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Biosystematics of the grylloblattodea Kamp, Joseph William 1973

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cl BIOSYSTEMATICS OF THE GRYLL03LA TTODEA by JOSEPH WILLIAM KAMP B.A. (1951), M.A. (1961) C a l i f o r n i a State University at Chico A Thesis Submitted i n P a r t i a l Fulfilment of the Requirements for the Degree of Doctor of Philosophy In the Department of Zoology We accept t h i s thesis as conforming to the required standard /7/f -2-THE UNIVERSITY OF BRITISH COLUMBIA AUGUST 1973. In presenting t h i s thesis i n p a r t i a l f ulfilment of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t freely available for reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. I t i s understood that copying or publication of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of The University of B r i t i s h Columbia Vancouver 8, Canada i i ABSTRACT The North American G r y l l o b l a t t i d a e are either hygro-p h i l i c occupants of the alpine-subalpine or f a c u l t a t i v e cavernicoles of lava ice caves. The'research indicates that the habitats preferred by G r y l l o b l a t t a must be cold and moist but not wet. The preferred alpine-subalpine hypolithion was found to be beneath stones 50 to 150 cm. buried 20 to 50 cm. deep i n the substrate. G r y l l o b l a t t a also temporarily occupied g l a c i e r s , snow f i e l d s , r o t t i n g logs and borders of g l a c i a l springs. The optimum micro-climate i n the alpine-subalpine habitats was within - 3 to -f 6° C. with a humidity above 70 per cent. Humidity evidently governs the occupancy of the hypolithion more than temperature. The insect w i l l not inhabit hypolithion with r e l a t i v e humidities le s s than 70 per cent, regardless of temperature. The cavernicolous habitat for G r y l l o b l a t t a i s a micro-environment l i m i t e d to a few ice caves i n lava f i e l d s . S u f f i c i e n t ice must be present to maintain a spring-to-f a l l temperature of - 3 to + 8° C. and over 80 per cent r e l a t i v e humidity. During the winter the Ice cave i s recharged with cold a i r below the tolerance of G r y l l o b l a t t a at which time the Insect inhabits the hypolithion. •The temperature preference of Gry l l o b l a t t a , estab-l i s h e d i n the laboratory, was between - 3.5 and + 5° C. at 90 to 99 per cent r e l a t i v e humidity, - 2.2 to + 4.5° C. at r e l a t i v e humidities between 70 and 90 per cent, and - 1.1 to + 1.6° C. at 50 to 70 per cent r e l a t i v e humidities. Temperature tolerance at humidities above 95 per cent ranges between - A and + 1 1° C. Four-hour exposure to 4 1 6° C. and one-hour exposure at - 5.5° C. produces 50 per cent mortality. Lethal extremes were - 8° C. and 4- 23° C. A l l stages of the insect were found to be active year around with no dormant period. The mean freezing point depression of the hemolymph was measured at - 0.98° C. therefore, the insect remains active i n a supercooled state. Twenty-six new populations were found during t h i s research, extending the d i s t r i b u t i o n from the Yukon-3ritish Columbia border to the southern Sierra Nevada of C a l i f o r n i a . Five new species and three new subspecies are here described The d i s t r i b u t i o n a l data indicate the presence of four divergent groups characterized by is o l a t e d endemic popula-tions or species. The present disjunct d i s t r i b u t i o n and zoogeography have been fundamentally influenced by the geologic and climatic events of the la t e Pleistocene. Further, regional and sometimes highly l o c a l i z e d volcanic a c t i v i t y during the post Pleistocene, the warm dry Hypsi-thermal period, and the r e - b i r t h of summit and cirque g l a c i e r s , commencing approximately 2500 years ago, have affected the d i s t r i b u t i o n a l patterns of G r y l l o b l a t t a . The recent zoogeography of the various species and populations i n western North America i s discussed. i v A comparative l i p i d analysis of G r y l l o b l a t t i d a e and six other insects from related orders, and with varying temperature preferences, shows differences i n fatty a c i d composition. The composition in Gr y l l o b l a t t a i s more l i k e that i n Dermaptera, but the a f f i n i t y i s as remote as i s demonstrated i n the numerical a n a l y s i s . Analysis of Gr y l l o b l a t t a shows 65.8 per cent of the t o t a l fatty acids are unsaturated, 91 per cent of which have melting points below i t s maximum tolerated temperature. These data c l e a r l y indicate the low temperature adaptation of G r y l l o b l a t t a ; such composition i s not seen i n the warmer tolerance forms. In a numerical analysis of 164 external and int e r n a l morphological characters i n G r y l l o b l a t t a and seven other related orthopteroid i n s e c t s , Dermaptera has the closest a f f i n i t y to the G r y l l o b l a t t a . The phenetic a f f i n i t i e s and r e l a t i o n s h i p s of the G r y l l o b l a t t a , as shown i n the a n a l y s i s , place the taxon at the ordinal l e v e l . The most acceptable systematic treatment of th i s group i s as the order Grylloblattodea. V TABLE OF CONTENTS ' ABSTRACT . . . . . . . . f . ... i i TABLE OF CONTENTS v LIST OF TABLES x LIST OF FIGURES xi ACKNOWLEDGMENTS xiv I. GENERAL INTRODUCTION ,. 1 I I . .. HABITAT AND BIOCLIMATIC REQUIREMENTS OF GRYLLOBLATTA SPECIES . . . . ; 6 Introduction 6 I. The Hypolithlon Habitat ' 6 I I . The Cavernlcolous Habitat 11 .The Microenvironment and Microclimate of Gr y l l o b l a t t a Habitats x3 I. Materials and Methods 13 I I . Results 1 5 A* The Alpine-Subalpine Habitat 15 1. Microhabitat 15 2. Microclimate ." 17 B. The Cave Habitat 20 1. Microhabitats of the Ice Cave ... 20 a . Entrance Zone 22 b. Twilight Zone 22 c. Upper Dark Zone 23 d. Lower'Dark Zone •... 24 • 2. M i c r o c l i m a t e o f t h e I c e Cave H a b i t a t s 24 a. Temperature . 26 b. I c e 28 c. H u m i d i t y 29 I I I . D i s c u s s i o n and Summary 31 A c t i v i t y o f G r y l l o b l a t t a i n A l p i n e and Cave H a b i t a t s 35 I . I n t r o d u c t i o n 35 I I . Methods 35 I I I . R e s u l t s 35 A. A c t i v i t y i n A l p i n e H a b i t a t s 35 1. W i n t e r Season 36 2. S p r i n g Season 37 3. Summer Season 39 4. F a l l Season 39 B. A c t i v i t y i n Cave H a b i t a t s 41 IV. D i s c u s s i o n and Summary 43 Temperature and H u m i d i t y P r e f e r e n c e o f G r y l l o b l a t t a 45 I . I n t r o d u c t i o n 45 J l . M a t e r i a l s and Methods 46 A. E x p e r i m e n t a l Chamber 47 B. T e s t Methods 51 I I I . R e s u l t s 53 I V . D i s c u s s i o n and Summary 55 v i i Temperature Tolerance and Lethal Limits of G r y l l o b l a t t a .... 61 I. Introduction 61 I I . Materials and Methods 61 I I I . Results 63 IV. Discussion and Summary 64 I I I . SYSTEMATICA, DISTRIBUTION AND ZOOGEOGRAPHY ... 71 Introduction . 71 , Materials and Methods 74 Results 74 The Nearctic Species 74 'The Nearctic D i s t r i b u t i o n 76 Zoogeography 81 I. The Rocky Mountain Co r d i l l e r a n Group ... 81 I I . Coast-Cascade Group 93 1. Northern Cascade Group 100 2. High Cascade Group 112 3. Southern Cascade Group 120 4. Modoc Plateau-Basin Range Group .... 124 5 a . Coast Range Group 125 5 b . Klamath Mountain Group 126 I I I . The Sierra Nevada Group 128 Discussion and Summary 133 IV. SYSTEMATIC POSITION OF GRYLLOBLATTODEA BY • NUMERICAL ANALYSIS . . 141 Introduction 141 Materials and Methods lZ<5 v i i i Results . l l*Q Discussion V. LIPID ANALYSIS . . I63 Introduction . 163 Materials and Methods 166 1. Extraction and P u r i f i c a t i o n 167 2. Separation of L i p i d Classes 168 3. Analysis of Fatty Acids by Gas Chromatography I69 Results and Comparisons 170 1. Order Thysanura (Pedetontus) 170 2.* Order B l a t t a r i a (Periplaneta) 174 3. Order Orthoptera (Gryllus) .... 176 4. Order Isoptera (Zootermopsls) 178 5. Order Dermaptera (Anlsolabls) 180 6. Order Grylloblattodea ( G r y l l o b l a t t a ) ..... I83 Discussion of Environmental and Systematic Relationships Demonstrated by L i p i d Analysis . 187 VI. DISCUSSION AND CONCLUSIONS 192 LITERATURE CITED 203 APPENDIX I 222 * G r y l l o b l a t t a s c u l l e n l s c u l l e n l Gurney 222 Gr y l l o b l a t t a skagltensis n. sp 225 Gr y l l o b l a t t a scudderl n. sp 229 i x G r y l l o b l a t t a paullnai n. sp 233 G r y l l o b l a t t a campodelformls athapa ska n.ssp, 239 G r y l l o b l a t t a campodelformis nahannl n. ssp 243 G r y l l o b l a t t a s c u l l e n l cryocola n. ssp 248 G r y l l o b l a t t a hoodalles n. sp 252 G r y l l o b l a t t a lava cola n. sp 256 G r y l l o b l a t t a o c c l d e n t a l l s S l l v e s t e r i , s t a t . nov.... 260 APPENDIX II . . . 262 X LIST OF TABLES IN TEXT Pages I. Temperature Tolerance Limits of Grylloblatta....66-67 II." L i s t of Species and Subspecies of G r y l l o b l a t t a . .75 I I I . Fatty Acid Composition of Glycerlde Classes Analyzed 171 IV. Total Per Cent Fatty Acids with M. P. above •f 10° C 189 V. Total Per Cent Unsaturated Fatty Acids 190 VI. Total Per Cent Unsaturated Fatty Acids with M. P. above * 10° C 190 VII. 84 External Characters of the Orthopteroids....262-268 VIII. 80 External and Internal Characters of the Orthopteroids • .269-275 x i LIST OF FIGURES IN TEXT Pages 1. Examples of various Gr y l l o b l a t t a habitats 16 2. Monthly atmospheric temperature of alpine-subalplne habitats of G r y l l o b l a t t a i n Oregon (1969-1971)..... 18 3. South Ice Cave, Oregon 21 4. South Ice Cave, Oregon. Monthly temperature p r o f i l e of rock and cave a i r 25 5.. Transparent view of temperature-humidity chamber... 49 6. Temperature response of adult G r y l l o b l a t t a  campodeiformis and G. lava cola to various controlled humidity ranges 54 7. Temperature tolerance and l e t h a l l i m i t s of adult G r y l l o b l a t t a campodeifor'mis and G. lava cola when exposed to 1° C. temperature change each 2-| minutes .65 8. 5000-foot elevation contour of the Cordillera of western North America 77 9. Species d i s t r i b u t i o n of Gryllobla tta l o c a l i t i e s 79-80 10. Maximum extent of late Pleistocene g l a e l a t i o n i n western North America 82 11. Quaternary to Recent volcanics of Coast-Cascade Range 96 12. Dendrogram showing d i s s i m i l a r i t y analysis of 84 external characters 149 13. Dendrogram showing, d i s s i m i l a r i t y analysis of 80 external and i n t e r n a l characters 152 x i i 14. Dendrogram showing d i s s i m i l a r i t y analysis of 164 characters 153 15. Dendrogram showing s i m i l a r i t y analysis of 84 external characters used In Figure 12 155 16. Dendrogram showing s i m i l a r i t y analysis of 80 external and Internal characters used i n Figure 12. 156 17. Dendrogram showing, s i m i l a r i t y analysis of 164 characters used i n Figure 14 158 18. Representative chromatograms of Pedetontus 172 19. Representative haseline-corrected chromatograms of Per Iplaneta amer i ca na 175 20. Representative baseline-corrected chromatograms of Gryllus a s s l m l l l s 177 21.. Representative baseline-corrected chromatograms of Zootermopsls angusticolli's 179 22. Representative baseline-corrected chromatograms of Anisola'ois maritima 181 23. Representative baseline-corrected chromatograms of Gr y l l o b l a t t a lava cola . . . : 184. 24. G r y l l o b l a t t a s c u l l e n l scullenl s t a t . nov 223 25. G r y l l o b l a t t a skapcitensis n. sp 226 26. G r y l l o b l a t t a scudderi n. sp 231 27. . G r y l l o b l a t t a p aulinal n. sp 235 28. G r y l l o b l a t t a camoodeiform!s athapa ska n. ssp 240 29. Gr y l l o b l a t t a campodelformls nahannl n. ssp 245 x i i i 30. G r y l l o b l a t t a s c u l l e n l cryocola n. ssp 249 31. G r y l l o b l a t t a hoodalies n. sp ...253 32. G r y l l o b l a t t a lava cola n. sp 258 x i v ACKNOWLEDGMENTS This study was carried out under the auspices of Dr. G. G. E. Scudder whose assistance and helpful discussions are g r a t e f u l l y acknowledged with sincere thanks. The research was financed by the National Research Council of Canada through grants to Dr. Scudder, and by the A r c t i c and Alpine Committee of the University of B r i t i s h Columbia. Drs. H. D. Fisher, W. H. Mathews, J . D. McPhail and J . E. P h i l l i p s c r i t i c a l l y read the manuscript and offered many helpful suggestions and valuable advice. Thanks are due to the following for providing G r y l l o -b l a t t a material used i n the systematic section: Dr. ¥ . F. Barr, University of Idaho, Dr. A. B. Gurney, U. S. National Museum, R. E. Leech, National Museum of Canada, and Miss K. Stuart, Spencer Entomological Museum, University of B r i t i s h Columbia. Living insects for the l i p i d comparison were kindly collected by Drs. T. H. Carefoot and G. G. E. Scudder, University of B r i t i s h Columbia, and by V. R. Vickery, Mcdonald College, Quebec, Special thanks are due my f i e l d companions, namely, Drs. G. G. E. Scudder, M. Topping, Messrs. L. B a r t l e t t , and A. T. Smith and family. Their willingness to work long hours, often under dangerous and uncomfortably wet and cold conditions, w i l l always be remembered with appreciation. XV Valuable assistance was provided i n other areas by-numerous persons and i s g r a t e f u l l y acknowledged. Mr. H. Fuhrer, Canadian National Park Service provided climbing advice and assistance for exploring ice walls and g l a c i e r s and also supplied weather data for Jasper National Park; Ranger E. Sloinker and Forester A. T. Smith, Deschutes National Forest, Oregon, supplied ice cave l o c a l i t i e s and specimens of G r y l l o b l a t t a ; P. F. Brogan, Bend, Oregon, provided weather data, cave l o c a l i t i e s and early h i s t o r y . F i n a l l y , my eternal thanks and gratitude to my wife, Mary, who has been a true source of i n s p i r a t i o n and patience, without whom t h i s thesis would not have been completed. XVI ADDENDUM. None of the new names used i n thi s thesis has any status i n zoological nomenclature u n t i l such time as they are published i n accordance with the provisions of the Inter-national Code of Zoological Nomenclature. They are used i n the thesis only for convenience. In f i n a l publication d i f f -erent names may be used. G r y l l o b l a t t a o c c i d . e n t a l i s o c c l d e n t a l l s a d u l t f e m a l e t h r e e t i m e s a d u l t s i z e 1 I. GENERAL INTRODUCTION Few insects discovered i n the 20th century have stimulated wider Interest among entomologists than the Grylloblattodea. The f i r s t two specimens discovered at Banff, A l b e r t a , by E. M. Walker i n 1913> were described by him the following year as Gry l l o b l a t t a campodei formis, l n a new family of Orthoptera, the Gr y l l o b l a t t i d a e (Walker, 1914). At that time the order Orthoptera was a "catch a l l " assemblage of insects of such diverse forms as the cockroaches, mantids, s t i c k i n s e c t s , earwigs, grasshoppers and c r i c k e t s . G r y l l o b l a t t a campodelformls i s the emblem of the Entomological Society of Canada and i s of special i n t e r e s t on three accounts, namely (1) i t s peculiar structure and hence taxonomic and evolutionary a f f i n i t i e s , (2) i t s peculiar habits and temperature tolerance, and (3) i t s peculiar zoogeographic d i s t r i b u t i o n . Walker (1914) described G r y l l o b l a t t a as a wingless insect with a head resembling that of an earwig, eyes l i k e those of a termite, and antennae l i k e those of Phasmida. Such a peculiar combination of morphological features to many suggests an ancient or primitive taxon. Thus, Walker (1937) ca l l e d G r y l l o b l a t t a a l i v i n g f o s s i l and the possible representative of some ancestral l i n e leading to modern orthopteroids, while Zeuner (1945) considered i t a recent 2 l i v i n g Protorthoptera. There has been much speculation and divergence of opinion concerning i t s taxonomic position and phylogenetic a f f i n i t i e s . Expert orthopterists have been unable to agree on i t s status and r e l a t i o n s h i p s , and individual authors have been troubled by i t s systematic placement over the years. Thus, Walker (1933) considered the Gry l l o b l a t t i d a e to have close a f f i n i t y to the Salt a t o r i a (grasshoppers), but by 1938 he placed i t between the B l a t t a r i a (cockroaches) and the Sal t a t o r i a , and l a t e r placed i t nearest to the l i v i n g Ensifera ( c r i c k e t s , katydids):, (Walker, 1943). Crampton (1915) considered that Gr y l l ob l a t t a occupied a position intermediate between the Dermaptera (earwigs) and Isoptera (termites), and believed i t to be the l i v i n g representative of the common ancestor of the Gryllidae and Tettigoniidae (long-horned grasshoppers). In the course of 16 papers by Crampton (1915-1938) on the a f f i n i t i e s of G r y l l o b l a t t a , he associated i t with l i t e r a l l y every order of the orthopteroids. Imms (1927) placed G r y l l o b l a t t a nearer to the Dictyoptera, while Snodgrass (1937) believed i t had a f f i n i t i e s with B l a t t a r i a and Orthoptera. One of the aims of t h i s study was to c l a r i f y the a f f i n i t i e s of the G r y l l o b l a t t i d a e . This was attempted by a numerical analysis of the information provided by mor-phological characters present i n the orthopteroids and by a study of the l i p i d chemical c h a r a c t e r i s t i c s . The a v a i l -a b i l i t y of l i v i n g G r y l l o b l a t t a material enabled me to id e n t i f y morphological structures not r e a d i l y d i s c e r n i b l e i n preserved materials. The most recent studies, using large numbers of morphological characters to assess evolutionary a f f i n i t i e s i n the orthopteroids, are those of Giles (I963) and B l a c k i t h and B l a c k i t h (1968). Based solely on drawings, B l a c k i t h and B l a c k i t h (1968) considered G r y l l o b l a t t a to be closest to the Ensifera (Orthoptera). Giles (1963), using a single specimen of G r y l l o b l a t t a , considered i t to be closest to the Dermaptera. Biochemical taxonomy i s now a favored area of study and many constituents of the body are available for t h i s research. Since l i p i d s have been used in the past i n thi s context (Fast, 1970) and since they are of importance i n determining certain physiological a t t r i b u t e s of insects, t h e i r study i n G r y l l o b l a t t a and other orthopteroids and lower Insects permits a two-fold use of the same data. The Grylloblattodea occur i n cold and moist habitats i n S i b e r i a , Japan and western North America. They are found on g l a c i e r s , beneath stones on the fringe of snow patches, on moist talus slopes, i n r o t t i n g logs and i n i c e caves. The d i s t r i b u t i o n s are very disjunct and the tax-onomy of the North American forms i s most confused. Gry l l o-b l a t t a has long been considered by entomologists to be one of the rarest of a l l insects. At the beginning of t h i s study fewer than 75 adult specimens (the majority being G r y l l o b l a t t a campodelformis campodelformls) were i n the major entomological c o l l e c t i o n s of the world. Only a few years ago the world's largest insect c o l l e c t i o n , i n the B r i t i s h Museum (Natural H i s t o r y ) , possessed a single specimen/ (G. _c. campodelf ormis) and i t was treated with the same care and "reverence" as the Museum's Dodo and Archaeopteryx. The discovery of many new populations of G r y l l o b l a t t a . and the c o l l e c t i o n of well over 500 adults and numerous larvae from many types of habitat, permitted a study of d i s t r i b u t i o n , h a bitats, and the biology i n a depth not previously a t t a i n a b l e . In order to undertake these studies l n depth, however, a thorough systematic study of a l l material was e s s e n t i a l . Only i n t h i s way could s a t i s f a c t o r y i d e n t i f i c a t i o n s be obtained and accurate comparisons of data be made. . Thus, i t was necessary i n the course of the research to describe f i v e new species, three new sub-species, and to change the taxonomic position of two species. This taxonomic work i s i n Appendix I. The discovery of these new taxa and many new popu-l a t i o n s made i t possible to describe and discuss the present d i s t r i b u t i o n as a d i r e c t r e s u l t of Pleistocene or Neoglacial a c t i v i t y . Therefore, I have discussed the Rocky Mountain "Cordillera, Coast-Cascade and Sierra Nevada d i s t r i b u t i o n , and the Implications of the Pleistocene- post Pleistocene climates on the present occurrence of G r y l l o b l a t t a . This 5 could only be completed, however, a f t e r data on (1) the hypolithion and cavernicolous habitats, (2) the b i o c l i m a t i c requirements of various species both In the microhabitats and i n the laboratory, and (3) the seasonal a c t i v i t y of G r y l l o b l a t t a in alpine and cave habitats had been obtained. F i n a l l y , the a v a i l a b i l i t y of a c r y o p h i l i c form made i t possible to investigate the r e l a t i o n s h i p between environ-mental temperature and the l i p i d composition of i n s e c t s . The l i t e r a t u r e reports on the e f f e c t s of temperature and degree of l i p i d saturation are i n c o n f l i c t . This seems to be due to the fact that a l l studies have been performed by acclimation of a species to d i f f e r e n t environmental temperature regimes. Therefore, a comparative l i p i d analysis was performed on six d i f f e r e n t orthopteroid orders: two orders with high temperature preference, three with a temperate though variable preference, and one with a cold preference. My analysis was also used as a tool to indicate evolutionary r e l a t i o n s h i p s among these six taxa. In summary then, t h i s thesis i s concerned with the biosystematics of the Grylloblattodea. It i s concerned with the habitats, the temperature and humidity preferences, the d i s t r i b u t i o n and zoogeography. The systematic a f f i n i t y of the taxon i s considered and the p e c u l i a r i t i e s of the l i p i d composition are discussed with reference both to the phylo-genetic r e l a t i o n s h i p s and the habitats occupied by the taxon. 6 I I . HABITAT AND 3I0CLIMATIC REQUIREMENTS OF GRYLLOBLATTA SPECIES Introduction The t y p i c a l environmental niche reported for the North American G r y l l o b l a t t a i s the alpine-subalpine l i f e zone (Walker, 1914; Caudell, 1923; Ford, 1926; S i l v e s t r i , 1931; Gurney, 1937, 1948, 196l), but It does occur at low eleva-tions (Campbell, 1949; Kamp, I963). The genus has also been reported as a f a c u l t a t i v e cavernicole (Kamp, 1970). The hypolithion and cavernicolous situations seem to be the resident areas, and the insects range only temporarily into other habitats w,hile foraging for food. I. The Hypolithion Habitat G r y l l o b l a t t a occurs i n the hypolithion of the alpine biome. The term alpine zone, as used i n this t h e s is, follows Swan (1968). He states that the alpine zone begins at.the forest l i n e , which i s the highest elevation where trees are the dominant f l o r a type either in continuous stands or i n groups, and that the upper l i m i t of the alpine zone Is the highest elevation where the biota are dependent upon l o c a l autotrophic plants as the base for n u t r i t i o n . Thus defined, the alpine zone i s the same as eualpine, high alpine, n i v a l , eunival, high a l t i t u d e , Canadian and Hudsonlan i n the l i t e r a t u r e . The lower l i m i t s of the alpine zone i n western North America can be le s s than 1000 feet i n elevation i n the Yukon, to over 10,000 feet i n the southern Sierra Nevada and Rocky Mountains. The a l t i t u d e depends upon a number of physical f a c t o r s : l a t i t u d e , massiveness of the mountain range, d i r e c t i o n of slope, gradient of slope, extent and duration of winter snow cover, temperature, humidity, type of s o i l , moisture and p r e v a i l i n g winds. The subalpine, as here used, i s a subdivision of the alpine zone extending from forest l i n e to tree l i n e . Tree l i n e i s the upper l i m i t of trees that are generally scattered over open ground cover. The subalpine i s a t r a n s i t i o n a l region of the alpine zone where the environ-ment changes from that of the forest to that, of the alpine The climatic factors i n t e r a c t i n g on organisms i n the alpine include : 1. High transparency of the a i r ; 2. Low atmospheric temperatures; 3. Reduced evaporation from exposed surfaces; 4. Reduced water vapor tension correlated with reduc atmospheric pressure; 5. Greatly increased i n t e n s i t y of the u l t r a v i o l e t r a d i a t i o n ; 6. Increased rates of i n s o l a t i o n and r a d i a t i o n ; 7. Wide differences i n atmospheric and ground temper tures r e s u l t i n g in large diurnal and nocturnal f l u c t u a t i o n s ; 8 8. Snow cover than insulates against a r i d i t y , low temperature, u l t r a v i o l e t r a d i a t i o n , and governs substrate moisture and temperature. Snow cover, p a r t i c u l a r l y the winter snow cover, i s perhaps the most important environmental condition that makes l i f e possible here on a year around b a s i s . The snow cover exerts an ameliorating influence on the extremes of atmospheric temperature f l u c t u a t i o n s . In winter the snow forms a protective blanket against low temperatures and de s i c c a t i o n , and prevents ground f r e e z i n g . A foot of snow (30 cm.) w i l l provide enough in s u l a t i o n from an a i r tempera-ture of - 12° C. to prevent freezing of the ground (Landsberg, 1962). The i n s u l a t i o n effectiveness increases as the density of the snow increases. In the la t e spring and early summer the atmospheric and s o i l temperatures do not r i s e as rapidly as expected under conditions of high transparency and a r i d i t y of the alpine atmosphere. Snow has a very low heat conductivity (0.01 cal/cm/degree) and a very high albedo (80-90 per cent r e f l e c t i o n ) (Sellers, 1967). With snow cover most of the sun's energy (heat) i s r e f l e c t e d back and, of the remainder, a large percentage i s l o s t as latent heat i n the melting of the snow. The slow melting of the snow allows a gradual increase In s o i l temperature rather than an abrupt increase, as experienced on bare s o i l s or rock. Snow counteracts the high i n s o l a t i o n and a r i d i t y of the 9 alpine by replacing s o i l moisture and increasing humidities. Without the presence of some snow cover i n the alpine zone and i t s ameliorating e f f e c t s on the atmospheric conditions, only the most cold-hardy arthropods could inhabit t h i s area. However, many Insects, not p a r t i c u l a r l y cold-hardy, do l i v e i n the alpine area. They can do t h i s by l a r g e l y avoiding the open exposed environments. They l i v e i n un-exposed and somewhat, protected areas such as under mats of vegetation, the the f i r s t few milimeters or centimeters of the s o i l , under stones or rocks or i n other underground c a v i t i e s . Within these special microhabitats, special micro-climates p r e v a i l , which, although largely a product of and dependent on the atmospheric conditions above, are much les s severe and more stable than-in the open alpine area. These special microhabitats, with t h e i r rather constant micro-climates, heavily dependent on the snow cover, provide a wide variety of niches for a great many d i f f e r e n t types of i n s e c t . It i s evident that i t i s the hypolithic microhabitat i n the alpine that i s the most favored by insects, since i t i s here that one finds the greatest d i v e r s i t y and the highest populations. By d e f i n i t i o n , the hypolithic habitat includes a l l the underground c a v i t i e s and narrow spaces under the many stones and boulders that are more or l e s s deeply buried i n the ground. 10 Schonborn (I96l) has described the s t r a t i f i c a t i o n of the hypolithion i n Europe and A s i a . He states that stones about 100 to 400 cm.2 and 20 cm. thick form the most stable h y p o l i t h i o n . The f i r s t layer i s on the undersurface of the stone and i s 5 to 10 mm. t h i c k ; the character species are predominantly zoophagous. The second layer i s that of the f l o o r of the substrate under the boulder. The depth of t h i s layer depends upon the nature of the s o i l , the percolation of melt-water and the vegetation of the l o c a l i t y . It i s usually 20 - 30 mm. deep. The arthropod community of t h i s layer i s usually feeding on d e t r i t u s . The t h i r d layer underneath the substrate contains more or less decomposed plant debris and ranges up to 40 mm. i n thickness. This t h i r d layer may be taken as the base of the food pyramid of the h y p o l i t h i c community and i s largely composed of phytophagous forms or d e t r i t u s feeders, dominated by Collembola, The f i r s t and second layers are also the r e s t i n g places for nocturnal Diptera that must contribute s u b s t a n t i a l l y to the d i e t of the carnivorous species. The hypolithion i s characterized by a conspicuous di s c o n t i n u i t y of d i s t r i b u t i o n . It i s only prevalent i n older consolidated slopes that have adequate s o i l to hold the stones or boulders s u f f i c i e n t l y long for the subsurface environment to evolve. The development of t h i s environment r e s u l t s from decay of vegetable matter, mechanical a c t i v i t y of animals and melt-water. At the elevations associated with t h i s habitat i n the alpine zone, there are 11 many areas of unconsolidated talus slopes with loosely l y i n g stones that often s l i d e or r o l l down. These areas a f f o r d a very temporary hypolithion which i s generally not occupied by other than transients. I I . The Cavernicolous Habitat The cavernicolous niche for G r y l l o b l a t t a i s a micro-environment l i m i t e d to the ice caves i n the lava plateaus of western North America. These caves occur i n Oregon, C a l i f o r n i a and Washington, and G r y l l o b l a t t a occupy them as t r o g l o p h i l i c inhabitants. Geologically, the lava caves i n western North America are l a t e Pliocene or Pleistocene i n age and are tubes or tunnels i n f i s s u r e flows of pahoehoe basalts (Williams, 1957; Walker, Peterson and Greene, 1967; Greeley;1971). The caves were formed either by superheated gases.that blew through lava that was s t i l l viscous and underlay the cooling crusted surfaces, or by cracks and f i s s u r e s that formed at the advancing front of the flows. Such fronts were followed by an outpouring or draining of l i q u i d veins or conduits which l e f t a hollow void or tube. It i s common to f i n d two or more caverns i n the same system, and a few systems are multi-leveled with as many as f i v e stories (Kamp, 1963; Greeley, 1971). The caverns range i n size from small grottos to c a v i t i e s more than two miles long, 16 m. i n height, and up to 26 m. wide (Halliday, 1959; Greeley, 1970). Not a l l caves contain i c e , and probably l e s s than 10 per cent house persistent i c e . Balch (1900) has suggested that the cave ice represents remnants of the Ice Age. E c o l o g i c a l l y , the western ice caves are located i n the Upper Sonoran with a dominant f l o r a of Ceanothus, Artemisia. and Juniperus, with scattered Pinus contorta and P. ponderosa that grade into thick stands of these pines i n the Transition. Total moisture from r a i n and snow r a r e l y exceeds 12 inches per year, with 50 per cent l o s t by evaporation ( S e l l e r s , 1967). The c l a s s i f i c a t i o n of cavernicole organisms i s based upon the fauna of the limestone caves i n Europe that have a homeothermic environment. The c l a s s i f i c a t i o n was estab-l i s h e d by Schiodte (1849) and Schiner (1853) and was i n t r o -duced to North America by Packard (1888). Today i n North America i t i s used with l i t t l e modification for the fauna of limestone and lava caves. Troglobites are animals found only i n caves and are so modified that rudimentations of structure r e s t r i c t them to a cave existence. Troglophiles are animals found f r e -quently i n caves, reproducing there, and completing t h e i r l i f e cycle underground but not necessarily i n the cave. Trogloxenes are animals often found i n caves, but not completing t h e i r whole l i f e cycle underground. 13 The Microenvironment and Microclimate of G r y l l o b l a t t a Habitats G r y l l o b l a t t a are not universally d i s t r i b u t e d through-out the alpine-subalpine areas of western North America. They occur i n only some of the ice caves associated with the volcanic regions of the C o r d i l l e r a n . These insects appear to have p a r t i c u l a r microclimatic requirements that are to be found i n only l i m i t e d areas. Since these micro-environments have not been studied i n d e t a i l i n the Nearctic, one of the aims of the present research was to obtain det a i l e d microclimatic data from the alpine-subalpine hypo-l i t h i o n and the cavernicolous habitat i n l o c a l i t i e s where G r y l l o b l a t t a are found. I . Materials and Methods The alpine-subalpine habitats at McKenzie Pass, Crater Lake, Mt. Hood, Bachelor Butte and Sunshine Shelter, a l l i n the Cascade Range, Oregon, and at Athabaska G l a c i e r , Jasper National Park, A l b e r t a , were selected for study. These s i t e s were chosen because a l l have populations of G r y l l o -b l a t t a . are centra l l y located i n the Cascade or Rocky C o r d i l l e r a , and government weather records covering at l e a s t 10 years are a v a i l a b l e . Temperatures were recorded using thermographs (Ryan Instruments, model D-45; J . P. Friez and Sons, model 594). 14 Maximum-minimum thermometers (Six-type, Taylor Instruments) were used i n areas where thermographs could not be reached to service throughout the year due to snow conditions. Additional substrate temperatures were obtained while c o l l e c t i n g G r y l l o b l a t t a specimens. These data were combined with government weather, bureau records to construct monthly p r o f i l e s . Humidity recordings were obtained by thermohygrographs ( J . P. Friez and Sons, model 594; Edney-Short Mason L t d . , model 215) and lithium chloride hygrosensors (Hygrodynamics, models 15-1810, 4-4821KW). Humidities were recorded beneath the rocks of the hypolithion with a micro-aspiration psychometer ( J . P. Friez and Sons). South Ice Cave and Edison Ice Cave, Deschutes National Forest, Oregon, were selected for study of the cavernicolous habitat since they are lava i c e caves with resident popula-tions of G r y l l o b l a t t a . Other caves i n C a l i f o r n i a , Oregon and Washington were also examined p e r i o d i c a l l y . Temperature and humidity records, l i m i t e d i n time but covering the seasons of the year, were obtained from South Ice Cave between 1963 and 1971. In a d d i t i o n , i s o l a t e d temperature and humidity measurements were made in the other i ce caves i n C a l i f o r n i a , Oregon and Washington. To obtain long term temperature data ten 45-day continuous record thermographs (Ryan Instrument, model D-45) were i n s t a l l e d i n the d i f f e r e n t speleological zones of 15 South Ice Cave. A i r temperatures were recorded 25 cm. above the rocks. Rock and a i r temperatures were recorded from June, 1970, to March, 1971. Spot data collected previously for A p r i l and May f i l l e d i n a year's record. Relative humidity measurements were obtained twice a month for each temperature s i t e with the help and cooperation of l o c a l forest service personnel. By the use of posted markers, the amounts of melting and formation of ice were recorded at monthly i n t e r v a l s throughout the year. Simul-taneously, l i g h t measurements were obtained using a Gossen Lunasix meter. I I . Results .A. The Alpine-Subalpine Habitat 1. Microhabitat The usual microhabitat of Gr y l l o b l a t t a i n the alpine-subalpine was found to be that of c a v i t i e s beneath stones and boulders which are more or less buried i n the ground. Most insects were found under isolated rocks 50 to 150 cm. in diameter and buried to a depth of about 30 cm. They occasionally were taken beneath small rocks 5 to 10 cm. i n diameter (scree), and i n talus slopes 50 or more feet deep that contain rocks of a l l sizes ( F i g . 1). In a l l cases 16 F i g u r e 1. Examples o f v a r i o u s G r y l l o b l a t t a h a b i t a t s : upper l e f t , a l p i n e - s u b a l p i n e h y p o l i t h i o n , Mt. Hood, Oregon; upper r i g h t , e n t r a n c e t o South I c e Cave, Oregon; l o w e r l e f t , a l p i n e h y p o l i t h i o n around g l a c i e r b o r d e r s , Athabaska G l a c i e r , J a s p e r N a t i o n a l P a r k ; lower r i g h t , t y p i c a l h y p o l i t h i c c a v i t y w i t h snow margin ( r u l e r 15 cm. l o n g ) , 5200 f e e t , Mt. B a k e r , Washington. 17 the substrate was subirrigated by melt water. On occasion G r y l l o b l a t t a was found to occupy the spaces beneath decaying bark and i n or beneath f a l l e n logs. In the alpine-subalpine G r y l l o b l a t t a were found to seasonally occupy d i f f e r e n t s i t e s . They inhabited the hypo l i t h i c spaces deep i n the talus or benea.th the largest boulders i n summer and winter and were found i n other areas only i n the late spring and f a l l . 2. Microclimate The microclimate i n the hypolithion was found to be more uniform than the surface climate. While the surface a i r temperatures were subject to wide cir c a d i a n , seasonal and yearly f l u c t u a t i o n s , the temperatures of the hypolithion occupied by the Gr y l l o b l a t t a fluctuated only s l i g h t l y . Figure 2 presents the yearly a i r temperature p r o f i l e for f i v e alpiner- subalpine s i t e s . Yearly surface a i r tempera-tures ranged from - 40° C. to + .40° C. and a circadian range in excess of 30 degrees was noted. In contrast to t h i s , the greatest yearly temperature range recorded i n the hypo-l i t h i o n preferred by the Gry l l o b l a t t a was 10 degrees. In the a l p i n e , atmospheric r e l a t i v e humidity ranged from less than 10 per cent to saturation. A summer alpine humidity fl u c t u a t i o n i n exces-s of 80 per cent frequently occurred i n a few hours. 