, (5), p. 55. northward to Lincoln Co., Nevada near Lyons, California and in California as far north as Lyons Arethaea brevicauda (Scudder) and the Inyo Mountains and as far west as (Figures 66, 67, 76; Table 46; Map 23) Cottonwod Station in the Mojave Desert and 1900. Dichopetala brevicauda Scudder, Canad. Palm Springs on the Western edge of the Colo- Entomologist, XXXII, p. 331. rado Desert. Southward distribution in Mexico unknown." (Rehn and Hebard, 1914). Distinctive Features. A small, Hght green

At the Nevada Test Site it was found species, with very long slender legs. The males wherever Larrea was present. are fully winged, the females with reduced alar

Table 45. Size variation of Insara coviUeae. Bmiciiam Vounc Univehsity Science Bulletin appt'iidages, much shorter than pronotiim. The vada Test Site it was found in onl\- two widely malt- tc'gmina has the stridnlating field very separated localities. strongly and narrowly produced at apex of stridulating \ein, as in Figure 76, with the pro- Habitats. These insects are difficult to ob- chiction at apex not equal to the width of the serve in their habitats of grasses or other low remaining portion of the field. The marginal plants. The onl\' male record from the Nevada field of the tegmina is normal. The eauda! mar- Test Site was found on Lycium pallidum. The gin of the pronotal disk is never sharply acute; \'egetation from which tJie female was taken the lateral lobes with the area of convex callosity was not recorded. This area, however, is all sometimes inflati'd. iMrrca-FranserUi. LaRivcrs (1948) reported these insects "associated with Insara covilleae and more abundant, being attracted to lights at

night in large numbers .... During the daw specimens were found hiding in the cooler depths of such plants as the omnipresent Larreo clivaricata, KramerUi cancscen.';. Frosopus jiiJiflora and Acacia

Seasonal Occurrence. There is no definite seasonal occurrence from the scant data. The Fig. 76. Arethaca brevicauda, male, pronotum and adults collec-ted pr().\imal tegmina .showing stridulating mechaiiLsm, two were on June 2 and July dorsal view. 15. No nymphs were collected.

Localities Represented. Specimens examin- Coloration. This insect is unicolorous, light ed (adults): 2. green, and resembles the vegetation upon which it is found. Study 5.\, 1 male, June 20, on Lijcium jxillidtim. Distribution. The type locality is Cahon Pass,

California. The species is found only in south- Forty Mile Canyon, 1 female, July 15, no ern California through southern Nevada to record of the vegetation upon which the speci- Crestline, near the Utah state line. At the Ne- men was taken.

Table 46. Size variation of Arethaea brevicauda.

s 3-= t U: T3 ~i s .3 p-^

1961 3.0 18.8 (f , 5A, June 20, 15.2 20.5 1.4 9.5 14.3" 9 , 40 Mile Canyon, July 15, 1960 4.0 3.1 21.7 1.7 0.0

Subfamily Tettigoniin.\e

Key to the Genera of Tettigoniinae^°

1. Wings non-functional, much shorter than pronotum (Fig. 77), not visible beyond the pronotum in females Atelaplus Scudder Wings functional, longer than abdomen 2

2. Prosternum armed with a pair of spines (Fig. 78); hind femora armed below on apical half with several distinct spines (Fig. 79) Capnobotes Scudder Prosternum unarmed; hind femora unarmed below Auoploclusa Caudell

"Female abdomen obviously shninkcn. ^'•AgUiothorax anniger Rehn and Mcbard was licstribtd from tlu- tree yucca {yucca l)rccifolm), the t\jx- locality being Lee Canyon, Nevada, not f;ir from the Nevada Test Site. Thi.s species was not collected at the test site, however. This shield-back katydid Ciui be distinguished from the other members of the subfamily by its very large pronotum, its wingless condition, and the corci of the male, which are without internal hook.s. iiOLOciCAL Series, Vol. 4, No. 3, Septembek, 1964

78 FIG. 77 FIG. FIG. 79

Figs. 77-79. 77, Ateloplus lutetts, male, pronotum and tegmina, dorsal view. 78, Capnobotes fuUginosus, male, prostemum showing spines, cephalo-ventral view. 79, C. fultginosus, female, caudal femur, lateral view.

Key to the Species of Capnobotes

Larger, at least 60 mm. total length; last dorsal segment of abdomen deeply divided apically, the angles forming attenuated prolongations e.xtending over the epiproct, almost or quite reaching the tip (Fig. 80); wings rather uniformly and deeply fuUginous C. fuUginosus (Thomas)

Smaller, under 50 mm. total length; last dorsal segment of abdomen less deeply divided apically, the angles forming prolongations scarely exceeding the middle of the epiproct; wings less fuliginous, at least in the posterior field C. occidentalis (Thomas)

Genus Capnobotes Scudder produced over the base of the abdomen. The tegmina and wings are fully developed, extend- 1897 Capnobotes Scudder, Canad. Entomolo- ing far beyond the tip of the abdomen in both gist, XXIX, p. 73. se.xes. The long, narrow cerci of the male have two apical internal hooks. The terminal abdomen appendages of the male are shown in Figure

80. The ovipositor of the female is distinctly shorter than the hind femora.

Coloration. The basic body color is mottled gray, brown, or occasionally greenish, or brownish mottled with gray. The tegmina have the FIG. 80 same general bodv color and the wings are uni-

Fig. 80. C. fuUginosus, male, apex of abdomen, dorso- formly and deeply fuliginous, darker on the lateral view. major veins.

Distribution. This species ranges from Calif- Capnobotes fuUginosus (Thomas) ornia, through Nevada and Utah, into Arizona (Figures 64, 78-80; Table 47; Map 23) and Mexico. At the Nevada Test Site it was 1872. Locusta fuUginosa Thomas, Ann. Rept. U. found widely distributed throughout many of S. Geo]. Surv. Terr., V, p. 443. the study areas.

Distinctive Features. A very large species, Habitats. This thamnophilous species appears the total length, including the tegmina and to be more nocturnal than diurnal in habit and wings, at least 60 mm. The pronotum is large, is often attracted to lights. At the Nevada Test

Table 47. Size variation of Capnobotes fuUginosus.

1 )

Biu(;iiA.\i VouNc; Univeksitv Science Bulleti.n

Siti- fhcv were qtiite rej^iilarlv found around tlie ing about the lights during this period of time. lights and often found inside the buildings that Study CM, Cane Springs, 7 adults, June 15 had been lighted and left open at night. They to June 20, on Atriplex canesceiis. frecpientlv flv for extended periods around the Miscellaneous collecting in rocky areas, higher lights. They are wary insects during the studies not identified, 3 adults, vegetation not be found hiding in of day and can some the recorded. dense shrubl)er\'. The host plant varies according to the desert habitat. A number of observa- ('apnolwtes oecideutalis ( Thomas tions were made of this insect, during the day, ( Table 18; Map 2.3) sitting near the top of a large shrub, such as Larrea or Atriplex. with the head downward. 1872. Locttsta oecideutalis Thomas, Ann. Rept. When disturbed thev jump into the center of U. S. Ck'ol. Surv. Terr., V, p. 444. the bush .:id frc(|uently escape. Established Synonomy. Capnohotes occi- When annoyed the insect often raises its teg- dentalis viridis Cockerell. mina and wings o\er its back in a defiant atti- tude, and if not handled properly will inflict a Comparative Features. The species is similar severe bite on the handler. Tliis is one of the to fidiginosus, but differs from that species by few species of Orthoptera found at the Nevada lacking the dark hind wings, the much smaller Test Site that will attempt to bite. size, and the longer o\ipositor in the female.

They are at least partialh- predacious on The male cercus is apicailv armed with a short other insects. internal spine with a large subapical internal prong proximally. Seasonal Occurrence. Nymphs were collect- Coloration. The species occurs as two dis- ed as earh' as March 23 and well into the month tinct phases, brown or green, described in the of June. The adults first appeared on May 27 literature as different subspecies. They should and were present until, at least, July 27. They not be recognized as distinct races, however, and were most numerous during the month of June. the green phase (viridis) is synonomized. Tlie

Localities Represented. Specimens e.xamin- brown phase is mottled with flecks of white, es- ed (nymphs and adults): 30. pecially on the tegmina; the green phase less Study TA, 3 nymphs and adults, March 23 maculate but with distinct pronotal markings of green. to June 23, probably on ArtemisUi tridentata. tan and The only specimen collected at the Nevada Study lOD, 3 adults, July 12, on Coleix^ifuc Test Site is representative of the green phase. riimosissima.

Area 1 (studies IB, IG), 3 adults, June 16 to Distribution. This is a Great Basin sjx-cies June 26, on Atriplex canesccns. found in the desert and jimiper-pinvon areas of Utah, northern Arizona, Nevada, California, New Study 5A, 1 adult, July IS, on lAirrea divaricata. Mexico, and southern Idaho. It is represented Study 5E, 2 adults, Jvuie 17, on Lyciuiii from the Nevada Test Site by one subadult fe- pallidum. male, with undeveloped tegmina and wings.

Mixed plant communities (studies JA, CR.\), Habitats. Tinkham (1944) listed juniper as 5 nymphs and adults, June 14 to July 27, on the only host of the species. LaRi\ers (1948)

Atriplex eanescens and Larrea divaricata. reported it as common to sagebrush in Nevada, Area MD, Mercury campsite, 3 adults, June with one specimen being taken on pin\'on pine. 14 to July 13, attracted to lights. This is a small He also reported the green phase on "two species percentage of the total specimens observed fly- of introduced weed, Salsoki hdi tcnuifoUa and

Table 48. Mea.surement.s of Capnohotes occidentalis.

B t Biological Series, Vol. 4, No. 3, September, 1964

an unidentified chenopodiac, whose vivid greens Nevada Test Site it is limited in distribution to the insect matched well." the Lower Sonoran life zone, the marginal areas The one specimen from the Nevada Test Site of Frenchman and Yucca playas. was swept from Oryzopsis hymenoides. Habitats. Anoplodusa arizonensis is adaptive- Seasonal Occurrence. The only specimen, a ly colored to the creosote bush, Larrea divari- subadult female, was collected on June 26. cata, but it is not restricted to that shrub. It has been observed on Dalea polijadenia, Franseria Localities Represented. Specimens examin- diimosu and Graijia spinosa. ed (subadult): 1. The insect is apparently limited to Larrea as Study 12A, Rainier Mesa disturbed area, 1 a source of food and is probably not carnivorous subadult female, June 26. This is a pinyon- in nature, as are its relatives, Capnobotes. It juniper area, but most of the trees have been will, however, attempt to bite when handled and destroyed as a result of an atomic explosion. can produce a rather severe bite. Although Tinkham's specimens (Tinkham Genus Anaplodusa Caudell 1942) were collected at night, at the Nevada Test Site the 1907. Anoplodusa Caudell, Proc. U. S. Nat. Mus., species was found only during the day, and searches at night failed XXXII, p. 319, fig. 25. to reveal its presence. This is perhaps due to its scarcity and

the large expanses of Larrea, making it difficult Anoplodusa arizonensis (Rehn) to encounter. (Figures 60, 63; Table 49; Map 24) Its very large size makes it readily discerni-

1904. Dnjmadtisa arizonensis Rehn, Proc. Acad. ble in a shrub upon which it rests. All the Nat. Sci. Philadelphia, LVI, p. 573. specimens were taken as a result of this visual study. It is a remarkable flier and after dis- Distinctive Features. The wings are very turbance has flown for nearly 200 yards. It ap- long as in the genus Capnobotes. The most dis- parently does not rely upon concealment for es- tinctive features, however, are given under cape, but upon the powers of flight. When dis- "Coloration." turbed it immediately flies at about 15 feet alti- Coloration. Greenish, occasionally slightly tude in a straight line, sometimes until nearly buff especially in dried condition, with ivory- out of sight, and immediately drops into another white markings on the entire body; pronotum, shrub. A second and third fhght is often en- especially the margins of the lateral lobes and countered in an attempt to capture specimens. the margins of metazona marked with white, Accordingly, it is one of the most difficult yellowish in pinned specimens; pronotum also orthopterans to collect. One specimen was first marked with brown. Tegmina of botli sexes observed flying in small circles about one green, with three distinct, or sometimes some- hundred feet from the ground. As soon as the what indistinct, rows of large circular nacreous author got out of the car it immediately flew in spots, the stridulation field of the male tegmen a straight line at that approximate altitude until reddish brown. Hind wings transparent. it had completely disappeared from sight.

Distribution. A member of the Lower Sonor- Seasonal Occurrence. Tinkham's report states an life zone, this species is found from the that this is an early spring form, persisting until Mohave Desert in California to Arizona. At the late summer. He reported specimens having

Tiible 49. Size variation of Anoplodusa arizonensis. BuiGiiAM Voi'Nc Univej

been collcxted as adults from April to early Aug- Morphological Variation. This is one of the ust. "Since the eggs arc laid in late July or early most \ariable species at the Nevada Test Site. August at about the lime the desert rains com- .\n obvious variation from the typical amdition mence, it appears highly probable that the ova is the spined nature of the fore tibiae. In the hatch iti the fall and the n\-mphs de\'elop dur- specimens from the test site there are three ing tilt' hill and late winter to mature in the spines instead of the usual one. early spring. This assumption is based on know- C;olorati()n. ledge of other Orthoptera in the region." Noth- In the series of specimens from ing has been pubhshed on the habits of nymphs, the .Nevada Test Site there is a remarkable dif- to date, however, and no discoveries were made ference in coloration and pattern. The females in the present study to reveal their habits. Speci- are tan with longitudinal stripes on the pro- mens from the .\e\ada Test Site were collected notum, while only one male is suggestive of this marking. CJenerally during June and Julv. the specimens have one iiKcIio-dorsal dark stripe extending from the examin- Localities Represented. Specimens anterior margin of the pronotum to the tip of ed (adults): 1.5. the abdomen, but the stripe mav be partially or Study 1BD2(), I adult female, July 12, on completely absent. The c;iudal tibiae in tvpical Gray id spinoso. specimens are light bluish-green. The veins of

Studv IC;, 1 adult female, June 19, on Larrea tile male tegmina are light, with diuk cells. lUvaricata. Distribution. The species is limited in distri- Study 4A, 1 adult, June 19. bution to (California and Nevada. At the Nevada Studies 5A and 5CQ, 11 adults, June 15 to Test Site it was widely distributed and found Franseria dumosa. July 14, on L. divaricala and in many collecting areas. Study lOD, 1 adult female, July 13, on L. divaricata. Habitats. These insects are frequently swept from low shrubs at the Nevada Test Site and were occasionallv found on upright stakes used Genus Ateloplus Scudder as markers. Heifer ( 19f)3) reported that they are 1894. Ateloplus Scudder, Canad. Entomologist, nocturnal; however, most specimens at the test XXVI, pp. 179, 182. site were collected during the day. A few specimens were found in can traps, but whether this was the result of a nocturnal or diurnal move- Alchiplii.s Uilcii.s Caudell ment could not be determined. ( Figure 77; Table 50; Map 24)

1907. Ateloplus luteits Caudell, Proc. U. S. Nat. Seasonal Occurrence. Nvinphs and adults were both collected in the nymphs from Museum, XXXII, p. 373. Mav — .May 10 into July; the adults from May 12 to Distinctive Features. Pronotum short, the September 1. The species was most abundant truncate, the lateral lobes posterior margin during June and Julv. shallow. Male tegmina visible and extending beyond pronotiun; female tegmina not extend- Localities Represented. Sjx'cimciis examin- ing beyond pronotum. ed (nymphs and adults): 36.