18 F i g u r e 2. Monthly atmospheric temperature of a l p i n e -subalpine h a b i t a t s of G r y l l o b l a t t a i n Oregon (1969 to 1971). 'wide bar = mean minimum and maximum temperatures; narrow bar = absolute minimum and maximum temperatures; s o l i d v e r t i c a l bar to r i g h t of °C. scale = pre-f e r r e d temperature range of G r y l l o b l a t t a . and narrow hatched bar - t o l e r a b l e temperature l i m i t s of G r y l l o b l a t t a . L o c a l i t y key to graph as f o l l o w s : £ I-icKenzie Pass 5,325 "ft. I Edison Ice Cave 5/200 f t . j Crater Lake 6,475 f t . I South Ice Cave 5/050 f t . • J Tirrbarlins, Mt. Hood 6,000ft. 9 , -_ Bacneior Butte/ Century Drive 6,450 f t . ^ Sunshine, Korth Sis-car 6,600 f t . !9 The microclimate i n the h y p o l i t h i c spaces occupied by the G r y l l o b l a t t a was found to vary and to depend on the size of the rock, substrate ground water, snow cover, surface temperature and exposure. The h y p o l i t h i c microclimate beneath boulders (50-150 cm. i n diameter and 20^ -50 cm. thick) p a r t i a l l y buried 30 cm. or more deep was the most uniform throughout the year. When t h i s type of hypolithion was inhabited by G r y l l o b l a t t a ' the yearly extreme range was - 3 to + 6° C. The hypolithion temperature was found to r i s e slowly from around 0° C. i n the summer to a maximum of + 6° C. i n l a t e September or early October. The temperature within the microhabitat then decreased ra p i d l y to the minimum, when the a i r tempera-ture went below 0° C. i n the f a l l . This cooling of the hypolithion continued u n t i l the snow cover provided i n s u l a -t i o n from further drop i n temperature. In the alplne-subalpine where Gr y l l o b l a t t a occurred the cooling ceased by mid-December. The h y p o l i t h i c temperature then remained . between - 3 and + 1° C. throughout the winter and spring. In the years when there was reduced or delayed snow cover, the temperature i n t h i s type of h y p o l i t h i c space f e l l below - 3° C. and the insects were found to move to the deeper hypolithion below the freezing zone. Only when the h y p o l i t h i c temperature reached between - 3 and 0° C. again did G r y l l o b l a t t a re-occupy the spaces. F i e l d records indicate that a humidity no lower than 20 70 per cent must be present i n combination with the - 3 to 4 6° C. temperature range for G r y l l o b l a t t a to inhabit the alpine hypolithion. The humidity i n the microclimate pre-ferred by the G r y l l o b l a t t a was found to fluctuate on a yearly cycle. Beneath the large boulders the humidity inhabited by G r y l l o b l a t t a ranged between 90 and 100 per cent during the spring season. . As the h y p o l i t h i c tempera-tures slowly rose during the summer and f a l l seasons the humidity gradually decreased. When the humidity was below 70 per cent the G r y l l o b l a t t a were found.to leave the hypo-l i t h i o n even i f the temperature remained below + 6° C. In the l a t e f a l l the h y p o l i t h i c humidity remained at the yearly minimum u n t i l the temperature decreased and the r a i n and f i r s t snows increased the moisture. The h y p o l i t h i c humidity then rose to at least 90 per cent and persisted at t h i s l e v e l through the winter. In response to the melting of the snow pack the maximum humidity i n the alpine hypolithion was found to occur i n the l a t e spring. B. The Cave Habitat 1. Microhabitats of the Ice Cave E c o l o g i c a l l y , the ice caves may be divided into four zones ( F i g . 3): 1) entrance zone, 2) t w i l i g h t zone, 3) upper dark zone, and 4) lower dark zone. The d i s t r i b u t i o n of the 21 Figure 3. South Ice Cave, Oregon. P r o f i l e to scale i n d i c a t i n g l i m i t s of entrance (E), tw i l i g h t ( T ) , upper dark (UD) and lower dark (LD) ecological zones. Two recorders i n each zone for a i r and rock temperatures. 22 fauna and the gradation of one zone into the other are t r a n s i t i o n a l and depend upon the season. a. Entrance Zone The entrance zone substrate i s composed of c e i l i n g and walls, spallated rocks and boulders interspaced by large amounts of wind- or water-borne s o i l and organic d e t r i t u s . B i o t i c a l l y , the entrance zone i s the r i c h e s t of the four zones. The walls and c e i l i n g are covered with lichens and the substrate i s l i t t e r e d with plant and animal material transported by E r l t h i z o n . C i t e l l u s . Neotoma, Peromyscus and a variety of b i r d s . The fauna of t h i s zone are f a c u l t a -t i v e troglophiles and w i l l also be found i n non-cave habitats. It contains, i n addition to the above rodents and b i r d s , an Invertebrate fauna of Arachnida, Diplopoda and Insecta. Examples are spiders, harvestmen, millipedes, moths, mosquitoes and Collembola. This zone grades into the t w i l i g h t zone that exhibits less organic material i n the substrate. This material decreases i n amount with increase i n distance from the entrance. b. Twilight Zone The t w i l i g h t zone receives only Indirect l i g h t of low i n t e n s i t y . B i o t i c a l l y , i t contains the nests and caches of rodents, a thin moss cover, and growth of fungi 23 on decaying organic m a t e r i a l . The fauna, i n addition to the same fauna of the entrance zone, contains Campodeidae, f a c u l t a t i v e G r y l l o b l a t t a . Machilidae and Coleoptera. This zone i s also used by many organisms as a retreat from extremes of surface temperature. During the hot summer and early f a l l the l o c a l b i r d and mammal fauna retreat here from midday u n t i l dusk and they u t i l i z e the small pools of melt or drip water present i n t h i s t w i l i g h t zone. In periods of very hot, dry weather Diptera and Lepidoptera invade t h i s zone as w e l l . The following two zones are spe l e o l o g i c a l l y described as one zone, the dark zone. B i o l o g i c a l l y , however, the dark zone i s d i v i s i b l e into two, as follows. c. Upper Dark Zone The upper dark zone may receive i n d i r e c t l i g h t of very low i n t e n s i t y , l e s s than .016 foot candles (.172 lumens/ sq. m.) for an hour or less during the early morning or near sunset. The length of l i g h t exposure depends on the entrance orientation and amount of c e i l i n g collapse i n the tw i l i g h t and entrance zones. B i o t i c a l l y , t h i s zone i s inhabited by small rodents. Rodent nests are always present and i n some caves very large nests and caches of twigs, pine cones and grass occur that are evidently deposited by pack r a t s , Douglas s q u i r r e l s 24 and porcupines. Plant f l o r a i s l i m i t e d to fungi growing on organic m a t e r i a l . The Arthropoda are the dominant fauna, composed of Campodeidae, Machilidae, Collembola, G r y l l o -b l a t t a . Diplopoda and Arachnida. During the f a l l there may be large numbers of dead Lepidoptera and Diptera k i l l e d by the cold and these are u t i l i z e d as food by the other Arthropoda. Chiroptera use t h i s zone during the diurnal period and may i n certain caves hibernate here during the winter. d. Lower Dark Zone This microhabitat i s a region of permanent darkness with l i t t l e atmospheric exchange throughout the year. Organic material i s very lim i t e d and i s r e s t r i c t e d to the droppings of wandering rodents. Fungi and molds are the exclusive f l o r a and are r e s t r i c t e d to the scat. The fauna i s l i m i t e d to Campodeidae, Diplopoda and Arachnida. 2. Microclimate of the Ice Cave Habitats The f i e l d data obtained (F i g s . 2 and 4) show that the microclimate of the caves i s i n general more stable than that of the surface climate. While the a i r tempera-tures and humidities i n the outer zones of the cave environ-ment fluctuated with the external conditions, the substrate 25 Figure 4. South Ice Cave, Oregon. Monthly temperature p r o f i l e of rock and cave a i r showing la t e n t response of rock temperature to that of a i r ( A p r i l , 1970, to March, 1971). S = surface, E = entrance zone, T = t w i l i g h t zone, UD = upper dark zone. , 2 6 and atmosphere 5 cm. above the substrate in the deeper zones were much more uniform. a. Temperature Figure 4 presents the mean monthly temperature pro-f i l e s obtained from South Ice Cave. Temperatures recorded at South Ice Cave of the different biospeleological habitats agree with those obtained from other ice caves in Cali-fornia, Oregon and Washington. I therefore believe the yearly temperature profile of South Ice Cave to be repre-sentative of the temperatures found in most western North American lava caves containing, ice that persists for a number of years. The atmospheric temperatures of the twilight and dark zones were below 0° C. from November to May. The rock temperatures of these zones were generally a few degrees colder than the air temperatures. Rock temperatures in these zones were - 15° C. in late winter. The rock did not remain at such temperatures but slowly rose in the spring and summer as i t lost cooling power to the slowly warming, atmosphere. By late summer and early f a l l , rock temperatures were between - 3 and + 3.5° C., depending on the distance from the cave entrance, proximity to ice, the mass of ice in the zone, and the heat flow through the ceiling and underlying rock. The temperatures in the entrance zone varied greatly, 27 both seasonally and d a i l y . This zone i s continually under the influence of the external climatic f l u c t u a t i o n s . From January through May t h i s zone was i n equilibrium with the temperature means of the external environment. From June through October the a i r temperatures of the entrance zone were generally higher than the mean maxima of the external temperatures. With the lowering of the external tempera-tures during November, the entrance rock temperature approached equilibrium with the external conditions. The mean a i r temperature i n t h i s zone dropped below the outside mean during November. In December there was a lag of the entrance zone rock, and a i r temperatures behind the external temperatures. Thus, there was a d e f i n i t e lag i n cave temperatures compared with surface temperatures. Seasonal values i n the cave responded as a dampened phase s h i f t of the annual surface temperature curves. In winter, when the external temperature f e l l below that of the cave atmosphere, colder, dense, a i r entered the cave and replaced the warmer cave a i r . Repeated influxes of colder a i r lowered the cave tempera-ture and gradually the outer ecological zones of the cave came into equilibrium with the mean outside winter tempera-ture. The deeper eco l o g i c a l zones f e l l below the outside mean and remained below the outside mean throughout the re s t of the winter. The repeated charges of cold that lower the t w i l i g h t and upper dark zone temperatures during the winter caused 28 those areas to remain r e l a t i v e l y stable during the spring and early summer. A gradual increase of temperature occurred during the warmest seasons and the microhabitat temperatures in the deeper zone of the cave rose to a maximum of 5° C. b. Ice The formation of new i c e i n the cave was dependent on the simultaneous presence of water and below-0° C. rock temperatures. The usual source of the water was seepage of r a i n and snow melt through the overlying rock; i n rare cases running water was the primary source. In winter the cave received l i t t l e or no seepage from the frozen surface and at t h i s time the r e l a t i v e humidity f e l l below 10 per cent. In general, during the winter the amount of i c e present represented the i c e and melt water that remained at the end of the f a l l season. However, wind-blown snow contributed to ice build-up i n the entrance zone during the winter i n some years. Ice accumulated in the lava caves only during the spring and early sumxer. Only during an exceptional year di d small amounts of ice accumulate during the la t e f a l l . A warm (above 0° C.) rainy period following an i n i t i a l below-0° C. cooling of the rock surface resulted i n seepage being transformed into i c e . 29 During the f a l l usually the dry cold c i r c u l a t i n g a i r caused evaporation of. most of the water formed by summer melting of the ice before i t could freeze. During the spring warmer external a i r c i r c u l a t i n g through these caves r a i s e d the rock temperature above 0° C. so rapidly that seepage from runoff did not always form i c e . In the spring, i c e formation was found to vary and t h i s depended upon surface temperatures, time of ground thaw, and the amounts of r a i n and melt water. Cave ice was found to vary from thin sheets a few mm. thick to gla c i e r e s 2 to 5 m. i n depth, up to 15 m. wide and 50 m. long. In some depressions,"lakes", ice was found and there was from 30 to 60 cm. of water on top during late summer and early f a l l . Ice i s not perpetual, but only t r a n s i t i o n a l , i n most of the lava caves studied i n the west. In caves with active c i r c u l a t i o n , ice only lasted 1 to 2 years and i t was only i n those caves with a s t a t i c or passive c i r c u l a t i o n that cave ice p e r s i s t s for years. The exact age of persistent ice i s unknown in the west beyond the 150 years of recorded h i s t o r y , since none of the current dating methods have been applied to western cave i c e . c. Humidity Ice cave humidities were also found to be seasonally 30 c y c l i c . In the t w i l i g h t and upper dark zones of South Ice Cave from November to March a l l water vapor was frozen by the below-0° C. temperatures and t h i s caused a drop i n humidity. At the onset of the winter season the humidity of these zones was usually above 80 per cent. It f e l l to 10 per cent or less by January and the microhabitat was very dry u n t i l the spring surface thaw. There was a rapid r i s e of humidity during the spring melt of A p r i l and May. The humidity was noted to r i s e i n a 2-week period from 10 per cent or less to over 80 per cent. The humidity then s t a b i l i z e d around 80 per cent for a short period as the seepage of melting snow diminished. From June to September seepage from spring and.summer r a i n and the gradual melting of ice caused the humidity to r i s e to 90 per cent or more. This same cycle was present i n other western lava i c e caves (personal observation). It must be re-stated that the temperature and humidity patterns presented here are for the rock and the bottom 25 cm. of the atmosphere. I took the climatic data from t h i s region for i t i s the active zone for almost a l l of the cavernicolous fauna. S t r a t i f i c a t i o n of temperature and humidity occurs i n the cave atmosphere, with tempera-tures increasing and humidities decreasing from f l o o r to c e i l i n g . 31 I I I . Discussion and Summary Gr y l l o b l a t t a has been found on g l a c i e r s (J. G. Edward, personal communication), at the borders of springs (Henson, 1957a), beneath stones on the fringe of snow patches (Gurney, 1937, 1948, 195 3, 1961), under stones (Ford, 1926; Gurney, 1937, 1961), boulders or logs firmly embedded i n the ground ( M i l l s and Pepper, 1937; Kamp, 1963), i n moist rock s l i d e s (Gurney, 1937, 1948; Campbell, 1949; Chapman, 1953), i n r o t t i n g logs (Henson, 1957a; Kamp, 1963; Gagne, 1968) and i n ice caves (Kamp, 1953, 1963, 1970). These insects have d e f i n i t e humidity and temperature require-ments ( M i l l s and Pepper, 1937). The present research indicates that the habitats preferred by G r y l l o b l a t t a must be cold, but not too cold, and moist, but not wet. Such cold and moist habitats, that are more or l e s s stable and permanent, occur i n Isolated regions of the alpine-subalpine. The hypolithion above timberline i s composed of a wide variety of microenvironments. Each type of micro-enviornment has a unique microclimate more or less d i r e c t l y influenced by the capricious alpine-subalpine climate. The c a v i t i e s beneath stones or boulders, 50 to 150 cm. i n diameter and 20 to 50 cm. thick, t y p i c a l l y provide the optimum conditions for G r y l l o b l a t t a i n the alpine-subalpine. The temperature-humidity conditions of such h y p o l i t h i c c a v i t i e s fluctuate minimally during the day, and seasonal 32 fluctuations beneath the larger boulders are neither abrupt nor extreme. This type of hypolithion has a yearly extreme tempera-ture range of - 3° to + 6° C. and a humidity above 70 per cent. Seasonal variations are hardly evident, with winter and spring temperatures between - 3° and + 1° C. The temperature slowly r i s e s during the summer, reaching the maximum by l a t e September or early October. The tempera-ture then decreases r a p i d l y to the minimum u n t i l the snow cover provides s u f f i c i e n t i n s u l a t i o n from further drop i n temperature. The most important physical factor of the hypolithion that makes possible habitation by G r y l l o b l a t t a i s snow cover. It provides i n s u l a t i o n against the intense winter atmospheric cold and a r i d i t y and i s the source for the gradual development of the hy p o l i t h i c cavity and climate. However, the humidity of the alpine-subalpine hypo-l i t h i o n microhabitat preferred by G r y l l o b l a t t a fluctuates on a yearly cycle. It ranges between 90 per cent and saturation during the spring. As the temperature slowly r i s e s during the summer and f a l l , the humidity gradually decreases, evidently reaching the minimum tolerated by Gr y l l o b l a t t a i n the early f a l l . The= f i r s t r a i n s and snow, coupled with decreasing temperatures during the l a t e f a l l , r a i s e the humidity to approximately 90 per cent where i t p e r s i s t s throughout the winter. Snow melt i n the spring then r a i s e s the humidity to the maximum l e v e l . 33 Humidity seems to govern the occupancy of the hypo-l i t h i o n by G r y l l o b l a t t a more than temperature. The Insect w i l l not inhabit any hypolithion with r e l a t i v e humidity l e s s than 70 per cent, regardless of the temperature. M i l l s and Pepper (1937) have e a r l i e r reported that the op-timum humidity i s around 100 per cent, and Henson (1957b) noted an almost saturated humidity i s required. Chapman (1953) has stated that i t i s moisture, rather than a l t i t u d e , that i s important to G r y l l o b l a t t a . The fact that these insects occur at Kamloops, B r i t i s h Columbia, i n talus slopes at 1400 feet (Campbell, 1949) and i n the talus at 3195 feet on the North Fork of the Feather River (Kamp, I963) would seem to bear t h i s out. Here, as i n many other l o c a l i t i e s , they can only be found i n the l a t e f a l l - early spring, when cool moist surface conditions p r e v a i l . While Campbell (1949) collected specimens out i n dry conditions i n the f a l l , t h i s i s most unusual. The cavernicolous habitat for G r y l l o b l a t t a i s a micro-environment li m i t e d to the ice caves i n the lava f i e l d s (Kamp, 1953, 1963, 1970). G r y l l o b l a t t a i s not found i n a l l i c e caves, but only i n those with s u f f i c i e n t quantity of persistent i ce to maintain a spring-to-fa11 temperature range of - 3° to + 5° C. and over 80 per cent r e l a t i v e humidity. These conditions are comparable to those pre-v a i l i n g i n the alpine-subalpine hypolithion c a v i t i e s . In contrast to the narrow yearly fluctuations of the h y p o l i t h i c microclimate, the cavernicolous habitat occupied 34 by G r y l l o b l a t t a undergoes a d r a s t i c change i n the winter. This winter change i s so great that i t renders the caverni-colous environment uninhabitable by G r y l l o b l a t t a . During the winter the cave atmosphere i s recharged with cold a i r and i t i s t h i s cold recharge which makes the cave an un-suitable environment for Gr y l l o b l a t t a at thi s time of year. The insects are forced out onto the surface, into a habitat which they cer t a i n l y cannot tole r a t e at any other time of the year. The present study thus confirms that G r y l l o b l a t t a has p a r t i c u l a r temperature and humidity requirements, which no doubt i n part account for the rather spotted d i s t r i b u t i o n of the genus i n western North America. (Gurney, 1937; Kamp, 1963). It i s l i k e l y that t h i s insect i s not as rare as often stated. Instead, the insect has p a r t i c u l a r environ-mental requirements that are either not often met with i n the f i e l d , or not recognized by entomologists searching for these insects. This, together with th e i r predaceous habit that tends to space them out ( M i l l s and Pepper, 1937; Campbell, 19^9; Gagne, 1968) and the fact that they appear photonegative and active mostly at night (Ford, 1926) has resulted,at times, i n l i t t l e success when searching for specimens. The insect i s active a l l winter and does not have a dormant period ( M i l l s and Pepper, 1937; Henson, 1957a). I t , thus, i s active also when most entomologists think there i s l i t t l e about to be coll e c t e d i n the f i e l d . 35 A c t i v i t y of G r y l l o b l a t t a i n Alpine and Cave Habitats I. Introduction The a c t i v i t y of G r y l l o b l a t t a i n the hyp o l i t h i c habitat has received l i t t l e attention i n the l i t e r a t u r e . What has been published i s Incidental to taxonomic descriptions or reports of laboratory cultures (Ford, 1926; Campbell, 1949; Gurney, 1953, 1961). The most, extensive description of f i e l d a c t i v i t y i s a report of the winter a c t i v i t y of G. c. campodeiformis from Kamloops, B r i t i s h Columbia (Campbell, 1949). A c t i v i t y i n the cavernicolous habitat has been l i m i t e d to the general accounts of Kamp (1963, 1970). I I . Methods Observations were made of the a c t i v i t y of the insects i n the s p e c i f i c habitats at seasonal i n t e r v a l s throughout the year. The v e r t i c a l and horizontal movement within the habitat was recorded as the seasonal macroclimate a f f e c t s the environmental conditions within the hypolithion and cave. I I I . Results A. A c t i v i t y i n Alpine Habitats 36 The seasonal h y p o l i t h i c a c t i v i t y of G r y l l o b l a t t a temporarily varied i n response to climatic conditions, extent and depth of hypolithion, l a t i t u d e , slope and e l e -v a t i o n . Winter snowfall and the spring and summer tempera-tures were the most important factors regulating yearly a c t i v i t y within the alpine-subalpine habitats. Most of the species and populations of G r y l l o b l a t t a d i s t r i b u t e d i n the C o r d i l l e r a were found to follow the same general yearly a c t i v i t y pattern. The insects were active throughout the year in d i f f e r e n t depths of the hypolithion i n response to the surface climate. 1. Winter Season Insects were found beneath rocks buried at shallow depths or on the surface beneath the snow pack. In certain l o c a l i t i e s , where the slope and exposure or below-normal snowfall tended to produce i s o l a t e d accumulations of snow, most of the G r y l l o b l a t t a population was active below the freezing zone i n the hypolithion. The insects did not tend to aggregate beneath the i s o l a t e d snow patches except i n a shale or sandstone substrate. This lack of aggregation i s not surprising since adult and older i n s t a r s were found to be extremely antagonistic toward other i n d i v i d u a l s and are c a n n i b a l i s t i c . Except for 5 pairs in. copulo, more than one in d i v i d u a l was never found under a rock or within \\ feet of another during c o l l e c t i o n s 37 of approximately 1000 i n d i v i d u a l s . On occasion, during d u l l overcaste days without wind and with temperatures between 0 and + 5° C . , i n d i v i d u a l s were taken on the snow surface. Two i s o l a t e d populations were found only during the l a t e f a l l and winter seasons. G r y l l o b l a t t a barberi (Feather River, Plumas Co., C a l i f o r n i a ) and G. c. campodei-formis (Mt. Paul, Kamloops, B r i t i s h Columbia) occurred at elevations below 2000 f e e t . F i e l d observations indicate that they both were active i n the upper 3 to 5 feet of the hypolithion between October and February. Temperature measurements and c o l l e c t i n g suggest that t h i s depth of habitat could not be tolerated by the insects during the rest of the year and they retreat into the substrate. The talus slopes of Mt. Paul receive l i t t l e winter snow and temperatures may be below - 30° C. Winter c o l l e c t i n g indicates that the G r y l l o b l a t t a population here migrates below the freezing l e v e l in the hypolithion and i s only active i n the upper portions where a i r temperatures are between - 5° C. and + 6° C. with no wind. 2. Spring Season In most years the winter snow pack persisted u n t i l l a t e spring at l o c a l i t i e s studied above 5000 fe e t . The G r y l l o b l a t t a were found to occupy the same regions of the hypolithion during t h i s period as they do i n the winter. 38 As atmospheric surface temperatures rose there seemed to "be increased snow surface a c t i v i t y . Y/ith the increased abundance of f l y i n g insects the G r y l l o b l a t t a became active foragers on the snow surface during the early evening. Nocturnal surface a c t i v i t y was r e s t r i c t e d to a i r tempera-tures between - 2 and + 5° C. and a humidity over 75 per cent. The insects retreated beneath the snow within minutes i f the temperature dropped a few degrees or a l i g h t wind came up to lower the humidity. In the subalpine, when the snow melted into i s o l a t e d banks or patches during May and June, G r y l l o b l a t t a was found i n the shallow hypolithion, in. f a l l e n logs, and under small stones and i n mosses at the snow edge. G r y l l o b l a t t a was not found under large rocks or i n shallow gravel embedded i n mud or running water. G r y l l o b l a t t a preferred a hypolithion that was moist but not wet. The early nymphal stage was found i n wetter conditions than were the larger, instars or adult forms. Short periods of contact with water did not have an adverse ef f e c t on these insects. Most stages of G r y l l o b l a t t a were observed to walk through shallow pools or melt r i v u l e t s , and even under water for short distances. Three hours of submergence have no apparent adverse e f f e c t on adults, but four hours of submergence appear to be l e t h a l . The penultimate instar can withstand 2 hours of submergence while 5 minutes of exposure beneath water appears l e t h a l to the f i r s t i n s t a r . 39 3. Summer Season In the alpine-subalpine habitats Gr y l l o b l a t t a occurred beneath the larger rocks deeply buried in subirrigated slopes, the smaller rocks at the snow edge, or deep within large r o t t i n g l o g s . A l l stages were found nocturnally active on persistent snow f i e l d s or g l a c i e r margins within the l i m i t s of tempera-ture and humidity stated previously. Nocturnal foraging began between 9'-30 and 1 0 : 0 0 P. M. and r a r e l y continued a f t e r 1:00 A. M. 4. F a l l Season The locations of the insects within the hypolithion during early f a l l were found to depend largely on the amount of winter snow pack and l a t e spring and summer temperatures. In normal years G r y l l o b l a t t a were usually found beneath rocks 2 to 3 feet i n diameter which were p a r t i a l l y buried at least 1-g- feet deep in subirrigated slopes. They were active between the smaller rocks that were buried beneath !•§• feet or more of the vegetation mat. During t h i s period nocturnal surface a c t i v i t y decreased because of the reduced amounts of snow and the shorter i n t e r v a l s of optimum temperatures and humidities. The substrate depth of a c t i v i t y during the l a t e f a l l In the hypolithion was found to depend solely on the time AO of the f i r s t persistent snow. In the normal years snow depth was s u f f i c i e n t to insulate the shallow regions of habitat against below-O0 C. temperatures and G r y l l o b l a t t a was active i n the same portions as during l a t e summer. With periods of below-O0 C. temperatures before the f i r s t snow the insects retreated deep into the habitat. With the a r r i v a l of the f i r s t snow G r y l l o b l a t t a were found beneath the smallest rock, above the surface i n tree stumps and i n damp moss i n the diurnal period. When r a i n or snowfall occurred with temperatures between - 3 and + 6° C., a l l stages of the insect were at times found active on the surface. Nocturnal a c t i v i t y was never observed i n l a t e November and early December. The f i r s t i n s t a r nymphs were only found a f t e r the f i r s t snow i n the f a l l . By mid-winter the second in s t a r was captured. This has also been reported by Ford (1926) and Walker (193T). The l i m i t e d presence of the f i r s t instar suggests that hatching of the eggs takes place a f t e r exposure to the f i r s t combination of below-0° C. temperature and snow. On occasion G r y l l o b l a t t a were found to be active beneath bark or small rocks that were covered with a trace of snow and when the atmospheric temperatures were down to - 10° C. These occasions were always preceded by higher a i r temperatures and some snow melt. 41 B. A c t i v i t y i n Cave Habitats In the Bend area, G r y l l o b l a t t a were found to be t r o g l o p h i l i c cavernicoles except in winter months. During the winter when there was below-freezing temperatures and low humidities i n the caves, the insects were not present i n t h i s habitat. As the temperatures and humidities f e l l , the arthropod fauna was found to migrate toward the surface, where they were found as t y p i c a l hypolithions under rocks below the f r o s t depth. During the spring, as the surface temperatures slowly rose, the insects migrated downward, eventually reaching the cave environment. Arachnida and Diplopoda were the f i r s t to re-occupy the t w i l i g h t and upper dark zones of the cave, usually by mid-April. Evidently these groups are not as r e s t r i c t e d by the humidities of the environment as Gr y l l o b l a t t a for they were also the l a s t to migrate out during the late f a l l . When humidities reached 80 per cent and percolating melt water-had moistened the rock surfaces, G r y l l o b l a t t a returned to the upper dark zone, usually i n la t e May or early June, where they remained u n t i l the next winter. Most years the hypolithion became very dry and warm by mid-April ( F i g . 4) and then was not inhabited by the G r y l l o b l a t t a . These insects invaded the various outer ecological zones of the cave i n sequence, moving to the next deeper as the previous one became uninhabitable. 42 By March the G r y l l o b l a t t a were active i n the moss and shallow hypolithion of the surface adjacent to the entrance. The length of time t h i s area was occupied appeared to depend on the temperature, r a i n f a l l and melt water runoff. In normal years t h i s region became very dry and warm by mid-April and the insects retreated deeper into the lower entrance zone. During the nocturnal period the G r y l l o b l a t t a were observed to search for food i n the upper regions of the entrance and on the surface, es p e c i a l l y on melting snow banks. This surface and entrance a c t i v i t y was evidently governed by humidity rather than by temperature, for night-time temperatures were usually around or below freezing at t h i s time of the year. The G r y l l o b l a t t a were only active on those nights when the humidity was above 75 per cent and i n s t i l l a i r . A s l i g h t breeze and drop of the humidity was found to induce the G r y l l o b l a t t a to retreat within minutes to the lower regions of the entrance zone. In May, when there was a sharp temperature r i s e in" the entrance zone ( F i g . 4), the insects moved into the t w i l i g h t and upper dark zones. These zones were occupied then u n t i l l a t e October or November, at which time the G r y l l o b l a t t a evacuated the cave environment r a p i d l y . They then became occupants of the hypolithion. During the winter months G r y l l o b l a t t a were active i n the hypolithion below the frost zone regardless of the surface temperature. Insects were observed and collected 4 A3 1 m. below the surface when the a i r temperature was - 12 to - 20° C. In mild periods of the winter, e s p e c i a l l y during and immediately after wet snow storms when tempera-tures were s l i g h t l y above zero, the insects were often active on the snow surface. The Gr y l l o b l a t t a ,found in the high lava plateaus of western North America, for most of the.year were thus true cavernicolous t r o g l o p h i l e s . When the cave environment became unfavorable, they then became hypolithions. IV. Discussion and Summary The h y p o l i t h i c a c t i v i t y of G r y l l o b l a t t a i n the alpine-subalpine of the western Co r d i l l e r a temporarily varies i n response to the e f f e c t s of the clima t i c conditions. The insect i s r e s t r i c t e d to certain parts of the hypolithion and shows l i m i t e d surface a c t i v i t y , probably because o f . i t s narrow temperature-humidity optimum. Snow depth and spring-summer temperatures are the most important factors regulating yearly a c t i v i t y . G r y l l o b l a t t a do not hibernate ( M i l l s and Pepper, 1937; Henson, 1957a, 1957b) and, hence, are dependent on adequate snow cover for protection against the atmos-pheric cold and low humidities of the winter season. When 4A snow depth i s s u f f i c i e n t to insulate against extreme dry-ness and cold the G r y l l o b l a t t a are active i n the shallow hypolithion and ground surface beneath the snow. Adequate snow pack during the spring and early summer provides the low temperatures and high humidities necessary to the insect and i t remains active i n the hypolithion and beneath the snow. As the snow melts during the summer, diurnal a c t i v i t y i s r e s t r i c t e d to the deeper hypolithion that i s subirrigated by melt water. Here, during the:, highest summer temperatures, 15 to 20 feet below the surface, high humidity, i c e , and a i r temperature around 0° C. are often present (Kamp, 1963). F a l l season i s the most c r i t i c a l for the G r y l l o b l a t t a . In the western C o r d i l l e r a the early f a l l i s generally the d r i e s t and warmest season. During t h i s period the insects w i l l only be active deep i n the hypolithion. Some snow i s necessary during the l a t e r f a l l as protection against the rapidly f a l l i n g temperatures. Surface a c t i v i t y may take place at any season of the year, but i s li m i t e d to a range of - 2 to + 5° C. and at least 75 per cent humidity, but i s usually r e s t r i c t e d to the nocturnal period. Winter snow surface a c t i v i t y has been reported by Pletsch (1947), Campbell (1949) and Gurney (1953)» hut they did not give d e t a i l s of the snow surface microclimatic conditions that permit such a c t i v i t y . Cavernicolous species are t y p i c a l h y p o l i t h i c Inhabitants during the winter when the cave environment i s too cold and dry. In the spring, when cave temperatures slowly r i s e , the insect re-enters the cavernicolous habitat, occupying each deeper zone as the more shallow one becomes too dry or warm. Temperature and Humidity Preference of G r y l l o b l a t t a I. Introduction It i s often stated i n the l i t e r a t u r e that the pre-ferred habitat of G r y l l o b l a t t a i s cool and damp. The few temperature-humidity preferenda and the optimum and l e t h a l l i m i t s reported are c o n f l i c t i n g . M i l l s and Pepper (1937) reported an optimum of + 3.7° C., a preferendum between + 0.1 and f 15.5° C., with l e t h a l l i m i t s about - 6.2° C. and + 27.8° C. A preference range between - 2.5° C. and + I I . 3° C. was stated by Edwards and Nutting (1950). Henson (1957b) published an optimum of + 1.0° C. and a preference range between - 3.0 and + 5.0° C. Humidity requirements are inferred to be 100 per cent. In the test procedures described by each there was no statement of the humidity at the time of t e s t i n g nor any control of the r e l a t i v e per cent. In the above studies the insect tested was G. _c. campodelformis, collect e d from G a l l a t i n Canyon, Montana. F i e l d data of temperatures at times of c o l l e c t i o n have been reported to be - 9.44 to + 15.56° C. by Campbell.(1949), - 1 .8 to + 7.22° C. by Kamp (1963), and he + 1.1° C. by Chapman (1953). These temperatures were from four d i f f e r e n t species of G r y l l o b l a t t a , ranging from B r i t i s h Columbia to Montana and C a l i f o r n i a . From these apparent contradictions, the questions a r i s e as to whether the differences are due to experimental conditions, a c c l i m a t i z a t i o n of test specimens, seasonal responses, n u t r i t i o n , age, or the varying preferences by d i f f e r e n t species and populations. In an attempt to answer these questions, I investigated the temperature-humidity responses under controlled laboratory conditions. I I . Materials and Methods The fundamental p r i n c i p l e underlying these studies i s that of attempting to simulate i n the laboratory the spectrum of natural conditions under which G r y l l o b l a t t a have been observed or have been reported to encounter i n th e i r natural habitats. Accordingly, four c r i t e r i a were established for these experimental procedures: 1) use of a multiple rather than a single factor approach, 2) use of gradients of temperatures and humidities comparable to those i n the habitat, 3) use of d i f f e r e n t populations and species, and 4) use of insects of known sex, length of time i n c a p t i v i t y , n u t r i t i o n , and holding temperatures and humidities. 47 A. Experimental Chamber I designed and b u i l t a multiple factor environmental gradient chamber. The chamber i s a highly modified and sp e c i a l i z e d design based i n part, i n terms of operating p r i n c i p l e s , on the c i r c u l a r gradient chamber of P r a t t , C o l l i n s and Witherspoon (1957). Pratt's chamber i s of ideal design for work on f l y i n g insects at ambient or higher temperatures. In working at temperatures ranging around 0° C. and humidities of 70 per cent and higher, many physical problems are encountered. The manufacturer's published data for transparent p l e x i -glass and i t s resistance to f l u i d s are not v a l i d i n the range of - 25 to + 15° C. The solutions generally used to e s t a b l i s h and main-t a i n r e l a t i v e humidity become toxic to the insects and to the sensors used to record humidity. In atmospheres of high r e l a t i v e humidity, as the dew point of the atmosphere i s reached by lowering the temperature, condensation, fog and r a i n form i n the chamber. The test chamber I used was a rectangular transparent ple x i g l a s s tube (15 x 15 x 138 cm. and 1.25 cm. thick) surrounded by separate heat exchangers lengthwise and at either.end for temperature c o n t r o l . Each heat exchanger had 48 separate i n l e t and outlet pipes ( F i g . 5 ). Three access ports were spaced equidistantly on the top for i n s e r t i o n of Insects and sensing elements. Solution boxes for humidity control were at either end. The entire test chamber was housed i n a 6 x 3 x 2 -foot r e f r i g e r a t e d cabinet. Three c i r c u l a t i n g pumps (Gebruber-Haake 2-model, FS 1-model HK) were outside the cabinet and connected to the heat exchangers by insulated tubing. Temperature gradients were established and maintained by the c i r c u l a t i o n of soya o i l and a saturated solution of NaCl through the two end heat exchangers. The temperature of these heat exchangers was higher and lower than the maximum and minimum temperature of the gradient for a given t e s t . The temperature of the l i q u i d i n the lengthwise heat exchanger was that of the mid-point of the gradient desired. The temperature of the r e f r i g e r a t o r cabinet was usually maintained at or below the mid-point of the gradient. By varying the choice of temperatures and the flow rates i n and out of the heat exchangers, i t was possible to e s t a b l i s h an atmospheric temperature gradient of 10 to 15° C. i n the inner chamber i n 4 to 10 hours. The lower the desired gradient was i n temperature, the-. longer the time necessary'to e s t a b l i s h the gradient. Once the gradient was established and stable i t held from 4 to 49 Figure 5. Transparent view of temperature-humidity chamber. Arrows indicate d i r e c t i o n of flow for temperature control into the three separate heat exchangers. Heat exchangers are separated by hatched p a r t i t i o n s i n diagram. Three access ports located on top for i n s e r t i o n of sensors and specimens. Scale i s 1/10 actual s i z e . 50 8 hours over a temperature spectrum of - 15 to + 20° C. The flow rate of the cold heat exchanger was 2 to 3 times that of the higher heat exchanger. The flow rate i n the middle heat exchanger was adjusted to be 1/3 to 1/4 that of the high exchanger. It was possible to maintain a gradient for 12 to 24 hours by periodic adjustment of temperature and flow rates of each heat exchanger, Temperatures were recorded along the gradient with 6 equally spaced YSI Type 4 thermistors calibrated to 0.2° C. Over the range of temperatures that were tolerated by G r y l l o b l a t t a , the regulation of r e l a t i v e humidity was the most d i f f i c u l t operation of the chamber. It was not hard to e s t a b l i s h and maintain a gradient of humidity of 10 to 15 per cent within the range of 40 to 80 per cent r e l a t i v e humidity at a temperature gradient above -j- 10° C. Below + 10° C., when the vapor pressure approached saturation (humidities of 95 per cent and higher), at the dew point temperature and below condensation, fog and rhime formation caused rapid s h i f t s i n the r e l a t i v e humidity i n the closed system. Mixtures of glycerol-water were found to be preferable to s a l t solutions i n establishing humidity gradients i n t h i s chamber. Glycerol-water mixtures maintained the de-si r e d humidity over the longer periods of time and were r e l a t i v e l y non-toxic to G r y l l o b l a t t a and the humidity sensors were not seriously affected by these chemicals. The concentrations of g l y c e r o l to water for a given humidity were found by t r i a l and er r o r . Data available regarding 51 concentrations of glycerol-water mixtures for a s p e c i f i c humidity were based on an a i r volume of 1 l i t e r , a very large surface-to-air r a t i o (Stokes and Robinson, 1 9 4 9 ; Winston and Bates, i 9 6 0 ) . For humidity gradients from 90 to 99 per cent r e l a -t i v e humidity, the bottom of the chamber was l i n e d with i - i n c h thick sponge rubber and saturated with d i s t i l l e d water. A gradient of 90 to 99 per cent was possible with the arrangement over a temperature range of - 5 to f 1 0 ° C. for periods of 2 hours. Lower humidity gradients of 20 per cent were possible for periods of four hours, using glycerol-water mixtures. Relative humidity was recorded In the chamber using 3 equally spaced narrow range hygro-sensors (Hygrodynamics Co., type H-3) i n d i v i d u a l l y c a l i -brated to - 1.5 per cent r e l a t i v e humidity from - 10 to + 60° C. B. Test Methods Specimens of adult G r y l l o b l a t t a campodelformis  campodelf ormis Walker were collected from Athabaska, Jasper National Park, Alberta, at an elevation of 7100 feet. Specimens of G r y l l o b l a t t a lava cola were obtained from McKenzie Pass, Cascade Mountains, Oregon. The specimens were transported to the laboratory i n i n d i v i d u a l containers at 95 + per cent humidity and at 0 to 4 4° C. They were then held under the same conditions i n environmental cabinets (Controlled Environment Ltd.) u n t i l needed and 52 also between the test periods. Specimens were tested a) within seven days of capture, and b) over a period of four months, for possible tempera-ture a c c l i m a t i z a t i o n . Individuals were exposed to the following temperature gradients: l ) - 8 to 4 5° C., 2) - 5 to + 10° C., 3) + 5 to 4 20° C. Each temperature gradient was tested with the following humidity gradients: 1) 50 to 70 per cent r e l a t i v e humidity, 2) 70 to 90 per cent r e l a t i v e humidity, 3) 90 to 99 per cent r e l a t i v e humidity. Each insect was exposed only to one combination per day and returned to the holding humidity and tempera-ture In the laboratory environmental chamber. For the experiments, the insects were placed i n the center of the test chamber, which i n most t r i a l s was the mid-point of the temperature gradient being tested. Since G r y l l o b l a t t a become very agitated and hyperactive upon handling, a ten minute period was allowed before recording position i n the gradient. During t h i s period the insect would rapidly explore the entire length of the test chamber regardless of the temperature-humidity gradient i t encoun-tered. The insect's p o s i t i o n i n the gradient was recorded every 10 minutes for 30 minutes. The insect was then made to move from the l a s t zone recorded and then i t s p o s i t i o n was recorded again a f t e r 10 minutes. These time periods were chosen for i t was observed that the insects would po s i t i o n themselves i n the most tolerable portion of the gradient within 10 to 20 minutes and then stay within that 53 narrow range. In an attempt to determine whether fatigue and/or habituation were influencing the i n i t i a l r est point, the insects were made to make another choice a f t e r 30 minutes. I I I . Results A t o t a l of 50 adult G r y l l o b l a t t a were tested i n each gradient of temperature-humidity. Twenty females and f i v e males were used of G. lava cola and twenty-two females and three males of G. _c. campodeiformls. At a temperature-humidity gradient of - 8 to + 5° C. and 90 to 99 per cent r e l a t i v e humidity, no ind i v i d u a l s preferred a temperature below - 3.5° C. Over the tempera-ture range of - 3.5 to 4 5° C., an approximately normal d i s t r i b u t i o n curve was recorded. Thirty-eight insects preferred a temperature range between 0 and + 2.22° C., with sixteen preferring about 1.67° C. Testing at the same temperature gradient and a humidity gradient between 70.and 90 per cent, the d i s t r i b u t i o n was compressed between - 2.22 to 4 4.50° C., with t h i r t y i n d i v i d u a l s preferring between 0 and + 1.67° C. At a humidity gradient of 50 to 70 per cent, only a narrow temperature range was chosen (between - 1.11 and + 1.67° C. ) by a l l indivi d u a l s ( F i g . 6). In a test temperature gradient of - 5 to 4 10° C. and at the above three humidity gradients, the insects showed very l i t t l e change from that exhibited at - 8 to 4 5° C. One female specimen of G. lava cola preferred a temperature 54 Figure 6. Temperature response of adult G r y l l o b l a t t a  campodei formi s and G. lava cola to various controlled r e l a t i v e humidity ranges. F i f t y t r i a l s were used for each range. 55 around 4 7° C. Over the other humidity gradient of 70^ to 90 and 50 to 70 per cent, no s t a t i s t i c a l changes were found. During t e s t i n g i n a temperature gradient of 4 5 to - 20° C., a l l i n d i v i d u a l s were recorded within + 5 to 4 6° C., regardless of the humidity gradient. Figure 6 i s a summary of t o t a l individuals tested at the various gradients. No ind i v i d u a l s were exposed to humidities below 50 per cent. I observed, both i n the f i e l d and i n the laboratory, that i n d i v i d u a l s exposed to lower r e l a t i v e humidities, even for very short periods, underwent rapid dessication and a 50 per cent mo r t a l i t y . With the sole exception of the female G. lava cola that chose a temperature of about 4 7° C., no difference was detectable between the two species or between sexes. G r y l l o b l a t t a campodelformls campodeiformls specimens were collected 685 miles north of G. lavacola and 2000 feet higher i n el e v a t i o n . One would possibly expect some differences i n temperature and humidity tolerances between these populations, yet test r e s u l t s were i d e n t i c a l . IV. Discussion and Summary Laboratory temperature preference data many times do not represent the actual habitat preferences of the organism. The apparent discrepancies of some reports are undoubtedly due to acclimation to pre-experimental conditions, wide range of preference, seasonal responses, n u t r i t i o n , age, 56 varying populations and species preferences, inadequate test chamber or methods, no control of humidity, and lack of f i e l d data for comparison. M i l l s and Pepper (1937), for example, did not take humidity into account i n t h e i r temperature gradient. An observation chamber in which one can control simultaneously gradients of temperature and humidity does present some problems of design and operation. However, the design I used functioned admirably over the ranges tested, that i s the ranges found i n the natural habitat. It has also been used with success by others testing amphibia and various orders of insects (Licht, unpubl.; Scudder, unpubl.). The parameters of age, sex, n u t r i t i o n and acclimation were c a r e f u l l y controlled i n the present experiments. The temperature preference of G r y l l o b l a t t a i n t h i s study, l i k e that observed i n e a r l i e r studies ( M i l l s and Pepper, 1937; Edwards and Nutting, 1950; Henson, 1957b) i s much lower than that found i n the majority of insects ( M i l l s and Pepper, 1937; Wigglesworth, 1965; Heath et a l , 1971). This i s not surprising i n an insect that i s active throughout the winter at high elevations. Both species of Gry l l o b l a t t a studied at 90 to 99 per cent r e l a t i v e humidity preferred a temperature between - 3 .5 and + 5° C., the optimum being + 1 . 6 7 ° C. These data agree with the temperature preference reported by Henson (1957b) with a saturated atmosphere. At no stage i n the experiments did the preferred temperature range expand s i g n i f i c a n t l y above t 5° C. Thus, when the gradient was + 5 to 4 20° C. and the r e l a t i v e humidities between 90 and 99 per cent, 70 and 90 per cent, or 50 and 70 per cent, the insects preferred the + 5 to t 6° C. area. Since the G r y l l o b l a t t a used by Henson (1957b) were G. c. campodeiformis, and those i n t h i s study were G. c. campodelformis from Athabaska, Alberta, and G. lava cola from McKenzie Pass, Oregon, and since a l l showed a similar temperature preference, i t would seem that the various species and populations of Gr y l l o b l a t t a i n western North America might have s i m i l a r requirements. Although one female G. lava cola showed a preference for + 7° C., t h i s i s s t i l l close to the r e s t of the data obtained. The findings of M i l l s and Pepper (1937) for G. c. campodeiformis from G a l l a t i n Canyon, Montana, d i f f e r s i g n i f i c a n t l y from the present' r e s u l t s . They found the optimum to be + 3.7° C. and the normal range of preference, from + 0.1 to + 15.5° C. While i t i s possible that they were using insects with quite d i f f e r e n t a t t r i b u t e s , i t i s also possible that t h e i r data are i n error because they did not control or evidently attempt to consider the humidity conditions i n t h e i r experimental set-up. This omission i s one that i s frequent i n much of the early work on insect temperature preference, and has been stated to be a factor producing erroneous r e s u l t s (White and Zar, 1968). 58 Edwards and Nutting (1950) stated the preference range i n G. _c. campodelf ormis (material obtained from Pepper) to be - 2.5 to + 11.3° C., since between these temperatures the insects remained quiet i n the respirometers used i n t h e i r experiments. While t h i s range i s somewhat closer to the range obtained i n the present study and by Henson (1957b), i t s t i l l seems rather high. Again, the insects may be d i f f e r e n t to those used herein, and humidity was evidently not taken into account. However, these data were not obtained i n gradient preference experiments and so should not be taken for close comparison with other f i n d i n g s . U n t i l more populations of G r y l l o b l a t t a are studied i n d e t a i l , with f u l l cognisance of the importance of the humidity-temperature i n t e r a c t i o n i n these i n s e c t s , i t i s not possible to state categorically that a l l G r y l l o b l a t t a have the same requirements. Indeed, i n situations where two species coexist, as i n Edison Ice Cave and McKenzie Pass, i t could well be that they d i f f e r i n t h e i r tempera-ture requirements, since one of the factors permitting coexistence i n closely r e l a t e d species can be a difference i n temperature preferences or optima (Heath et a l , 1971; Jamieson, 1973). The most recent data suggest that G r y l l o b l a t t a i n western North America do have s i m i l a r temperature-humidity preferences, and i t would be i n s t r u c t i v e to study coexisting species and other populations, such as those at Kamloops, B r i t i s h Columbia, where they occur 59 at 1400 feet In an area- that i s normally quite hot .and dry. Unfortunately, material from such populations was not avai l a b l e for i n c l u s i o n i n the present research. The preference range of - 3.5 to + 5° C. obtained i n the laboratory seems to coincide almost exactly with the range of temperatures normally encountered i n the hypo-l i t h i o n and cavernicolous microhabitats that G r y l l o b l a t t a occupy. The f i e l d data showed the hypolithion to remain over the year between - 3 and + 6° C. with a humidity over 70 per cent, and the cavernicolous habitats, when occupied i n the spring, summer and f a l l , had the same range. Chap-man (1953), from f i e l d c o l l e c t i o n s , considered that the optimum was + 1.1° C. and Kamp (1963) has previously reported the range as between - 1.8 and +• 7.22° C. i n the f i e l d . The figures given by Campbell (1949), namely - 9.44 to + 15.56° C., would appear to be too wide and perhaps based on general environmental temperatures rather than on the microenvironment i n which the insects l i v e . F i n a l l y of note i s the narrowing of the preference temperature range as the humidity decreases. This same phenomenon has been found i n various species of woodlice (Tracheoniscus) (White and Zar, 1968) and i s evidently a response that can lead to a reduction i n water loss by evaporation. The integument of G r y l l o b l a t t a i s a t h i n ' c u t i c l e with' very extensive membraneous areas. F i e l d and laboratory observations confirm that G r y l l o b l a t t a i s 60 extremely susceptible to dehydration,, even over the preferred temperature range, when the humidity f a l l s below 70 per cent. While warmer a i r can hold more water vapor than colder a i r , the vapor pressure i s also higher for warmer a i r . For example, maximum vapor pressure (saturation) at + 5° C. i s 6.54 mm. Hg. and i s 4.58 mm. Hg. at 0° C. (Landsberg, 1962). When the a i r i s lower than saturation, the d i f f u s i o n grad-ient would be from the insect to the a i r and the organism would lose water. G r y l l o b l a t t a , by moving to a lower temperature when exposed to humidities below 70 per cent, decreases the vapor pressure d e f i c i t (saturation d e f i c i t ) and an equilibrium r e l a t i v e humidity condition i s found higher than the atmospheric humidity around the insect by the process of vapor pressure s t a b i l i z a t i o n ( S e l l e r s , 1967). For conservation of water at low atmospheric humidities, the saturation d e f i c i t at a given temperature i s of greater importance to sur v i v a l than r e l a t i v e humidity alone. Therefore, G r y l l o b l a t t a decreases the range of temperature preference as humidity decreases and t h i s changes the saturation d e f i c i t which reduces water loss from the body of the insect. 61 Temperature Tolerance and Lethal Limits of G r y l l o b l a t t a I. Introduction The temperature-humidity experiments showed that two species of G r y l l o b l a t t a have a temperature preference between about - 4 and + 7° C. and a humidity preference of greater than 90 per cent. These experiments, which allowed free choice of temperature and humidity, do not demonstrate the f u l l range of temperature i n which G r y l l o -b l a t t a can survive. Previous studies have set the upper l e t h a l l i m i t i n G. c. campodeiformis as + 27.8 _ 1° C., and the lower l i m i t as - 6.2° C. ( M i l l s and Pepper, 1937). However, Edwards and Nutting (1950) set the upper l i m i t at + 20.5° C. and Campbell (1949) states that they freeze at - 3.5° C. ( + 26° F . ) . There i s obviously some disagreement. Since i t has been, suggested that the temperature preferences reported i n the l i t e r a t u r e may be i n e r r o r , the extremes of temperature survival need v e r i f i c a t i o n . This i s important because the extremes of high or low temperature may govern t h e i r d i s t r i b u t i o n to a considerable degree. I I . Materials and Methods In an attempt to e s t a b l i s h parameters of tolerance 62 to temperatures and l e t h a l l i m i t s , the following procedure was employed. The test equipment u t i l i z e d a Gebruder-Haake constant temperature c i r c u l a t o r and a small transparent chamber immersed i n the c i r c u l a t o r bath. The c i r c u l a t o r was con-t r o l l e d by a variable thermoregulator accurate to 0.05° C. and operated over a range of - 35 to •+ 100° C. A saturated solution of NaCl was used i n the bath as the heat exchanger The test chamber for the insects was constructed of 25 mil milar p l a s t i c 1^ x 3 x 5 inches and was sealed to maintain humidity. This chamber was l i n e d with i - i n c h sponge rubber on the bottom and 1-inch on a l l sides. The sponge rubber, when saturated with d i s t i l l e d water, pro-vided a humidity of 95 Pe r cent and also insulated the insects from contact with temperatures d i f f e r i n g from the chamber atmosphere. A YSI thermistor, calibrated to 0.05° C from - 10 to -v 25° C. with a 10-second response time, monitored the chamber a i r temperature. The G r y l l o b l a t t a were chosen at random from the series used i n the preference experiments. I believe the choice t be v a l i d for I found no change i n response, at l e a s t over the range tested, to have taken place during the four-month test period. Prior to a given test run, the atmospheric temperature of the immersed test chamber was adjusted to the same temperature as the housing container for the insect ( •+ 2 to + 4° C.). The insect was allowed to explore and to become adjusted to the chamber for 10 minutes a f t e r being placed i n i t . The atmospheric temperature of the chamber was changed at a constant rate of 1° C. per 2.5 minutes. Temperatures and reactions of the insect were recorded every minute. "Normal a c t i v i t y " of G r y l l o b l a t t a was recorded when the insect exhibited the following behavior: body s l i g h t l y elevated above substrate, slow measured walking, antennae moving and touching some object as i f exploring surrounding cerci straight or with s l i g h t downward curve. "Decreased a c t i v i t y " was t y p i f i e d by: sternum on substrate, antennae limp or not moving, cer c i relaxed. "Increased a c t i v i t y " was defined as: rapid non-directional movement over sub-s t r a t e , sternum greatly elevated, rapid waving of antennae. "Hyperactivity" was defined as being an i n t e n s i f i e d pattern of the increased a c t i v i t y with an upward arching of the entire abdomen. Paralysis was shown by: l i t t l e or no response to an external stimulus, entire sternal region on substrate, legs l a t e r a l l y extended. Spasm or tetany was defined as an uncontrolled jerking and twitching, of legs and antennae. Death was preceded by a tetany i n which the insect assumed almost normal posture and then slowly f e l l over on the pleural region, with r i g i d legs and no recovery i f removed from that temperature. I I I . Results Under the test parameters as the temperature increased the insects exhibited normal a c t i v i t y u n t i l a temperature of + 6.5 - 0.2° C. was reached. From + 6.8 to + 11° C. 0.1° C, the a c t i v i t y was reduced. Increased a c t i v i t y began at + 11.5 ± 0.3° C. and progressively i n t e n s i f i e d when i t became hyperactive at + 16° C. The insects were hyperactive over the temperature range of + 16 to 4 21.0 - 0.5° C. At 4 22° C. the insects were paralyzed and death took place between 4 23.0° C. and 4 23.3° C. ( F i g . 7 ) . Shortage of specimens prevented extensive study of short term acclimation i n these i n s e c t s . The insects exposed to decreasing temperatures remained normally active to - 2.2° C. A c t i v i t y was reduced between - 2.2 and - 4° C. £ 0.5° C. The insects become increasingly active during the period of temperature between - 4.0 and - 5.5° C. i 0.5° C. The insects experienced a sudden para l y s i s at - 5.5° C. and a tetany and spasms between - 6.5 and - 7° C. with death following between - 7.5 and - 8.0° C. Insects were also exposed to various tempera-tures for long term survival i n the zone of f a t a l high temperature. The r e s u l t s are presented i n Table I and Figure 7. IV. Discussion and Summary Sudden changes of climatic condition i n the alpine-subalpine are common. A thirty-degree change i n tempera-ture within an hour i s not unusual and the humidity can also change as r a p i d l y . Hence, G r y l l o b l a t t a can be exposed to temperatures and humidities beyond the preferenda 65 Figure 7. Temperature tolerance and l e t h a l l i m i t s of adult G r y l l o b l a t t a campodelformis and G. lava cola when exposed to 1° C. temperature change each 2-g- minutes. Solid bar = length of t o l e r a b l e exposure; oblique hatched bar = occurrence of 50 per cent mortality; narrow bar = longevity from 50 to 100 per cent mortality; v e r t i c a l hatched bar = sur v i v a l through test period. Alphabetical l e t t e r s indicate the a c t i v i t y behavior and temperature range shown i n Table 1. 66 TABLE I Temperature Tolerance and Limits of G r y l l o b l a t t a Tolerance and Lethal Limits* ( 1° C. shift/2.5 minutes) A Normal a c t i v i t y -2.2 to -6.8° C. Lonpc Term Exposure (mortality i n per cent) 4 10° C. for 12 hours - 50$ for 24 hours - 100% B~ Decreased a c t i v i t y -2.2 to - 4° C. + 16° C. for 4 hours - 50% for 7 hours - 100$ C" Increased a c t i v i t y - 4 to - 5.5° C. 4 18° C. for 1| hours - 50$ for 2k hours - 100$ D~ Paralysis - 5.5 to - 6° C, 4 20° C. for 40 min. - 50$ for 2 hours - 100$ E~ Spasm tetany - 6.5 to - 7° C 4 22u C. for 1 min. - 75$ for 10 min. - 100$ F~ Death - 7.5 to - 8° C. 4 23.5° C.for 1 min. - 100$ B Decreased a c t i v i t y 4 6.8 to 4 11 C ^Alphabetical l e t t e r s r e f e r to those i n Figure 7. 67 C Increased a c t i v i t y 4 11 to 4 16° C. D Hyperactivity 4 16 to +. 21° C. E Paralysis 4 22° C. F Death 4 23 to 4 24° C. 68 should they venture out from th e i r regular microhabitats. This they do quite often for they come out, for example, to forage on snow f i e l d s or on g l a c i e r s at night, where they can capture prey numbed by the cold (Ford, 1926). They are then often caught by a sudden change i n tempera-ture and p e r i s h . G r y l l o b l a t t a can maintain a c t i v i t y over a wider range of temperature than the preferred range. In humidities of 95 + per cent the range of a c t i v i t y was between - 4 and 4- 11° C. i n contrast to the preferred choice of - 3.5 to + 5° C. These r e s u l t s are i n agreement with the data obtained by Edwards and Nutting (1950). A c t i v i t y was reduced at the extremes, but G r y l l o b l a t t a can survive at + 10° C. and at - 4° C. for at least six hours without mor t a l i t y . A c t i v i t y increased between + 11 and + 16° C. and also between - 4 and - 5.5° C. This increased a c t i v i t y exhibited i n the small test chamber was undoubtedly a searching mechanism for a more tolerable temperature. There i s evidence for th i s hypothesis i n that an exposure of four hours at + 16° C. caused 50 per cent mortality and increased to 100 per cent within seven hours. The time of exposure to - 5.5° C. that caused mortality was shorter than the higher temperature with 50 per cent mortality within one hour. Paralysis from exposures to below-zero temperatures i s r e v e r s i b l e i f the insect can reach - 4° C. within f i v e 69 minutes. G r y l l o b l a t t a . paralyzed by exposure to high temperatures ( i e . + 22° C.),cannot recover and a one-minute exposure i s l e t h a l to most i n d i v i d u a l s . Thus, the short term su r v i v a l data suggest an upper l i m i t of about + 23° C. and a lower l i m i t of - 8° C. These r e s u l t s are not greatly d i f f e r e n t from those obtained by M i l l s and Pepper (1937), although t h e i r upper l i m i t would appear to be too high. The r e s u l t s i n the present thesis were obtained with a 95 per cent r e l a t i v e humidity; the humidity i n the M i l l s and Pepper (1937) experiments i s unknown. The upper l i m i t of + 20.5° C., stated by Edwards and Nutting (1950),was based on a l^-hour exposure period and so cannot be compared with the short term exposure r e s u l t s of the present study. However, the long term exposure data obtained i n the present research agree almost exactly with the above figure, for at •+ 20° C. G r y l l o b l a t t a survived only for about two hours. There would thus seem to be no great discrepancy i n the general temperature extremes said to be tolerated by these i n s e c t s : the experimental methods each time must be taken into account. These data indicate that G r y l l o b l a t t a i s a c r y o p h i l i c form with a narrow range of temperature and humidity tolerance. The fact that the hemolymph has a freezing point depression of 0.98° C. (unpublished data) bears t h i s out. However, the depression i s not as great as i n such cold-hardy insects as 3ra con cephl (Salt, 1959). Further, 70 i t i s of i n t e r e s t to note that, while G r y l l o b l a t t a shows a r e s p i r a t i o n maximum at + 20° C., there i s no metabolic adjustment to temperature i n t h i s Insect. The slope of the temperature-metabolism curve i s the same as for Thermobia domestica which has a temperature optimum of + 37.5° C. (Edwards and Nutting, 1950). 71 I I I . SYSTEMATICA, DISTRIBUTION AND ZOOGEOGRAPHY Introduction Knowledge of the family G r y l l o b l a t t i d a e has grown slowly since the o r i g i n a l d e s c r i p t i o n . I t now contains three genera, namely, G r y l l o b l a t t a Walker,;19l4, Gryllo-b l a t t i n a Bei-Bienko,1951, and Galloisiana Caudell,1924, and f i f t e e n species. The early described forms, such as G r y l l o b l a t t a  campodelformis campodelformis Walker 919l4, G r y l l o b l a t t a  barberi Caudell^1924, Galloisiana n o t a b i l i s S i l v e r t r i t 1 9 2 7 , G r y l l o b l a t t a campodeiformis occidentalis S i l v e s t r i , 1 9 3 1 , and G r y l l o b l a t t a s c u l l e n l Gurney >1937» were based on immature stages or adult females as type material. Such species and d e s c r i p t i o n s , based on a few females or immature specimens, i s not surprising since G r y l l o b l a t t i d a e have long been considered ' among the rarest of the i n s e c t s . The description of species and subspecies from female and immature specimens has caused some confusion regarding the v a l i d i t y of some populations, for topotype adults and es p e c i a l l y adult males are s t i l l unknown for some. Immature stages of most species are exceedingly d i f f i c u l t , i f not Impossible, to d i f f e r e n t i a t e from each other. In a d d i t i o n , adult females possess few taxonomic characters of discrimina-tory value at the species l e v e l . It has been increasingly 72 evident that taxonomy and systematics must be based on males since the male g e n i t a l i a are of great importance having many s p e c i f i c characters (Gurney, 1961; Kamp, 1 9 6 3 ) . The known d i s t r i b u t i o n of Gry l l o b l a t t i d a e suggests that they are l i m i t e d to the eastern Palaearctic and western Nearctic faunal regions of the world. A more l i m i t i n g descriptive faunal region would be the Cordilleran complex of the P a c i f i c rim. It i s , of course, not impossible that G r y l l o b l a t t i d a e may eventually be found i n some other parts of the world. The discovery of Gry l l o b l a t t i d a e elsewhere would be surpri s i n g , since there has been widespread i n t e r e s t i n the group, and many competent entomologists have searched most areas of the world f o r these insects. Mani ( 1 9 6 8 ) and his associates have found no evidence of the group i n the Co r d i l l e r a of the Himalaya-Pamir complex. Bei-Bienko ( 1 9 5 1 ) and Sharov ( 1 9 6 8 ) have l i t e r a l l y combed the U. S. S. R. without finding a single population except for one form found i n the Maritime-Siberian region of the P a c i f i c . I and many others have searched the alps and ice. caves of Europe and the a r c t i c of North America and others have looked during expeditions i n South America without finding a single specimen. In addition, the fact that no insect at a l l nearly related to the group has ever been found i s evidence of the r e s t r i c t e d and scattered d i s t r i -bution, and i s suggestive of the fact that these insects 73 must be the l a s t survivors of an e c o l o g i c a l l y highly s p e c i a l i z e d nearly extinct group. The separation of the Palaearctic and Nearctic genera was probably not a very recent event (Gurney, I 9 6 I ) . There has been intermittent land connection between North America and Asia throughout the geological ages (Simpson, 1 9 ^ 7 ) and i n the absence of f o s s i l evidence, i t i s impossible to say at what time the range of the family was continuous. I f the family i s confined to mountainous hypolithion and lava ice cave habitat, as the present evidence indicates, and i s not present i n the regions of continuous permafrost of the a r c t i c or subarctic low elevations, i t i s somewhat Improbable that the group crossed the land bridges connecting Alaska and Siberia at various times during the Pliocene and Pleistocene epochs. Before t h i s study knowledge of the family showed that there were nine species of G r y l l o b l a t t a i n the western Nearc-t i c C o r d i l l e r a , a single species, Gryllobla t t i n a d.jakonovi, i n the Primuryo T e r r i t o r y , 90 miles east of Vladivostok, Mariames, S i b e r i a , U. S. S. R., and six species of G a l l o i s i a n a on the Japan Archipelago. Since the Grylloblattodea are unknown i n the f o s s i l record, any attempt to decipher the d i s t r i b u t i o n a l history of the genus G r y l l o b l a t t a cannot be based on paleontological data. The present d i s t r i b u t i o n and zoogeography 'can only be understood by reference to the c l i m a t i c , geographic 74 and geological changes that have taken place i n the past. Since i t has been noted i n the previous section of the thesis that t h i s requires very d e t a i l e d study of these factors for each population and since I have had f i r s t -hand experience only i n western North America, only the Nearctic d i s t r i b u t i o n can be adequately considered herein. Materials and Methods The d i s t r i b u t i o n of a l l populations of G r y l l o b l a t t a i n western North America was mapped. Material from as many these l o c a l i t i e s as possible was then -obtained and studied taxonomically. Further c o l l e c t i n g then i n these same l o c a l i t i e s and at l i k e l y intermediate stations was then undertaken. From these studies, the taxonomy of the Nearctic G r y l l o b l a t t a was c l a r i f i e d and the possible systematic r e l a t i o n s h i p s were established. Detailed d i s t r i b u t i o n s for each species could then be determined. Results The Nearctic Species Fi f t e e n species and f i v e subspecies have been recog-nized i n the present research (Table II ). From morpho-l o g i c a l features, such as s i z e , number of antennal segments, TABLE II L i s t of Species and Subspecies of G r y l l o b l a t t a G r y l l o b l a t t a campodeiform!s campodeiforrols Walker 1914 Gry l l o b l a t t a campodeiformls athapa ska ssp.n. G r y l l o b l a t t a campodeiformis nahannl ssp.n. G r y l l o b l a t t a scudderi sp.n. G r y l l o b l a t t a o c c i d e n t a l i s S i l v e s t r i 1931 Gry l l o b l a t t a skagitensis sp.n. G r y l l o b l a t t a c hi rug ica Gurney I96L. G r y l l o b l a t t a hoodalles sp.n. G r y l l o b l a t t a s c u l l e n l s c u l l e n i Gurney 1937 Gry l l o b l a t t a s c u l l e n i cryocola ssp.n. G r y l l o b l a t t a lava cola sp.n. G r y l l o b l a t t a paulinai sp.n. G r y l l o b l a t t a r o t h i Gurney 1953 Gry l l o b l a t t a gurneyi Kamp 1963 Gry l l o b l a t t a chandleri Kamp 1963 Gry l l o b l a t t a barberl Caudell 1924 Gry l l o b l a t t a washoa Gurney I 9 6 I G r y l l o b l a t t a b i f r a c t r i l e c t a Gurney 1953 76 shape of pronotum, length and shape of ovipositor, and symmetry of male g e n i t a l i a , the taxa can he separated in t o groups as follows: 1) G. occidentalls and G. scudderi, 2) G. skagitensis and the Mt. Rainier population, 3) G. c h i r u g i c a , A) G. hoodalles and G. r o t h i , 5) G. paulina, G. lava cola and the Mary's Peak population, 6) G. s c u l l e n l  s c u l l e n i and G. _s. cryocola . 7) G. gurneyi f G. barberi f G. chandler!. Mt. Shasta, Mt. Elwell and Sierra Butte populations, 8) G. washoa, G. b i f r a c t r i l e c t a , Yosemite, Convict Basin, and Sequoia populations, and 9) G. campodei-formis campodelformis, G. c. athapa ska and G. c. nahannl. The Nearctic D i s t r i b u t i o n The necessity of a specialized habitat has produced a scattered d i s t r i b u t i o n of the Nearctic G r y l l o b l a t t i d a e . l i m i t e d to western North America. Gryllobla tta are usually found above 5000 feet elevation i n the Rocky Mountain and Cascade Co r d i l l e r a of Canada and the United States and i n the Cascade and Sierra Nevada Cor d i l l e r a and i t s assoc-iated plateaus of V/ashington, Oregon and C a l i f o r n i a (Fig. 8 ) . D i s t r i b u t i o n data for Western Hemisphere G r y l l o b l a t t a points to the presence of three divergent groups char-acterized by i s o l a t e d endemic populations or species. 77 Figure 8. 5000-foot elevation contour of the C o r d i l l e r a of western North America i s indicated by stip p l e d areas. Solid c i r c l e s = h y p o l i t h i c l o c a l i t i e s of G r y l l o b l a t t a ; s o l i d t r i a n g l e s = cavernicolous l o c a l i t i e s of G r y l l o b l a t t a . 78 The genus i s d i s t r i b u t e d from the Upper Convict Basin i n the Sierra Nevada of C a l i f o r n i a , along the major ranges to the Cassiar Range on the Yukon-British Columbia border, and north from Yellowstone National Park, Wyoming, to the end of the Canadian Rocky Mountains (Fig. 9). -Individual populations or species d i s t r i b u t i o n may be very l i m i t e d . The d i s t r i b u t i o n may be as small as a single cave system or one side of a mountain peak or canyon, or may be large and occupy a chain along a mountain range. One species group i s found i n the Rocky Mountain C o r d i l l e r a of Canada and Montana, northern Wyoming and eastern Idaho. The Coast-Cascade C o r d i l l e r a group of species extends from Mt. G a r i b a l d i , B r i t i s h Columbia, south to Crater Lake, Oregon, and east-southeast into the Modoc Plateau of the Basin Range province i n C a l i f o r n i a . This group occupies rocks mainly composed of various types of b a s a l t s . The ranges are predominantly composed of volcanlcs of l a t e Tertiary (Miocene and Pliocene) and Pleistocene age (Peck, I960). The t h i r d group of species occurs in the Sierra Nevada of C a l i f o r n i a from Mt. Elwell i n the north to the Sequoia National Park in the southern portion of the range. This group of species occurs c h i e f l y i n substrate of g r a n i t i c o r i g i n and the present range i s from diastrophism of upper Miocene to present (King, 1958, 1 9 5 9 ) . Figure 9 . Species d i s t r i b u t i o n of G r y l l o b l a t t a l o c a l i t i e s . Insert A = Oregon Cascade Mountain and high desert; i n s e r t B = Canadian Rocky Mountain l o c a l i t i e s . Key to G r y l l o b l a t t a species and l o c a l i t i e s : 1. G. sp., May's Hole (Cave), Sequoia National Park, C a l i f o r n i a . 