Table .50. Size variation of Ateloplus lulcua. Biological Series, \"ol. 4, No. 3, Sepiembek, 1964

Studies IB and IG, 5 specimens, May 10 to are commonly elongate, flexible, and covered August 9. with long, erect tactile hairs (Hubbell 1936). Studies 5A and 5CQ, 12 specimens, May 12 to July 18. Subfamily Stenopelmatin.ae Study 5E, 3 specimens. May 17 to June 6. The members of this group are called Study 6A, 3 specimens, June 18 to August 14. Jerusalem crickets or sand crickets. They are lOD, 5 specimens, 21 to Septem- Study June completely wingless witli strong, spiny legs, aud ber 4. a large inflated head with powerful jaws, with Study CBA, 1 specimen, June 13. which they can bite severely. Tliey have a Study JA, 1 specimen, June 3. reputation for being actually poisonous. They Studies TA, Midvalley, and ECB, Target live in the soil or under rocks and other objects Rock area, 6 specimens, June 22 to August 18. and are able to burrow rather rapidly. Their food consists, to a large extent, of other insects. Additional Remarks. Because of the diversity The family is represented by only one species species, specimens were submitted to Dr. of the at the Nevada Test Site. A. B. Gumey of the U. S. National Museum, who made the determination. Genus Stetwpclmattis Burmeister One female (ECB, August 11, 1961) is atypical with narrower pronotum and reduced 1838. Stenopclmatus Burmeister, Handb. Ent. spines on the ventral margins of the caudal fe- II, p. 720. mora. It perhaps is a morphological aberrant, or may represent an undescribed group. Stenopclmatus fuscus Haldeman (Figure 81; Table 51; Map 25) Family Gryllacbxdidae 1853. Stenopelmattis fuscus Haldeman, Appendix Representatives of the Gryllacrididae at the C in Stansbury, Exploration of Great Salt Lake, Nevada Test Site belong to two subfamilies, each p. 372. distinct from the other. They all have four- segmented tarsi, the tegmina and wings are com- Established Synonomy. Stenopclmatus ce- pletely absent, and the general features are more phalotes Walker; Stenopclmatus fasciatus or less grylloid. The female ovipositor is com- Thomas; Stenopclmatus oculatus Scudder; Steno- pressed, of the tettigonioid type; the male cerci pclmatus hijdrocephalus Bruner; Stenopclmatus

Key to the Subfamilies of Gryllacrididae

Antennal bases widely separated, by more than twice the length of the eye (Fig. 81); tarsi with puivilli; head large; cephalic coxa unarmed laterad Subfamily Stenopelmatinae, page 9i

Antennal bases very close together (Fig. 82); tarsi without puivilli; head smaller; cephalic margin of cephalic coxa armed with a spine (Fig. 83) Subfamily Rhaphidophorinae, page 94

FIG. 82 FIG. 83

FIG. 81

Figs. 81-83. 81, Stenopelmatus fuscus, male, head, facial view, 82, Ceuthophilus fossor, female, head, facial view. 83, C. fossor, female, cephalic coxa showing spine 92 BituaiANf V(^UNC University Science Bulletin JioLOGicAL Series, Vol. 4, No. 3, Septemher, 1964 93 comanchus Saussure and Pictet; Stenopelmatus Coloration. The general body color of the in- terrenus Rehn (?). sect is yellowish marked \\ith dark bro\vn. Some specimens show a particular barring on the ab- Distinctive Features. The most striking char- domen. In most of the heavily spined species acter of the species is the very large head, en- of orthopterans the spines are very dark. This larged out of all proportions to the body, par- is particularly true with Stenopelmatus, and the ticularly in the occipital region. Tliis character ver\' robust insect appears light yellow or tan alone is sufficient for the recognition of the and dark brown, almost black. The dorsal sur- species. In addition, the characters given in the faces are very shiny. key can be used to distinguish it from its near relatives. Distribution. This species has a wide distribution in the United States from the eastern Morphological Variation. This species was edge of the Great Plains to the Sierras of west- described early in the work on the Orthoptera ern North America. The specimens from the Ne- and has been known by a number of common vada Test Site represent collections from near names as well as having been described by dir- its western limits. Here, it is well distributed ferent workers. The synonomy, of course, indi- throughout most of the Sixndy areas of tlie test cates the variable characters of the species. site, particularly on and around Yucca Flat and The genital structures have been most use- on Rainier Mesa. ful in studies on orthopteran speciation. How- ever, differential genitalic characters apparently Habitats. This subterranean insect is largely e.xist group. According to Hebard do not for this nocturnal. Individuals can frecjuently be found has a small stout incurved chi- (1916) the male wandering about the desert in late afternoon, tinous hook on each side of the epiproct just especially when the humidity is relatively high pro.ximad of the cerci. The epiproct and sub- and the temperatures lower. Because of its noc- plates other specialization and genital show no turnal habits, most of the specimens collected general similar. The stuctures within the are in from the Nevada Test Site were taken in the can genital chamber are soft, unmodified and shrivel traps. A few specimens were picked up from drying. female epiproct and subgenital in The the ground or as a result of over-turning rocks plates show specialization. The ovipositor is no in search for fossorial orthopterans. short and simple. Because of these genitalic characters it is Seasonal Occurrence. Nymphs were collect- very difficult to separate the adults from the ed as early as March 10 and were still present in subadults, or individuals in the last instars pre- September. Adults were first collected in April ceding maturity. Tlie complete absence of wings and were taken as late as November 21. They leaves the abdomen and the genital structures were collected throughout most of the year, but as being the criteria for determining adult speci- were most numerous in June and July. An in- mens, and both the external and internal genital creased number of specimens were again found structures are unspecialized. The only apparent in May, August, and October. For some reason difference, then, is the general heaviness and fewer specimens were collected during the solidity of the limbs of the true adult. month of September. The series collected repre- See "Additional Remarks" for further com- sents all ages, from first nymphal instars to ments. adults.

Table 51. Size variation of Stenopelmatus ftiscus.

t t>\

(^, 1GB7, October 2, 1961 33.3 cT, 1FA5, August 2, 1961 October 1961 § , 12EC10, 23, 1961 9 , 12EC2, October 23, October 1961 § , 1BD20, 24, .

94 Bhicham Vounc University Science Buixetin

Localities Hi-prfsciited. Spccimons pxamin- Those specimens from the lower elevations t-d (nymplis and adidts): 12n. ar(! \ery hairy compared to the practically glabrous appearance of die otliers. Studies IH. Ki, and 4A {all in C.raijui-Lijcium If genitalic characters could be relied upon communities), 74 si)ecimcns March 10 to No- it miglit mean a separation of these two popu- \cmlxT 2 1 lations. In (he absence of such characters it Study IK, 2 spedmciis, August 2 and Novem- might be possible to carry out breeding c.\peri- ber 1. ments to see w hether or not they are distinct or

Study lOD. 1 spociiiun, July 13. the same. SurcK' lesser differences exist in the genus CculUopliilus, which group has highly Study lOS, 36 specimens, June 19 to July 5. developed genitalic structures and species can be (Tliis study area was opened lati- in tiie re- separated on this basis. A complete revision of search, and a complete evaluation nl the area the genus Stenopelmatits may warrant separation has not been made.) of tiiese forms. Area 10, miscellaneous collections, 1 specimen, September Ifi. Subfamily Ridvphidophorinae Studies 12A and 12E, Rainier Mesa, 12 specimens, Jime 15 to October 27. More specimens The members of this group are commonly were collected in the undisturbed area than the known as the cave or camel crickets. They are disturbed area. tan to brown, wingless, with curved or humped body, rather than flattened as in the true Additional Hemarks. Some variation exists crickets. Although very common they are not within the coniines of the Nevada Test Site. often noticed, since they are nocturnal in habit Those specimens taken at higher elevations. Area or live in caves or other cavities in the ground. 12 on Rainier Mesa, differ somewhat in the sub- The system of special can traps used in the genital plate, in the ovipositors, and by the pile studies at the Nevada Test Site resulted in a far on the body, which latter difference could be larger collection of these crickets than any a result of activity and nonnal wear. In these other orthopteran group. After extensive obser- specimens the dorsal valves of the female ovi- vations throughout this area during all seasons positor are slightly less than twice the length of the year it has been determined that this of the ventral valves and the subgenital plate is group is the most common of all the Orthoptera. rounded. In the specimens taken at lower elevations the dorsal valves are only one-fifth long- The subfamily is represented by two genera, er than the dorsal valves and the subgenital the PrisioceutJiophilus being found only at high-

plate is acute. er elevations.

Key to the Genera of RHAPumopnoRiN.vE

.Male with styles (Fig. S4); ventral ovipositor valves of female armed distad with crenulations or low serrations (Fig. 85); tarsal claw witli a distinct ventro-proximal sensory seta (Fig. ggv PrkioccuthitpJiilii.s Rehn

four (in Nevada Male without styles; vcnlral (i\ i|i()silor vaKes of Irinale armed distad with Test Site specimens) Iriaiigulai or atieular terlli in addition to the tenninal decurved hook- Cculliophilus Scudder like apex (Fig. 87); tarsal claws without sen.sorv setae

FIG. 84 iloiso-lalcral view. 85, female, distal Figs. 84-87. Prisloci-uthophilus piicijkiis. 84, m.\V, ;ipo\ iii.n, claws and sen.sory setae, valves of ovipositor, hitcnil view. 86, male, (list;)! si meiit ot eaiidal tarsus showing view. 87, Ccuthophilus UwuUipis. fiiiialc, distal v:ilv< of ovipositor, lateral Biological Series, Vol. 4, No. 3, September, 1964 95

Genus Ceuthophilus Scudder

1862. Ceuthophilus Scudder, Ciuiadian Nat. & Geol., VII, p. 284.

Key to the Species of Ceuthophilus (Adapted from Hubbell, 1936)

1. Subgenital plate of male divided into lateral halves by a percurrent median fold or sulcus (Fig. 88); ovipositor of female very long, the ovipositor/pronotum ratio 2.0 or more. Form less compact; legs longer and more slender, caudal femur rhaphidophoroid, spines of ventro-

cephalic carina fewer and separated bv distinct inter\als . .- C. nevadcims Barnum, new species

Subgenital plate of male with proximal portion undivided by a median sulcus, but dis- tolateral portions often prolonged and separated by a cleft or notch extending in from

FIG. 93 FIG. 95 FIG. 97

FIG. 98 FIG. 96 FIG. 99 Figs. 88-99. 88, C. nevadensis, male holotype, subgenital plate, caudal 89, C. fossor, m-Ac, subgenital plate, caudal view. 90, C. fossor, male, cephalic margin of cephalic femur, lateral view. 91, C. hehurdi, male, cephalic margin of cephalic femur, latcr;J view. 92, liimellipes, male, cephalic margin of caudal femur, lateral view. 93, C. lamcllipcs, female, distal end of cephalic margin of caudal femur, lateral view. 94, C. deserticolti, male holotype, caudal tarsus, laeral view. 95, C. descrticoUi, distal abdominal tergites, dorsal vew. 96, C. deserticohi, male holotype, subgenital plate, caudal view. 97, C. hehurdi, male, caudal tarsus, lateral view. 98, C. hebardi, male, distal abdominal tergites, dorsal view. 99, C. hebardi, male, subgenital plate, caudal view. 96 Bhigiiam Young Univehsity Science Bulletin

apex of free margin (Fig. 89); ovipositor of female short, the ovipositor/pronotiim ratio

less than 2.0. Fonii compact, robust, with sliort, stout legs; caudal femur grylloid, its ventrocephaUc carina usually with numerous, closely-spaced denticulations; often unicolorous or nearly so 2

2. Ventrocephalic carina of cephalic femur ornamented with a row of numerous nodules or denticulations in addition to the movable di.stal spurs (Fig. 90) C. fossur Hubbell

Ventroci'phalic c;iriiia of cephalic lemur not nodulose, either unarmed or bearing one or more movable spurs (Fig. 91) 3

3. Ventral carinae of caudal femur strongly laminate-explaiiatc, breadth of carinae increasing distad, each terminated just pro.ximad of base of genicular lobes by deep excision of margin, end of lamella forming an acute trigonal process; margins of carinae closely denticulate (figs. 92, 93) C/lamellipes Rehn

Ventral carinae of caudal femur not as in alternative 4

4. Ventral carina of caudal metatarsus armed with a row of short, bristle-like setae extending ail the way to apex (Fig. 94); 8th abdominal tergite of male not produced aiudad, the 9th tergite being much the wider (Fig. 95); subgenital plate of male suborbicular, separated for most of its distance (but never proximally) by a median sulcus (Fig. 96) C. deserticola Bamum, new species

Ventral carina of caudal metatarsus glabrous except for a proximal group of setae (Fig. 97); Sth abdominal tergite of male greatly produced caudad, concealing most of the 9th tergite (Fig. 98); subgenital plate of male elongate, terminating in a pair of large, widely separated rounded protuberances (Fig. 99) C. hebardi Hubbell

The descriptions of the species of Cetithophi- Allotype Female, same localitv', July 16, 1961. lus, as indicated by indices and measurements, Both specimens are deposited in the U. S. Na- are the same as those employed by Hubbell tional Museum. (1936) in his monographic revision of the genus. This species belongs to the subgenus Ceu- All measurements were made with a micrometer thopfiilus, {Uiherisis series, and Pditcisjniiosus

in one ocular of a binocular microscope. In group of Hubbell. Within the group it shows cases of doubt as to the identification of the a greater id^finity to C. t/avajxii Hubbell, but dif- species the male internal genital structures fers from this and all otlier species by the form and/or subgenital plate should be compared of the terminal abdominal appendages of the with the indicated drawings. male, particularK'. In thi- distribution the species

The worker may find it diflicult to separate is closely allied to utahaisis but differs remark- the adults from the larger nymphs or subadults. ably in the terminal abdominal appendanges of The females of the species can be told by the both male and female. Superficially it is per- fully developed teeth on the ventral valves of haps indistinguishable from the closely related the ovipositor, these being very acicular or at species. As is indicated by Hubbell, it is more least sharp-tipped in the adult, mere rounded difficult to distinguish the females. protuberances in the subadult and larger The general body form is elongate with nymphs. The adult males can be told by the moderately long, slender legs and short spurs, scleroti/afion of epiphallus pseudosternite calcars, and claws. the ( of llubbcll) and associated structures. In the Description of Ilololvpe Male. Body very iiN'mpIis and subadults these structures are in no elongate and slender, length 16.8 mm. Dorsum, way sclerotizcd. including abdomen, weakly polished, subglabrous with very scattered minute setae. Cculliopliihis ncvadcn.fui Rarnum, lllvVD. F,ves moderate in size, length 0.7 mm., New Species width 0.55 mm.; interocular distance 1.3 mm.; infraocular distance 1.45 mm.; clypeal sutiu-e (Ki^piris 8S. 110-120; Table 52; Map 26) 1.9 mm.; iuitennae long, iipproxiniati'ly two and nolot\'])e Male, Nevada, Nye Co., Nevada one-half times bocK' length; distal segment of Test Site, Studs TK (Tippipah Springs), Aug- ma\illar\- palpi 2.2 mm.; fastigiinn more prom-

ust 2-1, 1961. inent than expanded mesal margins of an- Biological Series, \'ol. 4, No. 3, Septemueh, 1964 tennal fossae, apex noticeably projected, very gite slightly concave, of tergite two to six trrm- bluntly rounded, subconical, sparsely setose. cate or subtruncate, of se\en rounded-angulate, THORAX. Pronotal length 3.3 mm.,' greatest of eight rather stronglv produced, the dorso- breadth 3.8 mm., depth, in side view 2.4 mm., caudal margin tumid; dorsal surface of ninth ter- in dorsal aspect subquadrate, cephalic margin gite greatly produced caudad, abruptly emargin- broadly emarginate, caudal margin truncate, ate mesially, verv tumid. Epiproct exposed dor- ventrolateral margins rather strongly arcuate, sally, the tenth tergite projecting beyond caudal slightly projected outward, broadest point just margins of ninth dorso-laterally; epiproct direct- caudad of midlength; mesonotal length 1.7 mm., ed ventrad, triangular, margins slightly elevated, greatest width 3.8 mm., caudal margin in dorsal straight and convergent to apex. Paraprocts aspect broadly conve.x; metanotal length 1.55 membranous, enlarged, produced beyond epi- mm., greatest width 3.5 mm., caudal margin in proct. Supraanal plate a small triangidar lobe dorsal aspect slightly emarginate. LEGS. Ceph- compressed between paraprocts, apex rounded. alic femur (Figure 117), length 5.5 mm., Cerci 3.1 mm., originating as a heavy appendage width 0.8 mm., equal in length and breadth to proximally, abruptly expanded to approximately middle femur, ventrocephalic carina with 1-1 twice the proximal diaineter, then gradually spurs, spur short 0.35 mm. long; middle coxa tapering to end. Epiphallus (Fig. 110) ex- with dorsocephalic carina not e.xplanate, dorso- posed, projecting dorsad of subgenital plate, cephalic angle obtuse, distal angle forming a heavily sclerotized, rami divergent to dorsum of very blunt projection directed cephalad; middle lu-ch, a straight horizontal bar, cephalic margin femur, ventrocephahc carina with 3-3 spurs, ven- turned under arch, junctions of rami and dor- trocaudal with 2-2 very small spurs. Caudal sum bearing large conspicuous, erect, laminate femur (Fig- 119) moderately elongate and auriculae, projected mesially as continuous with slender, length 11.5 mm., greatest breadth 2.6 dorsum and sub-horizontal to rami, projected mm. occurring at the proximal one-sixth, then ventrad one-half the distance of the sclerotized gently tapering distad to base of genicular lobes, rami, the auriculae rounded protuberances both ventrocephalic carina with 23 somewhat irregu- \entrad and dorsad. Mesa! margins of mem- larly spaced subequal minute spinulose denticu- branous areas within arch forming two bro\\'nish, lations on the distal two-thirds of carina, the heavily sclerotized tumid folds with laminate largest of these being restricted to the distal surfaces. Subgenital plate similar to C. pima, one-half, ventrocaudal carina with 45 similar rather heavilv corneous, including distal margin, but somewhat smaller denticulations, these ap- except a small proximal area on either side of pearing in one series on caudal side of carina, ex- mesal groove, completely divided into lateral cept for eight minute denticulations on cephalic halves by a small groove, terminating distally as side of carina in proximal one-half, dorsal denti- two rounded lobes. culations about 40, scattered on distal half of fe- Description of Allotype Female. General mur, but more restricted to cephalic surface, characteristics as given for male holotype. Body caudal genicular lobe distinctly spinulose. length 17.8 mm., including ovipositor, 27.6 mm. Caudal tibia straight, 11.6 mm. in length, dorsal HEAD. Eyes, length 0.85 mm., width 0.65 min., spurs (Fig. 120) moderately slender and elon- interocular distance 1.4 mm., infraocular dis- gate, subdistal spur of cephalic carina 0.75 tance 1.5 mm., clypeal suture 2.1 mm., distal mm. long, spurs gently curved, apex minutely segment of maxillary palpi 2.3 mm. THORAX. hooked, dorsal face bicarinate, outer face sparse- Pronotal length 3.8 mm., greatest breadth 4.6 ly setose, the setae varying on different spurs, mm., depth in side view 2.85 mm.; mesonotal spines of carinae variously arranged according length 2.75 mm., greatest breadth 4.8 mm. in to distance between spurs, averaging 13 between dorsal aspect; metanotal length 2.3 mm., greatest proximal tibia and first spur, then averaging 8 breadth 4.4 mm. LEGS. Cephalic femur, length between other spurs; subdistal ventral spurs 1-1; 5.9 mm., breadth 1.1 mm., ventrocephalic carina calcars short, dorsocephalic calcar 0.75 mm. in with 2-3 spurs, the distal spur 0.4 mm. long; midlength, the other calcars being 1.45 mm., 0.6 dle femur, ventrocephalic carina with 4-3 spurs, mm., 0.4 mm. respectively. Metatarsus 2.1 mm. ventrocaudal with 1-2 spurs. Caudal femur long, 2nd segment of tarsus 0.9 mm. long, 0.5 length 11.9 mm., greatest breadth 3.2 mm., ven- mm. wide, 4th segment 1.3 mm. long, claws 0.6 trocephalic carina with 29 spinulose denticula- mm. long, ventral carina of tarsal segments with- tions, \entrocaudal carina with .30 denticulations, out setae except a small group at proximal end. dorsal denticulations 55. Caudal tibia 12.3 mm. Terminal Abdominal Structures. (Figures 113- in length, subdistal spur of cephalic carina 0.95 116). Dorsocaudal margin of first abdominal ter- mm. long, spines of carinae averaging as in male; Bhiciiam Young Univkhsity Science Bulletin stibdistul ventral spurs 1-1; dorsoceplialic calcar gineil with brown along ventrolateral as well as 0.4 mm. in length, tiie other caicars being 0.6 cephalic and caudal margins, the disk with a mm., 1.45 mm., 0.75 mm. respectively. .Metatar- pair of admesal brown bands separated bv a sus 2.1 mni. long, 2nd segment of tarsus 1.05 narrow Ncllouish line and b\' a pair of caudal mm. long, 0..5.5 mm. uide, 4tli segment 1.5 mm. admesal triangles t'lnbracing the caudal end of long, claws O.S mm. long. till- pair of admesal bands and separated from Temiinal Abdominal Structures. Cerci 2.3 them b\- a U-shaped light area connecting the mm. long, slender, not expanded and modified rcniform areas of each side, these mottled with as in male. Subgenital plate simple. Ovipositor brownish spots and lines. Meso- and iiietanotum largely brownish, with extensive yellowish spots (Figure 111 ) 10. S mm. long, gradually tapering througliout, \erv sliglith' upturned, dorsal valves near cephalic margin, leaving the caudal and terminating in a slender point, sub-aciculati', ventrolateral margins solid brownish, .\bdominal longer than ventral \'al\es, five aciculate teeth tergites margined broadlv with brownish, ce- of ventral valves short, separated etjuidistantlv, plialic portions light, giving a trans\erse-band- restricted to the distal one-fifth of valve. ('(! ap]U'araiicc which disappears caudad due to crowding of tergites and concealment of paler Coloration. Similar to utahen.^Ls and its re- areas. Caudal femur with usual scalariform pat- lated species, the following color description is tern indicated bv darker brown, the remainder modified from Hubbeil for that species. General impression of dorsum \ell()wish brown, with of legs iargelv without maculations or markings, weakly contrasted pattern on pronotum and being unicolorous. All spines, caicars, spurs, transverse banding on abdomen. Pronotum mar- and tubercles tipped with darker brown.