2. G. sp., Convict Basin, C a l i f o r n i a 3. G. b i f r a c t r i l e c t a , Sonora Pass, C a l i f o r n i a 4. G. washoa. Echo Summit, C a l i f o r n i a 5. G. sp., Sierra Buttes, C a l i f o r n i a 6. G. sp., Mt. E l w e l l , C a l i f o r n i a 7. G. b a r b e r i . North Fork Feather River, C a l i f o r n i a 8. G. sp., Ice Cave, Plumas Co., C a l i f o r n i a 9. G. sp., Ice Cave, G r i f f i t h Meadows, C a l i f o r n i a 10. G. chandleri. Ice Cave, Eagle Lake, C a l i f o r n i a 11. G. sp., Ice Cave, Siskiyou Co., C a l i f o r n i a 12. G. sp., Mt. Shasta, C a l i f o r n i a 13. G. sp., Ice Cave, Siskiyou Co., C a l i f o r n i a 14. G. sp., Blue Lake, C a l i f o r n i a 15. G. gurneyi, Ice Cave,Lava Beds National Monument, Ca l i f o r n i a 16. G. sp., Mt. Ashland, Oregon 17. G. r o t h i . Crater Lake, Oregon 18. G. sp., Mary's Peak, Coast Range, Oregon 19. G. hoodalles n. sp., Mt. Hood, Oregon 20. G. chirugi c a, Ape Cave, Skamania Co., Washington 21. G. sp., Mt. R a i n i e r , Washington Figure 9 continued 22. G. skagitensls n. sp., Glacier Peak, Washington 23. G. o c c i d e n t a l i s , Mt. Baker, Washington 24. G. o c c i d e n t a l i s . Hanagen Peak, Washington 25. G. sp., Timberline V a l l e y , Manning Park, B r i t i s h Columbia 26. Report from Grouse Mountain, B r i t i s h Columbia (doubt-f u l ) 27. G. scudderi n. sp, Mt. .Garibaldi, B r i t i s h Columbia 28. G. scudderi n. sp. , Wedge Peak, B r i t i s h Columbia 29. Report from Forbidden Plateau, Vancouver Island, B r i t i s h Columbia (doubtful) 30-37. G. campodeiformis campodeiformis 38. G. campodeiformis athapaska n. ssp., Mt. St. Paul, B r i t i s h Columbia _.. ,39. G. campodeiformis nahanni n. ssp., Cassiar, B r i t i s h Columbia 40-54. G. campodeiformis campodeiformis Insert A 1. G. lava cola n. sp., McKenzie Pass, Oregon 2. G. s c u l l e n l s c u l l e n i s t a t . nov., North S i s t e r , Oregon 3. G. s c u l l e n i cryo'cola n. ssp., Edison Ice Cave, Oregon 4. G. r o t h i , Bachelor Butte, Oregon 5. G. sp., Ice Cave, Deschutes Co., Oregon 6 . G. paulinai n. sp., South Ice Cave, Oregon Insert B 1-12. G. campodeiformis campodeiformis 81 I have no evidence that the d i f f e r e n t groups are sel e c t i v e f o r habitats,,in. substrate of different, geologic o r i g i n . It i s most l i k e l y that the d i f f e r e n t groups occupy these ranges because of h i s t o r i c a l reasons and because now these areas are the only tolerable .habitats present. The rock types occupied indicate no more than the geologic o r i g i n of the i n d i v i d u a l ranges. Zoogeography I. The Rocky Mountain Cordilleran Group The entire d i s t r i b u t i o n of the Canadian population of the Rocky Mountain Cord i l l e r a n group l i e s within the area covered by the l a t e Pleistocene Cordilleran Glacier complex ( F l i n t , 1947).. The Co r d i l l e r a n Ice Sheet formed through coalescence of piedmont and valley g l a c i e r s i n the Rocky Mountains and Cascade Range of Canada and flowed south into the United States, extending south as three major lobes to about 47° 30 l a t i t u d e ( F i g . 1 0 ) (Miller, 1958; Williams, 1961; Richmond, 1964). Nunataks projected a few hundred feet above i t . The surface sloped 15 to 50 feet/mile (3 to 10 meters/km) ( F l i n t , 1935) from an a l t i t u d e of over 8500 feet i n central B r i t i s h Columbia to 7300 feet (2200 m) at the Canada-United States border and the B r i t i s h Columbia-Yukon boundary. At the Canada-United States border ice was 82 Figure 10. Maximum extent of Late Pleistocene g l a e l a t i o n i n western North America, modified i n part from F l i n t (1957) and Canadian Geological Survey (1958). Large st i p p l e s = extent of C o r d i l l e r a n ice sheet; v e r t i c a l hatch -extent of Keewatin ice sheet; small stipples = extent of mountain g l a c i a t i o n ; R = possible i c e ^ f r e e r e f u g i a ; s o l i d c i r c l e s = h y p o l i t h i c l o c a l i t i e s of G r y l l o b l a t t a ; s o l i d t r i a n g l e s = cavernicolous l o c a l i t i e s of G r y l l o b l a t t a . ,9®.' 0 * 0 83 3500-5000 feet (1000-15000 m) thick over major v a l l e y s , to an a l t i t u d e of about 2000 feet ( 6 0 0 m) with i t s southern l i m i t s in Montana and Washington ( F l i n t , 1957; Canadian Geological Survey, 1958). Four or fi v e major periods of g l a c i a t i o n separated by i n t e r g l a c i a t i o n s are recognized for the whole Pleistocene i n t h i s region. The l a t e Pleistocene (la s t g l a c i a t i o n ) i s sub-divided into at le a s t three stades, or minor advances, separa-ted by b r i e f interstades (Alden, 1953; Richmond, 1965). Numerous i s o l a t e d populations of Gr y l l o b l a t t a campodei-formis campodeiformis occur along the Cordilleran crest with the type l o c a l i t y being Sulphur Mt., Banff National Park, Alberta ( F i g . 9 B). The known populations scattered along the mountain chain 'might possibly r e f l e c t incomplete sampling of the habitat: there may be many undiscovered populations. There are many areas that are remote and only accessible a f t e r long hikes or a great deal of technical climbing. A scattered d i s t r i b u t i o n a l pattern i s , however, probably the true picture of G. c. campodeiformis in the Rocky Mountain Cordilleran complex. Over the past f i v e years I and others (R. E. Leech, E. R. MacDonald, L. B a r t l e t t and J . Gordon Edwards) have v i s i t e d most of the areas between known l o c a l i -t i e s . The combined e f f o r t s have yielded less than six new populations and the spotty d i s t r i b u t i o n seems to be a f a c t . The range of in d i v i d u a l populations varies from l e s s than 100 yards to approximately §-mile area. In no instance have longer continuous d i s t r i b u t i o n s been discovered. 84 I believe that t h i s "pocket" d i s t r i b u t i o n i s due to l o c a l or micro-topography and i t s e f f e c t s on the micro-climate conditions. Around such centers of populations within a short distance other areas of apparently suitable habitat may be present, yet the insect w i l l not be found i n them. Ecological or geographical.features such as a d i f f e r e n t face of a slope, a g l a c i a l outwash stream, a moraine, f l a t open area of a l p i n e , lack of s o i l between rocks, seem to function as e f f e c t i v e b a r r i e r s to colonization of adjacent suitable habitat. These b a r r i e r s which may be. only a few yards i n breadth seem to be b a r r i e r s to dispersal and colonization to t h i s apterous, photonegative, cryobionic G. c. campodeiformis. The populations appear to be more or l e s s e f f e c t i v e l y Isolated from each other. Such "pocket" d i s t r i b u t i o n i s not unique among the Insecta. A l l the various orders found i n the d i f f e r e n t mountainous regions of the world have forms exhibiting s i m i l a r d i s c o n t i n u i t y of d i s t r i b u t i o n . It reaches i t s highest development i n the Himalayas where 98 per cent of the Dermaptera, 95 per cent of the Coleoptera, and over 80 per cent of the Orthoptera exhibit such I s o l a t i o n (Mani, 1968). Much the same Is o l a t i o n i n very l i m i t e d ranges i s also known for f l i g h t l e s s grasshoppers along the west coast of North America (Cohn and C a n t r a l l , pers. comm.). Populations a few miles apart maintain species and subspecies i n t e g r i t y . 85 I have not been able to obtain any f i e l d evidence for the presence or absence of gene flow between populations of G. _c. campodeiformis occurring within a short distance of one another. Further, the size of populations i s not known; too few specimens are avail a b l e for mark and recapture techniques and the animals' habits are not conducive to sequential samp-l i n g . The state of the "art" of discovering populations and the c o l l e c t i o n s of i n d i v i d u a l s did not 'develop early enough i n th i s project to permit f i e l d studies on movement and in t e r -breeding. It i s now possible to conduct such long term study and I hope to i n s t i g a t e t h i s i n the near future. From geologic evidence the regions currently occupied by the Canadian Rocky G. c. campodeiformls were subjected to successive massive g l a c i a t i o n during the Pleistocene. From t h i s evidence the following questions a r i s e . Are the present-day G r y l l o b l a t t a descendants of pre-Pleistocene inhabitants and did they survive the l a s t Cordilleran g l a c i a t i o n , or are they new a r r i v a l s following the ice recession? Four hypotheses are immediately a v a i l a b l e . Each hypo-thesis has i t s proponents when applied to the present d i s t r i -bution of the various f l o r a and fauna of the Rocky Mountains. (1) The populations have continuously occupied the region during Pleistocene g l a c i a t i o n and the present d i s t r i b u t i o n i s l i t t l e changed. (2) The populations survived the Pleistocene g l a c i a l episodes i n some l o c a l i c e - f r e e refugia and invaded the present habitats as the ice receded. (3) The 86 d i f f e r e n t populations retreated to l o c a l nunataks that projected above the i c e sheet during g l a c i a l periods and l a t e r spread out from them. (A) The fauna retreated north or south beyond the ice sheet borders and the present d i s t r i -bution i s of post Pleistocene origin.;" We can consider each of these i n turn. (1) Continuous habitation of the region by the G. c. campodeiformis i s t o t a l l y i n v a l i d . G r y l l o b l a t t a do occupy the hypolithion under snow f i e l d s and under favorable conditions may feed on the peripheral surface of g l a c i e r s , but long term survival under or on an i ce sheet would be impossible. The movement of g l a c i e r ice abrades to bedrock or below and removes a l l unconsolidated substrate. The action of valley and cirque g l a c i e r . i c e deposits . material as l a t e r a l or terminal moraines. The continual movement and b u i l d i n g of moraines by active g l a c i e r s make the substrate uninhabitable for G r y l l o b l a t t a . While G r y l l o b l a t t a do a c t i v e l y search for food for short periods under favorable conditions on the margins of small g l a c i e r s , continual occupation i s not possible. The circadian temperature and humidity fluctuations are beyond the extremes tolerated by the organisms (see other sections of t h e s i s ) . Nocturnal foraging by G r y l l o b l a t t a on small snow f i e l d s a few hundred feet i n diameter has been noted, but any rapid fluctuations of temperature and humidity can be l e t h a l . I have observed on numerous occasions while c o l l e c t i n g on snow f i e l d s at night that a few degree f a l l i n temperature or s l i g h t wind movement w i l l cause the G r y l l o b l a t t a to retreat 87 within 15-20 minutes to the hypolithion. During such condi-tions i n d i v i d u a l s far from the snow f i e l d margins become s t u p i f i e d and d i e . It i s usually possible on large snow f i e l d s where G r y l l o b l a t t a forage to find a few dead i n d i v i d u a l s next morning that did not safely retreat to the hypolithion. I f present free a i r temperature gradients can be extra-polated to paleoclimates the mean summer temperatures during the l a t e Pleistocene g l a c i a l maxima were up to 9° C. (17.5° F.) cooler than the present temperature of the region (Heusser,1964; Richmond, 1965). Since the present mean for the region at 5000 feet i s approximately 2° C. (35.6° F.) (Meteorological D i v i s i o n , Department of Transport, 1970) continual g l a c i e r surface survival by G r y l l o b l a t t a seems highly improbable. It can be hypothesized that l a t e Pleistocene G r y l l o b l a t t a had a lower temperature-humidity tolerance than the present forms and could survive g l a c i a l maxima temperatures. It seems most l i k e l y that i f such a lowered tolerance were present during the Pleistocene, natural selection would have been for retention of t h i s a ttribute rather than against i t , fo r i n the present habitat occupied by these insects i t would s t i l l be an advantageous physiological feature. (2) Late Pleistocene g l a c i a t i o n survival i n i c e - f r e e refugia i s a p o s s i b i l i t y for at le a s t a few of the Canadian Rocky Mountain G r y l l o b l a t t a . However, survival i n ice refugia close to or matching the present d i s t r i b u t i o n Is highly improbable. The Keewatin and Cordilleran ice sheets are generally considered to have been at times in continuous contact throughout most of the region east of the Cordilleran crest. Extensive reworking of t i l l s , outwashes, and moraines has made i t d i f f i c u l t to determine the l i m i t s and absolute zones of contact. There i s some evidence for at least two and possibly as many as six refugia l y i n g between the two ice sheet complexes (Canadian Geological Association, 1958). A small refugium i s thought to have existed during the l a t e Pleistocene i n the Pincher Creek-Macleod (Porcupine H i l l s ) , A l b e r t a , region (Canadian Geological Association, 1958) ( F i g . Southeast of Edmonton i n the Buffalo Lake Region, Bretz (1943) has mapped end moraines whose arrangement suggests that a deglaelated zone existed between the C o r d i l l -eran and Keewatin ice sheets. Halllday and Brown (1943), i n t h e i r study of the present d i s t r i b u t i o n of pine trees i n Canada, speculate that t h i s region could have Deen a refugium for the western pine species. Hansen (1949), from analysis of bog pollen p r o f i l e s , has also recognized the p o s s i b i l i t y that pre-late Pleistocene forests existed i n an ice- f r e e b e l t i n the same area. The Fernie population of G. _c. campodeiformis may possibly have inhabited the Pincher Creek-Macleod (Porcupine H i l l s ) refugium during late Pleistocene. However, the same species i s known from Jasper National Park, A l b e r t a , to Yellowstone National Park, V/yoming. I suggest that while the Fernie population may have occupied t h i s refugium during l a t e Pleistocene, i t i s unlikely to be the nucleus or parent popula 89 t i o n from which the present t o t a l d i s t r i b u t i o n of the species has been derived. Further, the p o s s i b i l i t y that t h i s Fernie population did not f i n d refuge i n the Pincher Creek-Macleod region must be considered. The region, even i f not g l a c i a t e d , was subjected to extensive outwash of t i l l and s i l t s from both i c e sheets (Horberg, 1952). Present populations of G r y l l  b l a t t a do not inhabit such mixtures of unconsolidated material The area,thought not to have been glaciated, i s l i m i t e d i n size with l i t t l e elevation change and i t i s probable that a suitable habitat did not e x i s t . The present s i t e of the population i s separated from the refugium area by the Flathead River valley which contained an extrusive southern lobe of the Continental Ice Sheet that projected approximately 100 miles into Montana. The ice sheet recessions at the beginning of the post Pleistocene are believed to have been by stagna-t i o n and melting instead of by f r o n t a l retreat (Rice, 1936; Nasmith, 1962). Stagnation melting freed the highland, but ice remained for possibly hundreds of years i n the valleys of the Flathead and S t i l l w a t e r drainages. In l i g h t of the present d i s t r i b u t i o n a l pattern and the systematica of the species, the Fernie population could well be an early post Pleistocene migrant along the Flathead and Whitefish ranges from a southern center of l a t e Pleistocene G. c. campodei-formls . The present d i s t r i b u t i o n pattern of the G. c. campodei-formls does not support l a t e Pleistocene occupation i n the questionably deglaciated Buffalo Lake region. I f t h i s area was the center of l a t e Pleistocene survival 9 0 and the Banff^Jasper populations are migrants from the region, i t would suggest the presence of populations up the only possible migration route, the Saskatchewan River. I t i s true that suitable climate and habitat exist above Nordegg, yet no populations of G. c. campodeiformls have been discovered i n t h i s drainage. Considering l a t e Pleistocene and post Pleistocene geological data, the present d i s t r i b u t i o n pattern, and the systematics of G. c. campodeif orml3, I suggest that the genus did not occupy the Buffalo Lake region. A l a t e Pleistocene habitation i n an ice - f r e e refugium i n the north i s most l i k e l y for the northern B r i t i s h Columbia • species of G r y l l o b l a t t a . The Summit Lake (Lat. 58° 45' N.) subspecies of G r y l l o b l a t t a i s found some 450 miles northwest of the G. _c. campodeiformls populations i n Jasper National Park. In t h i s context i t should be noted that the main Canadian Rocky Mountain Cor d i l l e r a decreases i n elevation to the north and i s interrupted by major r i v e r drainages, such as the Fraser, Parsnip, and especially the Peace River: the extensive i n t e r i o r valleys present major geographical b a r r i e r s . The Cassiar- Mt. McDame subspecies of G. campodeiformls, found on the east slopes of the Cassiar Range, i s presently Isolated from the Summit Lake species. The climatic condi-tions that exist i n the Liard P l a i n , the Dease and L i a r d r i v e r s and t r i b u t a r i e s , and the Rocky Mountains, tend to maintain t h i s i s o l a t i o n . 91 Two large, unglaciated refugia existed west of the Mackenzie River during l a t e Pleistocene (Canadian Geological Association, 1958). Topography and geological evidence suggest that the southern unglaciated area of the Li a r d and Nahanni ranges, extending north to the Canyon Range west of Fort L i a r d and Fort Simpson, was the most l i k e l y refugium for these populations during the late Pleistocene. (Fig. 10). As the i c e sheets receded from the highlands along the L i a r d and Fort Nelson r i v e r s migration routes would have been a v a i l a b l e , permitting refugium populations to reach the present l o c a l i t i e s of Cassiar and Summit Lake. Morpho-l o g i c a l features indicate a close r e l a t i o n s h i p of these populations to G. c. campodeiformls. The topography of the t e r r a i n and the distance separating the northern group from G. _c. campodeiformls predicates against post Pleistocene migration of the northern forms from the south. The close r e l a t i o n s h i p , however, does suggest a p're-l a t e Pleistocene common o r i g i n with a disjunct i s o l a t i o n of the northern group during l a t e Pleistocene i n the refugium . A future discovery of G r y l l o b l a t t a from the L i a r d or Nahanni ranges would substantiate t h i s hypothesis. Based on observa-tions and c o l l e c t i o n s I have made during t r i p s into the Fort Nelson and Peace River v a l l e y , I doubt that G r y l l o b l a t t a w i l l be discovered from the area. (3) Survival during Pleistocene g l a c i a t i o n on nunataks projecting above the ice sheets i s frequently suggested as an explanation of present d i s t r i b u t i o n patterns for the f l o r a and fauna of western Canada (see McCabe and Cowan, 1945; Calder and Savile, 1959; Mathias and Constance, 1959). The present climatic conditions that exist on nunataks above the small ice f i e l d s , Wapta and Columbia (Jasper-Banff National Parks),coupled with f i e l d observations suggest that only those organisms most r e s i s t a n t to extreme cold and dessica-t i o n could survive any extended period of time. The bare rock substrate on such projections i s frozen throughout the year except for a few small areas on the west-southwest face that are sheltered from wind and thaw during July and August. A i r temperature may r i s e a few degrees above 0° C. during these months, but the peaks are buffeted by cold katabalic wind which counteracts the above zero temperatures. During July and August, 1969, I climbed Mt. Columbia, Mt. Athabaska, and the Twin mountains, nunataks above the Columbia Ice F i e l d , and found no animal l i f e and observed only the rare crusteous l i c h e n s . One would assume that present conditions-, on such nunataks are ameliorated from those that existed during massive Cord i l l e r a n and Continental g l a c i a t i o n s . I f l i f e i s marginal today on such areas i t was even more so during the P l e i s t o -cene. Even i f conditions were less severe during the Pleistocene than present, evidence from the narrow range of temperature and humidity tolerated by G r y l l o b l a t t a would indicate again that these insects would not be present. In addition, the widespread d i s t r i b u t i o n of the species l n the Rocky Mountain C o r d i l l e r a does not suggest l a t e Pleistocene s u r v i v a l on nunatak re f u g i a . 1 am i n c l i n e d to believe that 93 those who propose Pleistocene nunatak survival have never studied the nature of the environment there. (4) A l a t e Pleistocene occupation of habitats south of the Cord i l l e r a n and Keewatin i c e fronts i n Montana and Idaho must be considered, with the present d i s t r i b u t i o n being established during or soon a f t e r ice recession (ca. 9 0 0 0 years ago). Nine populations of G. c. campodeiformls are known from l o c a l i t i e s south of the maximum Pleistocene ice advances i n Montana and Wyoming ( F i g . 1 0 ) . Five other popula-tions of the species occur i n regions that had a p e r l g l a c i a l climate on the margins of the Flathead and Lewis lobes of the Cor d i l l e r a n Ice Sheet i n northern Montana and Idaho. The present d i s t r i b u t i o n of the species along the crest of the Rocky Mountain C o r d i l l e r a from Yellowstone, Wyoming, to Jasper, A l b e r t a , and the concentration of present populations south of the maximum ice advance, suggest that Montana was the center of the la t e Pleistocene G. c. campodeiformls d i s t r i b u t i o n . Thus, the present d i s t r i b u t i o n i s a r e s u l t of migrations along i c e - f r e e highlands during g l a c i a l recession some 9000 years ago. I I . Coast-Cascade Group The d i s t r i b u t i o n of Gr y l l o b l a t t i d a e i n the Coast- 5 Cascade C o r d i l l e r a i s r e s t r i c t e d , with few exceptions, to the major stratovolcanoes or ice caves i n plateau lava f i e l d s . Each inhabited peak and cave system has a character-i s t i c species or subspecies usually a l l o p a t r i c with other G r y l l o b l a t t a . They are separated by climatic or geographic b a r r i e r s , such as elevation, r i v e r canyons and lava f i e l d s . Every known species now occurs i n a l o c a l i t y which was within the maximum extent of l a t e Pleistocene g l a c i a t i o n or the p e r i g l a c i a l zone surrounding such g l a c i a t i o n (Figs. 9 A and 10). The Coast-Cascade C o r d i l l e r a , though usually treated as a single physiographic unit, can be considered as f i v e major subdividions. These correlate with the known d i s t r i -bution and systematics of the G r y l l o b l a t t a and show i n d i v i d u a l regional Pleistocene - post Pleistocene geological and cli m a t i c changes. These subdivisions are: 1. The Northern Cascades, extending from Garibaldi-Wedge Peak, B r i t i s h Columbia, south to Mt. St. Helens, Washington; 2, The High Cascades, a narrow b e l t of summit volcanoes beginning with Mt. Hood and including the Three Sisters-Bachelor Peak complex; 3. The Southern Cascades, comprising Crater Lake, Mt. Shasta and Mt. Lassen; 4. Plateau and Basin ranges, l y i n g east of the Cascades and including the Columbia, High Desert and Modoc plateaus; 5. Secondary ranges, including portions of the Coast and Klamath ranges. The Quaternary (Pleistocene-post Pleistocene) history of the C o r d i l l e r a i s exceedingly varied and includes such events as repeated alpine g l a c i a l advances, invasion of the Northern Cascades by the Cordilleran Ice Sheet, catastrophic 95 floods released by glacier-dammed lakes and covering wide expanses, and the bui l d i n g of numerous stratovolcanoes and huge f i s s u r e flows of lava covering hundreds of square miles ( F i g . 11). The Cascade Range i s 100-160 km. wide and extensive areas along the crest are 1200 m. (4000 f t . ) i n elevation. Towering 1000-4000 m. (3000-10,000 ft.) above the average c r e s t l i n e are 12 large stratovolcanoes. Most of these volcanoes l i e west of the crest and form the headwaters of the many major r i v e r s that drain the east and west slopes of the mountains. These major peaks, because of t h e i r elevation above the average c r e s t l i n e and the east-west di s s e c t i o n of the t o t a l range by the major drainage systems, function as evolutionary islands for the G r y l l o b l a t t i d a e . The most extensive record of Pleistocene g l a c i a t i o n i s recognized in Washington, where g l a c i a l strata representing at l e a s t four major g l a c i a t i o n s , are interbedded with non-g l a c i a l deposits. The stratographic evidence for each major g l a c i a t i o n includes records of at least two g l a c i e r fluctuations for each period (Armstrong et a l , 1965; Crandell, 1965). The records of Pleistocene g l a c i a t i o n and cli m a t i c changes are not as well documented for the southern portion of the range. Nevertheless, geological formations indicate Pleistocene g l a c i a t i o n along the entire range. This consisted of the Cord i l l e r a n Ice Sheet, with i t s o r i g i n i n the Coast Mountains of B r i t i s h Columbia'*, invading northern Washington on either side of the Cascades; there were broad valley and piedmont g l a c i e r s flowing down the major drainage with the 96 Figure .11. Quaternary to Recent volcanica of Coast-Cascade Range indicated by stippled areas; s o l i d c i r c l e s = h y p o l i t h i c l o c a l i t i e s of G r y l l o -blatta ; s o l i d t r i a n g l e s = cavernicolous l o c a l i t i e s of G r y l l o b l a t t a . 97 c r e s t l i n e covered by expansive ice f i e l d s and i c e caps. The early and middle Pleistocene i s recorded to have had two major g l a c i a t i o n s , each with two or more advances of the Puget Lobe of the Cordilleran Ice Sheet, extending approximately 170 miles south of the B r i t i s h Columbia-Washington border to the v i c i n i t y of Mt. St. Helens (Crandell, 1963; Snavely et a l , 1958). Each of these major advances was contemporary with the formation of ice f i e l d s at higher elevations. Three g l a c i a l episodes occurred during the l a t e Pleistocene. The l a s t , the Fraser G l a c i a t i o n , occurred between 25,000 and 11,000 years ago. During the e a r l i e r two episodes of the l a t e Pleistocene, large valley and piedmont g l a c i e r s covered the lowland south of the Puget Lobe and were fed by vast ice f i e l d s and ice caps that mantled the entire length of the Cascade Range. Mountain g l a c i a t i o n i s thought to have been of lesser extent during the Fraser period, although ice f i e l d s and ice caps again covered the areas now occupied by the G r y l l o b l a t t i d a e . During the Fraser Glaciation alpine g l a c i e r s in B r i t i s h Columbia formed the Puget Lobe which invaded Washington about 22,000 years ago and reached t h e i r maximum advance 50 miles south of Seattle between 15,000 and 13,000 years B. P. (Crandell, 1963; Heusser, 1964). This g l a c i a l episode, the Vashon Stade, was followed by the Everson nonglacial i n t e r v a l that terminated about 11,500 years ago and had a climate cooler and more moist thanvthe 98 present. A sh o r t - l i v e d Sumas Stade followed with the re-advance of the Cordilleran Ice Sheet and the r e b i r t h of mountain g l a c i e r s and ended with the disappearance of the ice sheet and mountain g l a c i a t i o n approximately 11,000 years B. P. At the time the Puget Lobe reached i t s maximum extent during the Vashon Stade, alpine g l a c i e r s i n the Cascades had greatly decreased in- size or had disappeared (Cary and Carlston, 1937). The recession of alpine g l a c i e r s i n the Cascade Range while the Cordilleran Ice Sheet continued to expand i s thought to have been the e f f e c t of a cli m a t i c change of regional d i s t r i b u t i o n p r i o r to the maximum expansion of the Puget Lobe (Mathews, 1951). Mathews (1951) suggested that t h i s change resulted i n increasing temperature and caused the wasting of the mountain g l a c i e r s . The increase of temperature could have resulted i n greater p r e c i p i t a t i o n on the ice sheet, because i t s size and elevation maintained a l o c a l cold climate on the ice surface and continued to expand. During the Sumas readvance of the Puget Lobe, about 11,500 years ago, there was a temporary return to cold c l i m a t i c conditions i n the mountains and the r e b i r t h of alpine and cirque g l a c i e r s . Sumas mountain g l a c i a t i o n i s indicated by morai-nal and t i l l features l y i n g on nonglacial deposits and covered by two d i s t i n c t i v e volcanic ash layers. The volcanic ash f a l l s are from Glacier Peak, Washington, with a tentative date about 11,000 years B. P. ( F r y x e l l , 1965) and from the widespread Mazama ash from Crater Lake, 99 Oregon, of 6600 years B. P. i n age (Powers and Wilcox, 1964). Two mountain g l a c i e r episodes are known for the post Pleistocene, the older advance during Neoglacial beginning about 3500 to 2000 years ago. During the younger advance, s t a r t i n g at l e a s t 700 years ago, various g l a c i e r s on the major peaks reached maximum stage, ranging from 600 to 100 years B. P. The Cascade g l a c i e r s have since been undergoing an inconsistent pattern of recession and advance and t h i s continued to the present day (Harrison, 1956a, 1956b; Sigafoos and Hendricks, 1961). The analysis of pollens from nonglacial deposits suggests that the i n t e r g l a c i a l climates were at le a s t 2° C. cooler (yearly mean) and were much moister than present (Heusser, 1964). Summer mean temperatures during the very l a t e Pleistocene (Fraser Glaciation) are considered to have been at least 6-9° C. below the present (Mullineaux , Waldron and Rubin, 1965). The present d i s t r i b u t i o n of G r y l l o b l a t t a i n the Coast-Cascade .Cordillera i s a d i r e c t r e s u l t of the various episodes i n the l a t e Pleistocene advances and recessions of mountain crest ice f i e l d s and ice caps. Late Pleistocene centers of G r y l l o b l a t t a survival were below the summit ice sheet and between the valley and piedmont g l a c i e r s fed from the C o r d i l l e r a . 100 1. Northern Cascade Group The G r y l l o b l a t t i d a e occurring i n the northern portion of the C o r d i l l e r a are: G r y l l o b l a t t a scudderi. Wedge Peak, B r i t i s h Columbia; G r y l l o b l a t t a o c c i d e n t a l i s , v i c i n i t y of Mt. Baker, Washington; G r y l l o b l a t t a skagltensls, G l a c i e r Peak, Washington; and G r y l l o b l a t t a chlruglca, Ape Cave, Skamania Co., Washington. In addition, frpm the north and east slopes of Mt. Rainier I discovered G r y l l o b l a t t a i n October, 1969. From the morphological characters of the adult females, I believe t h i s population to be a new species. I have not as yet named i t as such for adult males are s t i l l unknown. There are l i t e r a t u r e reports of G r y l l o b l a t t a species occurring on the Forbidden Plateau, Vancouver Island (Buckell, 1930; Spencer, 1945). Spencer (1945) also reports that the genus has been found on Grouse Mountain, north of Vancouver, B r i t i s h Columbia. A single female i s reported from Garibaldi Peak, B r i t i s h Columbia (Walker, 1937). In the B. C. P r o v i n c i a l Museum c o l l e c t i o n there i s a single adult female G r y l l o b l a t t a from Timberline Valley, Manning Park. The condition of the specimen does not allow compari-son with G r y l l o b l a t t a o c c i d e n t a l i s or G. scudderi. Except for the Garibaldi and Manning Park specimens, there i s reason to believe that these are hearsay records for they have not been substantiated by material. However, I believe that eventually G r y l l o b l a t t a w i l l be discovered 101 from more l o c a l i t i e s i n southwestern B r i t i s h Columbia. The Northern Cascade species of G r y l l o b l a t t a , except for G. chl r u g l c a , occur i n l o c a l i t i e s which were covered by the Cordilleran Ice Sheet, ice f i e l d s and crest ice caps during Fraser G l a c i a t i o n about 25,000 to 10,000 years ago. Ape Cave on the slope of Mt. St. Helens (type l o c a l i t y of G. chlruplca) has been the scene of repeated post Pleistocene l a v a , ash and mud flows. Some of t h i s a c t i v i t y i s as recent as 1802, 1831, 1842 and 1854 (Lawrence, 1954; Heusser, I960). These and e a r l i e r eruptions have reworked or obliterated a l l trace of l a t e Pleistocene g l a c i a t i o n to the extent that only Neoglacial and recent features are known. Gry l l o b l a t t a scudderi occurs' i n the alpine zone about 6000 feet on the north and east slopes of Wedge Peak, Garibaldi Mt., B r i t i s h Columbia. The returns from six d i f f e r e n t c o l l e c t i n g t r i p s by Topping, B a r t l e t t and myself indicate that t h i s i s a very sparse population. Late Pleistocene g l a c i a l evidence indicates that G r y l l o b l a t t a scudderi could not have occurred i n the region p r i o r to the Sumas Stade about 11,500 to 11,000 years B.P. The present population i s undoubtedly from stock that survived the extensive l a t e Pleistocene g l a c i a t i o n south of the ice f r o n t . The very small population would te n t a t i v e l y suggest post Pleistocene migration during the Neoglacial period about 3500 to 2000 years ago from parent stock i n the v i c i n i t y of Mt. Baker. The broad expanse of the Fraser River and delta has been an e f f e c t i v e topogeographical bar r i e r for many species. This 102 major b a r r i e r tends to rule out a p o s t g l a c i a l migration d i r e c t l y from the southern regions of the range, unless Neoglacial climate was much colder than present records i n d i c a t e . A post Pleistocene dispersal route from the south to the present l o c a l i t y i s suggested. The elevations through the Manning Park Cascades, the L i l l o o e t Range, are such that suitable c l i m a t i c conditions must have existed along the coast during Neoglacial times. Any future discovery of G r y l l o b l a t t a populations from either the Grouse Mountain-Mt. Seymour area or the Manning-Lillooet region could greatly contribute to our understanding of the d i s t r i b u t i o n and systematic r e l a t i o n s h i p of G r y l l o b l a t t a scudderi. Irrespective of l a t e or post Pleistocene o r i g i n , the species did not occur i n i t s present l o c a l i t y during the greater portion of the Fraser G l a c i a t i o n . Nunatak sur v i v a l for the genus above the Cordilleran Ice Sheet i s not considered possible for reasons presented e a r l i e r . G r y l l o b l a t t a o c c i d e n t a l i s i s known from Mt. Baker and from surrounding mountains within a v i c i n i t y of 15 miles. Populations of l i m i t e d d i s t r i b u t i o n are known from Mts. Winchester, Goat, Table, Herman, Hannegan, Shuksan and Baker. Morphological examination of a l i m i t e d series indicates that the species i s composed of three subspecies that are Isolated from each other by topographical features. The populations i n the Skagit Range, which occur on Winchester Mountain, S e r f r i t Mountain, Goat Mountain and Hannegan Peak, are found above 5200 feet and form one 103 subspecies group. The Skagit Range i s a small subsidiary chain of the above peaks branching northwesterly o f f the main Cascade c r e s t l i n e . . The populations i n the Skagit Range are topographically i s o l a t e d from other populations i n the region by the Nooksack River flowing to the west and the Chilliwack River draining the eastern slopes. The canyons cut by these r i v e r s and t h e i r t r i b u t a r i e s almost i s o l a t h i s part of the Cascade crest. The elevations of the ridges, low mountains and f o o t h i l l s between the Skagit Range sub-species and the populations i n the Mt. Shuksan complex, average below 3000 feet with the bottom of the r i v e r canyons between 1000-2000 feet. The lower elevations are much too dry and the temperature too high to be tolerated by Gryll o-b l a t t a during most of a yearly season. Suitable hypolithion habitat i s scarce and generally covered by erosional s i l t s and persistent snow does not ex i s t . While the Skagit Range and Mt. Shuksan populations are only 7 miles apart, the cli m a t i c and topogeographical b a r r i e r s seem to be e f f e c t i v e and are.presumed to have been i n existence long enough for morphological d i f f e r e n t i a t i o n to have taken place. The Mt. Shuksan populations occurring on Mt. Herman, Table Mountain and across the Kulshan Ridge to Mt. Shuksan constitute a further subspecies. These populations are morphologically d i s t i n c t from those on the 6000-8000 foot slope of Mt. Baker which are another subspecies. The topography to the south and west of the Shuksan 104 complex i s at an elevation that the climatic conditions most of the year would be a possible e f f e c t i v e b a r r i e r for interchange with the Mt. Baker subspecies. From Table Mountain, l y i n g in. the center of the Shuksan subspecies range, Ptarmigan Ridge leads southwest d i r e c t l y to Mt. Baker. Ptarmigan Ridge has small patches of snow which p e r s i s t throughout the year and the lower portions toward Table Mountain are composed of substrate which appears to be id e a l habitat for G r y l l o b l a t t a . Yet G. occidentalis i s not found anywhere along the 5 miles between Table Mountain and Mt. Baker. The ancestral population of G. occidentalis most l i k e l y survived the late Pleistocene west or southwest of Mt. Baker between mountain ice and the Fraser Lobe. The present Mt. Baker subspecies populations are found only on the western and northwestern facing slopes. Flowing down from the summit ice f i e l d of Mt. Baker to the northeast along the upper por-t i o n of Ptarmigan Ridge are Manning, Rainbow and Park G l a c i e r s . Below the g l a c i e r s the upper portion of the ridge i s composed of e a r l i e r moraines and t i l l which are unconsolidated and sub-ject to avalanche. This loose, moving rock evidently i s not a suitable habitat and acts as a b a r r i e r between the two subspecies. I believe that these three morphologically d i s t i n c t population groups are subspecies, although I have no evidence that gene flow does not occur between them. I t i s possible that during exceptional years of much greater snow f a l l and a long cool spring, the peripheral populations of each sub-105 species may come in contact. A long cool spring preceded by very heavy snow could allow the snow cover to exist across the lower elevations well into the normal summer conditions. Under such c l i m a t i c conditions l o c a l populations could spread out and meet; short distances only separate the three groups. The Mt. Baker G. o c c i d e n t a l i s are presently i s o l a t e d from G. scudderi by the cli m a t i c and major topographical b a r r i e r s previously discussed. They are also separated by 50 miles from G. skapitensis on Glacier Peak. Both of these stratovolcanoes are 20 to 40 miles west of the Cascade crest-l i n e . Most of the topography i n the 50 miles of separation i s below 3000 feet i n elevation and dry during the l a t e spring through f a l l seasons. The wide valleys of the Skagit, Sauk, Stehekin and S u i a t t l e r i v e r s , with eleva-tions below 1500 f e e t , areL the major topographical b a r r i e r s . The heads of the valley and r i v e r drainages lack s u i t a b l e , moist, cool hypolithion. The 50-mile separation with a lack of suitable hypolithion and the present temperature-humidity conditions i n the lowlands would appear to be an e f f e c t i v e b a r r i e r between the two species, i f habitat .and temperature-humidity tolerance i s constant throughout the genus. G r y l l o b l a t t a skagltensls occurs i n the compound cirque basin between Kennedy G l a c i e r , Glacier Ridge and the head-waters of the White Chuck River. The basin between 6000-9000 feet i n elevation contains large permanent snow f i e l d s through the summer and f a l l . The entire basin substrate i s a hypolithion i n material of Neoglacial or younger age. 106 The basin was ice-covered during the Fraser G l a c i a t i o n and the present g l a c i e r s and morainal features are a l l of Neoglacial age (ca. 3500-2000 years B. P.). Glacier Peak experienced a major eruption of pumice and ash near the time of the Sumas Stade ice r e t r e a t . This pumice and ash covered the slopes to a depth of a meter or more and are recorded as f a r east as Montana (Wilcox, 1965). I f alpine g l a c i e r s disappeared about 11,500 years ago, as suggested by Mathews (1951) and Crandell (1963), any hypolithion occu-pied then by G. skagitensls would have been destroyed. The hypolithion inhabited by the insects i s thus l e s s than 3500 years o l d , the maximum age of the current g l a c i e r s . The upper elevations of the basin are presently the s i t e s of much avalanche a c t i v i t y of snow, ice and rock. The sequence of volcanism, Fraser advance and recession, and Neoglacial events suggests recent invasion of the present habitat. The pioneering stock of G. skagltensls certainly1 did not come from the west for that region was either under an i c e sheet or too hot and dry during the l a t e r Hypsithermal Interval (ca. 9000-3500 years ago. ). The only possible areas of survival during Fraser G l a c i a t i o n and Hypsithermal times are to the east or south near Mt. R a i n i e r . Discovery of G r y l l o b l a t t a from the Cascade crest east of Glacier Peak would help to c l a r i f y the o r i g i n of the present populations. Th e discovery of more males and females from Mt. Rainier would certainly contribute to the knowledge of the r e l a t i o n s h i p and phylogeny of G. skagltensla. 107 I speculate that populations of G r y l l o b l a t t a occur above Holden i n the Chilon mountains and i n the eastern Glacier Peak Wilderness Area. Numerous independent t r i p s by Mr. B a r t l e t t and myself, searching for G r y l l o b l a t t a i n the mountains between Glacier Peak and Mt. Rainier,have f a i l e d to produce any populations. Glacier Peak i s well i s o l a t e d by distance and topography from both G. oc c i d e n t a l i s to the north and the undescribed Mt. Rainier populations. Between Glacier Peak and Mt. R a i n i e r , a 90-mile expanse, the Cascade Range decreases i n elevation and breadth. The highest peaks are less than 6000 feet except for three near Mt. Rainier that exceed t h i s e l evation. This portion of the Cascade Range 'Is divided by two major drainage systems: the Skykomish River flowing west through Stevens Pass at 4000 foot elevation and the Wenatchee River draining the eastern slopes of the range. The headwater canyons overlap i n Stevens Pass at 4061-foot elevation. Further south toward Mt. R a i n i e r , the Cascade Range i s divided by an even lower pass, the Snoqualmie, 3010 feet above sea l e v e l . In between these low passes a dozen other r i v e r systems have dissected both slopes with deep canyons. The combination of l a t e Pleistocene g l a c i a l evidence, the lower elevations of the dissected intervening portion of the Cascade Range, the i n f e r r e d climate during Hypsi-thermal and Neoglacial periods, and f i e l d data, strongly suggest separation between Glacier Peak and the Mt. Rainier G r y l l o b l a t t a for at l e a s t 40,000 years. 108 The Mt. Rainier G r y l l o b l a t t a are found on the northern and eastern slopes i n Cayuse and Chinook pass hypolithion habitats above 4600 feet. They occur up to 7500 feet eleva-t i o n on the older moraines of Emmons Glacier . This occurrence at elevations below those from which they are generally known from l o c a l i t i e s i n the ranges, i s undoubtedly due to the much cooler more moist summer climate of the north and east slopes of Mt. Rainier. This l o c a l i z e d cool summer climate seems to be the e f f e c t of the topography of the mountain and of many g l a c i e r s extending to lower elevations. Mt. Rainier was continually glaciated during l a t e Pleistocene. While the Puget Lobe during Fraser G l a c i a t i o n did not overrun the western slopes, the e a r l i e r Salmon Springs Stade (ca. 35,000 years ago) did meet the ice cap exis t i n g on the mountain. The regional snowline (the elevation above which snow accumulated and forms i c e f i e l d s ) (Matthes, 1940) i n the mountains near Rainier was approximately 3100 feet (1000 m.) during Fraser G l a c i a t i o n about 25,000 to 9000 years ago. Glaciers from the ice f i e l d s of Mt. Rainier extended to lower elevations i n the r i v e r valleys and covered the regions now inhabited by the G r y l l o b l a t t a . The extent of g l a c i a t i o n on Mt. Rainier decreased during the middle portion of the Fraser episode. A rigorous cold climate returned to Mt. Rainier before the end of the Sumas Stade with the formation of cirque and valley g l a c i e r s extending below 4000 feet around 11,000 years ago (Crandell et a_, 1962). 109 Glaciers and persistent snow f i e l d s disappeared during the Hypsithermal i n t e r v a l on Mt. R a i n i e r . The present large g l a c i e r s are of Neoglacial o r i g i n and not older than 3500 years. G l a c i a l a c t i v i t y has continued to the present with major advances occurring during the 14th and 15th centuries and again i n the 1850's, and then reached or covered the present habitat of the insects (Harrison, 1956a; Sigafoos and Hendricks, 1961). Post Pleistocene volcanic a c t i v i t y of Mt. Rainier i s recognized hy ash deposits and mud flows i n the l o c a l i t y of the G r y l l o b l a t t a between 2300-2000 years ago (Crandell et a l 1962). Extensive mud flows are recorded i n modern time covering the habitat (Fiske, Hopson, and Waters, 1963). The populations on Mt. Rainier must thus be very recent inhabitants of the present habitats. The parent stock could have survived at l e a s t the l a s t portion of l a t e Pleistocene g l a c i a t i o n on the western slopes i n an area between the summit ice f i e l d s and the Cordilleran Puget Lobe i n the coastal p l a i n . Continual survival during the entire l a t e Pleistocene i n the upper region of the American River east of Chinook Pass i s a p o s s i b i l i t y . The present d i s t r i b u t i o n would support the existence of Pleistocene refugia on the eastern slope of Mt. R a i n i e r . G r y l l o b l a t t a c h l r u g l c a . known from Ape Cave and other i c e caves on the southern slopes of Mt. St. Helens, occurs i n habitats no older than the l a t e Pleistocene (Verhoogen, 1937). In f a c t , most of the present cone and surrounding 110 lava f i e l d s appear to be of p o s t g l a c i a l o r i g i n and le s s than 12,000 years old (Lawrence and Lawrence, 1959; Mullineaux, 1964)., Mt. St. Helens has been the most active of the nor-thern Cascade stratovolcanoes during the post Pleistocene. The l a s t 1000 years have been a period of violent a c t i v i t y with major eruptions of lava, pumice, and mud as recently as 1854. The entire area of the lava f i e l d s surrounding the peak are mantled with pumice varying i n depth from 1 to 6 feet. The eruption of 1843 deposited ash |-lnch deep 93 miles away at The Dalles, Oregon. c Between Mt. Rainier and the Columbia River the Cascade Range reaches i t s lowest crest elevation. The entire region i s covered with g l a c i a l outwash or extensive lava f i e l d s and possesses a climate too dry and hot for post P l e i s t o -cene habitation by G. chirup,!ca. The region was not extensively glaciated during the la t e Pleistocene for g l a c i e r i z a t i o n was r e s t r i c t e d to Mt. Adams, Mt. St. Helens, and the c r e s t l i n e . The entire region c e r t a i n l y must have had a p e r i g l a c i a l climate under the influence of the Cascade ice f i e l d s to the north and south. During l a t e Pleistocene, 40,000 to 9000 years ago, cli m a t i c conditions were probably i d e a l anywhere i n the area for G r y l l o b l a t t a . However, at the termination of the l a s t g l a c i a t i o n and the return for a 5000-6000 year period of much warmer dry conditions (the Hypsithermal), G r y l l o b l a t t a  chlruglca were undoubtedly forced to retreat to the only habitat t o l e r a b l e , namely, the ice caves. Occupation of I l l the higher elevations and alpine on Mt. St. Helens was not possible owing to continual volcanic a c t i v i t y . " G r y l l o b l a t t a chirugica, or ancestral species, has undoubtedly existed i n the region since at le a s t the l a t e Pleistocene. Geological and paleobotanical information suggest much the same cycles of clim a t i c conditions for the entire Quaternary 1,000,000+ years (Em i l i a n i , 1955; F l i n t , 1957; Martin and H a r r e l l , 1957). Therefore, the present stock may have inhabited the general region for a considerable length of time. The topographical b a r r i e r of the Columbia River and the nature of the insect's morphology show a long i s o l a t i o n and no r e l a t i o n s h i p to Gr y l l o b l a t t a on Mt. Rainier or Mt. Hood. The present d i s t r i b u t i o n of the Northern Cascade species of G r y l l o b l a t t a i s of a post Pleistocene o r i g i n and no older than 9000 years. The present populations of Gr y l l o b l a t t a  scudderi. G r y l l o b l a t t a o c c i d e n t a l i s and Gry l l o b l a t t a skagitensia are derived from ancestral stock which l i v e d during the late Pleistocene south or east of the ice fronts and reached the present l o c a l i t i e s as the ice receded at the beginning of post Pleistocene (ca.10,000 years ago). The present d i s t r i b u t i o n of G. scudderi may be as recent as the Neoglacial (ca. 3500-2000 years ago). 112 2. High Cascade Group The Mt. Hood G r y l l o b l a t t a are geographically i s o l a t e d by the Columbia River Gorge from G, chiruglca found i n the lava caves on the slopes of Mt. St. Helens. The Columbia River has functioned as an e f f e c t i v e b a r r i e r to northern migration to many animal species(e.g» Amphibia, see Stebbins, 19^9; Lowe, 1950). The Columbia River Gorge b a r r i e r evidently has been i n e f f e c t throughout the Pleistocene. Certainly i t was a very e f f e c t i v e b a r r i e r during periods of l a t e Pleistocene when the gorge was repeatedly swept rim to rim by catastro-phic floods a r i s i n g i n eastern Idaho and western Montana (Richmond, 1965). The population of G r y l l o b l a t t a hoodalles i s presently i s o l a t e d from G. r o t h i . G. lavacola and G. s. s c u l l e n l i n the Three Si s t e r s complex by the Warm Springs Plateau and the lowlands between Mt. Hood and Mt. Jefferson. Other b a r r i e r s to population interchange e x i s t , such as the deep canyons of the -Salmon, Clackamas and Matolius r i v e r s . The present temperature i n these lower areas i s such that G r y l l o b l a t t a could not survive except during the short periods of mid-winter. This has been an e f f e c t i v e b a r r i e r to the taxon since at least the beginning of the post Pleistocene (ca. 8500 B. P.). Even during the coldest of 113 Neoglaelation, temperature means were not appreciably lower than today. Populations could exist i n t h i s area i f they inhabited lava ice caves, but currently none has been found. Mt, Hood was extensively glaciated during the Fraser G l a c i a t i o n of the l a t e Pleistocene. The severity of the g l a c i a t i o n i s exemplified by the possible plucking of at l e a s t 600 v e r t i c a l feet from the summit and possibly more from the slope (Wise, 1964a, 1964b). The mountain was almost completely i c e covered with terminal moraines below 4500 feet (Crandell, 1965; Wise, 1966). ..Ice completely covered the High-Cascades from O l l a l i e Butte jU3t south of Mt. Hood to Mt. McLoughlin near the Oregon-California border during Fraser Glaciation (Thayer, 1939; Crandell, 1965) (Fig. 10) . During that time the present b a r r i e r between the more southern Cascade species of G r y l l o b l a t t a and the Mt. Hood species did not e x i s t . This region would have experienced a p e r i g l a c i a l climate during Fraser G l a c i a t i o n since i t i s between the extensive ice cap of the Cascades and that of Mt. Hood. In fact t h i s southern peripheral zone may well have been of post Pleistocene o r i g i n for the present species on Mt. Hood. As the ice retreated toward Mt. Hood the Gry l l o-b l a t t a may have followed the front, the - populati on -even-t u a l l y then finding the only tolerable habitat at the begin-ning of the Hypsithermal on the slopes of Mt. Hood. Evidence 114 for t h i s southern o r i g i n of the present population i s i t s close r e l a t i o n s h i p with G. r o t h l found at south Broken Top i n the High Cascades. The G. hoodalles at Timberline Lodge, 6000-foot e l e -v a t i o n , Mt. Hood, has occupied t h i s l o c a l i t y for le s s than 1700 years. The southeast flank of Mt. Hood experienced a. huge mud and water flow mixed with eruptive hornblende andesite debris about 1670 B. P. - 200 years (Lawrence and Lawrence, 1959; Wise, 1966). This mud-andesite flow com-pl e t e l y covered the whole southwest flank of Mt. Hood, being many feet deep as far as the 3800-foot elevation on Zigzag River. The source of t h i s material i s believed to have been the l a s t eruptive phase of Mt. Hood and the breaching of a crater lake of l/3-mlle diameter and 800 feet deep. The flow completely buried large trees and denuded the upper slopes. The present habitat of the G r y l l o b l a t t a i s i n t h i s unsorted debris beneath rotten logs and large.buried rocks. Search by myself and b i o l o g i s t s of the U. S. Forest Service has f a i l e d to locate any other populations of G. hoodalles on the mountain. I do not believe that the genus migrated to the mountain before 1670 B. P. The present population must have moved to the unsorted material from some other region of the mountain since 1670 - 200 years B. P. The unsorted material of g l a c i a l flows and boulders of a l l sizes create an ideal hypolithion not found in other areas of the south 115 slopes. Extensive erosion on the other flanks of the mountain of Recent o r i g i n may have destroyed the o r i g i n a l post Pleistocene habitat. I seriously doubt that t h i s i s the only population of G. hoodalles on Mt. Hood. I f e e l that investigations during the summer w i l l turn up at le a s t one more at Cloud Cap on the east slope about 6800-foot elevation. Should , additional populations be found i n the future, they w i l l be geographically i s o l a t e d from each other by the unstable canyons present on a l l flanks. The G r y l l o b l a t t a that occur i n the Three Si s t e r s region are the most concentrated group of species of the entire genus i n North America. Within an area of sixty by t h i r t y miles are found G. s. s c u l l e n i . G. r o t h i . G.. lava cola, G. s. cryocola and G. pa u l l n a l , plus two cavernicolous popu-l a t i o n s of uncertain systematic r e l a t i o n s h i p s . G r y l l o b l a t t a s. s c u l l e n i . G. ro t h i and G. lava cola are sympatric i n the Three S i s t e r s , McKenzie Pass-Belknap lava f i e l d s region. G r y l l o b l a t t a paulinai i s an a l l o p a t r i c and t r o g l o p h i l i c cavernicolous form. A peripheral population of G. lava cola i s sympatric with G. s. cryocola i n the Edison Ice Cave system for at least a portion of the yearly climate cycle and i s thought to exist sympatrically with G. r o t h i . The Three Sisters complex i s composed of extinct i n d i v i d u a l volcanic cones that range i n elevation from 7045 to 10,854 feet with such picturesque names as North S i s t e r , Middle S i s t e r , South S i s t e r , L i t t l e Brother, Husband, Wife, Bachelor and Broken Top. Hodge (1925), i n describing the 116 geology of the region, v i s u a l i z e d these volcanic cones to be Pleistocene remnants of a single huge mountain over 18,000 feet high which he called Mt. Multnomah. In l a t e r more det a i l e d work, Williams (1957) considered these to be of i n d i v i d u a l volcanic o r i g i n of Pleistocene or younger age. Today most geologists follow Williams' theory. In any event, there has been almost continuous volcanic a c t i v i t y on or around the peaks throughout the entire span of the P l e i s t o -cene, and th i s continued during the post Pleistocene u n t i l l e s s than 2000 years ago (Williams, 1957; Peck, I960). Evidence of early and middle Pleistocene g l a c i a t i o n i s unknown for the High Cascades. The sequence of events, documented for the Northern Cascades and the Sierra Nevada to the south, i s i n f e r r e d to have taken place i n the Three Sisters region (Williams, 1942, 1944; F l i n t , 1947; Peck, i960). Late Pleistocene g l a c i a t i o n i s well documented with g l a c i a l stades correlated i n age with the Northern Cascades and the Sierra Nevada (Heusser, 1966). The Fraser G l a c i a t i o n i n the High Cascades was widespread. The entire range was covered with summit ice f i e l d s 500-1000 feet thick at an elevation of 5000 feet and deeper over the high peaks (Thayer, 1939; Williams, 1942, 1957). Lobes from the summit ice f i e l d s extended far down the east slopes transporting boulder e r r a t i c s of crest o r i g i n over 80 miles into the present high desert (Williams, 1957; A l l i s o n , 1966). Summit ice f i e l d lobes extended down the valleys on the western slopes to the Willamette lowlands around 2000 to 2500 feet i n elevation. E a r l i e r l a t e Pleistocene 117 g l a c i a t i o n entered the lowlands, transporting er r a t i c s , across the v a l l e y as far west as Monmouth and C o r v a l l i s ( A l l i s o n , 1 9 3 5 , 1 9 3 6 ) . There are f i f t e e n major g l a c i e r s and numerous snow f i e l d s of Neoglacial or Recent o r i g i n flowing from subsummit ice f i e l d s on North S i s t e r , Middle S i s t e r , South S i s t e r , Bachelor and Broken Top. These peaks have populations of G r y l l o b l a t t a occurring on the lower slopes which range up to the present terminal moraines of the g l a c i e r s between 7500 and 8000 f e e t . Most traces of l a t e Pleistocene g l a c i a t i o n have been obl i t e r a t e d on these peaks by the extensive post Pleistocene volcanism and Neoglacial ice movement. North Sister i s the oldest of the complex and i s considered to be early or middle Pleistocene i n age, having l o s t one-fourth of i t s mass during l a t e Fraser G l a c i a t i o n . Middle S i s t e r , South S i s t e r , Broken Top and Bachelor are thought to be perhaps middle but most l i k e l y . l a t e Pleistocene i n age, with growth extending well into the post Pleistocene. Summit craters exist on a l l of these.peaks and the slopes show erosion attri b u t e d only to the present g l a c i e r s (Williams, 1942, 1 9 5 7 ) . McKenzie Pass, immediately north of the Three S i s t e r s , i s composed of extensive lava f i e l d s of Neoglacial or younger age and the area i s dominated by Belknap Crater. The 6800-foot high Belknap Crater i s the primary source of the extensive lava flow that covered some 50 square miles and which flowed down the pass to 4C00 feet and covered the lower 118 slopes of North Sist e r to an elevation of 6500 f e e t . This flow, hundreds of feet t h i c k , completely covered the l a t e Pleistocene headwaters of the McKenzie River and i s dated 3000 years B. P. (Baldwin, 1964). There are younger i s o l a t e d f i s s u r e flows that form steptoes (islands) i n the 3000-year old Belknap Crater flow. G r y l l o b l a t t a s_. s c u l l e n i occurs i n the basin between North S i s t e r and Middle S i s t e r . The type l o c a l i t y at Scott Camp, 6600 f e e t , i s i n consolidated old talus of morainal hypo-l i t h i o n below Renfrew G l a c i e r . This species has been taken from over 8000 feet i n t h i s basin north, down the slopes of North S i s t e r , to the Belknap lava f i e l d s where i t occurs sympatrically with G. r o t h i and G. lavacola T G r y l l o b l a t t a lavacola inhabits the Belknap lava f i e l d s and i s d i s t r i b u t e d along the southeastern margins of the flows from Windy Point (4909..feet) to Condon Butte (5030 feet) and up the lavas covering the slope of North S i s t e r . The type"1 l o c a l i t y of G r y l l o b l a t t a r o t h i i s Happy Valley (6450 feet) located below Crook Glacier on Broken Top. This i s a species of wide d i s t r i b u t i o n , occurring at 7000 feet on 3achelor 6 miles south and through the Three Sisters to McKenzie Pass, 15 miles southwest of Broken Top. Gry l l o b l a t t a s. cryocola i s a cavernicolous subspecies from the Edison Ice Cave system, 5 miles southeast of Bachelor. The Edison Ice Cave system formed i n a pressure ridge on a post Pleistocene b a s a l t i c f i s s u r e flow from the southeastern flank of Bachelor. A 6888 year B. P. age i s assigned to Bachelor and the Edison flow i s possibly the 119 same age (Taylor, 1965) . G r y l l o b l a t t a a . cryocola i s sea-sonally t r o g l o p h i l i c i n the caves or hypolithion i n the substrate above and around the caverns. It occurs sympatri-c a l l y with G. lava cola i n the t w i l i g h t and entrance zone portions of the cave system. Gryllobla tta s. cryocola i s r e s t r i c t e d to the cave area but G. lava cola appears to be a disjunct population i n t h i s area, being unknown from Bachelor, Broken Top or Middle S i s t e r . The present l o c a l i t i e s of G r y l l o b l a t t a i n the Three Si s t e r s complex were covered by l a t e Pleistocene (23,000-9000 years B. P.) ice f i e l d s or g l a c i e r s . The present ranges are a l l located i n post Pleistocene and younger lava f i e l d s or i n hypolithion of Neoglacial to Recent o r i g i n . The ancestral populations of these species were undoubted-l y forced out o f the region by late Pleistocene ice advance. The present d i s t r i b u t i o n suggests a retreat both to the east and west beyond the ice fronts and i n the p e r i g l a c i a l zone. As late Pleistocene ice receded i n the High Cascades, approximately 11,000 to 9000 years ago, the parent stocks probably followed the. retreating ice and peripheral zone back to the higher elevations near the v i c i n i t y of the Three S i s t e r s . An eastern retreat and subsequent return i s evidenced by G r y l l o b l a t t a paullnai from South Ice Cave, located i n the high desert lavas to the east. A western l a t e Pleistocene retreat and early post Pleistocene return i s documented by the discovery of a new species of G r y l l o b l a t t a on Mary's 120 Peak near the western borders of the Willamette lowlands and Coast Range. Both South Ice Cave and Mary's Peak are located within the l a t e Pleistocene p e r i g l a c i a l zone, as indicated by f o s s i l ice wedges i n t h e i r v i c i n i t y . The present d i s t r i b u t i o n s of the species of G r y l l o b l a t t a i n t h i s region of the High Cascades must have formed after the post-volcanic and Neoglaelation periods that began approximately 3500 years ago. Based on the present sympatric d i s t r i b u t i o n and the fact that no hybrid populations have been discovered, I hypothesize that the parental populations were the same species a s t ound today. 3. Southern Cascade Group The Southern Cascade G r y l l o b l a t t i d a e are known only from Crater Lake, Oregon, Mt. Shasta and the v i c i n i t y of Mt. Lassen, C a l i f o r n i a . The mountains, i s o l a t e d from the main range by extensive low elevation lava f i e l d s , are the southern extension of the major volcanic peaks which dominate the Cascade Range. Major volcanic a c t i v i t y took place throughout the Pleistocene and has continued to Recent time, with Mt. Lassen erupting i n 1914 and active u n t i l 1917 (Williams, 1932a, 1942; Anderson, 1941). In a d d i t i o n , each was the s i t e of l o c a l ice caps and g l a c i e r s during the various g l a c i a t i o n s of the Pleistocene and Neoglacial. Mt. Shasta (14,164 feet) has currently active g l a c i e r s and 121 both Mt. Lassen (10,466 feet) and Crater Lake (6000 to 8300 feet) have large snow f i e l d s which l a s t throughout the year. G r y l l o b l a t t a r o t h l was reported by Gurney (1953) from Crater Lake. Two specimens, tentatively i d e n t i f i e d as G r y l l o b l a t t a prurneyi, are known from Mt. Shasta. G r y l l o - b l a t t a barberi and G r y l l o b l a t t a chandlerl.. while not occurring on Mt. Lassen, are found within the p e r i g l a c i a l zone that existed around the Mt. Lassen ice shield during Tioga (Fraser) G l a c i a t i o n (Williams, 1932b; Anderson, 1941). Eagle Lake, Lassen Co., the type l o c a l i t y for G. chandler!, i s 30 miles east of Mt. Lassen. The type l o c a l i t y of G. barberi i s 20 miles southeast of Lassen i n the north fork of the Feather River. An i s o l a t e d population of G r y l l o b l a t t a occurs i n Wilson Ice Cave located within the southern boundary^ of Tioga G l a c i a t i o n on Mt. Lassen. The present Crater Lake i s situated i n a caldera formed when the top of approximately 12,500-foot Mt. Mazama collapsed 6453 - 250 years ago (Powers and Wilcox, 1964). Mt. Mazama was mantled by extensive ice f i e l d s during the l a t e P l e i s t o -cene, volcanic a c t i v i t y also occuring at t h i s time (Williams, 1942). It i s thought that just p r i o r to the mountain collapse which formed the lake caldera, a violent explosion took place, depositing pumice 30-40 feet deep as far as 30-45 miles to the north and east. Some of t h i s ash and pumice was wind-borne to central Montana and Alberta (Williams, 1942; Wilcox, 1965). The l o c a l i t i e s around the Crater Lake rim where G r y l l o -b l a t t a i s now found were covered many feet deep by the hot 122 pumice and huge avalanches of mud and lava r e s u l t i n g from the r a p i d l y melting g l a c i e r s during the explosion. Before the Hypsithermal period, during which the crater formed, the l o c a l i t i e s were extensively glaciated and ice-covered during Fraser G l a c i a t i o n (ca. 2 3 , 0 0 0 - 1 2 , 0 0 0 years ago). It seems unl i k e l y that G r y l l o b l a t t a inhabited Mt. Mazama during the l a t e Pleistocene i n l i g h t of the widespread occurrence of ice f i e l d s and g l a c i e r s and the periodic volcanic a c t i v i t y . At least once during the l a t e P l e i s t o -cene Mt. Mazama was covered by the i c e sheet which mantled the entire Oregon Cascades as far south as Mt. McLaughlin (Crandell, 1965) ( F i g . 10) The p o s s i b i l i t y that G r y l l o b l a t t a inhabited some areas on Mt. Mazama i n the time between ice recession and about 6600 years ago must be considered. The almost continuous volcanic a c t i v i t y during the period tends to negate t h i s p o s s i b i l i t y . I f G r y l l o b l a t t a did not occupy Mt. Mazama during the l a t e Pleistocene and 6600 years ago, where did the parental population survive t h i s period? Other species of G r y l l o -b l a t t a are known from the south in the v i c i n i t y of Mt. Shasta, the Lava Beds National Monument and Medicine Lake Highlands. These numerous northern C a l i f o r n i a populations, of which many are cavernicolous, are not closely related systematically to G. r o t h i . The northern C a l i f o r n i a species form a d i s t i n c t group seemingly r e s t r i c t e d to the lava i n the Modoc Plateau and southern terminus of the Cascades. The Crater Lake G r y l l o b l a t t a . probably subspecific with 123 G_. r o t h l , undoubtedly came from the same ancestral stock. The parental stocks of each modern form most l i k e l y inhabited the high desert plateau east of the ice f i e l d s covering the Three Sisters complex and Mt. Mazama. They then returned, following the receding p e r i g l a c i a l zone, into the High and Southern Cascades. I believe the Crater Lake populations migrated to th e i r present l o c a l i t i e s sometime af t e r the caldera formation from the northeast and most l i k e l y during Neoglaelation (ca. 3500-2000 years ago). The discovery of additional populations on Mt. Thielson, Diamond Peak, the Twins and Maiden Peak, between the Three Sisters and Crater Lake, would substantiate t h i s hypothesis. The Crater Lake G r y l l o b l a t t a are presently i s o l a t e d from the species i n the Modoc Plateau by the Klamath Marsh and dry rabbit brush scrublands. This Crater Lake population i s also rather e f f e c t i v e l y i s o l a t e d from the Klamath Mountain population to the southwest by the low elevations associated with the Rogue River drainage and the Grants Pass-Medford V a l l e y . The drainage canyons of the Willamette, Deschutes and Umpqua r i v e r s v i r t u a l l y divide the low crest of the Cascade Range between Crater Lake and the High Cascade system. In a d d i t i o n , Just north of the Crater Lake rim exists a pumice desert some 10 miles wide which, u n t i l h i s t o r i c a l times, was apparently devoid of plant l i f e . The i d e n t i f i c a t i o n of the Crater Lake population as G. r o t h i by Gurney (1953) was based on a single female which was declared the a l l o t y p e . The holotype i s from Broken Top, 124 over 90 miles north. This association of holotype and a l l o -type was based on the shortness of the cereal segments, these being the only specimens then known. I have since collect e d G r y l l o b l a t t a of both sexes from various l o c a l i t i e s on the rim of Crater Lake and believe that these populations are not t y p i c a l G. r o t h i . Any future discovery of G r y l l o -b l a t t a between Crater Lake and Broken Top would be highly important, since i t might lead to a decision regarding the exact i d e n t i t y of the Crater Lake G r y l l o b l a t t a . The parental populations of both the Mt. Shasta and Mt. Lassen species undoubtedly inhabited regions within the p e r i g l a c i a l zone and possibly occupied a l l the Modoc Plateau and Medicine Lake Highlands. They are conspeciflc with, or closely related t o , other species now occurring within i c e caves of the plateau lava f i e l d s (Kamp, 1963). 4 . Modoc Plateau-Basin Range Group The G r y l l o b l a t t a i n the Modoc Plateau-Basin Ranges are cavernicolous species found i n the numerous ice caves of the lava f i e l d s . G r y l l o b l a t t a chandlerl. with type l o c a l i t y as Eagle Lake, ranges into the ice caves on the eastern slope of Mt. Lassen. Almost every ice cave explored between Eagle Lake and the Medicine Lake Highlands contains populations of G r y l l o b l a t t a . North of these highlands, G. gurneyl occurs i n many of the ice caves i n the Lava Beds National Monument. Ge o l o g i c a l l y , the Modoc Plateau i s a b a s a l t i c block 125 3000-5000 feet high, surrounded by numerous volcanoes 1000-2000 feet higher. On the western edge i t borders the Southern Cascades. During the l a t e Pleistocene Tioga Stade of g l a c i a t i o n , l o c a l ice caps and mountain g l a c i e r s were present on the highest peaks, especially the Medicine Lake Highlands between the Lava Beds National Monument and Eagle Lake-Mt. Lassen region (Stearns, 1929; Blackwelder, 1931; Peacock, 1931; Williams, 1932a, 1932b). A discussion of the l a t e Pleistocene-post Pleistocene d i s t r i b u t i o n and phylogenetic relationships of the species i n t h i s area i s given by. Kamp (1963). 5 a . Coast Range Group 1 The only known G r y l l o b l a t t a from the Coast Range i s a new species which occurs on the upper elevations of Mary's Peak, 14 miles west of C o r v a l l i s , Oregon. In the Coast Range of western Washington and Oregon the c r e s t l i n e i s below 3000 f e e t , but a few summits, such as Mary's Peak, are over 4000 feet i n elevation. Gl a c i a t i o n during the Fraser Stade of l a t e Pleistocene was apparently r e s t r i c t e d to cirque g l a c i e r s on the protected eastern side of the highest peaks (Baldwin and Roberts, 1952). Mary's Peak was the site- of such l o c a l g l a c i a t i o n during the l a t e Pleistocene. The eastern region of the Coast Range around Mary's Peak was also apparently affected by the climate present i n the p e r i g l a c i a l zone and the valley g l a c i e r s which carried e r r a t i c s derived from the High Cascades to a few miles 126 west of C o r v a l l i s . This i s o l a t e d species of G r y l l o b l a t t a i s most probably a remnant of G r y l l o b l a t t a displaced to the west by the extensive ice f i e l d s and valley g l a c i e r s from the High Cascades to the east. As ice receded with the advent of the Hypsithermal i n t e r v a l , a segment of the population may have retreated to the present l o c a l i t y of Mary's Peak. The present d i s t r i b u t i o n of t h i s species i s r e s t r i c t e d to the eastern slopes between 3000 feet and the 4100-foot summit. A regional, storm pattern deposits larger depths of snow on the east side of the Peak where i t may p e r s i s t i n sheltered areas well into the f a l l . 5b. Klamath Mountain Group , G r y l l o b l a t t a i s known from the Klamath Mountains near the Oregon-California boundary. A single specimen was found on the Siskiyou Summit, 4464 f e e t , 14 miles south of Ashland, Oregon, i n a t y p i c a l hypolithion s i t u a t i o n . The Klamath Mountains are a very rugged region somewhat higher i n elevation than the Coast Range. Clusters of high peaks, 6000-9000 feet i n a l t i t u d e , r i s e above the general l e v e l s which are between 2700 and 4000 feet i n e l e v a t i o n . The narrow canyon f l o o r s of the many r i v e r s are less than 900-2200 feet above sea l e v e l , even i n the center of the range, giving the mountains a precipitous r e l i e f of 3200-6600 f e e t . 