FIG. 112

FIG. 117

FIG. 115

FIG. 114 FIG. 118

FIG 116

FIG. 119

Figs. 110-120. Ceitlhotthilus neiiidt-nxvi. 110. male par;ilypc opipluilliis. Ill, femali- allotype, distal ViJvcs of ovipositor, lateral view. 112. ft-malc partaypc. distar \alvcs of o\i|5<>sitor, l.itiral viow. 11.3, male holotype, apex of alidomin. Literal view. 114, male holotype, subjjenital plate, eaudal \ie\v. 11.5, male holotype, epi- proef. 116, male holotype, distal abdominal tergites, dorsal view. 117, m.ile holotype, eephalic margin of oephalie fimiir, later.i! view. 118, male paratype, eephalic margin of c<^phalic femur, lateral view. 119, male holotype, (aiid.d femur, lateral view. 120, male holotype, eaud.il t.usus, latenil \iew. Biological Series, Vol. 4, No. 3, September, 1964

Variation. Sizes of the minimum, maximum On Rainier Mesa, Study 12A had many and average of the se.xes, as well as the holo- large cracks and fissures as a result of an atomic type male and allotype female are indicated in explosion nearby. The insects were collected the following table. The general remarks of C. onlv in the can traps, probably as a result of nevadcnsis hold true for this species, as well as their nocturnal movement from the fissures. At for all specie^ of Ceuthopliilus, seemingly. Vari- this higher altitude the fissures are undoubtedly ations in the spurs of the ventrocephalic carina of quite humid, similar to the environment of the cephalic femur, which count was taken as an Tippipah Springs. indication for the variation of the spinulose con- Tippipah Springs was visited on only three dition of the insect, are as follows: the general occasions during the course of this study, one condition is with two spurs, one larger distal spur trip in June, July and August. The insects were and a smaller more pro.ximal spur, this occurring very numerous at each visit, and no correlation in 70 per cent of the individuals. Twenty per cent was made according to the appearance of of the specimens had only one larger distal spur, nymphs, adults, or sex. Collections were made and three spurs were least common. One speci- as follows at Tippipah Springs: men was .observed with a series of five spurs, the two distal spurs being large, the three proximal ones minute.

Habitat. This insect is known only from t\\'0 areas on the Nevada Test Site, in an abandoned tunnel known as Tippipah Springs and in the disturbed area on Rainier Mesa. The tunnel, dug in clay and shale, has many cracks and fissures so that the ceiling frequently sloughs off. It has a perennial water supply, keeping the en\ironment humid and cool, a typical habitat for a cave- dwelling camel cricket. The insects are found on the ceiling of the tumiel, never on the sides, as is typical with other such cavernicolous species, and they escape readily into the cracks and fissures when tlie beam of hght is shined on them. They are never found near the en- trance and are not subjected to light from the opening, as they ;vre back in a darkened area of the tunnel. This characteristic has been observed with other species of this group in similar habitats. This species was not found in any of the abandoned mine tunnels in the area and has not been located in any of the caves, perhaps due to the aridity of those areas. 100 Bricham Young University Science Bulletin

Nevada Test Site. It could conceivably be found ately polished, subglabrous, notal setae minute in other areas similar to tlie type locality and and scattered, abdominal setae much more Rainier Mesa, but siidi favorable areas are not immcrous and evenly spaced. HF.VD. Eyes common throughout the arid Great Basin. small in size, length 0.7 mm., breadth 0..5 mm.; mterocular dLstance 1.3 mm., infraocular distance Localities Represented. Specimens examin- 1.4 mm., clypeal suture 1.7 mm.; antennae \'ery ed (nymphs and adults): 119. long, approximately three times body length; Tippipah Springs (Study TC), 113 speci- distal segment of maxillary palpus 1.8 nun.; mens: 24 male adults, 15 female adults; 16 male lastigium as in fusiformis, strongly declivent, subadults, 28 h-niale suliadults; 15 male n\niphs, tlatleiied above, apex in side view slightly more 15 female nymphs, ai cording to the al)o\e ilatcs. prominent than margins of antenna! fossae, Rainier Mesa (Study 12A), 6 adults: 12.\C6, bluntK' rounded-obtuse, with erect setae. THOR- 2 females, August II and August 24, 1961; AX. Length of pronotum 3.5 mm., greiitest 12AC8, 1 male, August 11, 1961; 12AC9, 1 fe- ])readth 4.1 mm., depth (in side view) 2.1 mm., male, August 14, 1961; 12AC1(), 2 males, July 25 in dorsal iispect sub(|uadrate, cephaUc margin and August 12, 1961. sliglitlv emaginate, caudal margin truncate, \en- Additional Remarks. The holotype male and trolateral margins rather strongly arcuate, allotype female, as noted previously, are de- moderately projected outward, broadest point positeid in the U. S. National Museum. Para- just caudad of midlength; mesonotal length 2.3 types are deposited at tlie Philadelphia Academy mm., greatest breadth 4.3 mm., caudal margin, of Natural Sciences; the California .\cademy of in dorsal aspect, broadly convex; metanotal Sciences; the Museum of Zoology, University of length 1.55 mm., greatest breadth 4.0 mm., Michigan; Brigham Young Universit)'; and the caudal margin in dorsal aspect truncate. LEGS. author's collection. Cephalic femur (Figure 127), length 4.5 mm., breadth 1.0 mm., shghtly stouter than middle Ceiithophilus deserticola Barnum femur, \'entrocephalic carina with 4-4 spurs, the New Species distal spur 0.55 mm. long, twice as long as other (Figures 94-96, 121-129; Tables 53, 54; Map 26) spurs; middle coxa with dorsocephalic carina explanate distad, distal angle u'cakly produced and Holotype Male, Nevada, Nye Co., Nevada forming a small projection; middle femur, ven- Test Site, Study ECB (foothills west of Yucca trocephalic carina with 4-3 spurs, ventroc-audal Flat), August 11, 1961. with 4-5 spurs. Caudal femur (Figure 128) Co., Allotype Female, Nevada, Nye Nevada short, stout, length 9.8 mm., greatest breadth 3.1 Flat), October Test Site, Study IBDll (Yucca' mm., tapering distad to base of genicular lobes, 13, 1960. Both specimens are deposited in tlie ventrocephalic carina with 45 closely set, some- U. S. National Museum. what irregularly spaced subequal spinulose This species belongs to the subgenus Geo- denticulations, ventrocaudal carina with 57 sim- of Ilubbell, but its re- tettix, Fiisifoiiiii.s series, ilar but somewhat smaller denticulations, these In lationship within that series is (juestionable. appearing mesially in two series on either side import- his monograph Hubbell emphasized the of carina, dorsal denticulations about 48, scat- ven- ance of the setose/non-setose natm"e of tlie tered on distal half of femur, both caudal and tral carinae of the tarsi. These setae, though cephalic genicular lobes distincth' spinulose. completely present in nymphal instars, may be Caudal tibia straight, 11.2 mm. in length, dorsal absent in adult specimens, the setae apparently spurs (Figure 129) moderately slender and decreasing in number with succeeding instars. elongate, subdistal spur of cephalic carina 1.1 northern The Ftmformi.'i group, principally of the mm, long, spurs gently curved, apex minutely Great Plains area but extending as far west as hooked, dorsal face bicarinate, outer face sparse- Scudder) is typi- northern Utah (C. fu.siformis Iv setose, the setae varying on different spurs, fied bv having the tarsal segments setose. The spines of carinae averaging 8 between spurs, western Cmukili group, on the other hand, more approximately equal in number between spurs; segments. in distribution, has non-setose tarsal subdistal ventral spurs 1-1: dorsocephalic calcar setose would belong The present species is and 0.95 mm. in length, the other calears being 1.7 the to the Fusijortitis group. .According to mm. and 1.05 mm., respectively. Metatarsus 2.0 species, characters given in llnbheirs keys to mm. long, 2nd segment of tarsus 0.6 mm. long, jusifonnis. (lescrtk'oJa is more closely relate

Terminal Abdominal Structures (Figures 121, 1.1 mm. long, metatarsus with a row of stout 123-126). Dorsocaudal margins of abdominal setae on ventral carina, proximal one-half of tergites 1 to 6 truncate, of 7th weakly rounded- 2iid tarsal segment \\ith setae. angulate, of 8th rather strongly produced, the Terminal Abdominal Structures. Gerci slen- caudal edge sHghtly convex upward, 9th tergite der, 4.0 mm. long; subgenital plate simple. Ovi- strongly exposed and thickened caudad, broadly positor (Figure 122) 6.2 mm. long, slightly up- emarginate, in lateral view 9th tergite stiongly curved, dorsal \alves terminating in elongate, convex upward, completely concealing phahic slenderly aciculate point; distal one-third of ven- complex and equidistant caudad to subgenital tral \'alves with five elongate, slenderly aciculate plate. Epiproct membranous, its distal portion teeth, the distal and subdistal teeth decidedly compressed between dorsal margins of para- curved and separated by rounded intervals, the procts, margins slightly elevated, straight and three proximal teeth separated by less rounded convergent to arcuate-emarginate apex. Supra- intervals. anal plate seemingly absent, bent beneath epiproct. Epiphallus narrow, sides of rami diver- Coloration. Entire body of both sexes uni- gent, rami narrow, separated by rather broad formly pale, the only contrast being the eyes, quadrate opening occupied by membrane; dor- the anterior articulations of the mandibles, the sum of arch bearing widely separated and well mandibles themselves at the clypeal sutures, and cliitinized paired auriculae curved along dorsal the spinulose denticulations. Tlie tips of the margin of epiphallus, cephalic lobe short, dorsal spines, the spurs and the calcars are dark. The curvature of epiphallus as a narrow, recurved auriculae of the epiphallus of the male are flange. Subgenital plate weakly sclerotized, darkened and the tips of the acicular teeth and suborbicular in ventrocaudal aspect, broadest terminal dorsal valves of the female are similar- proximally, apices rounded, together forming ly darkened. The tibiae and tarsi are only slight- rounded termination with narrow mesal cleft, ly darker than the general body coloration. the terminal portion less sclerotized, separated Variation. The usual variation in size is from basal portion by distinct sulcations angulate noted in Table 53 on page 102. In addi- from the mesal cleft. tion, there appears such obvious variants as tlie Description of Allotype Female. General spurs of the ventrocephalic carina of the cephalic characteristics as given for male holotype. Body femur. Occasionally there are only two spurs, length 21.2 mm., ovipositor length 6.2 mm. frequently three, but generally four. These are HEAD. Eyes, length 0.7 mm., breadth 0.6 mm., noticeably spaced differently, probably due to interocular distance 1.3 mm., infraocular dis- the loss of a spur during growth of the indi- tance 1.35 mm., clypeal suture 1.7 mm., distal vidual. Where the spur is missing from the segment of ma.xillary palpus 1.9 mm. THOR-AX. carina there is always a space, indicating that Pronotal length 3.5 mm., greatest breadth 4.4 it has been broken off. There is even a variation mm., depth, in side view, 2.3 mm.; mesonotal of the spurs on the left and right femora of the length 2.15 mm., greatest breadth 4.8 mm. in same insect. Characteristics of the caiinae of the dorsal aspect; metanotal length 1.7 mm., great- tibiae were not observed as regularly, but the est breadth 4.7 mm. LEGS. Cephalic femur, usual variations in these conditions could be ex- length 4.6 mm., breadth 1.0 mm., ventroceplialic pected. The spinulose denticulations of the carina with 3-3 spurs, the distal spur 0.5 mm. caudal femora vary within recognized limits, long; middle femur, ventrocephalic carina with not only as to size but also to number. Conditions 3-4 ventrocaudal carina with 4-4 spurs. spurs, of the terminal abdominal appendages of both Caudal femur length 9.8 mm., greatest breadth male and female are remarkably consistent in 3.1 mm., ventrocephalic carina with 41 spinulose the observed specimens. denticulations, ventrocaudal carina with 56 denticulations, dorsal denticulations 41, both Habitat. This insect has been collected under caudal and cephalic genicular lobes distinctly extreme conditions of aridity, surrounding the spinulose. Caudal tibia 10.0 mm. in length, sub- plava lakes of both Frenchman and Yucca flats. distal spur of cephalic carina 1.1 mm. long, spines Owing to the absence of large rocks or other of carinae averaging 8 between spurs; subdistal suitable ground cover, it undoubtedly is an in- ventral spurs 1-1; dorsocephalic calcar 0.95 mm. habitant of rodent burrows which are so com- in length, the other calcars being 1.75 mm., 1.0 mon throughout these areas. It is present, also, mm., and 0.55 mm., respectively; metatarsus 2.3 along the bajadas and into the lower foothills mm. long, 2nd segment of tarsus 0.7 mm. long, surrounding the mountains. All the specimens 0.6 mm. wide, 4th segment 1.3 mm. long, claws were captured in regularly maintained can traps BitK.iiAM VoiMc Univkhsitv Sciente Bulletin

FIG. 121

FIG. 123

Figs. 121-129. Ceuthophilus deserticola. 121, male paratype, epiphuUu.s . 122, female allotype, di.stal v;dves of ovipositor, lateral view. 123, male holotype, ape.x of abdomen, lateral view. 124, male holotype, .subgenital plate, caudal view. 12.5, male holotype, epiproct. 126, male holotype, distal abdominal tergites, dorsal view. 127, male holotype, cephalic marj^in of cephalic femur, lateral view. 128, male holotype, caudal femur, lateral view. 129, male holotype, caudal tarsus, lateral view.

Table 53. Size variation of Ceuthophilus deserticola.