127 Evidence of widespread l a t e Pleistocene Tioga G l a c i a t i o n i s known from above 3000 feet for the Klamath and T r i n i t y Ranges ( F l i n t , 1947; Wells, Hotz and Carter, 1949). T h l s g l a c i a t i o n of cirques, valley g l a c i e r s and summit ice f i e l d s may have extended east to the Sacramento River Canyon. This i s near the western border of the g l a c i a t i o n present i n the Mt. Shasta region of the Southern Cascades ( C a l i f o r n i a State Water Resources, 1959; Sharp, I960). I have not seen the specimeni.in the U. S. National Museum c o l l e c t i o n and A. B. Gurney has not been able to determine the species, nor i t s r e l a t i o n s h i p (Gurney, pers. comm.) It i s not surprising that G r y l l o b l a t t a occurs i n the Klamath Mountains considering the proximity to the range of the northern C a l i f o r n i a and southern Oregon species. The Klamath-Trinity mountains are i n an area of heavy snow-f a l l which p e r s i s t s throughout the year i n the highest elevations and even i n summer i s under the influence of northern P a c i f i c storm systems. Dr. Gurney, members of the C a l i f o r n i a Department of Entomology, and I searched i n the region of G r i z z l y Peak i n 1964. This region, of high alpine and large cirque basins, contains the only g l a c i e r in the mountain range. We d i d not discover G r y l l o b l a t t a at that time, though we did f i n d new forms of high a l t i t u d e Orthoptera such as Boona c r l s . Melanoplus a'hd Napaia. We f e e l confident that G r y l l o b l a t t a occurs in the region and south on higher peaks of the T r i n i t y Range. The Siskiyou Summit material i s presently i s o l a t e d 128 both topographically and c l i m a t i c a l l y from other species. Additional material w i l l probably e s t a b l i s h a close r e l a -tionship to the extreme southern Cascade-Modoc Plateau species. I I I . The Sierra Nevada Group From the standpoint of systematlcs and d i s t r i b u t i o n , the Sierra Nevada G r y l l o b l a t t a are a diverse fauna. Seven areas containing G r y l l o b l a t t a are known for the Sierra Nevada, scattered i n l o c a l "pockets" from the northern terminus south along the crest below Sequoia National Park, a distance of 300 miles. G r y l l o b l a t t a washoa i s described from 7382 feet on Echo Pass Summit, Eldorado Co., south of Lake Tahoe. G r y l l o b l a t t a b i f r a t r i l e c t a i s known from the 9000-10,000 foot l e v e l of Sonora Pass, 50 miles southeast of Lake Tahoe. A new species (Gurney, pers. comm.) i s found on the Badger Pass-Glacier Peak region of Yosemite National ^Park between 6000 and 7500 feet i n a l t i t u d e . A single female has been found at 12,000 feet i n the Upper Convict Basin near Devil's Postpile National Monument, 7 0 miles southeast of Sonora Pass. The southernmost popula-t i o n i s from May's Hole, Sequoia. National Park, Tulare Co. From the northern Sierra Nevada I discovered two new species, one from 8587-foot Mt. E l w e l l , 60 miles south-east of Mt. Lassen, and another from the 6000-7000 foot elevation i n the Sierra Butte, 10 miles south of 129 Yuba Pass located approximately halfway between Mt. Lassen and Lake Tahoe. The apparently disjunct d i s t r i b u t i o n of t h i s group along the Sierra Nevada i s probably a consequence of incomplete c o l l e c t i n g . There are few accessible passes through the high elevations of the range and there has been much searching perhaps i n the wrong habitat. In f a c t , the lack of habitat knowledge impeded my collecting•success during the two weeks I spent i n the region i n 1967. A number of entomologists i n the past 10 years have looked for G r y l l o b l a t t a during t h e i r other c o l l e c t i n g . These records were based on the descriptions of either the Rocky Mountain habitat or that of G. s c u l l e n i i n loose talus at the foot of g l a c i e r s . I firmly believe that G r y l l o b l a t t a populations w i l l be found the length of the range as far south as V/alker Pass, west of Death V a l l e y , and possibly in the San Gorgonio-San Jacinto ranges of southern C a l i f o r n i a . Present systematic data suggest that possibly two groups of species are d i s t r i b u t e d i n the Sierra Nevada: a northern group of large s i z e , long, many-segmented antennae, very long legs and symmetrical male g e n i t a l i a , occurring from Lake Tahoe to the v i c i n i t y of the closely related Modoc Plateau-Southern Cascades species, and a southern group of small s i z e , short, l e s s segmented antennae and moderately asymmetrical male g e n i t a l i a , from Lake Tahoe to Kings Canyon National Park. The Sierra Nevada i s a gigantic westward-tilted f a u l t 130 block up to 14,000 feet i n elevation. Its eastern slope Is an abrupt wall of 15-35° slope r i s i n g i n 5 miles from 2800 to 11,000 feet above the basin and range province. I t has an average c r e s t l i n e a l t i t u d e of over 7500 f e e t . The western slope descends from the crest to an a l t i t u d e of 300-600 feet at i t s foot i n a distance of 50-60 miles. The numerous streams and r i v e r s have cut narrow gorges 2000-7000 feet deep. A region 35 miles wide along the 350-mile crest was intensely glaciated during the P l e i s t o -cene (Blackwelder, 19315- A v e r i l l , 1937; Wahrhaftig and Birman, 1965). Six major g l a c i a t i o n s have been recognized i n the Sierra Nevada for the Pleistocene, and three smaller advances are known for the Neoglacial period. The Tioga G l a c i a t i o n i s correlated for the late Pleistocene and i s equal to the Pinedale G l a c i a t i o n i n the Rocky Mountains and the Fraser G l a c i a t i o n i n the Coast-Cascade ranges (Matthes, 1930; Blackwelder, 1931; Birman, 1957, 1964; Thompson and White, 1964). The Sierra Nevada at the present time contains only a few cirque g l a c i e r s on the sheltered north slope above 10,500 feet near Lake Tahoe and above 13,000 feet between Sequoia-King's Canyon National Parks. Climatic snowline i s 14,000 feet i n the Sequoia-King's Canyon area and 7000 to 9000 feet i n the northern portions ( F l i n t , 1957). The climate i s very seasonal, with 90 per cent of the .precipi-t a t i o n i n the winter, and i s controlled more by topography than by l a t i t u d e . 131 P r e c i p i t a t i o n ranges from 90 inches i n the north to only 55 inches i n the southern regions. P r e c i p i t a t i o n increases 2 to 4 inches for each 300-foot r i s e , reaching a maximum at about 5000-6000 feet i n the central part of the range. At higher elevations 86 per cent of the pre-c i p i t a t i o n f a l l s as snow, and t h i s reaches an average depth of 34 feet on Donner Summit. The east side of the Sierra Nevada i s i n a r a i n shadow and has a high desert condition of l i t t l e r a i n and snow. Summer temperatures range from 15-100° F. and during the winter from -30° to about 55° F. (U. S. Weather Bureau, 1964). The season's snow, which begins i n October, has usually disappeared by August except for small banks and f i e l d s at the highest elevations (Dale, 1959). In the l a t e Pleistocene a f a i r l y continuous complex of ice f i e l d s formed along the crest, with g l a c i e r s descending both slopes. The d i r e c t i o n of the descending g l a c i e r s was closely controlled by the present drainage patterns and general topography which had been previously established. Snowline was approximately 2000 feet lower during the g l a c i a l maxima than i t i s at present. During Tioga G l a c i a t i o n i ce f i e l d s were thicker in the southern and central regions between 37-38° l a t i t u d e than elsewhere i n the range. To the north the. lower elevations of the crest resulted i n a smaller volume, and at the southern terminus, higher temperatures kept ice volumes r e s t r i c t e d to the highest elevations ( F l i n t , 1957). The larger g l a c i e r s that descended from the crest ice 132 f i e l d s were only about 10 miles long on the eastern slope and as much as 65 miles long on the western side of the range. In the southern Lake Tahoe region the g l a c i e r s descended to a l t i t u d e s of 3000-4000 feet on the east (the f l o o r of the Basin Range) and 1800-3000 feet to the west (Blackwelder, 1931; Putnam, I960; Birman, 1964). In the north g l a c i e r s seldom exceeded 25 miles i n length and ra r e l y descended below 3200 f e e t . However, the northern i ce flowed into the middle fork of the Feather River, 10 miles north of Mt. Elwell ( A v e r i l l , 1937). The known l o c a l i t i e s of the Sierra Nevada G r y l l o b l a t t a are r e s t r i c t e d t,o the higher summits and is o l a t e d peaks which r i s e well above the c r e s t l i n e . Each species i s i s o -lated from others by the climate of the intervening elevations, the r i v e r canyons and glaciated topography. As an example, between G. washoa and G_ b l f r a t r i l e c t a the range i s divided by three passes and four major r i v e r s . The new species In Yosemite i s separated from G. b l f r a t r i l e c t a by the Yosemlte Valley and the 3000-foot deep Merced River Canyon. The southern G r y l l o b l a t t a species are well i s o l a t e d topographically from the others by the Kern Canyon and King's River Gorge, each 6000-7000 feet deep. The northern two populations are separated from each other by the much lower elevation of the c r e s t l i n e . The d r i e r , warmer conditions of the low elevations i s o l a t e t h i s northern group from the other Sierra Nevada species. The ancestors of the present Sierra Nevada species 133 probably inhabited the western slopes of the range below the ice f i e l d s and g l a c i e r fronts during the late P l e i s t o -cene. A post Pleistocene return to the v i c i n i t y of the present l o c a l i t i e s probably occurred, being r e s t r i c t e d by the existing drainage and topography and valley g l a c i e r s which persisted longer than the g l a c i e r s on the intervening highlands. Their persistence well into the post Pleistocene probably channeled migration between the canyons to the present l o c a l i t i e s , for these a l l l i e between the major passes and canyons. The northern forms may have retreated either to the north or west and then returned during ice recession. The close systematic r e l a t i o n s h i p s between these northern Sierra Nevada populations and those of the Modoc Plateau-Southern Cascades suggest a possible common Pleistocene o r i g i n . Discussion and Summary The i n t e r p r e t a t i o n of present d i s t r i b u t i o n a l patterns of G r y l l o b l a t t a i s hindered by the complete void i n the f o s s i l record. As i n other such studies (fleirne, 1952), the conclusions arrived at on the o r i g i n and history of the Grylloblattodea i n western North America are those that appear the most probable when a l l known aspects of the biology, d i s t r i b u t i o n and taxonomy of the animals, and a l l available information on past c l i m a t i c , vegetational and geographical changes i n the area are taken into account. 134 These conclusions are, however, a matter of personal opinion and so may be somewhat co n t r o v e r s i a l . The reconstructions are based almost completely on deductive evidence. Such evidence i s often capable of several i n t e r p r e t a t i o n s , but the conclusions drawn herein are those that appear most l i k e l y with the present data. The further we r e t r e a t into past h i s t o r i e s of d i s t r i -bution, the more tenuous and incomplete i n d i r e c t evidence becomes, especially i n the western C o r d i l l e r a . The P l e i s t o -cene i s represented by f i v e major ice ages i n the west and the l a s t major advance has obscured most evidences of e a r l i e r stades. Hence, most i n d i r e c t evidence i s avail a b l e for only the l a s t 25,000 to 40,000 years. The l a t e Pleistocene and post Pleistocene geologic and climatic events have been found to be useful i n the analysis of present patterns of d i s t r i b u t i o n and speciation of other t e r r e s t r i a l organisms. It has been recognized for many years that the Pleistocene g l a c i a t i o n played an important role i n the present d i s t r i b u t i o n of animals. In western North America, for example, Hubbs and M i l l e r (1948) used such data for the understanding of the d i s t r i b u t i o n of f i s h i n the Great Basin. B l a i r (1958, 1959, 1963) and Thurlow (1961) found Pleistocene and post Pleistocene events had pronounced influence on the d i s t r i b u t i o n and speciation of the Amphibia. The Pleistocene- post Pleistocene has been found to have had an important influence on the present d i s t r i b u t i o n 135 of western R e p t i l i a (Smith, 1957), birds (Mengel, 1964) and mammals (Peterson, 1955). The same periods have influenced the present d i s t r i b u t i o n of invertebrates, especially the Insecta (Ricker, 1964; Howden, 1966). The basic argument i n the above papers i s that g l a c i a t i o n was the most important influence. The premise i s that g l a c i a l advances were accompanied by c l i m a t i c effects far south of continental g l a c i a l borders and at lower elevations than evidence from terminal moraines would indicate for cirque and mountain g l a c i a t i o n . The clima t i c effects produced the cold region, or zone, known as the P e r i g l a c i a l (Boreal) of F l i n t (1957) and Brum-schwieler (1962). The degree of displacement has been argued pro and con,with some plant ecologists (e.g. 3raun, 1955) arguing strongly against any s i g n i f i c a n t displacement. However, evidence of widespread displacement i n the west can be documented from botanical work. Wendorf ( I 9 6 I ) , using pollen a n a l y s i s , indicates a boreal woodland of Pinus and P i c i a 15,000 to 22,500 years ago for eastern New Mexico and western Texas; these areas today are xero-phytic desert shrub and c a c t i . Martin (1963) finds the Mohave Desert of C a l i f o r n i a with a Junlperus and Pinus forest and the southeastern region of Arizona containing a forest of Pinus ponderosa and Abies; these regions are presently a r i d desert grass and shrub f l o r a . 136 When the continental ice sheets of the Pleistocene disappeared, i t i s usually assumed that the deglaciated areas were repopulated by species previously p e r s i s t i n g north (Yukon-Mackenzie valleys) or south of the i c e . This was undoubtedly true for highly mobile species where success for dispersal was not influenced by the vagaries of the post-g l a c i a l t e r r a i n and climate. This theory poses problems when applied to such an organism as Gr y l l o b l a t t a which i s apterous, has a very slow growth r a t e , and i s r e s t r i c t e d to a narrow range of temperature and humidity. Pleistocene survival i n refugia has been proposed many times for species with low dispersal powers. Such a theory would seem very applicable to G r y l l o b l a t t a . Evidences of refugia within and between the Cordilleran and Keewatin ice sheets are incomplete and i n many cases contradictory. Some of the proposed refugia are mantled with g l a c i a l outwash which would seemingly r e s t r i c t the amounts of suitable h y p o l i t h i c habitat for the in s e c t . Nunatak sur-v i v a l during the Pleistocene was not possible for G r y l l o -bla t t a . It i s reasonable to assume that the present environmental conditions on nunataks projecting above small ice f i e l d s are no more severe today than during the massive Pleistocene g l a c i a t i o n . Thus, while two theories have been advanced to explain the d i s t r i b u t i o n of organisms since the Pleistocene, namely, 1) survival north and south of the ice sheets and post 137 Pleistocene invasion into areas that were ice-covered, and 2) survival within the glaciated areas i n ice-fr e e r e f u g i a , either can be applied to explain the present d i s t r i b u t i o n of those species of Gryllobla tta which occur within the boundaries of continental g l a c i a t i o n . The Coast-Cascade and Sierra Nevada d i s t r i b u t i o n s of Gryllobla tta can be explained by l a t e Pleistocene survival at elevations below the borders of the summit ice sheets. The presence of a single species of Gry l l o b l a t t a occurring throughout the .Rocky Mountain C o r d i l l e r a , and the present d i s t r i b u t i o n of many populations south of the margins of maximum ice advance, indicate l a t e Pleistocene s u r v i v a l of G. c. campodeiformis in Montana and Wyoming. The present d i s t r i b u t i o n of G. c. campodelformis along the crest of the Rocky Mountain Cor d i l l e r a regions north of the boundaries of continental g l a c i a t i o n suggests migra-tion along highlands during g l a c i a l recession some 9000 years ago. The d i s t r i b u t i o n pattern and present i s o l a t i o n suggest la t e Pleistocene survival of G. c. at ha pa ska and G. c. nahanni in the refugia of the Liard and Nahanni ranges. The close r e l a t i o n s h i p of these subspecies further suggests a pre-late Pleistocene common o r i g i n with i s o l a t i o n of G. c. atha pa ska and G. c. nahanni during late Pleistocene. The present d i s t r i b u t i o n and taxonomic relationships of G. scudderi and G. occidentalis plus geologic evidence indicate a l a t e Pleistocene survival south of the Cordilleran 138 ic e sheet and between the Fraser lobe and summit ice f i e l d s . The very small population of G. scudderi and the assumed subspecies d i s t r i b u t i o n of G. occidentalis t e n t a t i v e l y suggest some disp e r s a l as recent as Neoglacial, beginning 3500 to 2000 years ago. The sequence of volcanism, Fraser advance and r e -cession, and Neoglacial events suggest a recent d i s t r i -bution of G. skagitensis that occurs i n the Glacier Peak region. The pioneering stock of the present population most l i k e l y survived the l a t e Pleistocene and Hypsithermal periods to the east of Glacier Peak or south toward Mt. Ra i n i e r . The continual g l a c i a t i o n since l a t e Pleistocene and the post Pleistocene volcanic a c t i v i t y in the l o c a l i t y of the Mt. Rainier G r y l l o b l a t t a suggest that these are very recent inhabitants i n the i r present l o c a l i t y . The current i s o l a t i o n of G. chirugica to the ice caves i n the lava f i e l d s of Mt. St. Helens and the past geologic and climatic history of the region indicate a l a t e P l e i s t o -cene d i s t r i b u t i o n approximately 8000 to 9000 years o l d . Gr y l l o b l a t t a were probably hypolithion inhabitants i n the region during the l a t e Pleistocene, 40,000 to 9000 years ago. However, with the termination of l a t e Pleistocene g l a c i a t i o n and the beginning of the Hypsithermal period, G. chirugica were forced to retreat to the ice caves. The d i s t r i b u t i o n of Gr y l l o b l a t t a species i n the High Cascades depicts a pattern of insul a r species and a center 139 of recent sympatric d i s t r i b u t i o n i n the hypolithion of the alpine-subalpine. The present d i s t r i b u t i o n must have formed after the volcanic and Neoglacial periods that began approximately 3500 years ago. The number of species and the presence of sympatric forms suggest systematic d i f f e r e n t i a t i o n well i n the past. The Southern Cascade and Modoc Plateau-Basin Range Gr y l l o b l a t t a are presently insular populations r e s t r i c t e d to the major volcanic peaks or ice caves of the lava deserts. The general cavernicolous d i s t r i b u t i o n predates the Hypsi-thermal i n t e r v a l with possibly some short distance di s p e r s a l during the Neoglacial. The i s o l a t i o n of the alpine-subalpine species was most certainly pre-Hypsithermal. The present d i s t r i b u t i o n of that species on the major peaks has taken place since the cessation of Hypsithermal-Neoglacia1 volcanic a c t i v i t y . The fact that many of the is o l a t e d populations are con-s p e c i f i c also indicates a rather recent dispersal to the present habitats. The d i s t r i b u t i o n patterns i n the Sierra Nevada shows scattered "pockets" of Gr y l l o b l a t t a existing the length of the range. Taxonomically, there are two d i s t i n c t groups i n the Cor d i l l e r a : a northern group more closely r e l a t e d to the Southern Cascade-Modoc-Basin Range species, and a southern species group. The northern populations most l i k e l y occupied the plateau regions to the north during 140 l a t e Pleistocene, the present d i s t r i b u t i o n being estab-l i s h e d on g l a c i a l recession and probably with no Neoglaci. re-adjustment. The topographical i s o l a t i o n of the d i f f e r e n t species suggests systematic d i f f e r e n t i a t i o n before the l a t e P l e i stocene. IV. SYSTEMATIC POSITION BY NUMERICAL OF GRYLLOBLATTODEA ANALYSIS 141 Introduction I t i s now generally agreed that the Grylloblattodea should be placed i n the orthopteroid group of insects with the Dictyoptera, Phasmida, Orthoptera, Isoptera, Plecoptera, Erabioptera and Zoraptera. These consitute a closely r e l a t e d group of exopterygotes, the Polyneoptera of Martynov (1938). The r e l a t i v e p o s i t i o n of the g r y l l o -b l a t t i d s within the orthopteroid complex has never been se t t l e d s a t i s f a c t o r i l y . From the phylogenetic standpoint, the G r y l l o b l a t t i d a e are of exceptional i n t e r e s t i n that they combine a mosaic of features found i n the other orthopteroids, as well as unique morphological characters that are viewed as primitive and r e s t r i c t e d • t o the taxon. For example, the head i s dermapteroid, the eyes are i s o p t e r o i d , and the antennae are l i k e Timea (Phasmida). The legs and t a r s i are si m i l a r to the Dictyoptera and Isoptera. The ovipositor resembles the Tettigoniidae (Orthoptera), but i s less developed, while the male g e n i t a l i a are asymmetrical and resemble i n part those of the Dictyoptera. The cerci of both sexes are l i k e those found only i n the Dictyoptera. Characters unique to the G r y l l o b l a t t i d a e include the lack of a 142 s u b g e n i t a l p l a t e i n t h e female and t h e o c c u r r e n c e o f t h r e e f r e e t h o r a c i c segments t h a t have r e t a i n e d t h e i r p r i m i t i v e m u s c u l a r c o n n e c t i o n s . The v e n t r a l n e r v e c o r d i s composed o f seven f r e e p a i r s o f abdominal g a n g l i a , as compared w i t h f i v e o r s i x i n t h e o t h e r o r t h o p t e r o i d s . The p r e s e n c e o f s t e r n a l s p i n a e f o l l o w i n g a l l t h r e e t h o r a c i c segments d i f f e r s - from a l l o t h e r l i v i n g , o r t h o p t e r o i d s i n w h i c h a t h i r d s p i n a i s a b s e n t . The p o s s e s s i o n o f a c o m b i n a t i o n o f c h a r a c t e r s t h a t a r e e x h i b i t e d a l s o i n o t h e r groups has made i t d i f f i c u l t t o s e t t l e the taxonomic p o s i t i o n o f t h e G r y l l o b l a t t i d a e . The p h e n e t i c and p h y l o g e n e t i c a f f i n i t i e s o f t h e t a x o n t o the o t h e r o r t h o p t e r o i d s have g i v e n r i s e t o much s p e c u l a t i o n and d i v e r g e n c e o f o p i n i o n . I n t h e o r i g i n a l d e s c r i p t i o n , Walker (1914) t r e a t e d G r y l l o b l a t t a as the t y p e genus o f a new f a m i l y w i t h i n t h e O r t h o p t e r a ( s e n . . l a t . ) . Crampton (1915) e l e v a t e d G r y l l o -b l a t t i d a e t o o r d i n a l s t a t u s as N o t o p t e r a . Other s u b o r d i n a l and o r d i n a l names proposed i n c l u d e G r y l l o b l a t t a r i a ( B r u n e r , 1 9 15), G r y l l o b l a t t o i d e a (Brues and M e l a n d e r , 1915), and G r y l l o b l a t t o d e a (Brues and Mel a n d e r , 1932). Much c o n f u s i o n e x i s t s i n the l i t e r a t u r e s i n c e a l l the above names a r e used i n v a r i o u s p a r t s o f t h e w o r l d . I n N o r t h A m e r i c a t h e G r y l l o b l a t t o d e a ^ i s u s u a l l y p l a c e d w i t h i n the O r t h o p t e r a G r y l l o b l a t t o d e a i s adopted here as t h e c o r r e c t name, f o l l o w i n g t h e s u g g e s t i o n o f Essig. (1942:105, f o o t n o t e ) 143 ( e.g. by Borror and DeLong, 1971), The evolutionary a f f i n i t i e s have been the subject of many pu b l i c a t i o n s . Walker (1933) considered G r y l l o b l a t t i d a e to have a close a f f i n i t y to the ancestor of the S a l t a t o r i a of the Orthoptera. After further morphological studies (Walker, 1938), he placed them between B l a t t a r i a (Dicty-optera) and the S a l t a t o r i a (Orthoptera), and l a t e r he placed them nearest to the l i v i n g Ensifera (sen. s t r . ) (Walker, 1943). Crampton (1915) considered that G r y l l o b l a t t a occupies a position intermediate between the Dermaptera and Isoptera and thought i t to be the nearest l i v i n g representative of the common ancestors of the Gryllidae and Tett i g o n i i d a e . Crampton (1917) l a t e r changed his mind and considered • Gr y l l o b l a t t a to be intermediate between the mantids and embiids as well as possibly related to the Orthoptera (sen. str . ) and Phasmida. A detailed discussion of Crampton's sixteen papers (1915-1938) on the a f f i n i t i e s of G r y l l o b l a t t a i s not necessary. In the course of his work he associated G r y l l o b l a t t a with l i t e r a l l y every order of orthopteroids. Imms (1927) placed G r y l l o b l a t t a nearer to the Dicty-optera than to the S a l t a t o r i a , while Snodgrass (1937) declared i t s a f f i n i t i e s to be with both the B l a t t a r i a and the Orthoptera. Zeuner (1939, 1945) went so far as to c a l l them "recent, l i v i n g Protorthoptera," and Walker (1937) called G r y l l o b l a t t a a " l i v i n g f o s s i l " a'nd a representative. 144 of some l i n e ancestral to modern orthopteroids. The orthopteroid f o s s i l record gives no i n d i c a t i o n of the a f f i n i t i e s of the G r y l l o b l a t t i d a e . The E n s i f e r a , B l a t t a r i a and Protorthoptera are known from the Carbon-i f e r o u s , the Caelifera and Phasmida from the T r i a s s i c , the Dermaptera from the Jurassic and the Mantodea from the Tertiary (Crowson et a l , 1 9 6 7). No f o s s i l record of the Grylloblattodea i s ' presently known. Considerable confusion exists about the taxonomic status and the a f f i n i t i e s of the taxon. Many of the opinions are purely speculative, and most of the above authors r e s t r i c t e d the basis of the i r opinions to the anatomical features of p a r t i c u l a r i n t e r e s t to them at the time of pu b l i c a t i o n ; often only a single character was considered. The h i s t o r i c a l confusion of the systematic position of the taxon has made i t necessary to attempt to c l a r i f y the a f f i n i t i e s of the Grylloblattodea. vfhat seemed to be needed was not so much another consensus of the opinions of e a r l i e r workers, but a synthesis of information provided by the morphological characters that are availab l e i n the orthopteroids. To compare various orthopteroid groups at the family, or higher, taxonomic l e v e l , using a wide range of anatomical characters, presents many d i f f i c u l t i e s , both p r a c t i c a l and t h e o r e t i c a l . Only the advent of high speed computers has 145 made i t possible to assess large masses of comparative data i n a reasonable length of time. Since the pioneering work of Bordas (I898), many authors have commented on the d e s i r a b i l i t y of basing com-parisons between various groups of the orthopteroid insects on a wide range of characters. To date, only two quanti-t a t i v e assessments of the a f f i n i t i e s of the higher orthop-teran taxa have been attempted ( G i l e s , 1963; B l a c k i t h and B l a c k i t h , 1968). Both .studies s u p e r f i c i a l l y discussed the a f f i n i t i e s of G r y l l o b l a t t i d a e . B l a c k i t h and B l a c k i t h (1968) had to r e l y upon drawings of G r y l l o b l a t t a , and from t h e i r numerical analysis considered the taxon to have greatest a f f i n i t y to the E n s i f e r a . G r y l l o b l a t t a was not included i n the cladograms and dendrograms by B l a c k i t h and B l a c k i t h (1968). In a l a t e r presentation of orthopteroid a f f i n i t i e s (Blackith and Reyment, 1971). G r y l l o b l a t t a was not discussed. Materials and Methods Detailed presentation of the various methods employed i n numerical taxonomic analysis can be found i n Sokal and Sneath (1963) and B l a c k i t h and Reyment (1971). At the subfamily l e v e l or below, there i s much to recommend the use of counts of d i s c r e t e variables and/or l i n e a r measure-ments ( B l a c k i t h , 1965). At the family l e v e l or above, the need i s mainly to make the measures somehow represent the 146 groups under comparison. Almost a l l the applications of such numerical techniques have been to comparisons at the generic l e v e l or belovr. The largest number of studies, i n which numerical taxonomic comparisons among insects have been made, use counts of d i s s i m i l a r i t i e s based on the methods of Sokal and Sneath (1963). The more recent approach by Camin and Sokal (1965), considered by the authors to be phylogenetic, requires that the inv e s t i g a t o r can so order, by weighting i f necessary, multi-state characters that his sequence of states follows the evolutionary sequence. I fin d the Camin-Sokal approach questionable because analysis must be preceded by judgments on the phylogeny that can influence the r e s u l t s . In my study a non-weighted analysis was employed. It was based i n part on that of Edwards and C a v a l l i - S f or za (1964, 1965), which u t i l i z e s a count of the number of d i s s i m i l a r i t i e s among the various groups i n the chosen suite of characters. This i s followed by a cluster analysis to y i e l d the shortest connection (generalized distance) between clusters (Prim, 1957), and the distance between clusters i s employed to construct the subdivisions of the suite of characters into the branches of the dendrogram. An alternate method was also used to check the r e -l i a b i l i t y of the analysis of d i s s i m i l a r i t i e s that show possible a f f i n i t y . This involved the development of a 147 s i m i l a r i t y index by cluster analysis of matching characters. This allowed the construction of a dendrogram i n which the lengths of the arms are equal to the s i m i l a r i t i e s and the branching equals the smallest distance c o e f f i c i e n t between a cluster p a i r . Such a dendrogram shows phenetic r e l a t i o n -ships based on degree of matching characters. The anatomical characters employed by Giles (1963) and B l a c k i t h and B l a c k i t h (1968) were used i n my a n a l y s i s , with modifications necessary to include G r y l l o b l a t t i d a e . A t o t a l of 80 a t t r i b u t e s were adapted from B l a c k i t h and B l a c k i t h (1968), and include features of the integument, nervous system, alimentary canal, circulatory system and musculature. Eighty-four of the 283 characters o r i g i n a l l y considered by Giles (1963) i n the analysis of the dermap-teran a f f i n i t i e s were chosen; these were li m i t e d to the external morphology (Appendix I I ) . Since the charac-ters used i n each suite were not chosen at random, and characters with a t t r i b u t e s shared by a l l of the groups would be of no discriminatory value, a p o s t e r i o r i weighting i s inevitable (Blackith and Reyment, 1971). I used fresh material wherever possible and supple-mented i t with descriptions from the l i t e r a t u r e . The complete delineation of a l l species i n any taxonomic hierarchy would require the examination of an impracticable number of organisms. I used one species as the "exemplar," the term employed by Sokal and Sneath (1963), of a higher 148 taxon. The r i s k that such "exemplars" might deviate widely from the means for t h e i r taxa must always be recog-nized. My analysis was based on material from the Mantodea (Stagmomantis C a r o l i n a ) , B l a t t a r i a (Perinlaneta amerlcana ), Dermaptera (Anisolabis maritima), Phasmida (Anlsomorpha sp . ) f Tettigoniidae (Ceuthophilus sp.), Gryllidae (Gryllus  a s s i m i l i s ) . Acrididae (Locusta mipratoria) and G r y l l o -blattodea ( G r y l l o b l a t t a campodeiformis). The suites of characters were transformed into Fortran IV by Mr. Borden and the basic data matrix computed on an IBM 1130. The program may be obtained from Mr. Borden, Computer Center, Zoology Department, University of B r i t i s h Columbia. Results Figure 12 presents the analysis of the number of d i s s i m i l a r i t i e s of 84 external morphological characters based on the shortest connection network between cl u s t e r s . The arrangement of the insects does not s i g n i f y l i n e a r phylogenetic r e l a t i o n s h i p s . The dendrograms are a two-dimensional representation of a multi-dimension model and the position of the branching i s the s i g n i f i c a n t feature i n showing degrees of phenetic r e l a t i o n s h i p s . For example, Figure 12 implies that the Acrididae has a greater s i m i l a r i t y to the Phasmida than to either the Tettigoniidae or the 149 Figure 1 2 . Dendrogram showing d i s s i m i l a r i t y analysis of 84 external characters, modified from Giles ( 1963) . Numbers at branchings equal generalized distance between clusters based on shortest connection network of Prim (1957). 50 G e n e r a l i z e d d i s t a n c e 40 30 i 20 10 4 1 30 3 3 3 0 2 0 2 0 2 6 D e r m a p t e r a G r y l l o b l a t t o d e a M a n t o d e a B l a t t a r i a t P h a s m i d a T e t t i g o n i i d a e G r y l l i d a e A c r i d i d a e P r i m : s h o r t e s t c o n n e c t i o n n e t w o r k ( 8 4 c h a r a c t e r s ) 150 G r y l l i d a e . The dendrogram also shows that the Phasmida and the Dictyoptera (Mantodea and B l a t t a r i a ) are more l i k e Grylloblattodea and Dermaptera than either i s to the Ensifera (Tettigoniidae and Gry l l i d a e ) or to the Caelifera (Acrididae). Therefore, the dendrogram i s a diagram of only phenetic r e l a t i o n s h i p s , and any attempt to read the branching as " e a r l i e r " or " l a t e r " evolutionary occurrences depends upon the assumption of equal, or nearly equal, evolutionary rates, evidences of which are unknown i n these taxa. The dendrogram agrees i n part with conventional c l a s s i f i c a t i o n of the groups (Imms, 1957). For example, the Tettigoniidae and Gryl l i d a e branch at a generalized distance of 20, which, i n the dendrogram, corresponds to the superfamily l e v e l . Modern c l a s s i f i c a t i o n places these two groups as•superfamilies of the suborder Ensifera (Imms, 1957). The generalized distance of the branching of the Caelifera and Ensifera corresponds to the suborder l e v e l . The generalized distance of the Dermaptera, Phasmida, Orthoptera and Dictyoptera equals the ordinal l e v e l i n th i s dendrogram. The presently c l a s s i f i e d suborders, Mantodea and B l a t t a r i a , are depicted at a d i s s i m i l a r i t y distance equal to the superfamily i n t h i s analysis of external characters. The Grylloblattodea, considered by by many to belong to the order Orthoptera, branches at the l e v e l of order. 151 Figure 13 represents an analysis of d i s s i m i l a r i t i e s , using a suite of 80 characters chosen from both external and i n t e r n a l anatomy. The Dictyoptera branch . from the other taxa at the ordinal l e v e l , but d i f f e r from Figure 12 by showing closer phenetic a f f i n i t i e s to the Dermaptera and Grylloblattodea. In the analysis shown i n Figure 13 the Acrididae (Caelifera) branch at the greatest general-ized distance and shows l i t t l e a f f i n i t y to the E n s i f e r a . The changes i n the degree of phenetic a f f i n i t y of the groups i n the two dendrograms graphically i l l u s t r a t e the danger of conferring relationships based on a single suite of features. : Figure 14 i s an analysis combining the suites of characters (164) used i n Figures 12andl3. This dendrogram depicts a compromise between the d i f f e r e n t degrees of a f f i n i t y suggested i n Figures 12 andl3. The generalized distances, equivalent to taxonomic ranking, do not change appreciably. The Dictyoptera branch ' at the greatest generalized distance, representing less phenetic a f f i n i t i e s to the other taxa than shown in Figures 12 andl3. In Figure 14, the Mantodea and B l a t t a r i a again remain-at the super-family l e v e l . The Acrididae (Caelifera) agrees i n the phenetic a f f i n i t y to the E n s i f e r a , as i n Figurel2. The greater generalized distance of ordinal l e v e l s i n Figure 14 suggests subordinal status for Grylloblattodea and Dermap-tera that i s equal to the Ensifera and Caelifera i n the Orthoptera. 152 Figure 13. Dendrogram showing d i s s i m i l a r i t y analysis of 80 external and i n t e r n a l characters, modified from B l a c k i t h and B l a c k i t h (1968). Same construction as i n Figure 12. Generalized distance 50 l 40 l 30 i — 20 l 10 _ J _ o _ l 30 3 4 27 1 9 16 2 Mantodea 6 Blattaria 5 Dermaptera 8 Grylloblattodea 4 Phasmida 1 Tettigoniidae 2 Gryl l idae 3 Acrididae Prim : shortest connection network (80 characters) 153 Figure 14. Dendrogram showing d i s s i m i l a r i t y analysis of 164 characters, combining those used i n Figures 12 and 13. Same construction as i n Figure 12. 50 Generalized distance 40 30 20 I l l 10 i o —I 37 35 2 8 30 1 9 18 Mantodea Blattaria Dermaptera Grylloblattodea Phasmida Tettigoniidae Gryllidae Acrididae Prim : shortest connection network (164 characters) 154 Figure 15 represents the phenetic a f f i n i t i e s obtained by a s i m i l a r i t y analysis of the 84 external characters used i n Figure 12. The Ensifera agree with Figurel2, which i s a d i s s i m i l a r i t y a n a l y s i s . The Acrididae,(. Caelifera) and Ensifera depict possible subordinal position i n Figurel5. In Figure 15 the Phasmida i s as i n Figurel2, i n that i t shows s i m i l a r i t y to the Orthoptera, but shows closer a f f i n i t i e s to the Grylloblattodea and the Dictyoptera, even though the common phenetic a f f i n i t i e s are the same i n each f i g u r e . The phenetic relat i o n s h i p of the Dictyoptera i s i n general agreement with Figure 12. The number of matching characters between the Mantodea and the B l a t t a r i a (78 per cent) hardly j u s t i f i e s subordinal status as now ranked i n conventional c l a s s i f i c a t i o n . In Figure 15 the Dermaptera shares only 13 per cent of the characters with the other orthopteroids. The Grylloblattodea departs r a d i c a l l y from the d i s s i m i l a r i t y analysis of FIgurel2, i n which the taxon branches from a base shared with the Dermaptera. Figure 15places the Grylloblattodea intermediate i n phenetic a f f i n i t y between the Dictyoptera and the Phasmida-Orthoptera branches, as suggested by Walker (1938). He l a t e r changed his mind (Walker, 1943) and placed i t nearer to the E n s i f e r a . Figure 16 i s a s i m i l a r i t y index of the 80 external and i n t e r n a l characters also used i n Figure 13. The Mantodea-B l a t t a r i a again share over 75 per cent of the characters. The a f f i n i t y of the Dictyoptera to the other groups i s i n 155 Figure 15. Dendrogram showing s i m i l a r i t y analysis of 84 external characters used i n Figure 12. Length of each arm equals number of s i m i l a r i t i e s and branching equals shortest distance c o e f f i c i e n t between a cluster p a i r . /JT* o Index Number of matching characters 10 i 20 30 40 i 50 i 60 i 70 80 90 Dermaptera 11 19 11 31 4 6 6 7 Mantodea Blat tar ia Gry l loblat todea Phasmida - Tettigoniidae 8 3 L Gry l l idae Acr id idae Simi lar i ty index (84 characters) 156 Figure 16. Dendrogram showing s i m i l a r i t y analysis of 80 external and in t e r n a l characters used i n Figure 13. 'Same construction as i n Figure 15. 