"^i 3 S !f^ !?^ ^ o r? 3 S to "S 2 S tbS = tcS M^ M-w "^-w Vd-S toa tc"? -si Vp« &1 Bog. a V 2 ^3 si-s- gogs- g3 sg g3 g| ss sis gs §s g;> P5 ^Q j:<(i: j£ JO JO mo JO jS jh can jh jh jo

Holotyjie 0.7 0..5 1.3 1.8 3.5 4.5 3.8 3.1 10.4 2.0 0.6 0.5 1.3 0.8 (^ ,

(-j* , Paratype, Minimum 0.65 0.4 1.1 1.7 3.2 4.3 9.0 2.75 9.8 2.0 0.6 0.5 1.2 0.8

(^ , Paratype, .^vcr.ige 0728 0.496 1.283 1.846 3.417 4.742 9.95 1.988 10.492 2.108 0.675 0.508 1.317 0.967

(^ , Paratype, Maximum 0.8 0.5 1.4 2.1 3.8 6.2 11.0 3.2 11.4 2.4 0.8 0.55 1.4 1.1

0.6 1.3 1.9 3.5 4.6 9.8 3.1 10.0 2.3 0.7 0.6 1.3 1.1 6.2 9 , Allotype 0.7

9 , I'ar.itype, Minimum 0.65 0.45 1.1 1.5 3.1 3.9 8.3 2.45 8.7 1.8 0.6 0.5 1.2 0.9 5.1

9 , Paratype, Average 0.7.37 0.53 1.267 1.81 3.467 4.553 J)..58 2.897 10.12 2.08 0.66 0.547 1.327 1.106 6.167

9 . P.iralvp.-, Maximum 0.9 0.6 1.5 2.1 4.1 5.3 11.6 3.3 12.0 2.3 0.7 0.6 1.5 1.2 7.1 JioLOCiCAL Series, Vol. 4, No. 3, September, 1964 with the exception of the holotype male which AfripJex-Kochia (study 6A), 5 specimens, was captured from one of the lower foothills in July 17 to October 13. the vicinity Flat. of Yucca The area is composed Coleogijnc (study lOD), 2 specimens, March of coarse sand from weathered the granitic rocks, 3 and June 12. and the one specimen was captured by the Mi.xed (studies ECA, NCB), 5 specimens, author from underneath one of the large rocks August 21 to November 27. on the west slope of a hill, while specifically looking for fossorial Orthoptera. The conceal- Study EM, 1 specimen, August 11. ment of the insect by the sand was remarkable. Study 5HL, 2 specimens, October 30 and Seasonal Occurrence. Late instar nymphs November 3. have been collected only in June, July and Aug- ust. This does not correlate with appearance the Cetithaphilus hehardi Hubbell of subadults and adults, however, and it is like- (Figure.s 91, 97-99, 1.30-137; Table 55; Map 26) ly that the early instars cannot be distinguished from those of C. fossor. Subadults were collected 1936. Ceuthophilus hehardi Hubbell, U. Florida in April, May, and June. Adults have been taken Biological Series, Vol. II, No. 1, pp. 457-460. during all months of the year except January. Distinctive Features. Superficially, all the The wide overlap in appearance of this insect is Ceuthophilus species at the Nevada Test Site undoubtedly due to the environment in which it resemble each other. They can best be told by is found, the winters being sufficiently mild that the conditions of the terminal abdominal ap- the insect can make periodic appearances dur- pendages. These differences can be recognized ing the winter months. See Table 54 for the by the illustrations and the key to the species. appearance of these insects. At the test site, hehardi most resembles deserti- Distribution. The present known distribution cola, but the differences are obvious as illus- of this insect is limited to the Nevada Test Site, trated by the figures. Nye County, Nevada. Insufficient collecting has Coloration. At the Nevada Test Site the been done in surrounding areas to arrive at any species is somewhat darker than species found at definite conclusions as to its total distribution. lower elevations. There is a correlation with en- It should be found through the more arid regions vironment in this respect. The soil on Rainier of the Great Basin, undoubtedly extending into Mesa is considerably darker than the soil around southeastern California. the playa lakes. The specimens are quite uni- Localities Represented. Specimens examin- formly colored, any infuscations and barring are ed (nymphs, subadults, and adults): 55, as fol- indistinct. lows: 7 male nymphs, 5 female nymphs, 4 sub- Distribution. The type locality of this species adult females, 20 adult males, 19 adult females. is at a high elevation ( 10,000 feet ) in Iron Coun- Salsola (studies IF, 5HQ), 6 specimens, June ty, Utah. It is known only from western Utah 16 to December 4. and southern Nevada, not necessarily at high Graijia-Lijcium (studies IB, IG, 4A), 34 elevations because of specimens taken at St. specimens, February 2 to December 5. George, Washington Co., Utah (Hubbell, 1936). BiiK.iiAM Vot'NC Univeiisity Science Bulletin

c;:^^ FIG. 135 FIG. 137 FIG. 134 FIG. 136

Figs. 130-137. Ceuthophilus hehardi. 130, miile, epiphallu.s. 131, female, distal valves of ovipositor, lateral view. 132, male, apex of abdomen, lateral view. 133, male, subgenital plate, caudal view. 134, male, distal abdominal tergltes, dorsal view. 135, male, cephalic margin of cephalic femur, laterjil view. 136, male, caudal femur, lateral view. 137, male, caudal tarus, lateral view.

Table 55. Size variation of Ceuthophilus hchurdi.

a -a I sl ^1 i r; o ^ — ~— -^— -S-. W)-C tloTJ '&'2 tc a ^3 ?-5 an toa c?r =5 c2 S-a - i2 SO J '-• Ph jo, JU JO JU jS JH caH cf , Minimum size, 12AC9, July 25, 1961 0.6 0.5 1.0 1.5 2.7 3.1 7.2 7.6 1.3 0.6 0.55 1.0 0.6 cf , Average 0.661 0.516 1.106 1.578 2.922 3.533 7.789 8.156 1.422 0.689 0.483 1.094 0.628 cf , Maximum size, 12EC3, July 25, 1961 0.7 0.55 1.15 1.8 3.2 4.2 9.3 9.5 1.8 0.8 0.5 1.3 0.7

9 , Minimum size, 12AC1, October 27, 1961 0.65 0.5 1.0 1.45 2.85 3.3 7.2 7.5 1.4 0.6 0.4 0.95 0.6 4.2 9, Average 0.7 0.571 1.107 1.65 3.236 3.9 8.243 8.343 1.493 0.693 0.479 1.114 0.729 4.971

9 , Maximum size, 12EC2, Augu.st 24, 1961 0.8 0.65 1.2 1.9 3.9 6.0 10.3 10.5 '2.3 0.8 0.55 1.5 1.1 6.4

At the Nevada Test Site it was found only in Seasonal Occurrence. Thriee male adults were two studies on Rainier Mesa. collected on April 10. The area was not visited until July, when both nymphs and adults were Habitats. Nothing is known of tlie habitats found. Both nvmphs and adults were present of this species, inasmuch as tlie specimens were until October 28. They were most numerous captured in can traps established in the studies. during the month of .\ugust. This species is ap- It can be suggested, however, that the species parently most active during summer and early Uves under rocks or in other holes in the ground. autumn. Of the total series, 72.2% were taken in the dis- Localities Represented. Specimens examin- turbed area, where it was found with C. iw- vadciisis and PristocetitJwphihis pacifims, ap- ed (nymphs and adults): 115. parently living in the fissures and under the Study 12A, 83 specimens, April 10 to Oc- loosened rocks caused by the nuclear explosion. tober 28. 105 Biological Series, Vol. 4, No. 3, September, 1964

Study 12E, 32 specimens, April 10 to Oc- darkened character is carried over in some tober 28. adults. The genicular area of the caudal femora and the caudal tibiae are generally darkened. Additional Remarks. In his description of C. fossor can be distinguished from C. la- the species, Hubbell described only the male, melUpes, the other very common gryllacridid at and gave measurements only on that sex. the Nevada Test Site, by the light maculations Measurements and drawings of the females of on that species. The nymphs of lamellipes are this species are here given for the first time. very maculate, while the nymphs of fossor are nearly unicolorous. Cetithophiliis fossor Hubbell Distribution. The type locality of this species (Figures 82, 83, 89, 90, 138-147; Tables 56, 57; Map 27) is near Tucson, Pima Co., Arizona. It ranges over the desert regions of Arizona, Nevada and 1936. Ceuthophilus fossor Hubbell, U. Florida California. At the Nevada Test Site it was col- Biological Series, Vol. H, No. 1, pp. 484-488. lected in all studies where can traps were main- Distinctive Features. This species can best tained, except at high elevations. be recognized by the row of numerous nodules Habitats. The distribution of this species or denticulations on the ventrocephalic carina of points to the fact that it is an inhabitant of the cephalic femur. These denticulations are rodent burrows, primarily. Like the other even present on the nymphs, although they are gryllacridids it is nocturnal and omnivorous. less conspicuous. The males can definitely be distinguished by the two finger-like lobes of Seasonal Occurrence. Nymphs and adults the subgenital plate. These lobes are present on have been collected throughout the year in all the male nymphs, more distinguishable, of months. It is primarily a spring insect, the adults course, with later instars. being most common from April to June. Adults were present in very few numbers from August Morphological Variation. As with any large to February. Nymphs began to appear more series of orthopterans, there is considerable vari- abundantly in October and declined in June. ation in the structures of this insect. A more See Table 57 for the occurence and distri- noticeable variation is the nature of the sub- bution of the species. distal spurs of the cephahc femur. The usual condition is with one spur. A very few speci- Localities Represented. Specimens examin- mens have two spines, and one female has one ed (nymphs and adults): 1,415. spine on one cephalic femur, two on the other. Salsola area (Study IF), 245 specimens, These differences have no appearance as being November to July. the result of an injury where the spines might Graijia-Lijcium area (studies IB, IG, 4A), have been broken off. 872 specimens, collected in all months.

Coloration. This species is very nearly uni- Larrea'Franseria area (studies 5A, 5CQ), 16 form light colored, with the tendency to darken- specimens, April 3 to November 7 (no specimens ed tibiae in the subadults of both sexes. This collected in July, August, or October).

Table 56. Size variation of Ceuthophilus fossor.

I 3

t log 50 -C n JPL, yjU 106 BiUGiiAM YouNc University Science Bulletin

FIG. 146

Figs. 138-147. Ceutlwphilus fossor. 138, male, cpiphallus. 139, female, distal valves of ovipositor, lateral \iew. 140, female, distal valves of ovipositor, lateral view. 141, male, apex of abdomen, lateral view. 142, male, suhnenita! plate, eaudal view. 143, male, epiproct. 144, male, distal alidominal tergites. dorsal view. 145, male, cephalie margin of cephalie femur, lateral view. 146, male, eaudal femur, lateral view. 147, male, c.iudal tarsus, lateral view. Biological Series, \'ol. 4, No. 3, September, 1964 107

Atriplex-Kochia area (Study 6A), 29 speci- on the caudal femur, the male nymphs more mens, February 2-3 to September 18. noticeably dentate. Coleogijne area (Study lOD), 73 specimens, Morphological Variation. The comments un- October to July (no specimens collected in Jan- der "Morphological Variation" of C fossor can uary or February). also be applied to this species. A considerable Artemisia area (Study TA), 2 specimens, degree of variation was found with respect to

April 15 and 20 . June the appendages, especially their spines and den- Lycium area (Study 5E), 33 specimens, No- ticulations. vember 28 to June (no specimens collected in Coloration. Color markings this December or January). of species have previously been discussed. This is the most Mixed areas (stvidies CBA, A, 12CJ), 134 J maculate member of the genus Ceuthophilus on specimens, September to June. the test site, resembling Pristoceuthophilus, Study lOS, 11 specimens, January 19 and found only at higher elevations, in that respect. and June July. These two insects may be confused by super- ficial study, but are quite different morpho- Ceuthophilus lamellipes Rehn logically.

(Figures 87, 92, 93, 148-154; Tables 58, 59; Map 28) Distribution. The type locality of this species 1907. Ceuthophilus lamellipes Rehn, Proc. Acad. is Phoenix, Maricopa Co., Arizona. It has pre- Nat. Sci. Philadelphia, LIX, pp. 78-80, figs. 19, viously been reported only from Arizona and 20. northwestern Utah in Tooele County. Its distri- Distinctive Features. This species can be bution is shown now to extend into western distinguished from the other ceuthophili found Nevada, and it should be found in eastern Cal- ifornia. At the Nevada Test Site it at the Nevada Test Site by the very large tooth was found in all areas except at higher altitudes. It is un- ( ventrocephalic carina strongly lamellate-e,x- planate) of the caudal femur of the male, the doubtedly limited to a desert environment. smaller tooth of the female, and by the numer- Habitats. The species is found in rodent bur- ous small denticulations on the dorsal and ce- rows, under rocks and debris, and, although not phalic surface of the caudal femur in both sexes, proven, undoubtedly burrows in the loose sand more numerous and stronger in the male. It according to the nigose nature of the append- can also be told by the purplish markings (es- ages. pecially in the nymphs and subadults) on the caudal femur and the dorsum of the thorax and Seasonal Occurrence. In contrast to the abdomen. The terminal abdominal structures of earlier occurrence of C. fossor, this species ap- the male, of course, are the most reliable criteria. pears later in the season. Nymphs were collect- The female nymphs have a slight indentation ed in all months of the year, although they were

Table 58. Size variation of Ceuthophilus lamellipes.

2 £ -5 -B^ -S^ t ei ti)'§ Mrt &"« "S S mS W)"" M) R, £-S cS^cg a a. as r-3 r'4H C£ ^.fc- Ck- C* Zi f .E- n JO JH mH JH JH

, size, 1BB30, (-f Minimum February 1, 1961 0.95 0.75 1.3 2.1 3.5 4.6 10.5 " 1.8 0.7 0.5 1.3 0.7 2.0 0.59 1.405 0.88 cf, Average 1.04 0.86 1.44 2.35 4.11 5.31 12.06 0.785

(j" , Maximum size, 6AL11, October 13, 1960 1.15 1.0 1.5 2.6 4.8 6.1 13.6 2.2 0.9 0.65 1.5 1.0 9 .Minimum size, 1BB5, February 9, 1961 0.95 0.8 1.1 2.0 3.7 4.1 9.1 9.5 1.6 0.6 0.5 1.1 0.7 4.4 3.8 4.47 9.78 10.08 1.76 0.69 0.545 1.2 0.79 4.62 9 , Average 1,015 0.83 1.2 2.115

9 , Maximum size, 1FL2, November 13, 1961 1.1 0.9 1.3 2.35 4.4 5.0 11.2 11.3 1.9 0.95 0.6 1.3 0.9 5.0

"Measurements of the length of the caudal tibia of the males were not made because of the normal curvature of that structure (See Figure 154). )

108 Bhiciiam Young University Science Bulletin

Figs. 148-154. C. kmrllipcs. 148, male, epiph;illus. 149, ft'nialc, distal valves of ovipositor, lateral view. 150,

male, apex of abdomen, lateral view. 151, male, subj^enital plate, laud.il \it'u . 152, male, distal abdominal tergites, dorsal view. 153, male, cephalie margin of cephalic femur, latir.il \iew. 154, male, caudal a|> pendage, lateral view.

not abundant from NovcmlxT to May. The 1,167 specimens, collected tliroughout the entire

adults made flicir fir.st appearance in August and year, in all months.

were present from then into May. They were I .arrca-Franscria area (studies 5A. 5CQ), most numerous from October to March. (See 67 specimens, from July to .May (no specimens Table 59 for occurrence. collected in June).

Localities Represented. Specimens examin- Atriplex-Kochia area (Study 6A), 188 speci- ed (nymphs and adults): 2,344. mens, collected throughout the year, in all Salsohi area (Study IF), 223 specimens, from months exc-ept January. July to April (no specimens collected in May Colcootjne area (Study lOD), 25S specimens, or June). from July to May (no specimens c-ollccted in Grai/ui-l.iiciiiin area (studies IB, IG, 4A), June). Biological Slhies, \'ol. 4, No. 3, Sepiember, 1964 Hkicmam V(junc Univehsity Science Bulletin Biological Series, Vol. 4, No. 3, September, 1964 111

FIG. 160 FIG. 157

Figs. 155-160. Pristoceuthopilus pacificus. 155, male, epiphallas. 156, female, distal valves of ovipositor, lateral view. 157, male, apex of abdomen, lateral view. 158, male, subgenital plate, caudal view. 159, male, distal abdominal tergites, dorsal view. 160, male, caudal femur, lateral view.

Table 60. Size variation of Pristoceuthophilus pacificus.

t 1 iltlitj 3q ^s2 S& BiiiciiAM VouNc Univehsity Science Bulletin

Scusonul Occurrence. 'I'lic coinplote seasonal antennae and, exc-ept for Mtjrmecopbila, auditory occurrence can nol he niven for tliis species in- organs on the front tibiae. The males have asmuch as the stndics in wliidi it is found were striilulatorv organs on the tegmina, except, not operated continuously as they were at lower again, for Mijrmccopliila which is completely elevations. Nytnphs were collected from July wingless. Crickets differ from the other Ensifera, 18 to November 2. .\dults were collected Irom the long-horned grasshoppers, in having three- August 11 to jannarv 10. The nvmphs were most segmented tarsi, an awl-like or needle-like ovi- common in .\ngust, the adults most coTiiinon in positor, and, when winged, tegmina which are October. fiat above and bent sharplv downward at the sides of the body. They are essentially nocturnal, Localities Kepresented. Specimens examin- but are also acti\'e to a considerable extent dur- ed (nvmphs and adults): 196. ing the day. Some are among the commonest

EC;il 1 adult male, January 10. Studv insects and are widely distributed; others are 4. Study TC.\I, 1 adult female, December rare and very local in distribution. Studies 12A and 12E, 194 nymphs and adults Crickets are more omnivorous than long- as follows: 70 nymph males, 81 nymph females, horned grasshoppers and will eat animal sub- 12 adults males, .31 adult females, from July IS stances and other insects. The tree-crickets, par- (nymphs) to November 9 (adult male). ticularly, feed largelv upon aphids.

life history, crickets agree, with ex- Additional Remarks. There is some <|uestion In few as to whether or not this series represents a dif- ceptions, with the majority of the Orthoptera in ferent species, or whether or not subspeciation hatching from the egg early in the season and might have taken place. No comparisons were developing to maturity during the summer. The made. eggs of field crickets are deposited in the soil; those of tree-crickets are placed in the bark or Family Gryllidae l^ithy stems of the plants among which they live, The crickets have long, delicately tapered in holes drilled bv the female.