0 Index Number of matching charac te rs 20 30 40 50 _J I I L_ 60 i 70 80 90 2 2 33 2 8 2 5 34 70 8 2 Mantodea Blattaria Dermaptera Phasmida Grylloblattodea Tettigoniidae Gryll idae Acrididae Similarity index (80 charac ters) 157 general agreement with the other dendrograms. The Acrididae shows few s i m i l a r i t i e s with the E n s i f e r a , and thi s agrees with F i g u r e l J . In l i g h t of present knowledge and the usually accepted c l a s s i f i c a t i o n of the rel a t i o n s h i p of the Ensifera and Caelifera as suborders of the Orthoptera, t h i s placement i s to be suspect. The position of Phasmida i s i n general agreement with the other dendrograms which use the same suite of characters and have close a f f i n i t i e s to the Dermaptera. Grylloblattodea show some a f f i n i t y to the E n s i f e r a , but have closer a f f i n i t i e s to the Dermaptera and Phasmida. Figure 17 presents a dendrogram of a s i m i l a r i t y analysis of the combined suites of characters as used i n Figure 14. The dendrogram of 164 characters i s again a compromise of the dendrograms for each separate s u i t e . The branches of the s i m i l a r i t y index are comparable to the generalized distances shown i n Figurel4. The Orthoptera branch from the common base which also leads to the Dermaptera, G r y l l o -blattodea, and Phasmida; whereas, i n Figure 15, the common a f f i n i t y i s with the Phasmida. In Figure 16 the Ensifera have the closest a f f i n i t y to the Grylloblattodea. In Figure 17the Dermaptera and Grylloblattodea have closer a f f i n i t i e s to each other than to the other taxa, and these agree with the relationships shown i n Figurel5. The small number of characters i n common would seem to warrant at least sub-ordinal rank. The a f f i n i t i e s of the Dictyoptera are i n 158 F i g u r e 17. Dendrogram showing s i m i l a r i t y a n a l y s i s of 164 c h a r a c t e r s used i n F i g u r e 14. Same c o n s t r u c t i o n as i n f i g u r e 15. o Index Number of matching charac te rs 10 _j 20 30 1_ 40 50 I 3 2 Mantodea Blattaria 1 0 16 «— 12 18 2 6 3 7 Dermaptera Grylloblattodea Phasmida Tettigon i idae Gryll idae Acrididae Similarity index (164 characters) 159 agreement with present knowledge of the group. The A c r i d -idae shows less phenetic a f f i n i t y to the Ensifera than would be expected from the current taxonomic groupings of the Orthoptera. Discussion An attempt has been made to determine the r e l a t i o n -ship of the Grylloblattodea to the other orthopteroid orders by an objective method using a large number of morphological characters. Although the dendrograms of t h i s study are not f u l l y consistent i n taxonomic hierarchy, they present a pattern of a f f i n i t i e s that some current opinions (e.g. Sharov, 1968) would support as a reasonable account of orthop-t e r o i d phenetic r e l a t i o n s h i p s . For example, the T e t t i -goniidae and Gr y l l i d a e show a high a f f i n i t y to each other i n a l l dendrograms. This implied, r e l a t i o n s h i p i s consistent with t h e i r p o s i t i o n i n several recent taxonomic works (Brues, Melander, and Carpenter, 1954; Borror and DeLong, 1971). The percentage of s i m i l a r i t i e s and the generalized distance for the Ensifera are at a l e v e l consistent with the accepted subordinal status of the taxon (Borror and DeLong, 1971). The phenetic r e l a t i o n s h i p of the Caelifera to the Ensifera i n Figures 14 and 17 might be at a higher taxonomic 160 l e v e l than would be acceptable by some authors (Ander, 1939; Rehn, 1952). In Figures 13 and 16 the remote a f f i n i t y of the Caelifera to the orthopteroid stem a t , or below, the ordinal l e v e l i s certainly not i n accordance with the current systematic concepts of the relationships between the Ensifera and C a e l i f e r a . Possibly the use of a suite of characters chosen from both the i n t e r n a l and external morphology has made the Caelifera more d i s t i n c t i v e than previously suspected. This tenuous a f f i n i t y was also found by BlacKith and B l a c k i t h (1968) i n t h e i r analysis of Australian orxhopteroids. In a study of redundance and non-selective (no change) characters, Le Quesne (1972) chose from the 92 characters used by B l a c k i t h and B l a c k i t h (1968) only those that behaved as i f they were uniquely derived. In an analysis based on Le Quesne's c r i t e r i a , 20 to 26 characters out of the o r i g i n a l suite were used by Le Quesne (1972) i n a re-assessment of phenetic r e l a -tionships i n the orthopteroids. The dendrogram that he constructed, did not d i f f e r in d e t a i l from the e a r l i e r analysis (Blackith. and B l a c k i t h , 1968). Le Quesne (1972) infe r r e d that there has been a considerable degree of r e p e t i t i o n in the evolution of the orthopteran group. These analyses suggest that a possible re-evaluation of the systematic r e l a t i o n s h i p s of the Ensifera and Caelifera i s needed. The close association of the Dictyoptera i n my dendro-grams does n o t agree w i t h Rehn (1951). The B l a t t a r i a and Mantodea show over 75 per cent s i m i l a r i t y o f c h a r a c t e r s ( F i g u r e s 15,16, a n d l 7 ) and a c l o s e g e n e r a l i z e d d i s t a n c e from each o t h e r i n t h e r e m a i n i n g dendrograms. One might r e g a r d them as more a p p r o p r i a t e l y t r e a t e d a t t h e super-f a m i l y l e v e l i n s t e a d o f a t the c u r r e n t s u b o r d i n a l r a n k . T h i s c l o s e agreement o f a f f i n i t i e s was a l s o found by B l a c k i t h and B l a c k i t h (1968) i n t h e i r study o f A u s t r a l i a n o r t h o p t e r o i d s . The c l o s e a s s o c i a t i o n o f the G r y l l o b l a t t o d e a w i t h t he Dermaptera i n my a n a l y s i s does not agree w i t h t h a t o f B o r r o r and DeLong (1971). I t i s not n e c e s s a r y , however, t o r e a d any p h y l o g e n e t i c i m p l i c a t i o n s i n t o t h i s p h e n e t i c ' S i m i l a r i t y . N e i t h e r t h e G r y l l o b l a t t o d e a nor t h e Dermaptera s h o u l d be c o n s i d e r e d as a n c e s t r a l t o the o t h e r , and whether t h e y shared c l o s e e v o l u t i o n a r y l i n e a g e i s d e b a t a b l e . A s u r p r i s i n g f e a t u r e i s the c l o s e p h e n e t i c a f f i n i t y o f the Phasmida, Dermaptera and G r y l l o b l a t t o d e a . T e n t a t i v e l y , t h e n , one c o u l d a s s i m i l a t e t he complex i n t o a. s i n g l e h i g h e r t a x o n . A t t he p r e s e n t time t h e r e i s no agreement among a u t h o r i t i e s about t h e h i g h e r c l a s s i f i c a t i o n o f t h e o r t h o p -t e r o i d s . I do n o t a d v o c a t e , a t p r e s e n t , any w h o l e s a l e u p g r a d i n g o f t a x a , but-am s i m p l y d r a w i n g a t t e n t i o n t o the use o f n u m e r i c a l methods o f i n v e s t i g a t i o n as a s t a r t f o r a c o n s i s t e n c y o f o r t h o p t e r o i d taxonomy. 162 Modern c l a s s i f i c a t i o n s of the orthopteroids consider Dictyoptera, Dermaptera, Orthoptera and Phasmida as orders. The phenetic a f f i n i t i e s and relationships of the G r y l l o -blattodea, as shown in the dendrograms here presented, place the taxon at the ord i n a l , rather than family, rank. In order to achieve some degree of consistency i n the hl e r a r c h i a l ranking among the orthopteroids, derived from morphological a f f i n i t i e s , I support the proposition that the most acceptable systematic rank of the Grylloblattodea Is that of an order. V. LIPID ANALYSIS 163 I. Introduction Composition of insect l i p i d s i s a topic of considera-ble current i n t e r e s t , and a number of excellent reviews of the subject have appeared i n the l a s t decade (Fast, 1964, 1970; Gilby, 1965; Gilmour, 1965, 1966). The l i p i d s of insects generally show a s i m i l a r i t y i n composition to those found i n other animals. The existing knowledge reveals that the insect l i p i d metabolism i s for the most part similar to that of vertebrates (Gilmour, 1966; K i n s e l l a , 1966a). The quantitative and q u a l i t l a t i v e composition of the fatt y acids i n insect l i p i d s have received extensive attention since g a s - l i q u i d chromatography became avail a b l e for microanalysis. Unfortunately, the majority of studies reported the fat t y acid composition of t o t a l or neutral and/or polar l i p i d f r a c t i o n s . The reports of s p e c i f i c l i p i d classes are predominantly for t r i g l y c e r i d e s and phospholipids . ( i e . Lambremont, Blum and Schrader, 1964; Fast, 1966; Harlow, Lumb and Wood, 1969; Beenakkers and Scheres, 1971). Nevertheless, certain generalizations have been put fo r t h : 1) there are s p e c i e s - s p e c i f i c differences, both / qu a l i t a t i v e and quantitative, i n the major l i p i d classes and i n fat t y acid composition; 2) while species differences 164 are apparent, there i s present a s p e c i f i c composition pattern of fatty acids for many higher taxa i n the Insecta; 3) many insects are unable to synthesize l i n o l e i c (18 ) and l i h o l e n i c (18^) acids and these are required i n the d i e t ( H i l d i t c h and Williams, 1964; House, 1965; Lambremont, Stein and Bennett, 1965); 4) the composition of fat t y acids i n insect l i p i d s varies depending on the fatty acids present i n the d i e t (Lambremont, Blum and Schrader, 1964; Nelson and Sukkestad, 1968; Schaefer, 1968; Vanderzant, 1968); 5) there i s a co r r e l a t i o n between environmental temperature and fa t t y a c i d composition. E a r l i e r works reviewed by Fast (1964) suggest the saturation of the fatty acids increases with temperature, or conversely, l i p i d s i n insects l i v i n g at, or acclimated to., lower temperatures have more unsaturated fa t t y acids. The l i t e r a t u r e on insects supports, with rare exceptions, s p e c i f i c species and higher taxa differences of l i p i d and fatt y acid composition. There are as many exceptions reported for dietary requirements, fatty acid- d i e t r e l a -tionship and ef f e c t s of temperature on fatty acid composition as there are substantiating studies. Major exceptions to ess e n t i a l dietary requirements of 18 and 18^ may be found i n the aphids (Strong, 1963; Bowie and Cameron, 1965), coccids (Tamaki and Kawai, 1968), certain Diptera and Lepidoptera (Chippendale, Beck and Strong, 1964; Fast, 1966; Vanderzant, 1968). Numerous reports suggesting that d i e t does not d i r e c t l y a f f e c t fatty acid composition are 165 known ( i e . Gilmour, 1961; Saha, Randell and Riegert, 196.6 ; Keith, 1967; Nakasone and Ito, 1967; Moore and Taft, 1970; L i p a i t z and McFarlane, 1971; Carter, Dinus and Smythe, 1972). In general, i t might be stated that i f dietary fatty acids do a f f e c t the fatty acid composition of the i n s e c t , i t i s usually only the t r i g l y c e r i d e f r a c t i o n j and synthesis can take place to supply essential fatty acids not present i n the d i e t . The e f f e c t s of environmental temperatures on f a t t y a c i d composition has been reviewed l n insects by Fast (1966) and on vertebrates by Knipprath and Mead (1968). The majority of studies confirm a c o r r e l a t i o n between temperature and degree of saturation of the fatty acids (Takata and Harwood, 1964; Keith, 1966; Knipprath and Mead, 1968; Zar, 1968; Baranska and Wlodawer, 1969; Schaefer and Washino, 1969). Van Handel (.1966), Buffington and Zar (1968) and Fast (1970) report exceptions to the correlation and object to the postulate that at low temperatures organisms attempt to maintain a constant l i q u i d i t y by use of lower melting unsaturated fatty acids. The insect studies cited have been conducted either on whole insects that overwinter aa adults i n hibernation, or eggs, or pupae, or on tissue containing t r i g l y c e r i d e s . In addition, the majority of studies have been to a c c l i m a t i z a t i o n to either high or low temperatures. To my knowledge, no insect has been i n v e s t i -gated that does not hibernate or does not pupate. Nor have there been any comparative studies between insect • 166 species which normally l i v e under d i f f e r e n t temperature conditions. When I considered the various hypotheses and counter-hypotheses, as discussed previously, G r y l l o b l a t t a seemed to be an i d e a l Insect to use i n the i n v e s t i g a t i o n of the various aspects presented. It i s generally accepted that there are s i m i l a r patterns of l i p i d and fatty acid compo-s i t i o n within a given family or order. As stated e a r l i e r , the systematic and phylogenetic p o s i t i o n of the G r y l l o b l a t t a i n the orthopteroid insects has been uncertain. I performed a comparative l i p i d and fa t t y acid analysis between G r y l l o b l a t t a , Thysanura, B l a t t a r i a and Orthoptera for the following reasons: 1) as a physiological comparison to augment my numerical morphological comparative study; 2) the l i p i d and fa t t y acid composition of Grylloblattodea has not been investigated, nor has that of any of the more "primitive" orthopteroids; 3) G r y l l o b l a t t a i s active through-out Its entire l i f e cycle within a narrow temperature range much lower than the majority of i n s e c t s . I I . Materials and Methods The insects used i n t h i s analysis are a l l females and are as follows: Isoptera (Zootermopsis a n g u s t i c o l l i s f adult alate primary reproductives, Galiano Island, B r i t i s h Columbia, collected by G. G. E. Scudder), Dictyoptera, B l a t t a r i a (Feriplaneta americana. one week old a d u l t s , 167 U. B. C. Department of Zoology c u l t u r e ) , Orthoptera, Ensifera (Gryllus a s s i m l l i s . a d u l t s , Macdonald College culture, V. R. Vickery, Quebec), Dermaptera (Anisolabis marltima. a d u l t , Mandarte Island, B r i t i s h Columbia, lan Robertson, c o l l e c t o r ) , Thysanura, iMachilidae (Pedetontus sp., Puget Sound, Washington, T. Carefoot, c o l l e c t o r ) , Grylloblattodea ( G r y l l o b l a t t a campodeiformis campodelformls, Jasper National Park, Alberta, and Gr y l l o b l a t t a lava c o l a . McKenzie Pass, Oregon, J . W. Kamp, c o l l e c t o r ) . P r i or to the extraction of the t o t a l l i p i d s , food was withheld from the specimens for 48 to 72 hours. Any foreign p a r t i c l e s were removed from the integument with compressed a i r . The specimens were then frozen i n a tared v i a l over s o l i d carbon dioxide and weighed. 1. Extraction and P u r i f i c a t i o n The extraction of the l i p i d s was by a modification of the Folch, Lees, Sloane-Stanley (1957) procedure which I found through experimentation to give the most sa t i s f a c t o r y y i e l d of the complex l i p i d s present i n i n s e c t s . A l l the reagents used were either chromatographic or pesticide grades and .checked for lipo-contaminants by gas chroma-tography. Solvents were evaporated under vacuum at temperatures less than + 5° C. The insects were homo-genized for 30 minutes at 0° C. with 20 volumes chloroform: methanol (2:1 v/v) and f i l t e r e d through a micropore funnel. The homogenate was then further homogenized and extracted 168 with 10 volumes chloroform: methanol (1:2 v/v) and 10 volumes chloroform: methanol (7:1 v/v). The f i l t r a t e s were combined and p a r t i t i o n e d by the addition of 1/5 the t o t a l f i l t r a t e volume of cold 0.9 per cent NaCl s o l u t i o n . The upper phase containing methanol, water, NaCl, proteins, amino acids and carbohydrates was removed. The i n t e r and lower phases were rinsed three times with chloroform: methanol: 0.9 per cent saline (3:47:48 v/v/v) and the chloroform-lipid phase extracted. To ensure complete removal of the saline and thus to prevent oxidation, the lipid-chloroform phase was desiccated with anhydrous magnesium sulfate under r e f r i g e r a t i o n , then evaporated under vacuum, dissolved i n 20 ml. chloroform and stored at - 10° C. i n nitrogen u n t i l further processed. 2. Separation of L i p i d Classes Three 0.5 ml aliquots of the t o t a l l i p i d extract were used for gravimetric determination of the l i p i d s extracted. The remaining extract was separated on a s i l i c i c acid column (1.5 x 14 cm.) into a neutral f r a c t i o n eluted with 200 ml. cold chloroform and a polar f r a c t i o n eluted with 200 ml. cold chloroform: methanol (1:1 v/v) (Morris, 1961). The eluates were then vacuum-evaporated to dryness, dissolved i n 20 ml. petroleum ether and three 0.5 ml. aliquots weighed for each f r a c t i o n . The neutral l i p i d f r a c t i o n was then separated into t r i g l y c e r i d e s , d i g l y c e r i d e s , raonoglycerides, free fatty a c i d s , hydro-carbons, esters and s t e r o l classes on a Florisi1, hydrated 7 per cent, column (1.5 x 15 cm.) according to the method of C a r r o l l (1961). The e f f i c i e n c y of separation into classes was monitored by thin-layer chromatography (Mangold, 1965). The s o l v e n t - l i p i d class mixtures were then refrigerated,while drying over magnesium s u l f a t e , i n a nitrogen atmosphere. The solvents of the l i p i d classes were vacuum-evaporated, l i p i d s dissolved i n 20 ml. of petroleum ether, aliquots taken for q u a n t i f i c a t i o n and stored at - 10° C. under nitrogen u n t i l s aponification and methylation. 3. Analysis of Fatty Acids by Gas Chromatography Aliquots of the t r i - , d i - and monoglyceride and free fatty acid classes were saponified with 0.5 N methanolic sodium hydroxide, e s t e r i f i e d with boron fluoride-methanol 14 per cent w/v using the procedure of Metcalfe, Schmitz and Pelka (1966). The fatty acid methyl esters were dissolved i n spectroanalyzed n-hexone for separation by gas chromatography. Qualitative and quantitative analyses of the f a t t y acid methyl esters present i n the four classes were performed on a Varian-Aerograph Model 1820 Gas Chromatograph equipped with dual d i f f e r e n t i a l flows Ionization detectors. Columns,6 or 12 feet long, 1/8-inch outside diameter, of s t a i n l e s s s t e e l , packed with 20 per cent DEGS (diethylene 170 g l y c o l succinate) on DMCS-chromosorb W, 60/80 mesh, main-tained at 4 200° C. and a c a r r i e r gas flow at 40 ml./ min. of Methyl esters of fatty a c i d standards for compara-ti v e purposes were purchased from Analabs, Inc., Conn-e c t i c u t , and Applied Science Laboratories, C a l i f o r n i a . Chromatograph peaks were i d e n t i f i e d by comparison with retention time of the standards. Unknown peaks were ten-t a t i v e l y i d e n t i f i e d by a plot of log r e l a t i v e retention time versus number of carbon atoms (James, 1959). Quanti-f i c a t i o n as per cent/weight were made with a Disc Integra-tor coupled to a recorder output. I I I . Results and Comparisons The fat t y acid composition of free fatty acids (FFA), mono-, d i - and t r i g l y c e r i d e s of the six insect taxa analyzed, are shown i n Table III and Figures 18, 19, 20, 21, 22 and 23. 1. border Thysanura (Pedetontus) ( F i g . 18) The f a t t y acids present i n the FFA and glycerides of Pedetontus (Thysanura) resemble those of Leplsma saccharins (Thysanura) ( K i n s e l l a , I 9 6 9 ) . Quantitatively, Pedetontus displays differences from those reported for Lepisma  saccharina and Thermobia domestlca (Fast, 1970). These quantitative differences are reasonable considering other reports of generic differences (Fast, 1964) and that the Thysanura sensu. l a t . u t i l i z e d i n t h i s t h e s i s . 17 li-5 5 -$ f 00 -*° • 5 5! • s o 5! • 5 • 55 55 55 a. . *. » «• -« c* 55555 55 51 55 55 $ • '5. o > 0 55 J S - 51 P. * 2 S - - S " 5 5 55| 5 5 5 | ... s a s s s 0 ~ 0 <» » 51 55 55 55 55 $ 5 5U 5 S 3 S S 3 § 55^555 ,551555 55 51 15 $ 55 51 s s s : c a 3 2 si si s -5 H 5 n : , g $ P S # 25 R ft 8 K .S S « » «, S g 2 2 5 S 5 5 5 5 5 * . 5 5^555 55 55 55 55 55 .55 555555 5 5 35? 5 555 : s s s s • «• ' * - ~ • 55 55 5| 55 • • 55 3 o ' 5 5 = G J! 0 55 55 55 5! .5 5 5 5 ^5 5S355S 55 55 55 55 55 55 5 6 S "> 2 2 P s s e s s a s 8 " s - ; s 3 ? s s s S S 8 S S 2 $ P £ P l i p 51S 5 5 s ° -•5'555 55 55 55 55 55 5 5 5 5 5 5 5 S S 3 2 S 55.555 t S S ! ! • d 0 0 v 55 55 55 55 55 55 s. s. ° * « 555555 S 3 S 8 t K =. S. * 6. K ? :55 55 55 55 55 55 I E ? , : ; ! c - -- o 5 5 5 555 ' ."; » 5 0 j - 55 51 55 55 5 5 5 •• 5 5- 5 55 • 5 55 55 55 55 55 5 5 5 5 5 5 5 5 . 5 5 5 55.555 5 5.555 5! 55 55 51 55 55 . 2 !• • 15 3 • 5 5 .... -55 55. 55 35 • 53 • 55 .55. 55.55. • 5 • !• § • 51 51 55 5! 2 • • 5 « . . . 0 2 . . . . . . . 5 . 5! 55 • - 5 • 172 Figure 18. Representative chromatograms of Fedetontus. F: free f a t t y acid f r a c t i o n , M: monoglyceride f r a c t i o n , D: diglyceride f r a c t i o n , T: t r i g l y -ceride f r a c t i o n . Baseline d r i f t corrected; attenuation i n t e r v a l s not indicated. Number equals carbon number of fatty acid and super-s c r i p t equals number of unsaturated bonds i n molecule. 173 fat t y acid content of the esters and phospholipids i n Pedetontus were not determined. Like the majority of the l i t e r a t u r e reports on fat t y acids (Fast, 1970), only the chain lengths of 14 to 18 carbons were quantified for Thermobla domestlca. The large proportion (52.9 %) of o l e i c (18:1) and palmitic (16:0) acid agrees with reports of other insects except the aphids and coccids (Fast, 1970). The major fatty acids present i n the Insecta : m y r i s t i c (14:0), palmitic (16:0), palmitoleic (16:1), s t e a r i c (18:0), o l e i c (18:1), l i n o l e i c (18:2) and l i n o l e n i c (18:3), comprised 83.6 % of the fatt y acid content of FFA and mono-, d i -and t r i g l y c e r i d e s i n Pedetontus. The unsaturated fa t t y acids make up 59.4 % of the t o t a l , with the unsaturated 18-carbon fatty acids predominating. In Pedetontus the unsaturated a c i d s , excluding the 18-carbon fatty a c i d s , represent 13.6 % of the FFA, with 7.2 % as my r i s t o l e i c (14:1), 12.9 % of the monoglycerides including 6.7 % palmitoleic (16:1), 19.2 % of the di g l y -cerides including 10.3 % as palmitoleic a c i d , and 11.35 % of the t r i g l y c e r i d e s mainly as myris t o l e i c (9.3 % ) . The concentration of my r i s t o l e i c and palmitoleic a c i d s , higher than those of most i n s e c t s , i s d i f f i c u l t to explain since l i t t l e i s known of the fatt y acid metabolism i n Thysanura. In f a c t , l i t t l e i s known about l i p i d metabolism and fatt y a c i d biosynthesis of Insecta; that reported i s i n d i r e c t evidence supporting hypotheses extrapolated from l i p i d research of microorganisms and vertebrates. The amounts of 14:1 and 16:1 approach those concentrations reported for some Diptera (Culicidae) (Van Handel, 1966) and Hemip-tera (Aphididae) (Tamaki and Kawai, 1968). 2. Order B l a t t a r i a (Perlplaneta) (Fig. 19) The f a t t y acid composition i n Perlplaneta amerlcana from the University of B r i t i s h Columbia culture closely resembles that of B l a t t e l l a germanlca (Krishnan, 1968) and the P. amerlcana used by K l n s e l l a (1966b). Quantification of my analysis d i f f e r s for the reasons discussed under Pedetontus sp. Oleic a c i d (18:1) was the largest com-ponent i n the l i p i d classes analyzed and accounts for 29.5 to 40.9 % (mean, 36.31 %) of the t o t a l f a t t y a c i d f r a c t i o n . The major f a t t y acids, m y r i s t i c , p a l m i t i c , p a l m i t o l e i c , s t e a r i c , o l e i c , l i n o l e i c and l i n o l e n i c , represent 89.6.% of the fat t y acids. Unsaturated acids were represented as a mean content of 60.7 % of the t o t a l , ranging from 53.7 % i n the monoglyceride f r a c t i o n to 64.8 % i n the t r i g l y c e r i d e s . The unsaturated 18-carbon f a t t y acids comprised 88.5 % of the t o t a l unsaturated fa t t y acids present i n the classes analyzed. The remaining unsaturated acids, represented by l i n d e r i c (12:1), myri s t o l e i c (14:1), palmitoleic (16:1), eicosenoic (20:1) and erucic (22:1), composed 22.1 % of the FFA, 9.9 % of the monoglycerides, 8.3 % of the digl y c e r i d e s and 5.7 % of the t r i g l y c e r i d e 175 Figure 19. Representative baseline-corrected chromatograms of Periplaneta americana. Legends as i n Figure 18. f r a c t i o n . The long chain fatty acids, 20:1 and 22:1, are not found i n the FFA f r a c t i o n and account for 1.83 % of the t o t a l fatty a c i d content. These r e s u l t s closely resemble those of 13 other species of B l a t t e l l a as l i s t e d i n Fast (1970). 3. Order Orthoptera (Gryllua) (Fig.:20) My fat t y acid analysis of Gryllus a s s i m i l i s resembles l i t e r a t u r e reports of other G r y l l i d a e (Fast, 1967; Young, 1967; Hut chins and Martin, 1968a). The major f a t t y acids present i n the four classes are myri s t i c , palmitic, palmito-l e i c , s t e a r i c , o l e i c , l i n o l e i c , and l i n o l e n i c , which average 97.5 % of the t o t a l composition, with l i n o l e i c representing 32.98 %. The mean percentage of unsaturated fatty acids present i n the four .classes i s 66.45 %, being the highest i n the six orders of insects I investigated. The 18-carbon unsaturated acids comprise 86.3 % of the t o t a l unsaturated f a t t y acids. Oleic (18:1), l i n o l e i c (18:2), and l i n o l e n i c (18:3) constitute 53.8 % of the t o t a l f a t t y acids present i n the FFA, 62.6 % of the monoglycerlde, 65.7 % of the d i g l y c e r i d e , and 57.2 % of the t r i g l y c e r i d e f r a c t i o n . Gryllus a s s i m i l i s has more of 18:1, 18:2, and 18:3~in the monoglyceride and d i g l y c e r i d e fr a c t i o n s than the other orders and the concentrations i n the FFA and t r i g l y c e r i d e f r a c t i o n s are only exceeded by Perlplaneta. Gryllus a s s i m i l i s d i f f e r s from the other orders investigated i n that l i n o l e i c 177 Figure 20. Representative baseline-corrected chromatograms of Gryllus a s s i m i l i s . Legends as i n Figure 18. acid (18:2) i s the predominant component of the FFA and glyceride f r a c t i o n s . The Gr y l l i d a e and Tettigoniidae seem to d i f f e r from the other Orthoptera i n respect to l i n o l e i c acid concentrations. Fast (1967) reported that Gryllus  bimaculatus contained greater amounts of l i n o l e i c a c i d than o l e i c and t h i s was found also i n the Tettig o n i i d a e , Scudderla furcata (Young, 1967). The high concentration of l i n o l e i c acid may be a possible r e f l e c t i o n of dietary intake or the metabolic need for a low melting p o i n t , highly mobile long chain fat t y a c i d . 4. Order Isoptera (Zootermopsis) ( F i g . 21) Twenty f a t t y acids were detected i n the isopteran, Zoo-termopsls a n g u s t l c o l l l s . The fatt y acid composition d i f f e r s from the subterranean termites, Reticulltermes f l a v l p e s (Young, 1967; Carter, Dinus and Smythe, 1972), and i s much l i k e Macrotermes f a l c i g e r . the African fungus-growing termite (Cmelik, 1972). The major components, m y r i s t i c (14:0), palmitic (16:0), palmitoleic (16:1), s t e a r i c (18:0), o l e i c (18:2) and l i n o l e i c . (18.2), make up 80.9 % of the t o t a l fatty acids present. The carbon-18 series of o l e i c and l i n o l e i c are the major unsaturated fatty acids present. The minor unsaturated components are l a u r o l e i c ? (12:1), m y r i s t o l e i c (14:1), palmitoleic (16:1), hexadecadienolc (16:2), l i n o l e i c (18:3) and eicosenoic (20:1), and make up 12.6 % of the FFA f r a c t i o n , 6.8 % of the monoglycerides, 13.3 % of the di g l y c e r l d e s and 12 % of the t r i g l y c e r i d e s . 179 Figure 21. Representative baseline-corrected chromatograms of Zootermopsis a n g u s t i c o l l i s . Legends as i n Figure. 18. The presence of hexadecadienoic acid (16:2) i n the FFA and d i g l y c e r i d e fractions i s d i f f i c u l t to explain. Cmelik . (1972) found 16:2 to be present i n a l l fr a c t i o n s of the fungus-feeding termite but offered no explanation. It i s possible that 16:2 i s a product of oxidation via a c e t y l CoA and the Malonyl pathway from l i n o l e i c a c i d (18:2). This has been suggested by Wakll (1961) to be the mechanism i n the r a t , and Keith (1967) believes t h i s i s also true for Drosophila. The presence of 16:2 i n both Ma crotermes and Zootermopsls, which either feed on fungi or on r o t t i n g wood containing fungal raycella, suggests to me that hirogenic acid i s of dietary o r i g i n . Since hirogenic acid i s found i n only the FFA and d l g l y c e r i d f r a c t i o n s , t h i s adds support for a dietary o r i g i n , for these two fra c t i o n s function primarily as transport and synthesis components. 5. Order Dermaptera (Anlsolabis) ( F i g . 22) Twenty-one straight chain fa t t y acids (C6 to C22:l) were i d e n t i f i e d from the dermapteran, Anisolabis marltlma. The presence of 6 other compounds was detected which may be i n the branched or hydroxy fatty a c i d s . The major fat t y acids present i n the combined fractio n s were my r l s t l c (14:0), palmi t i c (16:0), palmitoleic (16:1), s t e a r i c (18:0), o l e i c (18:1), l i n o l e i c (18:2) and l i n o l e n i c (18:3) and account for 85.8 % of the t o t a l fatty acid composition. Unsaturated fat t y acids average 65.3 %, with 71.3 % i n the 181 Figure 22. Representative baseline-. corrected chromatograms of Anisolabis maritima. Legends as i n Figure 18. 182 t r i g l y c e r i d e s , 67.6 % i n the monogly cerides, 56.0 % i n the'' dig l y c e r i d e s and 54.8 % i n the FFA f r a c t i o n . The amounts of unsaturated acids present i s exceeded i n t h i s comparative analysis hy the Gryllus that contain 66.45 % and G r y l l o -b l a t t a that contain 65.8 % of the t o t a l f r a c t i o n . The unsaturated 18-carbon chain fatty acids average 40.4 % of the t o t a l f r a c t i o n s and are present i n lower concentrations than i n the other orders. Of note are the small amounts of the polyunsaturated 18 carbons which are necessary for growth and metamorphosis. The low amounts of 18:2 and 18:3 are r e f l e c t e d by the large concentrations of p a l m i t o l e i c acid (16:1) present i n An-i s o l a b i s FFA and glyceride f r a c t i o n s . The condition of low 1.8:2 and 18:3 and high 16:1 i s c h a r a c t e r i s t i c of the Diptera (Schaefer and Washino, 1969; Fast, 1970) and has been observed i n the cockroach (McGuire and Gussin, 1967) and i n moths (Schaefer, 1968). The significance of t h i s condition i s not certain and there are c o n f l i c t i n g hypo-theses that i t i s a dietary and a non-dietary fatty acid synthesis phenomenon (Fast, 1970; Madariaga et a l . 1972). Since nothing i s known of the fatty acid compo-s i t i o n of the Anisolabis d i e t , no conclusions can be drawn. Nevertheless, I suggest a tentative hypothesis for the condition i n A n i s o l a b i s . Municlo et aJL (1971) have shown that certain Diptera are capable of synthesizing saturated and mono-saturated fatty acids independently of the fa t t y acid composition of the d i e t . In Anisolabis 16:1 compensates 183 for a lack of 18-polyunsaturated acids and palmitoleic acid i s a synthesis product of mitochondria or oxidation of a fatty alcohol C14 :0 (Lambremont, 1972) and i s de-saturated to 16:0 and then elongated to 18:0 or 18:1 as Keith (1967) found to happen i n Drosophila. This method would then account for the elevated concentrations of-16:0 (palmitic acid) present i n A n l s o l a b i s . The high con-centration of 16:1 and 18:1 may be of environmental s i g -n i f i c a n c e to the genus as w i l l be discussed l a t e r . 6 . Order Grylloblattodea (Grylloblatta) ( F i g . 23) Twenty-four fatty acids were i d e n t i f i e d i n G r y l l o -b l a t t a lava cola and range i n chain length from 6 to 24 carbons. Six other un i d e n t i f i e d compounds were present and most l i k e l y are branched, hydroxy fatty acids or fat t y a l c o h o l s . The composition of the major fat t y acids i n the four fractio n s d i f f e r s from the other orders analyzed. M y r i s t i c (14:0), palmi t i c (16:0), palmitoleic (16 : l ) , s t e a r i c (18:0), o l e i c (18:1), l i n o l e i c (18:2) and l i n o -l e n i c (18:3) account for only 69.1 % of the t o t a l f a t t y acid composition In contrast to the 82 4 to 97 % found i n the other orders tested i n t h i s study. The other major fatt y acids present are m y r i s t o l e l c (14:1) and the unusual occurrence of eicosenoic (gandoic) (20:1) and l i g n o c e r l c (24:0). Unsaturated f a t t y acids make up 65.8 % of the t o t a l present i n the four f r a c t i o n s and t h i s percentage 184 Figure 23. Representative baseline-corrected chromatograms of G r y l l o b l a t t a lava cola. Legends as i n Figure 18. 185 i s exceeded only by the G r y l l u s . The unsaturated 18-carbon fa t t y acids average 41.1 % of the t o t a l acid composition, with concentrations of 19.6 % FFA, 37.9 % of t r i g l y c e r i d e s , 48.8 % of monoglycerides and 58.4 % of the d i g l y c e r i d e . f r a c t i o n . Lignoceric (24:0) has not been reported pre-viously as occurring i n insects and i t s presence i s d i f f i c u l t to explain from current knowledge of l i p i d synthesis and metabolism i n the Insecta. There i s an i n t r i g u i n g p o s s i b i l i t y which warrants further i n v e s t i g a t i o n . Faurot-Bouchet and Michel (1964) discovered i n Coccidae natural occurring esters of fa t t y acids with carbon chain lengths of 16 to 34. They i d e n t i f i e d these esters and fatty alcohols (24 to 30 carbons) from cut i c u l a r waxes. Shikata (i960) and Amin (i960) showed that lepidopteran waxes consisted of a mixture of fa t t y acid esters and alcohols of chain lengths C16 to C34. Lignoceric acid (C24) i n the G r y l l o b l a t t a i s found only i n the FFA f r a c t i o n and therefore may be an oxidation product of a cuticular waxy ester or a precursor by mitochondrial synthesis to cut i c u l a r wax components. There i s also the suggestion (Hutchins and Martin, 1968b) that a C24 could be a cleavage of an alkan or o l e f i n i c hydrocarbon or a possible precursor to the formation of such compounds. My temperature-humidity experiments show that as environmental temperature r i s e s a higher humidity i s sought by the G r y l l o b l a t t a . The integument of G r y l l o b l a t t a has few areas of heavy s c l e r i t i -zation usually associated with i n s e c t s , being l i m i t e d to 186 appendages, head capsule and the tergal segments. The remaining c u t i c l e i s t h i n and i n areas transparent. The fatty acids present i n G r y l l o b l a t t a . such as erucic (22:1), behenlc (22:0), eicosenoic (20:1), heptadecanoic (17:0), pentadecanoic (15:0), undecanoic (11:0) and nonanoic (9:0), are not generally reported as occurring i n Insecta. The few exceptions where these fatty acids are reported (Lindsay and Barlow, 1970; Yendol, 1970; Blomquist et a l , 1972; Thompson and Barlow, 1972) may well prove to be the r u l e . Recent studies of insect cutic u l a r waxes and hydrocarbons (Bursell and Clements, 1967; Hutchins and Martin, 1968b; Blomquist et a l , 1972) suggest that these compounds are most l i k e l y associated with the synthesis of waxes, hydrocarbons and esters In the c u t i c l e . As shown by Thompson and Barlow (1972), undecanoic (11:0) and nonanoic (9:0) acids are formed by s t e a r i c (18:0) and acetate, elongated to 20:0, attenu-ated to 20:1, and oxidated to C11:0 and C9:0. Erucic (22:1), behenic (22:0) and heptadecanoic (17:0) acids have been reported to be associated with secondary alcohols and wax esters i n the orthopteran, Melanplus (Blomquist et a l , 1972). In G r y l l o b l a t t a these compounds are formed at much higher concentrations than reported for other i n s e c t s . Few insects are active at the temperatures pre-ferred by the G r y l l o b l a t t a . The higher concentrations of these compounds i n G r y l l o b l a t t a suggest that they may be ess e n t i a l components of the c u t i c l e to combat water loss 187 at temperatures around 0° C. IV. Discussion of Environmental and Systematic Relationships Demonstrated by L i p i d Analysis s The c o n f l i c t i n g evidence of the effects of environ-mental temperature on fatty acid composition i n insects may be due to the fact that the studies were concerned with changes from acclimation to d i f f e r e n t varying temperatures. I know of no studies where the saturation or unsaturation of the fatty acid composition has been analyzed from insects with d i f f e r e n t habitat temperatures. The insect orders used i n t h i s study have, i n general, d i f f e r e n t temperature preferences. Periplaneta americana i s a warm-preferring form and i s indigenous from subtropical America. It i s also able to survive i n the colder temperate regions i n a r t i f i c i a l habitats such as human habitations. The thysanuran, Pedetontus. occurs i n a temperate maritime climate, but i s only found i n the cracks and crevices of the shore c l i f f s during the warmer seasons of the year when habitat temperatures are In the range of + 25 to + 5 0° C. The maritime earwig, Anisola bis marltlma. inhabits ocean beaches just above high tide mark and escapes the lower temperatures of the northern coastal climate by re t r e a t i n g under stones and beneath decaying vegetation. I found, when maintaining cultures i n the laboratory, that -f 1 0° C. i s the approximate lower l i m i t of tolerance. Zootermopsls  a n p u s t i c o l l l s . the P a c i f i c coast coniferous dampwood, or "rottenwood", termite, occurs from B r i t i s h Columbia to Baja C a l i f o r n i a . While often found at high elevations and occurring i n areas of wide temperature v a r i a t i o n , i t i s , nevertheless, a warm-adapted species. The heat of decay within the wood maintains a rather uniform temperature between 4 15 and 4 25° C. Gryllus a s s i m i l i s , the f i e l d c r i c k e t , i s a wide-ranging species, occurring from the middle of South America and.extends throughout most of North America. This genus seems to be of wide temperature tolerance, but does escape freezing temperatures by over-wintering as nymphs or retr e a t i n g into burrows or vegetation. G r y l l o b l a t t a i s a c r y o p h i l i c form r e s t r i c t e d to an optimum temperature range of - 3 to + 6° C. The insect does not hibernate and i s active at temperatures that are generally l e t h a l to most insects. I f lower temperatures are correlated with higher concentrations of the unsaturated fa t t y acids that have lower melting points than saturated ones, i t should be evident i n t h i s series of insects. In addition, the melting points of the t o t a l fatty acids present should also r e f l e c t to some degree temperature preferences or tolerances. For G r y l l o b l a t t a 4 10° C. i s the maximum l i m i t for any long term s u r v i v a l . Therefore, when considering the l i q u i d i t y of the fatty acids, I chose the 4 10° C. melting point as the reference point. The t o t a l per'cent of fatty acids present i n the four classes analyzed with a melting point above 4 10° C. i s as follows: TABLE IV ' 189 Total Per Cent Fatty Acids with M.P. above- + 10° C. Pedetontus 76.0 S.E.0.11 Zootermopsis ; '70.4 S..E.0.02 Perlplaneta 75.0 0.02 Gryllus 58.4 0.02 Anlsolabis 71.5 0.10 G r y l l o b l a t t a 43.1 o . l l . The percentages above + 10° C. are i n excellent agree-ment with habitat temperatures. Pedetontus and Perlplaneta are the two most thermophilic of the orders and form a group d i s t i n c t from the others. Anisolabis and Zootermopsis are both maritime forms and are exposed to a wider range of moderate habitat temperatures. These, too, form a pair with a somewhat lower per cent of above + 10° C. melting point fatty acids. Gryllus forms a d i s t i n c t e n t i t y , as might be predicted by i t s wide range and exposure to varying temperature f l u c t u a t i o n s . The only s t r i c t l y c r y o p h i l i c form of t h i s group, G r y l l o b l a t t a . contains merely 43.1 % of the fatty acids that have a melting point at + 10° C. or higher. This i s consistent with the habitat temperatures i f G r y l l o b l a t t a i s to remain highly a c t i v e . The percentage of unsaturated fatty acids present i n the d i f f e r e n t orders tested do not present the same d i s t i n c t pattern as found i n the melting point of fatty acids present. The t o t a l per cent of unsaturated fa t t y acids i s as follows: 190 TABLE V Total Per Cent Unsaturated Fatty Acids Pedetontus 59.4 0.01 Anisolabis 65.3 0.05 Periplaneta 61.0 0.02 G r y l l o b l a t t a 65.8 0610 Zootermopsis 63.0 0.05 Gryllus 66.4 0.01 Pedetontus and Periplaneta again form a group and i s what would be expected i f higher environmental temperatures are correlated with higher percentages of saturated fatty acids. Zootermopsis i s also consistent with a moderately temperate form. The pattern changes here i n that Aniso-l a b i s , Gryllus and G r y l l o b l a t t a a l l form a group, yet have widely d i f f e r i n g temperature preferences. Environmental temperatures are correlated with the degree of saturation of the fatty acids present i n the insects i f one considers the melting points of the un-saturated f a t t y acids. The t o t a l per cent of unsaturated fa t t y acids with a melting point above + 10° C. i s as follows: TABLE VI Total Per Cent Unsaturated Fatty Acids With M. P. Above + 10° C. Periplaneta 36.3 0.05 Zootermopsis 35.3 0.01. Pedetontus 35.3 0.01 Anisolabis 33.2 0.05 Gryllus 25.0 o.Ol G r y l l o b l a t t a 8.9 0.02 191 The order groupings are once again consistent with habitat temperatures. From these data the environmental temperatures do appear to a f f e c t the degree or percentages of saturated-unsaturated fatty acids present i n the organism. The c o n f l i c t i n g data of previous reports might be i n agree-ment i f the temperature tolerances of the d i f f e r e n t insects tested were taken into consideration. The o v e r a l l quantitative and q u a l i t a t i v e patterns of f a t t y acids i n the orders analyzed agree with the fa t t y a c i d composition found i n other insects. Each order has i t s own d i s t i n c t composition as has been found i n other studies. Nevertheless, G r y l l o b l a t t a and Anlsolabis, while each possesses a d i s t i n c t composition, more closely resemble each other than the other orders analyzed. This closer r e l a t i o n s h i p i s consistent with the morphological r e l a t i o n -ships presented earli e r . 