Ke\- to the Subfamilies of (aivLLro.AE

1. Caudal tibiae armed with rows of long .spines (Fig. 100) 2

Caudal tibiae without rows of long spines, but with rows of short teeth (Fig. 101); body covered with scales Subfamily .Mogoplistinae, page 113

2. Completely wingless; iiind femora ovate, enormously enlarged (Fig. 102); eyes small; size minute, less than 4 mm. in total body length. Inhabitants of ants nests. '_ Subfamily Myrmecopliilinae, page 116

Winged, at least in the adult male; hind femora elongate; eyes not small; size well over 4 mm., medium to large in.sects 3

horizontal; fonn 3. Caudal tibia with minute teeth between the spines (figs. 103, 104); head slender. Cncnisli. Subfamily OEcanthinae, page 115 or Caudal tibia i.u king niiiiute teeth between the sjiines; head \t-rtieal; form robust. Brown Gr\'llinae, 113 l3la(_.l( Subfamily page

FIG. 103 FIG. 104

cotnprc- 100-104, 100, Arlu'la iixsimilis, ft-male, caiulai (ibiii aiul tarsus, latc-ral xii'w. 101, CijcloptHum. male, caudal ap- Itcncleiui fortior. male, caudal tibia and tarsus, lateral view. 102, MijrmccopMu manni. appendage, lateral view. 104, pendage, lateral view. 10.3, OEcunthus citlifomicus c-iliforiticus. male, caud.il OE. c. californicus, male, detail of caudal tibia, lateral view. Biological Series, \'ol. 4, No. 3, September, 1964

Subfamily Mogoplistinae da Test Site is was found in only three areas, near Cane Springs and Jackass approach. The bush-crickets are small (approximately 10 mm. in total body length), flat, slender- Habitats. This cricket inhabits the desert bodied insects, brown in color and covered by regions and is present over most of tlie Lower translucent scales. The hind tibiae have two Sonoran zone throughout its range. The insect rows of short teeth but no true spines, which is nocturnal, and was collected only in the can character will distinguish them all from other traps established in the studies. It apparently species in this family. moves at night but is secretive during the day. The song is reported as a high-pitched trilling which is continued over a considerable period. Genus Ctjcloptilum Scudder Seasonal Occurrence. Specimens were col- 1869. Cijcloptilum Scudder, Proc. Boston Soc. lected only in September and the insect can Nat. Hist., XII, 142. p. be found at the test site probably in late summer and early fall.

Cijcloptilum comprehemlens fortior Localities Represented. Specimens examin-

Hebaid ed ( adults ) : 4. (Figure 101; Table 61; Map 30) Stiidy CBA, 1 adult male, 2 adult females, September 2 and 14. 1931. Ctjcloptilum comprehemlens fortior He- bard, Trans. Amer. Entom. Soc, LVII, pp. 153- Study JA, 1 adult female, September 2. 157. .\dditional Remarks. This insect is probably Distinctive Features. This tliamnophilous more common than would be indicated by the cricket is distinctive and will not be confused citations. Some areas were swept for the insect, with any other insect from the Nevada Test Site. however, without success. It should be searched It is characterized by the body covering of trans- for in the evenings when it first becomes active. lucent scales and the absence of spines on the The specimens were compared with Cijclo- caudal tibiae. ptilum comprehendem interior Hebard from the type locality in Washington County, Utah, and Coloration. The head and pronotum of tliis they are distinctly of the fortior group. No valu- insect are reddish brown, the short wings of the ation or intergradation could be detected by the male are light brown, the dorsal abdomen of few specimens. the female dark brown to blackish.

Distribution. The tvpe locality of this species Subfamily Gryllinae was designated as Ajo, Pima Co., Arizona. The species, itself, extends from the Great Plains This subfamily includes tlie common crickets states south to Texas and west to California. The which need no introduction to anyone aware of race fortior is the southwestern representative, the insect world. They are robust, brown or being found from western Texas to southeastern black, and easily recognized morphologically by California and south into Mexico. At the Neva- the rows of fixed spines on the caudal tibiae.

Table 61. Size variation of Cycloptilum comprehemlens fortior.

t c Bhiciiam Vounc UiNivEnsiTY Science Bulletin

Ck-nus Acluia Faliriciiis Cloioration. The color of the field cricket ranges from light brown through dark brown 1775. Achrta Fabriciiis, Systema Entoinologiai'. and almost black. There is considerable c-olor Flcnslnirgi cl Lipsiac, p. 279. Variation in the series from the test site. Speci- mens from higher ele\ations tend to be darker Acheta assimilis Fabricius and smaller; specimens from more sandy areas (Kigurt-s 77. 78. 116; Table 62; Map 30) tend to be larger and much hghter. Intermediate forms can also be found. They are all generally 1775. Acheta assimilis Fabricus, Syst. Ent., p. quite unicolorous. 280. Distribution. In North America the field Synonomy. Tlic svnonoim- of this .species is cricket can be found nearly everywhere except still in (jiiestion. More than 45 names Iiave been at high altitudes and in the far north. It ap- applied to the American field crickets. Rehn parently cannot tolerate too much moisture as and Ilebiird (1915) conckided that only one it is not found in wet areas. It is found nearly liiglily variable species was represented, and everywhere o\er the Nevada Test Site, but was since that time ail of the native field crickets not collected on Rainier Mesa. have been discussed under the name of assimilis-. Tliis common insect has been discussed in liter- Habitats. Most of the insects were captured ature under the generic names of Gryllus and in can traps where they were collected as a re- Grtjlhdus. sult of their nocturnal wanderings. Tliey seek Alexander (1957) recognizes five species of refuge during the dav under anv debris or under field crickets in eastern United States in addi- rocks on the ground. They are particularly active tion to the house cricket, Acheta clomesticus after a summer's rain. The nymphs are more (Linnaeus). Even though tlie type locality of active during dry periods. In July, 1961, the assimilis is in Jamaica, the western populations adults were taken in the can traps only after a of the field cricket are recognized as that species rain, although the nvmphs were previouslv col- until a complete revision is made of the Ameri- lected from the same areas. can field crickets. Seasonal Occurrence. Nvmphs were collect-

Distinctive Features. Tliis is the very com- ed throughout the entire year, except no speci- mon thick-bodied cricket which is so widely dis- mens were taken during February and Decem- tributed. No other insect at the Nevada Test ber, and only one in January. Adults were col- Site even resembles the species. There are cer- lected from .April through September. They tain dark crickets, smaller in size, throughout were most common during the month of May. the west (genus Nemobius) which resemble Localities Represented. Specimens examin- assimilis, but which can be recognized by the ed (nymphs and adults): 113. large movable spines of the caudal tibiae. In Acheta the spines are not movable. Salsola area (Study IF), 6 specimens, July 5 to November 24. Morphological Variation. Probably no other Cmtjia-Lijcium area (studies IB, IG, 4A), 59 species of Orthoptera has as much variation as specimens, January 17 to October 24. this insect. Extreme differences can be found in a series collected at the same time in one area, Larrea-Franseria area (Study 5A), I speci- in the development of the wings and tegmina, men, October 16. the distribution of hairs on the body, and even Afriplcx-Kochia area (Study 6A), 4 speci- the development of the appendages. mens. May 3 to September 6.

Table 62. Size variation of Acheta assimilis.

Length Length Length Extension of Length Breadth Length Body" Pronotum Tegmcn Wiiig Beyond Caudal Caudal Ovipositoj Tegmen Femur Femur

(^, Minimum Size 24.6 2.9

(j' , MiUiimum Size 31.7

9 , Nfinimum Size 27.3 31.7 9 , Maximum Size

9 , CBA5, July 1, 1960 34.5 Biological Series, V'ol. 4, No. 3, Septemder, 1964 115

Coleogtjne area (Study lOD), 1 specimen, Distinctive Features. The tree-crickets are July 5. distinctive insects and can be recognized by the Lijcium area (Study 5E), 2 specimens, July characters given in the keys. There is no other 18 and August 13. insect found at the Nevada Test Site with which they can be confused. The two species found in Yucca-Coleogijne area (Area 6), 10 speci- the fauna at the test site can be distinguished mens, August 28. by color and markings on the antennal segments. Mixed areas (studies CBA, JA), 27 specimens. May 16 to November 24. Coloration. This insect is ivory to

Distribution. It is limited to the western Subfamily OEcanthinae United States where it is widespread. At the

The tree crickets are small, delicate crea- Nevada Test Site it was very limited in distri- tures, whitish in color, usually shaded with green bution and probably never appears commonly. or brown. The wings are fully developed in both Habitats. This tree-cricket is sexes, in the male the tegmina broadly expand- most commonly found on the larger shrubs and, in certain ed and paddle-shaped, flat on the back, in the female the tegmina narrow and wrapped closely areas, in trees. The only recorded vegetation from the Nevada Test Site was from Atriplex about the body. The ovipositor of die female is rod-shaped. canescens. Tree crickets are found on trees and shrubs, Seasonal Occurrence. The incidence of the or on vegetation generally, only accidentally on insect at tiie test site is unknown. It was col- the ground. Their songs are loud and are among lected only in August and September, and the most noticeable night noises. These insects throughout most of its range is a late summer are most active at night, but may be encounter- and early fall insect. ed during the day. They feed not only on leaves, flowers, fungi, and fruit, but consume large num- Localities Represented. Specimens examin- bers of small insects, such as aphids and scales. ed (nymphs and adults): 4. The eggs are deposited in the bark of trees or in the pithy center of plants, in holes made Area 4, miscellaneous collection, 1 adult by the ovipositor of the female. During their male, August 26, no record of the vegetation early nymphal existence they possess pronounced upon which it was taken. predaceous habits. As they approach maturity they become more vegetarian.

Genus OEcanthtts Audinet-Serville

1831. OEcanthus Audinet-ServiUe, Ann. Sci. Nat., XXII, p. 134.

FIG. 105 FIG. 106 OEcanthus californicus califoniiciis Saussure (Figures 103-105; Map 31) Figs. 105-106. 105, OEcanthus californicus californicus, male, proximal antennal segments, cephalic view. 1874. OEcanthus califarnictis Saussure, Miss. 106, OE. nigricornui quadripunctatus, male, Scient. Mex., Rech. Zool. VI, p. 462. proximal antennal segments, cephalic view.

Key to the Species of OEcanthus

Front side of first antennal segment never ornamented with more than a narrow black line along inner edge (Fig. 105); subgenital plate of female with a notch half as broad as the widest part of the plate; tegmina of male plainly colored. OE. californicus californicus Saussure

Front side of first two antennal segments ornamented with two black marks, the first segment with black line and dot which are narrow and well separated (Fig. 106); female subgenital plate with a narrow notch OE. nigricornis quadripunctatus Beutenmuller )

116 Bkiciiam Vol'nc Unmvkhsity Science Bulletin

Area 5, miscdlaneous collection, 1 adult Tlie one female, however, has more the typical male, September 26, no record of the vegetation. antennal markings of iptadripunclatiis to the Area CM, Cane Springs, 2 female nymphs, east (through Utah and Colorado), and is less August 10 and 22, on Atriplex canesccns. distincti\i' of the heavier markings of O. m

1894. OEcarUhus ijuadripunctatus Beutenmuller, Subfamily Myrmecophilinae Bull. Amer. Mus. Nat. Mist., VI, p. 271. Genus Mijrmecophila Latreille Distinctive F"eatures. Tiic features by which 1829. Mi/rniC(oplul(i Latreille, Regno Anim., this tree-cricket can be distinguished from cali- (ed. 2), V, p. 183. foniictis are the markings on the antennae. The The ant-io\ing crickets can be recognized by first and second antcnnal segments each have their extremely small size and the fact that, with two black marks on the under (front) side, few exceptions, they are found as commensals of the inner mark on the first segment is linear, ants. (Specimens have been found b\' the autlior straight, the distal end with a tendency to curve under rocks where no ants coidd be located. outward toward the outer spot which is small Colonies of ants may have, at one time, been and round. located imder the rocks, but none were there

Coloration. Compared to tiie other tree- when the crickets were captured. ) The %'arious cricket from the test site, (juadripunctotus is species of crickets can be determined by the pale greenish-white, becoming yellowish when number and proportionate length of the spines dried. and spurs of the caudal tibia, and the spines and spinulae of the caudal metatarsus. Tlie dor- Distribution. This insect has a wide distri- sal margins of the caudal tibia are armed with bution over the United States from Canada to one external and three or four internal spines; Texas and from the Great Plains westward. It the distal extreinit\' is armed with three pairs of was found only at Cane Springs at the Nevada spms, tlie \entrai pair being minute. Test Site.

Habitats. Low, heavy vegetation is the usual Mytmecophila nmnni Schimmer habitat of this group. At the Nevada Test Site (Figure 102; Table 64; Map 32) it was collected on Eltjmus cinereus. 1911. Mijrmecophila mantii Schimmer, Deutsch. Seasonal Occurrence. The specimens were Ent. Seitschr., 1911, p. 443. collected in June, but no information can be Distinctive Features. This species can be re- given as to its seasonal occurrence at the test cognized by the characters discussed under the site. It is, however, an earlier insect tlian culif- genus above. Specifically, the dorso-internal ornicus. margin of the caudal tibiae are armed with four Localities Represented. Specimens examin- spines, alternating in length. In his revision of ed ( nymphs and adults ) : 3. the genus, Hebard (1920b) commented tliat Study CM, Cane Springs, 3 specimens, 1 male rarely in manni one spine is missing. subadult, 1 female subadult, 1 female adult, C^oloration. The general coloration of the June 15 and 24, on Elymus- cinereus. spec ies is pale, yellowish brown or slightly dark-

.Additional Remarks. It is rather difficult to er, except the eyes which are blackish-brown determine with any certainty a race of Ortho- and the distal portion of the female ovipositor ptera on the basis of just one adult specimen, es- which is shining dark reddish-brown. The ab- pecially where color markings are key factors. dominal segments are sometimes margined

Table 63. Measurements of OEcanthus nigricornis qwidripttnctatus.

Length igth Length Length Bre;iflth Length Body oluni 'I'egmen Clau'dal Cand;J Ovipositor Femur Femur

18.2 2.0 11.7 8.0 0.9 4.6 9 , CM, June 24. 1961 Biological Series, Vol. 4, No. 3, September, 1964

Table 64. Size variation of Mymiecophila nuinni.

•s-^ ^1 A '« -= f.S t

1) ;- papa J £ rati; ju m u ju J ^ JO JO

0.5 0.88 cT, 12E, July 24, 1961 3.4 1.78 0.88 1.65 1.6 0.95 1.18 1.3 0.85 1.2 1.4 1.3 0.9 1.43 0.9 cf, 12A, July 24 ,1961 4.0 2.1 0.93 1.85 1.88 0.98 0.63 2.1 1.15 2.03 1.98 1.03 1.43 1..35 1.03 0.65 1.45 0.98 1.3 9 , 12AC9, AugiLst 12, 1961 3.53 14, 1961 4.25 2.28 1.08 2.08 1.93 1.05 1.45 1.4 1.0 0.5 1.53 0.9 1.4 § , 12AC9, August

caudad with a slightly darker shade, giving these able object, stone or board, the crickets would individuals a banded appearance. All of the im- be clinging to the object removed. This has also mature specimens are noticeably light tan in been the habit of Mnjmecophila aregonetisis color. Bruner collected by the author in Oregon. In an effort to determine the habits of this Distribution. This species is typically an in- species of cricket the author maintained a colony habitant of the semi-arid and arid regions of the of ants, crickets and other mrymecophilous western United States, ranging from southern arthropods in the laboratory and a number of Washington to the Mexican border. The insect significant observations were made. These speci- was collected at the Nevada Test Site in two mens were not from the Nevada Test Site, but extreme areas: one immature specimen was col- from the vicinity of St. George, Utah, and while lected at the edge of the playa at French- the artificial conditions may be different and man Flat; the other habitat, one where Mijrmcco- the crickets react differently under laboratory phihi was common, was at a higher elevation in conditions, the information may contribute to the pinyon-juniper areas on Rainier Mesa. a general knowledge of the group. The first ob-

Habitats. This species is a commensal of servations were made from October 28 to No- ants, the host species at the Nevada Test Site vember 25. The material was originally col- being Formica integroicles pkinipilLs- Creighton, lected from the ant's nest, including soil, grass Formica fusca Linnaeus, Formica lasioides and roots, and different arthropods including

Emery, and Camponotus vicinus Mayr. ( De- ants and their eggs, pseudoscorpions, small tene- terminations of ants were made by Dr. Arthur brionid beetles, hister beetles, collembola, and C. Cole, who has been associated with the radi- fifteen crickets in various stages of development. ation ecology project at Mercury.) The one Thev were introduced into a glass jar for obser- nymph from Study 5E was captured in a can vations. trap. In a subsequent visit to this station a The adult crickets almost immediately seem- colony of Pogonomijnnex calijormcus (Buckley) ed to establish territories and would not ap- was found within a few feet of the trap, and al- proach other adults. They would extend their though the tunnels and nest were carefully ex- legs to maximum extensions (standing up) in posed no other specimens were collected. One order to see over a larger area and ward off any significant generalization can be made from the intruder. This territorialism may account for the capture: the crickets do leave the protective cus- fact that very few specimens are foimd in any tody of the ant colonies and wander about on one colony of ants, at least in those areas ex- the ground, probably being nocturnal. amined. On Rainier Mesa some of the specimens Tlie following day cracked wheat and grass introduced the colony and the were captured with ants in can traps established seeds were to crickets fed this material, or at least they in the area, but more frecjuently they were cap- upon tured by turning over rocks, under which ant were attracted by it. colonies would be found. In these situations the During the first week of observation the crickets were always, without exception, cling- crickets spent most of their time on the exposed ing to the rocks rather than being on the ground. grass roots and stems and had not apparently in- This habit was observed by the author in many vaded the ant chambers which had been dug. collections in widely separated areas in Wash- They were extremely active over the surface ington and Kane counties, Utah. Whenever the area at night, only moderately active by day. crickets and ants were found under any move- Some of the ants had died off during the first BitioiiAM Vot'Sf: Univehsity Scienck Bui.uirriN

wi'fk, but tlu- t()llcrnl)()la, I'spi-cially, wore aci- In the special chamber where the under- justed to till- laboratory conditions in that their ground acti\'ity could be observed, whenever numbers luid increased. a cricket would emerge into a timnel, it would

After this colony iiad been maintained for immediately jump and remain on the ceiling two weeks nearly all the ants had died off. Ad- wliene\'er an ant would come through the tunnel, ditional ants, along with dirt anil debris, were returning to the bottom after the ant had gone. collected from the same natural colony and re- Mi/niircophilus seems to breed at any sea- established in tlu' laboratory. Three crickets son inasmuch as nymphs have been found were also introtluced with this second addition. throughout the year. 11 both the crickets and the On \o\ember Seasonal Occurrence. .No complete seasonal collembola had multiplied, as there were many data can be given for this species because of the first instar n\'mph crickets among the grass. No few trips and records made from Rainier Mesa, cricket eggs had been obseryed, but they may the area where the insects were found. Nymphs have been present in one or botli of tiie intro- .uitl adults were collected onh' during the ductions. months of July and August, with most of the The crickets were seldom seen in the ant captures of both ininphs and adults being in burrows by day or night, but apparently pre- July. ferred the grass. Localities Represented. Specimens examin- This original experiment was terminated he- ed (nymphs and adults): 11. cause the plant materials molded and all the in- Study 5E.\7, Frenchman Flat, 1 male nymph, sect life eventually died off. July 15, commensal of Pogonoimjrmex californi- Another colony was established in February cus (?). in a special narrow chamber, glassed on both Study 12A, disturbed area on Rainier Mesa, sides, so any digging and acti\'ity underground 7 n\mphs and adults as follows: 3 nymph males, could be noted. .\nts and crickets were placed July 24 to August 24, commensal with Formica in the chamber and after the establishment of inlcgroides planipilis (2 specimens) and Far- tunnels some table sugar was placed on the sur- tnica kisioides (1 specimen); 1 adult male and face. Some of the crickets went directly to the 3 adult females, July 24 to August 14, commensal sugar and began eating it, but avoided the ants. with Formica jtisca (1 specimen) and Formica It was observed that the crickets stood up as intcoroides plampili.f (3 specimens). frequently and in so doing high as possible \ery Study 12E, undisturbed area on Rainier the females would thrust the ovipositor forward Mesa, 3 nymphs and adults as follows: 1 mmph, to be cleaned by the mouthparts. The hind tarsi sex not determined, and 1 female nymph, July tibiae were also cleaned in this same man- and 24, commensal with Formica fusca: I adult male, ner. The crickets always avoided the ants and July 24, commensal with Componotus vicitms. never came close enough to them to feed on the oily secretions on the surface of the body .Additional Remarks. It is interesting to note of the 'ants, as stated by Wheeler (1900). Ob- that the comparatise numbers of specimens is the servations made by various authors indicate that same as the species of Ceiitliopliihi.s and Pristo- the food of these crickets is largely the secre- ceulhophilus, in that most of the specimens have tions which lubricate the ants' bodies and which lx;en collected from the disturbed area on Rain- are left on the walls of their passage-ways, tiiis ier Mesa where the rocks have been loosened being partly the food of the ants, also. This and fissiues occur in the ground. could not be contradictetl nor substantiated in .\lthough the genus Mijrmecophila was re- the observations made. Ilebard {I92()b) stated vised in 1920, additional work is needed to re- that Mt/ivH'cophild are wiiolly dependent upon define the \'arious species and to designate their the host for the type of nourishment re(|uired. distributions. It was found, however, in all colonies examined that whenever protection was offered, as in the Suborder Pii asm.\toptkh.\ case of a rock or board over the ant nest, the Superfamily Pn.\SM.\T<)n)K.\ crickets emerged from the nest and found shel- Family Phasm.\tid.\e ter under the cover and away from the ants. They retreated for tlie protective custody of the The walking-stick is among the curiosities ant nest when molested, but would not enter un- of the insect world. It has an elongate, slender, exserted heail. Tlu' til they could do so without running into the and i\liudrical body with an ants. prothorax is \ery short, the meso- and meta- Biological Series, \'dl. 4, \o. 3, September. 1964 119

Key to the Subfamilies of Phasmatidae

Antennae not more tlian one-half as long as the anterior femora Subfamilv Pachymorphinae, page 119

Antennae distinctly longer than the anterior femora Subfamih' Heteronomiinae, page 119 thorax elongate. The legs are slender and all A striking broad band of brown is present on the alike in form. Teginina and wings are lacking head and thorax but becomes weak on the ab- in all of the United States species. A large domen. arolium is present between the claws at the end Distribution. The range of this species is of the five-segmented tarsus. Tlie ovipositor of from Oregon and California east to Utah and the female is concealed by the subgenital plate Arizona. At the Nevada Test Site it was found and the cerci are not segmented. only in Area 6. The walking-sticks are remarkable for their resemblance to twigs of plants or to dead grass. Habitats. This species is largely found on They are protected effectively by their habit of range grasses, but has been reported from rab- moving very slowly and deliberateh' and of bit brush, burroweed and other desert perenni- remaining motionless for long periods of time, als. It was found in a Yucca-Coleogijne area at which makes them very difficult to observe. The the Nevada Test Site, but no record of the legs, if lost, may under certain circumstances, vegetation was made upon which the insect be regenerated, and individuals exhibiting ap- was found. pendages in this process are not infrc(juently Seasonal Occurrence. The only collection of seen. These regenerated appendages may be dis- the insect at the test site was in August. No tinguished bv the absence of one tarsal segment. other data can be gi\cn as to its seasonal occur- Tlie eggs closely resemble seeds of plants and rence. are dropped on the ground at random. All of the walking-sticks feed on the leaves of plants. Localities Represented. Specimens examin-

They are herbivorous and are usually found on ed (adult): 1. shrubs and trees or among grasses. Area 6, miscellaneous collection, 1 specimen, The insects can best be collected by sweep- August 28. ing the vegetation inasmuch as they arc difficult to see before capture.

Subfamih- 1 Ieteronenhinae Genus Psciulosermijlc Caudell Subfamily Pachymorphinae Genus Parahacillus Caudell 1903. Pscudosermijle Caudell, Proc. U.S. Nat. Mus., XXVI, p. 867. 1903. PambaciUm Caudell, Entom. News, XIV, p. 314.

Psciidoscniiiile siramincus ( Scudder) PamhaciUus licspertis Hebard (Tabic 65; Map 33) (M;,p 33) 1902. Bacuncuhis stmmineus Scudder, Proc. 1934. Parahacillus hesperus Hebard, Trans. Davenport Acad. Sci., IX, p. 20. Amer. Entom. Soc, XL, 286-290. pp. Established Synonomy. Pseudosermyle trtin-

Distinctive Features. Tiiis is a medium or cata Caudell; Pseudosermyle tenuis Rehn and small and extremely slender walking-stick. The Hebard. antennae are short, less than three times the Distinctive Features. This species can be dis- length of the head. Tlie surface is smooth, with- tinguished from the other walking-stick found out tubercles, but with a prominent medio- longi- on the Nevada Test Site by the longer antennae, tudinal carina on the pronotum and with coarse distinctly longer than the anterior femora in both low sub-marginal longitudinal carinae on each sexes. The surface is sub-rugose and the head side. The limbs are very slender, unarmed. The has two pairs of prominent carinae. The males female is considerably longer and more robust differ from the females in being entirely smooth than the male. except for two main carinae on the anterior part

C;oloration. The coloration is typically straw- of the head between the eyes, and in being yellow, but may vary from light to dark brown. smaller and more slender. 120 HiiioiiAM ^ouNc Univehsity Scienck Bulletin Biological Sebies, Vol. 4, No. 3, September, 1964

Table 65. Size variation of Pseudosermyle stramineus.

o -I Sh

J', IGDl, July 17, 1961 40.5 1.8 9.8 6.8 1.2 J', TCB, June 22, 1961 33.7 1.7 7.5 6.3 6.4 9 , CM, June 13, 1961 37.8 2.1 8.4 122 Bkiciiam Yoi'NC Univkksitv Scienck Bvi.lktin

Lilancutria minor (Seuddcr) Localities Represented. Specimens e,\amin- (T.il.K- m. Map 34) etl( nymphs and adults): 88.

1872. Htd'^mdloplcra minor Sciiddcr, U. S. Ct-ol. Saiwla area (Study IF), 2 specimens, June Surv. Nebraska, Final Heporf, Part 3, p. 251. UJ to 2.3. Graijia'Ltjcium Established Synonoiny. Lilancutria ocularis area (studies IB, IG. 4.\) 15 specimens. 5 to October .30. Saiissiire; Lilancutria ohscura Scudder; Lilaneu- May tria pacifica Scuddor; Litanculria nkinncri Rehn; Larrca-Framcria area (studies .5.\, 5C(^), 9 Lilanculria lon

Distribution. This is a verv widespread spe- mens, September 26 and October 3. cies of the west, occurring from the Creat Plains Area 6, miscellaneous collecting, 1 speci- westward and from IJritish (.'oiiunbia south into men, July 15. Mexico. It is distributed throughout most of the -Mercury area, 3 specimens, .August 24. Nevada Test Site, but was not found at higher Camp elevations. .Additional Remarks. One nymph (5CQ23, Habitats. This small, elongated insect can July 11, I9(>1 ), 5 mm. long, has the eyes distinct- be detected running rapidly about on the ground 1\' pointcil abo\e, suggestive of Yermniops. No in the desert areas and will only occasionally re- other nymphs show this condition and no adults of that collected sort to vegetation in an effort to escape capture. genus were at the Nevada Test Site. insect referred is therefore The terrestrial habits of the insect resulted in The to considered an abberant form of Lilancutria. its fre(juent capture in can traps.

Seasonal Occurrence. Collections were made from April 29 to October 30. Nymphs were pre- SubfamiK M \n ri:iNAK sent into September; adults were first collected Genus Sta^j^niouuiiifis Saussure in June. It would appear from the occurrence of the n\ui[)hs that part of the eggs laid during the 1869. Sla<^momantis Saussure, Mitth. Schweiz. summer hatch into nymphs that same season. Ent. Ges. Ill, pp. .56, (i5.

Table 66. Size variation of LiUineutria mhtor. Biological Series, Vol. 4, No. 3, September, 1964 123

Stagmonuinfis californicus Studies CBA and JA, 3 specimens, July 15 Rehn and Hebard to September 2, vegetation not recorded from which the (Table 67; .Map .34) specimens were taken. Study CM, Cane Springs, 3 specimens, July 1909. ^tagmomantis californicus Rehn and He- 15 to September 30, no vegetation records. bard, Proc. Acad. Nat. Sci. Pliila., LXI. pp. 409- Study TCB, 2 specimens, 16, no vege- 483. July tation records. Distinctive Features. This insect can be Area 6, miscellaneous collecting, 2 speci- distinguished Li- readily recognized and from mens, August 28, no vegetation records. taiiciitiia by its large size, its coloration, and its Camp iMercurv area, 1 specimen, Septem- tliamnophilous habits. ber 10.

Coloration. Tlie bodv of this insect is usual- .Additional Remarks. Thomas (1875) report- !)• green, but occasional yellowish or brown ed three specimens belonging to the family specimens are found. The brown specimens are Manteidae that were collected by members of most frequently with light maculations. The the \Mieeler Survey, one of which was new and hind wings are usually brown, often marked with was described at that time as follows: ashy blotches, or frequently purple, red, or even orange-yellow. The wings of the specimens col- "Mantis wlicclcrii, sp. nov. lected at the test site were red, but in observed specimens they changed to brown within four "The specimen is dry, and is so badly dam- hours after death. The first four dorsal ab- aged that it is impossible to determine positively dominal segments of the male are broadly edged the genus to which it belongs, or to do more with darker brown. than indicate some of its leading specific characters. Distribution This species is common "Female. Head flat, transverse, triangular in throughout the Larrea-Franseria deserts of the — front. Occiput short, reduced to a transverse southwest. It distribution extends from Calif- ridge. Vertex transverse, directed downward and ornia east to Colorado and Texas. This mantid backward toward the face, with four slight is not common at the Ne\'ada Test Site even longitudinal depressions. Ocelli distinct and though it was found in a nimiber of different prominent. The face transversely quadrilateral; areas. the upper carinate margin bent upward be- Habitats. Cenerallv (|uite common through- tween the antennae. The antennae wanting. out its distribution, this insect is found on shrubs Prothorax about twice the length of the rest of and low vegetation, and is frequently attracted the thorax; the margins minutely serrate, slight- to lights at night. Its most common occurrence ly emarginate, scarcely expanding posteriorly at the test site was on Larreci divaricata. near the transverse incision. Anterior femora denticulate on the exterior carina. Abdomen en- Seasonal Occurrence. Nymphs were collected larged, fusiform. Middle and posterior legs from July 10 to September 30. The only adult wanting, and but a remnant of the wings re- occurrence was in September. Tlie adults im- maining. doubtedly are found into October, at least, al- "Color. Yellow, probably faded from a pale though no specimens were collected. — green. The abdominal segments with a piceous Localities Represented. Specimens examin- black fascia or ring on the posterior margin of ed (nymphs and adults): 15. each. The remnants of the wings carneous-red. Study 5A, 4 specimens, July 10 to IS, on "The specimen is too much injured to give Lanca divaricata. any very acurate measurements; but the follow-

Tablc 67. Mea.surements of Stagmomantis californicus.

3 3 5 S bO >^ A C 13

, 14.8 12.6 10.9 J' CM, September 30, 1961 124 Briciiam Young University Science Bulletin

Key to the Genera of Polyphagidae

Middle and caiidal ti-iiiora with dorsal genicular spine at apex (Fig. 107) Arenivaga Rehn

Middle and caudal femora without dorsal genicular spine at apex Eremoblatta Rehn ing approximations will indicate the size: — Length, 2.2 inches; prothorax, 1.0 inch; anterior femora, 0.5 incli; anterior tibiae, 0.6 inch." Tliis species was subsequently placed into c::^ synonomy with ^tagmoniantis carolitm (Johann- son), but the c-olor markings arc as in S. californ- FIG. 107 icus. Eidier S. calijornicus of Rehn and Hebard I'^ig. 107. Arenivaga erratica, female, caudal femur of S. Carolina should be is a color variation and slxowiug distal spine, lateral view. placed into synonomy, or, if it is a good species, it should be relegated to a synonomic position The number of generations of these insects of Stagmomantis wheelerii (Thomas), and tlie per year appears to differ with the species. The latter reestablished as tlie scientific name of this native species produce apparently a single brood insect. per year, but those ad\entive species, commonly Superfamily Bl.\ttode.\ found in houses, may produce several broods per year. Tlie eggs are laid enclosed in ootheca, Family Polyphagidae which shows their true relationships to the The orthopterans with strongly depressed, mantids, which are carried about for several days more or less oval, bodies are readily referred to protruding from the body of the female before the superfamily Blattodea. Other distinguishing being finaliv dropped, appareiitlN' at random. characteristics separate them from the other The native roaches live under or within ob- Orthoptera. The head is concealed beneath the jects and are commonly found in rodent nests, pronotum, tlie face is ventral, the mouth poster- especially of the rat Neotanui. No attempt has ior, and tlie antennae long and filiform. The been made to include in this discussion those legs are slender, similar, and compressed, with roaches which might have become established the coxae long and free. When fully developed, as adventives, and which are found only in tlie the tegmina are parchment-like and overlapping, residences and buildings. and the wings membranous, with a large anal The males of the native species are long- area. Both tegmina and wings are often rudi- winged, buff with brown markings, and are mentary or wanting in the female and some- often seen because they are attracted to lights times in both sexes. The Nevada Test Site fe- in large numbers. The females are round and males are all apterous. wingless. Most of the specimens taken at the The sexes may be distinguished without diffi- Nevada Test Site were collected in can traps as culty, although there is no visible ovipositor. a result of their foraging at night. The males are characterized, in addition to tlie conspicuous cerci, by the presence of a pair of Genus Arenivaga Rehn styles at the sides of the caudal margin of the 1903. Arenivaga Rehn, Proc. Acad. Nat. Sd. last ventral segment of the abdomen. 181. These insects are commonly known as roach- Phila., 1903, p. es. They are nocturnal and remain in darkened In his revisionan,' studies of the genus. He- places during the day. At night they run about bard (1920c) stated: "So much indi\idual \ari- seeking food, and attack everything edible. ation occurs in tlie species of tliis genus, in the

Key to the Species of Arenivaga (Modified from Hebard)

Right, ventral genital plate of male without projections, the right dorsal genital plate vertically broad, with margins rounded and surface smooth (Fig. 108). Limbs of female more elongate and slender; dorsal surface of abdomen normally maculate A. erratica Rehn

Right ventral genital plate of male with projections; right dorsal genital plate large and lobi- internal fomi, proiluced inward from its left distal portion in an elongate heavy spike, the margin beyond the base of this spike armi'd with two small teeth (Fig. 109). Limbs of female shorter and stouter; segments of abdomen nonnally immaculate A. apacha (Saussure) Biological Series, \"ol. 4, Xo. 3. Septemulh. 1964 125

Arenivaga found at the test site. It is western in distribution, being found from California to western Texas and from southern Utah and southern California to Mexico. It has a wide distribution over most of the Nevada Test Site.

Habitats. Like other roaches, the species is FIG. 108 FIG. 109 nocturnal, the males being attracted to lights at Figs. 108-109. 108, Arenivaga erratica, male, con- night. They were most commonh' collected in cealed genital structure. 109, A. apacha, male, the can traps as a result of tlieir nocturnal concealed genital structures. wanderings. Tliey are frequently found in rodent nests, or in the tunnels associated witli the nests. features normally used for specific separation, They are especially common in sandy areas. that we feel it is imperative for the student to Seasonal Occurrence. This roach has been examine the concealed genitalia of all males to collected from April 4 to October 23. be recorded. The other features which we con- Botli nymphs and adults were present sider of some diagnostic value, and the degree from April through October. They are of variation known, are discussed under the most common during the month of August. species." Localities Represented. Specimens examined (nymphs and adults): 99. Arenivaga erratica Rehn Salsola area (Study IF), 1 specimen, (Figures 107, 108; Table 68; .Map 35) Aug- ust 4. 1903. Homoeogamia {Arenivaga) erratica Rehn, Graijia-Ltjcium areas (studies IB, IG, 4A), Proc. Acad. Nat. Sci. Phila., 1903, p. 187. 47 specimens, April 6 to October 12.

Distinctive Features. Several different spe- Larrea-Franseria areas (studies 5A, 5CQ), 3 cies of Arenivaga are found in the southwestern specimens. May 22 to September 2. desert areas. The two species found at the Ne- Atriplex-Kochia area (Study 6A), 4 speci- vada Test Site are difficult to differentiate, but mens, June 26 to September 15. may be recognized by an examination of the Coleogijne area (Study lOD), 8 specimens, external genital characters of the male. The fe- June 2 to September 15. males are more problematical in their differ- Lycium area (Study 5E), 8 specimens, May entiation, as no reliance can be placed on the 4 to August 26. dorsal markings. Mixed vegetation areas (studies CBA, JA), Coloration. The males of the species ar€ 52 specimens, April 4 to October 23. light buff in coloration with tegmina and wings Study lOS, 13 specimens, June 19 to 29. of approximately the same color. The females Additional typically have some darker maculations on the Remarks. The concealed genital characters of entire series dorsal surface. The ground color of the females the of male specimens of Arenivaga collected at the Nevada Test Site is usually darker than the males. were checked. In addition, measurements were

Distribution. This species is more numerous made on sufficient female adults to show no and has a wider distribution than the other statistical differences in the population, so the

Table 68. Size variation of Arenivagu erratica. B)\i(;iiAM VoiNc Univebsitv Science Bulletin

entire series of feniales is referred to crratica Habitats. The information given for crratica on the basis of numbers. As compared to fe- also applies to apaclia. The latter species, how- males of apaclia from California, there is notli- ever, is freijuentK' found on sand dunes through- ing in the collection from this area to indicate out its range, and can be recognized and col- the presence of apaclia females. Some male lected, particniarh' at night, by tlie small mole- sf>ecimens had the abdomen broken off and like burrows they make immediately under the were referred to crratica only on the basis of surface of the ground. The males, too are at- numbers of specimens. tracted to lights at night. A tinv spine is present on the right ventral Seasonal Occurrence. The imlv specimen as- genital plate of some males. This condition is signed to this species was collected in .\Iav. slightb' atvpicai to comparative specimens to the There is no reason to belie\e, howe\'er, that it east, and the Nevada Test Site specimens may occurs aii\ earlier than crratica. be an undescribed group, intermediate bet\veen apacha and crratica, and may indicate that these Localities Represented. Specimens examin- two species are the same with the presence of ed (adult): 1. subspeciation over their ranges. .\ complete re- Study 5E, 1 adult. May 4. vision of the genus will have to be made. .\dditional Remarks. This specimen was compared to a series of males from Riverside Arenivaga apacha (Saussure) County, California, and more nearly resembles (Figure 109; Table 69; Map 35) this species than it does a large series of 1893. (Homoeogamia) apacha Saussure, Rew crratica from a number of localities through-

Suisse Zool., I, Fasc. 2. p. 296. out that species range. For further comments see "Additional Remarks" of that species. Established Synonomy. (Homoeogamia) apacha infuscafa Caudell. Genus Ercmoblatta Rehn Distinctive Features. This species am be distingnislu'd from crratica with any assurance 190.3. F.rcmohlalla Helm, Proc. Acad. Nat. Sci. onlv bv the external genital structures of the Pliila., 1903, p. 181. male, and even these structures are subject to variation througliout its range. Ercmoblatta suhdiaphomi (Scudder)

7(1: Coloration. Both sexes are so similar to I r.il)l,. Map 36) distinction be based on crratica that little can 1902. Homoeogamia suhdiaphana Scudder, Proc. Generally the males are slightly coloration. Da\enport Acad. Sci., L\. p. 19. darker, although light individuals are also although found, and the females are reddish brown, as Distinctive Features. This species, Arenivaga, is quite dis- are the females of crratica. In apacha, liowever, superficiallv resembling is moderately the females have no dorsal maculations. tinct morphologically. The body ccnered with \ellowish hairs, with the middle occurs o\'er i)art Distribution. This species and caudal femora very hairy, and lacking the range of crratica, in that it is found from of the tlistal spine of Arenivaga. Seven spines are (California, through extreme soutiiern southern IouikI at tlu> distal end of the cephalic tibiae. south into Mexico, Nevada, into Arizona, and rhc males arc tulK winged, the females wing- locaiitv being in the state of Chihuaiiua. the t\'pe less. Onh' one specimen from the Nevada Test Site, tan colored species, collected near Frenchman Playa, was assigned Coloration. I'liis is a light brown in some specimens. to the species. ranging to a medium

Table 69. Measurement.s of Arenimga apacha.

X 5 2 5 -Sl j=^ -S^-gji ;5ji §)? lr% c 2 JH mt^ -j£ o5£ •J" CO'-' -^"^ M*^ JO nu

14.5 4.7 5.0 3.5 2.4 0.55 3.0 0.7 3.1 0.8 (^, 5EA7, May 4, 1961 Biological Series, \'ol. 4, No. 3, September, 1964 127

Table 70. Size variation of Eremohlatta suhdiaphana.

3 1 m 3 •55 5, -3 g ^5 ^ S c £- 5> g 1 i ?f

CQ H J CL, « ft. a'a Rkiciiam VoiNc Univeksity Science Bulletin

distribution of each species is given, overall and plete destruction due to nuclear explosions few for tile Nevada Test Site, botii geogra]>hical and orfhopti-rans are found, perhaps due to the ecological, and the habits and habitats are pre- scarcity of vegetation, particularly in tJie areas sented where known. The seasonal occmrence of short grasses and their associated \egctation. given is not necessarily the earliest or latest for To the other extreme, an obvious benefit was the species at the test site, but the dates when noted as a result of an explosion on Rainier the insects were collected in nymph or adult Mesa. In the disturbed area, v\liere the rocks form. Tile distribution of each species is pre- had been loosened and fissures of \'arious sizes sented for all localities, including the total num- occurred in the ground, and the soil had general- ber of specimens examined. .\ map plots tlie 1\' been loosened, there were large numbers of collection sites for each group. most orthopterans, even though the dominant xegetatiou had been killed b\- the force. Few An apparent scarcity of specimens for most numbers were found in the comparative area species was apparent at the test site as com- wiiere no nuclear effects could be noted. pared to other desert and semi-desert areas. The The Orthoptera, as a group, are ideal indi- reasons for this scarcity can only be speculated. cator animals for radiation studies as lia\e been It might be due to the normal fluctuation of and ma\- be conducted at the Xe\ada Test Site. numbers due to natural parasites, predators, or In addition to adecjuate numbers being present, en\ironmental conditions (i.e., wet seasons as both uvmphs and adults arc present at any sea- compared to dr\' seasons). On the other hand, .son of the vear. Most importantlv, perhaps, the it could be due to radiation effects on the tossorial types, such as the species of Ceu- animals. thophiliis. PiLstoceuthophihts. or Stcnopehnatus. With reference to possible radiation effects, max be protected in their underground environ- no statistical data were maintained to determine ments, wliile species of Tiimerofropis, Cibohicris. whether or not there is a high incidence of and Litanciitria may be more exposed in their aberrant forms. There is a great deal of di\ersit\' terrestrial environment. In addition the winged in all orthopteran groups and the aberrant in- species might actually migrate from one place dividuals may not occur with any greater de- to another. Finally, the thamnophilous forms, gree at the test site than elsewhere. Radiation such as Rootettix. are never found on the ground effects on a smaller [wpulation could concei\ablv and ma\' react (juite differently to or be ex{X>sed produce the same results of normal speciation to different intensities of radiation. or subspeciation as over a complete range of dis- Before any cximplete evaluation can be made tribution and in time. with reference to numbers or radiation effects An obvious environmental difference has on the Orthoptera, a follow-up study should be been noted at the test site. In areas of com- made after a lapse of time.

LITERATURE CITED

Albrecht, F. O. 1953. 'Ilir Anatomy of the Migratory Barnum, A. H. 1959. Th- Phallic Complex in the Locust. University of I.dikIom, Tlu- Athlone Press, OEdipodinae (Orthoptera. .\crididae). Unpuh- 118 pp. hshed Ph. D. Thesis deposited at Iowa State Uni- versity, Alexander, R. D. 1957. The 'ra.vonomy of tlie Field 220 pp.

Crickets of tlie Eastern United States (Orthoptera; Cantrall, I. j. 1943. The Ecology of the Orthoptera C'.rvllidae; Aclictu). Annals Enfom. Soe. Americ.i, and Oermaptera of the George Reserve, Michigan. Vol. ,50 (6), pp. .584-602. Misc. Puhl., Museum Zoology, U. Michigan, No. Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963. .54, pp. 1-182. Biotic Communities of the Nevada Test Site. Brig- Caudell, A. N. 1908. Notes on some Western Or- ham Young Univ. Sci. Bull., Biol. Ser., Vol. II, No. tiioptera; with the Description of One New Species. 2, .52 pp. Proc. U. S. N;.t. Mus., Vol. 34, pp. 71-81. Ball, E. D., E. R. Tinkham, R. Flock, and C. T. Vor- Dirsh. \'. M. 19.55. Tanaoceridae and .Xyronotidae: hies. 1942. The C.rasshoppers and Other Or- two new families of Acridoidea (Orthoptera). Viin. thoptera of Arizona. Arizona Agric. Exp. Sta. Tech. Mag. \at. Hist., (12) 8, pp. 285-288. Bull. 93, pp. 2.5.5-373. Oirsli. \'. M, 19.56. The Ph.dlic Complex in Acriil- Banium, A. H. 1952. The Ta.xononiy of Utah Or- oide.i (Orthoptera) in Relation to Taxouonn Tr.uis. thoptera with Notes on Distril)uti(>n. Uiipuhlished Royal Ent. Soc. London. \ol. 108 (7). pp. 2J3-3.56. Master's Thesis deposited at the Brigham Young Essig, E. O. 1926. Iiiscits of Western North Ameiic.i. University, 236 pp. Ihc \l,ic\lill,ni Company, New York. 1035 pp. .

Biological Series, \'ol. 4, \o. 3, September, 1964

1944. the significance of localized Hebard, M. 1916. A Study of the Species of the Rehn, J. A. G. On Genus Stcnopclmcitus found in the United States. coloration in the Creosote Bush Locust (Bootettix) Journ. \. V. Entom. Soc., Vol. 24, pp. 70-86. (Orthoptera; Acrididae; Acridinae). Entom. News, Hebard, M. 1920a. New Genera and Species of \'ol. .55, pp. 1.58-164. A. Grant, 1957. The Genus Melanopli found within the United States (Part Rehn, J. G. and H. J. Jr. III). Trans. Amer. Entom. Soc, Vol. 46, pp. .3.55- Pcirtitettix as found in North America (Orthoptera; 403. Acridoidea; Tetrigidae). Proc Acad. Nat. Sci. Hebard, M. 192(lb. A Revision of the North Ameri- Phila., Vol. 109, pp. 247-319. A. Grant, 1958. Revision can Species of the Genus Mi/rmecophila (Or- Rehn, J. G. and H. J. Jr. A

) Morsca ( Orthoptera; Acridoidca; thoptera; GryUidae; MyrmecophiHnae . Trans. of the Genus

Amer. Entom. Soc, Vol. 46, pp. 91-111. Eumastacidae ) . Trans. Amer. Entom. Soc, Vol. Hebard, M. 1920c. Revisionarv Studies in the Genus 84, pp. 217-259.

Areinvaga (Orthoptera, Blattidae, Polyphaginae ) Grant, 1959a. Review Rehn, J. A. G. and H. J. Jr. A Trans. Amer. Entom. Soc, \'ol. 46, pp. 197-217. of the Genera Fsijchomastax and Eumorsea (Or- Hebard, M. 1929. The Orthoptera of Colorado. Proc thoptera; Acridoidea; Eumastacidae). Trans. Amer. Acad. Nat. Sci. Phila., Vol. 81, pp. 30.3-425. Entom. Soc, Vol. 84, pp. 273-302. of the United Hebard. M. 1931. Th- Magoplistinae Relin, A. Grant, Review J. G. and H. J. Jr. 1959b. A Vol. State^. Trans. Amer. Entom. Soc, 57, pp. of the Romalcinae (Oithoptera; Acrididae) Found 135-160. in America North of Me.\ico. Proc. Acad. Nat. Sci. Hebard, M. 1934. Studies in Orthoptera which occur Phila., Vol. 109, pp. 109-271. in North America North of the Mexican Boundary. Rehn, A. G. and H.J. Grant, Jr., 1960. A New Con- V. The Pachymorphinae of the United States J. cept Involving the Subfamily Acridinae (Ortho- ( Phismidae). Trails. Amer. Entom. Soc, Vol. 60, ptera: Acridoidea). Trans. Amer. Entom. Soc, Vol. pp. 284-294. 86, pp. 173-185. Hebard, M. 1937. Studies in Orthoptera which occur Grant, 1961. Mono- Rehn, J. A. G. and H. J. Jr. A in North America North of the Me.\ican Boundary. graph of the Orthoptera of North America (North IX. On Arphia conspersa, notes and a new species of Mexico), X'olume I. Monographs Acad. Nat. Sci. of Sphamgemon, a new Genus and its Races of the Phila., No. 12, 2.57 pp. group Heliasti and a new generic name for the Rehn, A. G. and M. Hebard. 1908. An Ortho- group Anconiae (OEdipodinae, Acrididae). Trans. J. pterological Reconnoissance of the Southwestern Amer. Entom, Soc, Vol. 63, pp. 361-379. United States. Part I: Arizona. Proc. Acad. Nat. Sci. Heifer, R. 1963. How to Know the Grasshoppers, J. Phil.L, Vol. 60, pp. 365-402. Cockroaches and Their Allies. W'm. C. Brown Com- Rehn, A. G. and M. Hebard. 1909. An Ortho- pany Publishers, 353 pp. J. pterological Reconnoissance of the Soutliwcstern Hubbell, T. H. 1922. The Dermaptera and Orthoptera United States. Part III: California and Nevada. of Berrien County, Michigan. Mus. Zool., Univ. Proc. Acad. Nat. Sci. Phila., Vol. 61, pp. 490-483. Mich. Occas. Papers. No. 116, pp. 1-77. Rehn, A. G. and M. Hebard. 1914. A Revision of Hubbell, T. H. 1936. A Monographic Revision of tlie J. the Orlhoptcrous Genus Insiira. Trans. Amer. En- Genus CcuthophUus ( Orthoptera, Gryllacrididae, tom. Soc, Vol. 40, pp. 37-184. Rhaphidophorinae). Univ. Fla. Publ. Biol. Sci. 2 Rehn, A. G. and M. Hebard. 1915. The Genus (1), 551 pp. J. Gnjllus (Orthoptera) as Found in America. Proc. Isely, F. B. 1937. Seasonal Succession, Soil Relations, Nat. Sci. Phila., Vol. 67, 293-322. Numbers, and Regional Distribution of North- Acad. pp. Color Nomen- eastern Te.xas Acridians. Ecol. Monog., Vol. 7, pp. Ridgway, R. 1912. Color Standards and 317-344. clature. Washington, D. C, 43 pp. 53 pis. Isely, F. B. 1938. The Relations of Texas Acrididae Strohecker, H. F. 1937. An Ecological Study of Some to Plants and Soils. Ecol. Monog., Vol. 8, pp. 551- Orthoptera of the Chicago Area. Ecology, Vol. 18 604. (2), pp. 231-250. LaRivers, I. 1948. A Synopsis of Nevada Orthoptera. Thomas, C. 1875. Report upon the Collections of Or- Amer. Midland Nat., Vol. 39, pp. 652-720. thoptera made in portions of Nevada, Utah, Cahf- Revision of the Orthopteran Genus ornia, Colorado, New Mexico, and Arizona, during McNeill, J. 1901. Chapter Trimerotropis. Proc. U. S. Nat. Mus., Vol. 23, pp. the Years 1871, 1872, 1873, and 1874. 393-449. XIII (in) Report upon Geog. & Geol. E,xpl. Surv. A. 1919. Descriptions of New and West One Hundredth Mer., G. M. Wheeler. Vol. V., Rehn, J. G. Critical Notes upon Previously Known Forms of Zoology, pp. 843-908. North American OEdipodinae (Orthoptera; Acrid- Tinkhim, E. R. 1938. Western Orthoptera Attracted idae). First Paper. Trans. Amer. Entom. Soc, Vol. to Lights. Journ. N. Y. Entom. Soc, Vol. 46, pp. 45, pp. 229-255. 339-353. A. 1923. North American Acrididae. A Tinkham, E. R. 1942. The Rediscovery of Anoplodusa Rehn, J. G. Study of the Ligurotettigi. Trans. Amer. Entom.. urizonensis (Orthoptera). Bull. Chicago Acad Sci., 221-227. Soc, Vol. 49, pp. 43-92. Vol. 6 (12), pp. Rehn, A. G. 1940. The South American Species Tinkham, E. R. 1944. Biological, Taxonomic and J. the of the OEdipodinc Genus Trimerotropis (Ortho- Faunistic Studies on the Shieldback Katydids of Vol. ptera: Acrididae). Trans Amer. Entom. Soc, \'ol. North American Deserts. Amer. Midland Nat., 257-328. 65, pp. 395-414. 31, pp. Ecological Stu- Rehn, A. G. 1942. On the Locust Genus Psoloessa Tinkh;im, E. R. 1948. Faunistic and J. of (Orthoptera; Acrididae; Acridinae). Trans. Amc. dies on the Orthoptera of the Big Bend Region Texas, with Especial Reference to the Entom. Soc, Vol. 68, pp. 167-237. Trans-Pecos 130 BiiiciiAM ^<)l'^•c Univkksity Science Bulletin

Orthopteran Zones and Kaunac of Midwestern pers in UrI.ition to Plant Association.s. Biol. Bull., North America. Amer. Midland Nat., Vol. 40 (.3), Vol. 25, pp. 141-180. pp. .521-663. Wallace, H. S. 19.5.5. Revision of the Cenu-s Aeolo- Unpihard, K. A. 1941. An Kcological Study of the plitk'n (Oilhoptera, Acrididae). Ann. Entom. Soc'. Sahatoria of Point I'elee, Ontario. Univ. Toronto Anur., \ol. 48 (6), pp. 4.5.3-480. Studies, Biol. Scr., No. 50. pp. 1-91. Wheler, W. M. 1900. The Habits of Mtjrmccophila Vestal, A. G. 1913. Local Distrihnlion of (;r,issliop- nrhru.icin.si.\ Bniner. Psyche, Vol. 9, pp. 111-115.

APPENDIX I

l)i:i'osiToiui;.s ok Spkc.imkns Collecikd in This Study

American Museum oi Natural ili.ston'. New Nevada Soutiicni University, Las Vegas, Ne- York City, New ^ork. \'ada. Brigliain Young University, Provo, Utali. Philaclclpiiia .\caclemv of Natural Sciences, California Acadeinv of Sciences, San Francisco, Philadelphia, Pennsylvania. California. University of Michigan, Ann .\rhor, Michigan. Chicago Natural llist()r\' .Museum, Chicago, Uni\'ersitv of Ne\ada, Reno, Ne\ada. Illinois. University of Utah, Salt Lake Cit)' Utah. Di.xie College, St. George, Utah. United States National Museum, Washington, Musuem of Comparative Zoology (Hanard), D. C. Cambridge, Massachusetts. Utah State Unixersit)-, l^ogan, Utah.

APPKXDI.X II

Notes on Collecting .\nu Pheserving Orthoptfr.v

Most Orthoptera are large and conspicuous providing sufficient fluid is present. A safe and may be collected with a minimum of equip- licjuid \oluine for adequate preservation would ment, but iit times with a great deal of eflorf. be ten times the volume of the insects. Some fossorial and terrestrial forms can best he No specimen killed in the cyanide bottle found by the use of special can pit-traps or should be retained in the bottle for more than by searching through rodent nests and tunnels, lour to six hours, as the cvanide gas discolors in caves, or under rocks. At least one species the specimens, turning them (juite reddish. .After is found associated with ants and re(|uires death, but still relaxed, they should be piimed special collecting. The thamnophilous ortho- bv forcing a No. 3 insect pin through the [pos- pterans are best collected b\' the use of a hea\y terior part of the pronotum immediatelv to the sweeping net, \\hile the rapid fliers may be cap- right of the mcxlian carina. .\s most orthopterans tured 1)\ dropping an aerial net ()\er them alter are heavy-bodied, the legs and abdomen tend to tlie\ lia\e alighted on the ground. Nocturnal sag. To correct this unsightlv condition they specimens may be collected at night, by looking should be allowed to drv for several davs, ac- for them with a light, oi' by locating them by cording to environmental conditions (i.e., humid- their calls. itv), by inserting the pin in a sheet of st\rofoam All winged specimens should be killed in a plastic covered with paper (to prevent their standard cvanide killing bottle, or b\- subjection tarsi from breaking off when they are removed). to another gas, as fluids have a tendency to The legs and antennae should be arranged in change body colors and make the wings un- the desired position before drving. suitable for ade(|uate study. Fossorial specimens, The left wing of acridids, especially the so- such as the species of CeuthopJiilus. Prlstoceu- called band-winged grasshoppers, those with thopliiliis, Miiiniccopliild. and SlcnofM'hiuiliis. as bright Iv colonel wings, should be spread on a well as the females of roac-hes (not the winged s|)reading board to show the color and pattern males), ma\' be killed in TO'J ethyl alcohol, in of till- wings, as these are important taxonomic which solution they can be permanently stored characters. Biological Series, Vol. 4, No. 3, Septemuer, 1964

Large specimens have a tendency to discolor All specimens must be completely labelled due to fluids, body especially fats. These speci- with exact locality, date of capture, and en\iron- mens siiould be eviscerated and the internal or- mental conditions. Specimens without complete gans replaced with cotton; otherwise the speci- data are of little scientific value. mens will turn dark and may decay. An incision Dried specimens must be kept free from dust should be made at the base of the ventral ab- and so-called museum pests. Fumigation may domen, the length of three segments and the be accomplished by keeping a supply of para- internal organs removed with forceps. A small dichlorobenzene and/or naphthaline roll of cotton, the size of the internal abdomen, flakes in the should be inserted to replace these organs. The box or case at all times. Specimens properly pre-

natural size and color of the specimen is thus served and fumigated may be retained in- retained. definitely.

APPENDIX III Glossary

(Modified fiom Torre-Bueno, J. R. de la, A Glossary of Entomology, The Science Press Printing Co., Lancaster, Pennsyhania, 1937)

acicular, needle-shaped; with a long slender carina (pi. carinae), an elevated ridge or keel, not point. necessarily high or acute. acute, pointed; terminating in or forming less carinate, keeled; having keels or carinae; with a, than a right angle. or several, longitudinal narrow raised lines. adventive, accidental; applied to exotics or in- carnivorous, feeding upon flesh food; an insect troduced species. preying on other insects or feeding on aedeagus, in male insects, the intromittent organ, their flesh. a part of the phallic complex and situated caudal, of or pertaining to the anal end of the beneath the pallium of the subgenital plate. insect body. alate, winged; as opposed to apterous. cephalic, belonging to or attached to the head; ambulatorial, fitted for walking. directed toward the head. annulus, a ring encircling a joint or segment. cercus (pi. cerci), an appendage (generally pair- apical, at, near, or pertaining to the apex of any ed) of the tenth abdominal segment, usually structure. slender, filamentous and segmented. apterous, without wings, wingless; see alate. cinereous, ash-colored; gray tinged with blackish. arcuate, arched; bow-like. clavate, clubbed; thickening gradually toward arolium, the terminal cushion-like pad between the tip. the claws of the tarsi. clypeus, that part of the head of the insect be- articulate, to connect bv a joint; jointed or seg- low the frons (front), to which labrum is at- mented. tached anteriorly. auditory, relating to the sense of hearing. coriaceous, leather-like; thick, tough, and some- auricula (pi. auriculae), an appendage resem- what rigid. bling a little ear. corneous, of a horny or chitinous substance; resembling horn in texture. brachypterous, with short or abbreviated wings. costa, any elevated ridge that is rounded at its bullate, blistered; a slightly swollen structure. crest; the thickened anterior margin of any wing, but usually of the forewings. calcar (pi. calcaria), a moveable .spur or spine- coxa (pi, coxae), the basal segment of the leg, by like process; specificallv the spines at the means of which it is articulated to the body. apex of the tibia. crenulate (crenulation), with small scallops, even- callosity, a thick swollen lump, harder than its ly rounded and rather deeply curved. surroundings; callus; also a rather flattened cristate, with a prominent carina or crest on the elevation not necessarily harder tlian the upper surface; crested. surrounding tissue. cuneifonn, wedge-shaped; elongate triangular. campestrian, inhabiting open areas (fields). cursorial, adapted for running. Bhicham Vounc University Science Bulletin

clecli\ant, slnping gracliiallv downward. geophilous, living on the ground; of S[x;cies, li\'- dc|)lanate, comprt-sst'd; flattened above and Ije- iiig on the surface or coming freely into con-

low. tact with it. dimorphism, a difference in form, color, etc., be- glabrous, smooth, hairless and with(jut punctures tween individuals of the same species, char- or structures. acterizing two distinct types; may be sea- glaucous, sea-green; pale bluish-green. sonal, se.\ual, or geographic. herbivorous, feeding upon plant tissue; discoidal, relating to the disk, or middle; shaped leaf fi'cder. like a round plate. hyaline, transparent or partK' so; waterlike in distal, near or toward the free end of any ap- color; pendage; that part of a segment farthest glassy. from the body. immaculate, destitute of spots or marks. diurnal, active or habitually fl>ing by day only. instar, the period or stage between molts in the dorsal, of or pertaining to the upper surface. larva, numbered to designate the various

ecdysis, the process of casting the skin; moulting. periods; e.g., the first instar is the stage be- ensiform, sword-shaped; two-edged, large at tween the egg and the first moult. interocular, the base and tapering to the point, between eyes. epiphallus, a sclerite in the floor of the genital labium, the second maxilla; the lower lip; a com- chamber proximal to the base of the pound structure which forms the floor of phallus; pseudosternite. the mouth in mandibulate insects, behind epiproct, the dorsal part of the eleventh segment the first maxilla and opposed to the labrum. of the abdomen; the supra-anal plate, labrum, the upper lip, which covers the base of explanate, spread out and flattened; applied to the mandible and forms the roof of the margin. a mouth. falcate, sickle-shaped; convexly curved. lamellate, sheet- or leaf-like; composed of or fascia (fasciation), a transverse band or broad covered with laminae or thin sheets.

line, especially when it crosses both tegmina laminate, formed of thin, flat hiyers or leaves. or femora. lateral, relating, pertaininng, or attached, to fastigium, the extreme point or front of vertex. the side. femur (pi. femora), the thigh; usually the stout- linguiform, tongue-shaped; linear, with the ex- est segment of the leg, articulated to the tremities obtuseh' rounded. bod\- through trochanter and coxa and bear- lobulate, divided into, or with many small holes ing the tibia at this distal end. or lobules. filiform, thread-like; slender and of equal maculate, spotted or marked with figures of any diameter. shape, of a color differing from the ground flavous, pure, clear yellow. color. fossa, a pit or deep sulcus. mandibles, the first pair of jaws, stout and tootli- fossorial, formed for or with the habit of digging hke. or burrowing. mandibulate, having biting jaws. foveola, (pi. foveolac), a deep depression with maxilla, (pi. maxillae), the second pair of jaws well-marked sides; a pit. in a mandibulate insect. frons, the unpaired sclerite of the head lying be- medial, referring to, or at the middle. tween the arms of the epicranial suture mesad, toward or in the direction of the median and bearing the median ocellus. plate of the insect body. furcula, a pair of backwardly directed append- mesially, at or to the middle. ages which o\erlie in a more or less forked mesonotum, the primiti\ely upper surface of the position the base of the epiproct. second or middle thoracic ring. fuscous, dark brown, approaching black; a plain mesosternum, the underside or breast of the mixture of black and red. mesothorax. ring gena (pi. gcnac), the cheek; the part of the hi-ad incsothorax, the second or middle thoracic on each side below the eyes, extending to which bears the middle legs and the an- the gular suture. terior wings, genicular, pertaining to the curved dark mark- metamorphosis, the series of changes through ings on the posterior knee-joint. which an insect passes in its growth from adult. genitalia, all the genital structures collectively. the egg tlirough the Biological Series, Vol. 4, No. 3, September, 1964 133

metanotum, the primitively upper surface of the proximal, that part of an appendage nearest the third or posterior thoracic ring. body. metasternum, the underside or breast of the prozona, the anterior part of the pronotum. metathorax. punctate, set with impressed points or punctures. metathorax, the third thoracic ring or segment, raptorial, adapted for seizing prey; predacious. which bears the hind legs and second pair reniform, kidnev-shaped. of wings. rostrum, in general, a snout-like prolongation of metazona, the dorsal surface of the prothtjrax the head. behind the principal sulcus. rugose, wrinkled. nacreous, pearly; resembling mother of pearl. saltatorial, adapted for leaping; having the nocturnal, of the night; applied to insects that power of leaping. fly or are active at night. saxicolous, frequenting rocky or stony areas. notum, the dorsal or upper part of a segment; scalariform, ladder-like; applied to venation tergum. when the veinlets between two longitudinal nymph, a young insect which quits ihe egg in \'eins are regularly arranged like the rungs a relatively advanced stage or morphological of a ladder. development, differing from the adult in sclerotized, of the insect integument, hardened having the wings and the genitalia present in definite areas by deposition or formation only in an incompletely developed condi- of other substances tlian chitin in the tion. cuticula. sellate, saddle-shaped. obtuse, not pointed; at an angle greater than a serrations, a tooth, as of a saw; a series of iuch right angle; opposed to acute. teeth. occiput, the hinder part of the epicranium be- serrulate, finely serrated; with minute teeth tween the vertex and the neck. or notches. ocellus, (pi. ocelli), tlie simple eye in adult in- seta, a slender hair-like appendage. sects, consisting of a single bead-like lens, setaceous, bristle-shaped; slender, gradually occurring singly or in srriall groups. tapering to a tip. omnivorous, feeding generally on animal or setose, furnished or covered with setae or stiff vegetable food, or on both. hairs. ootheca, the covering or case over an egg mass. spatulate, rounded and broad at the top; slender ovate, egg-shaped in outline. or drawn out at base. ovipositor, the tubular or valved structure by spine, a multicellular more or less thorn-like means of which the eggs are placed; usually process or outgrowth of the cuticula not somewhat concealed, but sometimes extend- separated from it by a joint; a large seta ed far beyond the end of the body. provided with a calyx or cup by wliich it is palpus, (pi. Palpi), a mouth feeler; a palp. articulated to the cuticula. paraproct, one of the two lobes formed by tlie spinifomi, in the form or shape of a spine. ventrolateral parts of tlie epiproct. spinule, a small spine. penultimate, next to the last. spiracle, a breathing pore; in the plural the later- phallic complex, the genital structures of the al openings on the segments of the insect male, especially the concealed structures. body through which air enters the tracheae. phallus, the intromittent genital organ of the spur, a spine-like appendage of the cuticula, male. connected to the bodyAvall by a joint. plantula, a lobe of the divided tarsal puhillus; stemite, the ventral piece in a ring or segment; one of the soles or climbing cushions of the a subdivision of a sternal plate, or any one foot. of the sclerotic components of a definitive pleuron (pi. pleura), the lateral region of any sternum. segment of the insect body, commonly of sternum (pi. sterna), the entire vential division the thoracic segments. of any segment; the underside of the insect pronotum, the upper or dorsal surface of the thorax, between the coxal cavities. prothorax. stria (pi. striae), any fine longitudinal impressed prostemum, the fore-breast; the sclerite between line. the fore-legs. stridulate, to make a creaking, grating or hissing or prothorax, the first thoracic ring or segment; it sound or noise, by rubbing two ridged bears the anterior legs but no wings. roughened surfaces against each other. BHiGHAAf Young University Science Bulletin style, stylus (pi. styli), small, usually pointed, ex- pecially when such part consists of a single articulate appendages, most freijiiently sclerite. found on the tenninal segments of (lie testaceous, bearing a test or hard covering; ;d)doluen. brow nisli-\ellow. subgciiital plate, the piati" or process underijing thaninuphiluus, living in thickets or dense shrub- the genital organs; the tenninal or distal bery. abdominal scleritc. thorax, the second or intermediate region of the subocular, beneath or below the e\i's. insect body bearing the tnie legs and wings, suborbicular, slightlv less than round and ll.it. made up of three rings, named in order, pro-, meso-, and metathorax. subterranean, underground, beneath the surtacc tibia (pi. tibiae), the fourth division of tlie leg, of the soil or ground. articulated at the proximal end to the femur sulcate, deepK' furrowed or grooved; .vith deep and bearing on the distal end the tarsi. grooves. trapezoidal, in the form of a four-sided figure sulcus, (pi. sulci), a furrow or groove; a groove- of which two sides are parallel and two like excavation. arc not. supra-anal, abo\e the anus; suranal; the epiproct. trigonal, triangular; an area boimded b\ a tri- suture, a scam or impressed line indicating the angle. division of the distinct parts of ihe body trochanter, a sclerite of the insect leg, sometimes wall. di\ided, between the coxa and femur. tarsus (pi. tarsi), the foot; the jointed appendage truncate, cut off squarely at tip. attached at the apc.x of the tibia, bearing the tubercle, a little solid pimple or small button. claws and pnlvilli; the distal part of the in- tympanum, any membrane stretched like the sect leg, consisting of from one to five seg- head of a drum, specificallv applied to the ments or joints. membrane covering the auditory organs. tectate, covered; concealed; tectiform. ultimate, last. tectiform, roof-like; sloping from a median ridge. undulate, wavy; obtusely wa\'cd in segments of tegmen (pi. tegmina), a covering; the hardened circles. leathery or homy forewing. teneral, the condition of the adult insect after ventral, pertaining to the under surface of the

the last moult when it is not entirely hard- abdomen. ened or fullv of the mature color. vertex, the top of the head between the eyes, tergite, a dorsal sclerite or part of a segment, es- frons and tx.'ciput.

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