192 VI. DISCUSSION AND CONCLUSIONS As noted in the general introduction to this t h e s i s , the Grylloblattodea are of interest owing to th e i r structure and systematic p o s i t i o n , t h e i r low temperature-high humidity preference and winter a c t i v i t y , and their disjunct d i s t r i -bution and zoogeography. These three aspects have been considered i n the present research, but much remains to be done. With respect to the climatic tolerance, my research-has shown that the two species of G r y l l o b l a t t a studied have a preferred temperature of - 3.5 to -f 5° C. at 90 to 99 per cent r e l a t i v e humidity, a r e s u l t i n general agree-ment with the r e s u l t s obtained by Kenson (1957b). The upper and lower l e t h a l l i m i t s are correspondingly low and i t i s clear that G r y l l o b l a t t a i s l i k e other winter-active insects such as Boreus (Mecoptera), Chionea (Diptera) (Chapman, 1954; Kagvar, 1971) and the carabid Pterostichus brevlcornis.(Coleoptera ) (Baust and M i l l e r , 1971, 1972) in t h i s respect. In such i n s e c t s , i t i s evident that there must have been evolutionary adaptations to temperature, permitting them to move, feed, grow and reproduce at temperatures much lower than those preferred by the majority of other i n s e c t s . In the present study, while the physiological basis of this low temperature-high humidity preference was not 193 selected as a main l i n e of inve s t i g a t i o n , the data on l i p i d composition have produced evidence relevant to environmental physiology and temperature preference of G r y l l o b l a t t a . My analysis shows that the unsaturated f a t t y acids account for 65.8 per cent of the t o t a l f a t t y acids in G r y l l o b l a t t a , of which only 8.9 per cent have melting points above + 10° C. In the temperate forms analyzed, unsaturated fa t t y acids with melting points above + 10° C. range from 25 to 36 per cent. The per-centage of fat t y acids that are l i q u i d at or below the maximum tolerated temperature suggests a phy s i o l o g i c a l mechanism i s present i n Gr y l l o b l a t t a that allows mobility of the insect and continued i n t r a - and i n t e r c e l l u l a r transport at low temperatures. The high concentration of l i q u i d fatty acids may also function as an "antifreeze" mechanism which retards freezing and nucleation of body water. In the future, i t would be worthwhile to examine the temperature preference and tolerance of coexisting species to see i f there are adaptations i n t h i s parameter per-mitting them to l i v e together, as documented i n other insects (Heath et a l , 1971; Jamieson, 1973). The immature stages should also be studied since d i f f e r e n t i n s t a r s i n a l i f e cycle may show d i f f e r e n t preferences or optima (Sherman and Watt, 1973). Many other aspects of the physiology would be worthwhile pursuing, g l y c e r o l content 194 and enzyme functions being two that seem appropriate. In general, i n insects cryoprotectants either increase cold resistance by greatly lowering freezing or supercooling points, without affording protection i n the event of ice formation, or they allow varying degrees of cold protection without necessarily profoundly lowering freezing or supercooling points (Baust and M i l l e r , 1972). The former, which frequently involves s p e c i f i c behavioral responses r e s u l t i n g i n the reduction of nucleation agents ( S a l t , 1961, 1968), seems not to occur i n G r y l l o b l a t t a , as t h i s Insect i s active and feeds during winter. Salt (1968) found cessation of feeding was an important, behavior reducing, for example, nucleation agents i n the gut, etc. In G r y l l o b l a t t a , as in Pterosti chus brevi cornls (Baust and M i l l e r , 1972), the cryoprotectant probably does not greatly influence the freezing and supercooling points. Here, the g l y c e r o l content and changes would be worthy of study, since these have been shown to have a pronounced cryoprotectant function ( S a l t , 1961; Sp'mme, 1964; Baust and M i l l e r , 1971, 1972). In t h i s context, one of the potential sources of g l y c e r o l might be the l i p i d s . The ready conversion of neutral fats to g l y c e r o l and t h e i r component fatty a c i d s , either by enzyme mediation or acid h y d r o l y s i s , could be important (Baust and M i l l e r , 1972). The adaptation to low-temperature living, by G r y l l o b l a t t a undoubtedly involves either quantitative changes i n enzymes 195 or changes i n the type of p a r t i c u l a r enzymes. Studies of such biochemical adaptations have received considerable attention i n cold-acclimated animals (Somero and Kochachka, 1971; Sjzfmme, 1972) and in fishes showing evolutionary cold adaptation (Somero, Giese and Wohlschlag, 1968), but to date similar studies i n evolutionary cold-adapted insects do not seem to have been undertaken. Gry l l o b l a t t a would be an i d e a l subject to study i n t h i s context, perhaps i n preference to Borens or Chionea because of i t s larger s i z e . Certainly, one might expect i n t h i s insect the production of enzymes which are p a r t i c u l a r l y well suited for function at the temperature to which they are adapted (see Somero and Hochachka, 1971). The narrow temperature-humidity tolerance of G r y l l o -b l a t t a seriously r e s t r i c t s the habitat that t h i s genus can occupy. Only the alpine-subalpine hypolithion and the ice cave environment seem to provide the required conditions on a more or less permanent basis. Even i n these habitats, however, the microenvironments are not universally a v a i l -able, for i n the alpine-subalpine, i t would seem that only under stones or boulders, 50-150 m. i n diameter and 30-50 cm. thick, do the required r e l a t i v e l y constant tempera-ture and high humidity occur. Such a l i m i t a t i o n and depen-dence upon p a r t i c u l a r stone size i s well known i n the hypolithion community (Hagvar and Ostbye, 1972). Likewise, i c e caves are highly variable (Halliday, 195^), but only 196 certain ones are known to contain populations of G r y l l o b l a t t a (Kamp, 1970). Here again, only under certain conditions and at certain times of the year, do the required tempera-ture-humidity conditions occur. With the narrow temperature-humidity tolerance of Gr y l l o b l a t t a documented, with the limited a v a i l a b i l i t y of permanent microhabitats evident, and the knowledge that much of the-present Nearctic range of Gr y l l o b l a t t a was glaciated during the Pleistocene (Hubbs and M i l l e r , 1948), i t i s obvious that the d i s t r i b u t i o n patterns i n western North America have been greatly changed i n the past. The present d i s t r i b u t i o n pattern has been fundamentally i n f l u -enced by the geologic and climatic events of the P l e i s t o -cene. Further, the regional, and sometimes, highly l o c a l i z e d , volcanic a c t i v i t y during the post Pleistocene, the warm dry period (Hypsithermal), and the r e b i r t h of summit and cirque g l a c i e r s , commencing approximately 2500 years ago, must have profoundly affected the d i s t r i b u t i o n patterns of G r y l l o b l a t t a . It would seem that the present d i s t r i b u t i o n of Gr y l l o -b l a t t a i n the Coast-Cascade Cordilleran has been strongly influenced by volcanism i n the post Pleistocene. The d i s t r i b u t i o n suggests two centers of dispersal for the northern Coast-Cascade species: one somewhere east of the Cordi l l e r a n crest between Mt. Baker and Glacier Peak, and the other east of the summit between Glacier Peak and Mt, Rainier. 197 The current d i s t r i b u t i o n of Gr y l l o b l a t t a and the events of the post Pleistocene indicate that there was no single center of post Pleistocene dispersal for species i n the High Cascade-Basin ranges or•the Sierra Nevada. The post Pleistocene pockets of dispersal were probably many and migration controlled by regional topography and volcanism. The early post Pleistocene center of d i s p e r s a l for Gryllobla tta campodei formis campodei formis was probably south of the Cordilleran-Keewatin ice sheet i n Montana, with subsequent northward migration to the b a r r i e r s of the Peace River region. The present d i s t r i b u t i o n pattern and systematics within G r y l l o b l a t t a suggest that the northern Great Basin and Columbia Basin have been e f f e c t i v e b a r r i e r s to d i s p e r s a l between the Rocky Mountains and the Cascades. These basins and ranges have evidently acted as b a r r i e r s since the beginning of the l a t e Pleistocene and possibly throughout the various episodes of the Quaternary periods. The e f f e c t i v e -ness of the b a r r i e r s i s r e f l e c t e d by the present d i s t r i b u t i o n of G r y l l o b l a t t a on the western fringe of the Columbia and Great Basins and its.absence elsewhere in the basins. G r y l l o b l a t t a i s absent east of the Rocky Mountain Co r d i l l e r a n . During the l a t e Pleistocene a zone of suitable climate south of the Keewatin Ice Sheet probably extended eastward across the Central Plains, but lack of suitable habitat there seems to have prevented dispersal to eastern mountain ranges. . 198 The d e f i c i t of Gr y l l o b l a t t a material south of Yellow-stone National Park along the Rocky Mountain crest i s , I bel i e v e , due to lack of investigation i n suitable areas Further research should concentrate i n thi s area. G r y l l o b l a t t a appear to be absent north of the southern Cassiar and Liard ranges. This may be due i n part to i n s u f f i c i e n t f i e l d i n v e s t i g a t i o n ; however, I believe the taxon w i l l not be found north of the - 5° C. isotherm or i n widespread permafrost. The permanently .frozen deeper substrate and the freezing and thawing of the surface layer of the permafrost would preclude occupancy of the avail a b l e hypolithion by G r y l l o b l a t t a . In the northern portions of i t s range, G. _c. campodeiformis does not occupy areas of r e l i c or discontinuous permafrost, even though suitably sized rock i s present. G r y l l o b l a t t a may occur i n the mountain ranges on the Gulf of Alaska, such as the Coast, St. E l i a s , Wrangell, Chugach and McKinley ranges, for these areas are influenced by the.maritime climate and permafrost i s not widespread. R e l i c populations are found at the B r i t i s h Columbia-Yukon border, not far from the more southern of these l o c a l i t i e s . Discovery of A'laskan populations may shed some l i g h t on the rel a t i o n s h i p of Gr y l l o b l a t t a to the Asian genera, G r y l l o b l a t t i n i a and G a l l o l s l a n a . Such r e l a -tionships have not yet been discussed i n d e t a i l , and the inte r p r e t a t i o n of the zoogeography of the Grylloblattodea as a whole i s also a task for the future. For t h i s , a f u l l e r knowledge of the phylogeny of the order, the orthop-199 teroids and the Insecta i s e s s e n t i a l . So f a r , the data here are too sparse. While early phylogenetic speculations related the Grylloblattodea to almost every order of the Orthopteroidea over the years, the recent phenetic analyses have suggested a f f i n i t i e s with the Dermaptera ( G i l e s , 1963) and the Ensifera (Blackith and B l a c k i t h , 1968). The analysis undertaken i n the present thesis indicate a phenetic a f f i n i t y of G r y l l o b l a t t a to the Dermaptera and Phasmida, but t h i s does not s e t t l e the phylogenetic problem. Perhaps not u n t i l f o s s i l s are discovered w i l l t h i s be f e a s i b l e . It i s possible that f o s s i l Grylloblattodea do exist and have been overlooked by .paleontologists. Most f o s s i l insects consist of fragments such as wing, s c l e r i t e and head capsule. Nymphs and adult males of Gr y l l o b l a t t a can eas i l y be mistaken for larvae of staphylinid or lamellicorn Coleoptera. For example, 7 specimens, adults of both sexes and nymphs, collected between 1906 and 1910 and i n the Rocky Mountain Park Museum and the Canadian National Collec-t i o n , were not recognized as Gryl l o b l a t t a u n t i l 1916 (Walker, 1919). The f i r s t specimens of Grylloblattodea discovered i n Russia were collected i n 1935 and were i d e n t i f i e d as coleopterous larvae. They were not recognized and described as Grylloblattodea u n t i l 1951 (Sharov, 1968). I have found over a dozen museum specimens of Gr y l l o b l a t t a which had not been recognized as g r y l l o b l a t t i d s . Hence, i f f o s s i l Grylloblattodea has been found, the id e n t i t y may be shrouded by lack of recognition. 200 There i s no doubt that the Grylloblattodea are among the most important of the insects for understanding the phylogeny of the Pterygota. They not only combine i n t h e i r morphology numerous "ancestral" features that occur i n six other orthopteroid orders, but they also possess a number of features not found i n other pterygote i n s e c t s . The oldest known insect f o s s i l s are the winged Archaeoptera of the Devonian (Rohdendorf, 1961) . Other pterygote f o s s i l s belong to the Protorthoptera, Proto-blattodea, Paleodictyoptera and Blattodea and occur i n the Carboniferous, but not u n t i l the Upper Carboniferous do apterygote f o s s i l s occur i n the form of the Monura (Rohden-dorf et a l , 1961). The modern Grylloblattodea are generally considered to be secondarily apterous or neotenic (Walker, 1914; Crampton, 1915; Caudell, 1923, 1924; Imms, 1957; Sharov, 1968), having supposedly evolved from some early winged form. In the examination of Gr y l l o b l a t t a for characters used in the- numerical a n a l y s i s , I found a number of morphological features i n the thorax that suggest a primarily apterous structure. The three thoracic terga are freely movable upon one another and are progressively shorter and less d i f f e r e n t i a t e d p o s t e r i o r l y . The intersegmental muscles are well developed and r e t a i n the•primitive connection to a single tergal antecosta. The meso- and metaterga show no evidence of being divided into a prescutum, scutum, scutellum and postnotum. These same stru c t u r a l conditions 201 are found i n the apterygote orders, Thysanura and Dlplura, "and are absent i n adult pterygote orders. The phragmata are absent i n the G r y l l o b l a t t a thorax, the l o n g i t u d i n a l dorsal muscles are f l a t and less developed than i n other ptery-gotes, tergo-sternal muscles are absent, and the tergo-p l e u r a l muscles are attached to the pleural areas. In., contrast, winged Pterygota have small pleuro-alar flexors attached to the p l e u r a l ridge. The thoracic pleura of G r y l l o b l a t t a are on a primitive plan and r e t a i n the apterygote condition. The pleura are t h i n l y s c l e r o t i z e d except along the p l e u r a l sutures and the point of o r i g i n of the pleural arms. The epimeron i s i n the o r i g i n a l p o s i t i o n of the apterygote anopleurite. The precoxal and postcoxal bridges, which are present i n the t y p i c a l wing-bearing segment, are absent i n G r y l l o b l a t t a . The p l e u r a l ridge bears a minute terminal expansion which may possibly be a precursor of a p l e u r a l a l a r process. The p l e u r a l arms (pleural apophyses) are not attached to the pleural ridge as i n other Pterygota. The sternal regions are l a r g e l y membraneous and each segment has a spinasternum. The presence of a t h i r d spinasternum i s unknown for other Pterygota. The sternal features resemble the Apterygota and are not suggestive of even feeble f l i e r s such as B l a t t a . The general structures of the thoracic s c l e r i t e s and t h e i r musculature indicate a primary apterous condition i n G r y l l o b l a t t a , unless regressive mutations and saltations 202 have o c c u r r e d t h r o u g h o u t the t h o r a x . Such l a r g e - s c a l e r e g r e s s i o n o f s t r u c t u r e s a s s o c i a t e d w i t h wings i s not known from secondary a p t e r o u s o r t h o p t e r o i d s . I t can be a r gued t h a t the G r y l l o b l a t t o d e a a r e neotenous and o n l y . a d u l t i n the g e n i t a l i a and i n t e r n a l r e p r o d u c t i v e system. However, the t h o r a c i c morphology s u g g e s t s t h a t the G r y l l o -b l a t t o d e a might be a modern r e l i c o f an a n c i e n t non-winged s t o c k . C e r t a i n l y , t h i s a s p e c t b e a r s f u r t h e r i n v e s t i g a t i o n i n the f u t u r e . N e v e r t h e l e s s , whether p r i m a r i l y a p t e r o u s or n o t , a common p h y l o g e n e t i c r e l a t i o n s h i p o f G r y l l o b l a t t o d e a w i t h o t h e r o r t h o p t e r o i d s must be remote, p r o b a b l y d a t i n g back to the p r e - C r e t a c e o u s . 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National Museum (1), Oregon State University C o l l e c t i o n (2), J . W. Kamp Colle c t i o n ( l ) . Topotype l o c a l i t y : Three Sisters (North S i s t e r ) , 6500 f e e t , Cascade Mountains, Lane Co., Oregon. Date: July 22, 1970. Co l l e c t o r : J . W. Kamp. Description of male topotype: Size large for males of genus; body and leg pubescence dark and conspicuous with many stout spine-like setae; antennae long, each 18 mm. and as long as t o t a l body-head length; 40 antennal segments l e f t , 32 right (broken); 3rd antennal segment les s than 1.2 times as long as 2nd. Pronotum ( F i g . 24-9) with l a t e r a l margins s l i g h t l y con-vergent as i n G. skacrltensis and G. chandler!, anterior margin bearing setae, two rows of setae on dorsum; legs long with many rows of spine-like dark brown setae; leg r a t i o s (length divided by.width): p r o t i b i a , 9.25; profemur, 4.04; Figure 24 . G r y l l o b l a t t a s c u l l e n i s c u l l e n i 224 metatibia, 13.64; metafemur, 7.12. Supra-anal plate ( F i g . 24-1) symmetrical to 10th abdominal tergum,- plate with l a t e r a l , margins nearly equal, antero-l a t e r a l corners normally concealed beneath tergum; ce r c l long, about one-half antennal length, d i s t a l t h i r d of each segment with stout setae; l e f t gonocoxa wider at base than long, with heavy fringe of weak setae, stylus double-tapered as i n G. chandler!; r i g h t gonocoxa ( F i g s . 24-2,3) with ventral margin nearly s t r a i g h t , antero-dorsal margin gently rounded to meet cla s p e r - l i k e process. P r i n c i p a l copulatory s c l e r i t e ( F i g s . 24-6,7) with dorso-lateral lobe extending l a t e r a l l y from main arm, dorsal processes and apical l i p short and more rounded than other species; f i r s t secondary accessory s c l e r i t e as i n Figure 24-5, second accessory s c l e r i t e as i n Figure 24-4, with t i p recurving Color: dorsum and head l i g h t tan-brown; legs and venter s l i g h t l y l i g h t e r approaching dry straw. Measurements (length i n mm.): pronotum, 3.30; eye, 0.60; p r o t i b i a , 3.70; profemur, 4.25; metatibia, 6.55; metafemur, 5.70. Measurements (width i n mm.): head, 3.15; pronotum, 2.65; p r o t i b i a , 0.40; profemur, 1.05; metatibia, 0.48; metafemur, 0.80. Collected at night on large persistent snowfleld on Belknap Crater lava f i e l d s , Scott Pass, south slope of North S i s t e r . The male of G. s. s c u l l e n i has been unknown u n t i l now. With t h i s the other material from the High Cascades can now be determined. 225 The discovery of another subspecies, G. s. cryocola. warrants recognition of the nominate form as G. s. s c u l l e n l . Additional material: Two males, col l e c t e d at night on McKenzie Pass, 2 miles west northwest, on July 18, 1971, by J . W. Kamp. Gr y l l o b l a t t a akagitensla n. ap. Type l o c a l i t y : G l a c i e r Basin, 5500 feet, Glacier Peak, Snokomish Co., Washington. Date: September 15, 1969. Co l l e c t o r : L. B a r t l e t t . Habitat: persistent snow f i e l d s . Holotype: adult male, unaccessioned, U. S. National Museum. All o t y p e : adult female, same data, unaccessioned, U. S. National Museum. Paratypes: 1 adult male and 9 adult females, unaccessioned, University of Idaho c o l l e c t i o n . Description of male holotype; Size medium for genus; f i n e l y pubescent; major body setae conspicuous as i n G. scudderi; l a t e r a l end of poster-i o r margins of abdominal terga with a single row of well developed setae (lacking i n G. scudderi); 32 antennal segments l e f t (broken), 37 right. Pronotum (Fig. 25-9) with l a t e r a l margins moticeably convergent p o s t e r i o r l y , more so than In G. scudderi. and post e r o - l a t e r a l angles d i s t i n c t l y obtuse; leg r a t i o s (length divided by width): p r o t i b l a , 5.85; profemur, 3.34; meta-t l b i a , 12.83; metafemur, 5.37. Supra-anal plate of the 10th abdominal tergum ( F i g . 25-1.) ( • • " 226 Figure 25 G r y l l o b l a t t a skagltensis rj. sp. 1 1-6 <*~ 227 borne symmetri c a l l y with postero-lateral margins weakly asymmetrical, r i g h t margin more obtuse-angulate than l e f t ; r i g h t cercus i n s e r t i o n anterior to l e f t , posterior margin of supra-anal plate s l i g h t l y obtuse emarginate; r i g h t gonocoxa shaped as In Figure 25-3, stylus 4 times as long as wide and attached to base; oblique view of r i g h t gonocoxa with arcuate margin below thumblike process forming acute angle with lobe. P r i n c i p a l copulatory s c l e r i t e (Figs. 25-6,7) with short copulatory process, apex of process mesoemarginate, dor s o l a t e r a l and ven t r o l a t e r a l lobes greatly expanded l a t e r a l l y from main body of s c l e r i t e ; secondary accessory s c l e r i t e 1 as i n Figure 25-5, as wide as long and approaching that of G. r o t h i ; secondary accessory s c l e r i t e 2 (Fig. 25-4) with beak almost straight and narrower than i n G. occiden-t a l i s . and larger than i n G. scudderi; cerci with dark major setae on distal portion each segment; s c l e r i t i z e d basal portion r i g h t phallomere as i n Figure 25-8. Color: medium suede brown; sterna, legs, c e r c i , antennae, l i g h t b u ff. Measurements (length In mm.): pronotum, 2.8; p r o t l b l a , 2.87; profemur, 3.18; metatibia, 4.75; metafemur, 4.41. Measurements (width i n mm.): head, 2.55; pronotum, 2.25; p r o t l b l a , 0.49; profemur, 0.95; metatibia, 0.37; metafemur, 0.83. Description of female a l l o t y p e : D i f f e r s from male l n usual sexual features and i s larger with more elongate legs; 36 antennal segments l e f t , ' 2 2 8 33 r i g h t ; 3rd antennal segment les s than 1.8 times as long as 2nd. Head wider than pronotum; pronotum with l a t e r a l margins s l i g h t l y convergent p o s t e r i o r l y , p o s t e r o - l a t e r a l margins almost acute, d i f f e r i n g from G. scudderi and G. oc c i d e n t a l l s . posterior margin doubly arcuate, each side with transverse sulcus; major body setae very conspicuous; leg r a t i o s (length divided by width): p r o t i b i a , 7.00; profemur, 3.60; metatibia, 13.78; metafemur, 5.62; legs with many strong setae. Ovipositor: ventral valves with short stout setae on d i s t a l t h i r d ; dorsal valves only s l i g h t l y arcuate. Color: as i n male. Measurements (length i n mm.): pronotum, 3.32; p r o t i b i a , 3.30; profemur, 3.78; metatibia, 5.65; metafemur, 5.01; ovipo s i t o r , 4.35. Measurements (width i n mm.): head, 3.10; pronotum, 2.92; p r o t i b i a , 0.47; profemur, 1.05; metatibia, 0.41; metafemur, 0.89. Description of paratypes: Collected at type l o c a l i t y between 5000-7500 feet. Paratype male d i f f e r s s l i g h t l y from holotype i n absolute measurements as follows: (length i n mm.): pronotum, 2.80; p r o t i b i a , 2.85; profemur, 3.22; metatibia, 4.78; metafemur, 4.40; (width i n mm.): head, 2.50; pronotum, 2.27; p r o t i b i a , 0.46; profemur, 0.98; metatibia, 0.40; metafemur, 0.85. Antennal segments of male 36, r i g h t and l e f t . Antennal segments of females range from 35 to 37. 229 General comments: Gr y l l o b l a t t a skagitensls resembles G. scudderi and G. o c c i d e n t a l l s upon gross examination. G r y l l o b l a t t a skagj-tensls males may be distinguished from G. scudderi and G. o c c i d e n t a l l s by the d e t a i l s of the primary and secondary copulatory s c l e r i t e s . It d i f f e r s from G. scudderi by having a shorter pronotum, d i f f e r e n t t i b i a r a t i o s , and i n general shape of supra-anal plate and r i g h t gonocoxa. The majority of specimens were collected at night, between 10 pm. and midnight, as they foraged on large persistent snow f i e l d s . Other material was obtained i n nearby hypolithion. The s p e c i f i c name r e f e r s to the Skagit Indian t r i b e which has occupied the Skagit River drainage for centuries. In some of the Skagit legends Glacier Peak i s refe r r e d to as a s p i r i t that rumbles and speaks, undoubtedly because of i t s volcanic a c t i v i t y . G r y l l o b l a t t a scudderi n. sp. Type l o c a l i t y : Whistler Mountain, 6400 feet, Gari-b a l d i P r o v i n c i a l Park, B r i t i s h Columbia. Date: July 13, 1970. C o l l e c t o r s : L. B a r t l e t t and J . W. Kamp. Habitat: alpine on snow. Holotype : adult male, unaccessioned, Canadian National C o l l e c t i o n . A l l o t y p e : adult female, same data, unaccessioned, Spencer Entomological Museum, University of B r i t i s h Columbia. Paratypes: none. 230 Description of male holotype; Larger than G. campodeiformis but equal l n size to G. skagltensls; body and legs densely clothed with fine tan pubescence; major body setae more conspicuous than i n G. campodeiformis. but about as i n G, skagltensls; 35 antennal segments l e f t ; 37 r i g h t . Pronotum '(Fig. 26-9). longer than head width, with l a t e r a l margins only s l i g h t l y convergent (as i n G. campodei- formis). p o s t e r o - l a t e r a l angle nearly acute; leg r a t i o s (length divided by width): p r o t l b l a , 7.62; profemur, 3.31; metatibia, 14.84; metafemur, 5.09; t i b i a proportionately longer than i n G. occi d e n t a l i s or G. skagltenala. Supra-anal plate symmetrical to the 10th abdominal tergum ( F i g . 26-1), l a t e r a l and posterior margina symmetri-c a l , l a t e r a l margins gently concave toward posterior; cerci i n s e r t i o n opposed; r i g h t gonocoxa shaped as i n Figure 26-3, stylus 3 times as long as wide and attached basally to gonocoxa; r i g h t gonocoxa l n l a t e r a l oblique view as i n Figure 26-2,.clasper not as wide as i n G. skagltensis or i n G. campodeiformlB.. - : • P r i n c i p a l copulatory s c l e r i t e (Figs. 26-6,7) d i f f e r i n g from that of G. akagitenais i n that apex of copulatory process i s not emarginate, but has apex more pronounced and dorsal cap more expanded than i n G. campodeiformis or G. o c c i d e n t a l i s . d o r s o - l a t e r a l lobe alatute; a p i c a l process of secondary accessory copulatory s c l e r i t e as in Figure 25-5; accessory s c l e r i t e 2 (F i g . 26-4) with apex more sharply 231 Figure 26. G r y l l o b l a t t a scudderi n. sp. 232 constricted than i n G. occi d e n t a l l s and G. skagltensis; basal s c l e r i t i z e d portion of r i g h t phallomere as i n Figure 26-8. , . Color: head and thorax grayish-tan; abdomen l i g h t gray; legs straw. Measurements.(length i n mm.): pronotum, 2.67; p r o t i b i a , 2.82; profemur, 3.15; metatibia, 4.75; metafemur, 3.82. Measurements (width i n mm.): head, 2.50; pronotum, 2.22; p r o t i b i a , 0.37; profemur, 0.95; metatibia, 0.32; metafemur, 0.75. Description of female a l l o t y p e : Size medium for genus but smaller than G. o c c i d e n t a l l s ; major body setae l e s s prominent than i n other species; 36 antennal segments l e f t and r i g h t ; antennal segments smaller than i n G. skagltensls or G. occidentalls but about as i n G. campodeiformls. Head as wide as pronotum length; pronotum with l a t e r a l margin l e s s emarginate than i n G. skagitensls; leg r a t i o s (length divided by width): p r o t i b i a , 7.67; profemur, 3.31; metatibia, 13.85; metafemur, 5.17. Ovipositor: short, dorsal valves reaching to d i s t a l margin of segment 4 of cercus; ventral valves moderately curved throughout length. Color: as i n male. Measurements (length i n mm.): pronotum, 2.90; p r o t i b i a , 2.80;" profemur, 3.15; metatibia, 4.85; metafemur, 4.40; ovipositor, 3.00. Measurements (width i n mm.): head, 2.67; 233 pronotum, 2.32; p r o t l b l a , 0.36; profemur, 0.95; metatibia, 0.35; metafemur, 0.85. General comments; Gr y l l o b l a t t a scudderi males can be distinguished from those of G. occ i d e n t a l i s and G. skagitensls by the shape and d e t a i l s of the supra-anal plate and r i g h t gonocoxa. It may also be separated from other species by d e t a i l s of the copulatory s c l e r i t e . Female G. scudderi may be separated from other species by the short ovipositor and pronotal shape. Holotype and allotype specimens were collected on a large persistent snow f i e l d l y i n g i n a cirque approximately 1000 feet below a g l a c i e r . The a i r temperature on the snow f i e l d was 5° C. colder than the a i r and snow f i e l d s outside the cirque. Much colder a i r from the higher elevations flowed o f f the g l a c i e r and was funnelled by the topography into the cirque over the snow f i e l d . Searching on other snow f i e l d s f a i l e d to f i n d any addit i o n a l specimens. Nine nymphs (3 males, 6 females) were collected from the hypo-l i t h i o n immediately adjacent to the cirque snow f i e l d . I t i s a pleasure to name thi s new species for Dr. G. G. E. Scudder, whose sustained i n t e r e s t and encourage-ment are deeply appreciated. G r y l l o b l a t t a paullnal n. ap. Gr y l l o b l a t t a sp. Kamp, 1970 (Annales de Speleologie, ' 234 25(1): 223-230). Type l o c a l i t y : South Ice Cave, 5000 feet (T. 23, S; R. 14E, Sec. 18, N.E. £; U. S. G. S. Newberry Crater Quadrangle), Deschutes National Forest, Lake Co.., Oregon. Date: November 21, 1968. Col l e c t o r : J . W. Kamp. Habitat: cavernicolous. Holotype: adult male, unaccessioned, U. S. National Museum. Allotype: adult female, same data except collected July 22, 1963, unaccessioned, U. S. National Museum. Paratypes: 4 adult males and 5 adult females, unaccessioned, 1 each sex, Oregon State University c o l l e c t i o n and remainder J . W. Kamp c o l l e c t i o n . Description of male holotype: Size large for genus; very fine dense body pubescence inconspicuous; major setae sparse except on legs; setae les s than i n G. skagltensls; antennae shorter than i n G. chirugica and about as long as thorax and abdomen combined; 35 antennal segments l e f t , 36 r i g h t . Head as wide as pronotum i s long and larger than i n G. skagltensls or G. r o t h i ; pronotum ( F i g . 27-9) with l a t e r a l margins converging less p o s t e r i o r l y than i n G. skagitensls or G. s c u l l e n i . p o s t e r o - l a t e r a l margin weakly obtuse, posterior margin s l i g h t l y arcuate between postero-l a t e r a l angle and mid-line; leg r a t i o s (length divided by width): p r o t i b i a , 5.88; profemur, 2.91; metatibia, 4.65; metafemur, 4.70. Supra-anal plate s l i g h t l y asymmetrical to 10th abdominal tergum (Fig. 27-1); supra-anal plate with l e f t posterior lobe almost acute to postero-lateral margin, lobe Figure 27. G r y l l o b l a t t a paullnal n. sp 23 f«-236 shorter and more acute than i n G. r o t h l ; r i g h t gonocoxa ~ ( F i g . 27-3) with anterior margin arcus hal f e l l i p t o i d ; clasper acute to dorsal margin, s l i g h t l y recurved at apex, and l a t e r a l l y expanded; ventral margin of gonocoxa straight i n l a t e r a l oblique view as i n Figure 27-2, narrow d i s t a l arm with wide clasper. P r i n c i p a l copulatory s c l e r i t e ( F i g s . 27-6,7) with dor s o - l a t e r a l lobe expanded and over hal f the length of l a t e r a l margin below dorsal cap; apex of dorsal process les s than half length of cap; ventro-lateral lobe short, s l i g h t l y pointed; secondary accessory s c l e r i t e s as i n Figures 27-4,5; accessory s c l e r i t e 2 ( F i g . 27-4) with short apex, shorter than i n other species; accessory s c l e r i t e ( F i g . 27-5) longer and more pointed than i n G. chlrugica; s c l e r i t i z e d basal portion of r i g h t phallomere as i n Figure 27-8. Color: dorsum and head l i g h t brown; legs and venter pale straw. Measurements (length i n mm.): pronotum, 3.00; p r o t i b i a , 3.00; profemur, 3.50. Measurements (width i n mm.): head, 3.00; pronotum, 2.42; p r o t i b i a , 0.51; profemur, 1.20; metatibia, 0.42; metafemur, 0.85. Description of female allotype : Size large for females of genus, l a r g e r , than holotype or paratype males; fewer major setae than i n G. s c u l l e n l or G. skagltensis, with very fine pubescence and sparse major setae; antennae with 35 segments each; t h i r d antennal 237 segment 1.25 times the length of second; second antennal segment longer than i n G. chlrugica , G. s c u l l e n i and G. r o t h i ; antennae as long as thorax and abdomen combined, shorter than i n G. s c u l l e n i . Head as wide as pronotum length; pronotum with l a t e r a l margins nearly straight as i n G. s c u l l e n i f but postero-l a t e r a l angle decidedly obtuse, d i f f e r i n g from G. s c u l l e n i and G. chlrugica; leg r a t i o s (length divided by width): p r o t i b i a , 8.33; profemur, 3.68; metatibia, 12.12; meta-femur, 6.71; r a t i o s proportionately greater than i n G. chlrugica and G. r o t h i ; pro- and metatibia proportionately longer than i n G. s c u l l e n i but pro-and metafemur shorter. Ovipositor: ventral valves bear short stout setae along entire length; valves long and tapering, reaching base of seventh cereal segment; ovipositor much longer than i n G. s c u l l e n i . G. chlruglca or G. skagltensls. Color: as i n male . Measurements (length i n mm.): pronotum, 3.00; pro-t i b i a , 3.00; profemur, 3.50; metatibia, A.85; metafemur, 4.70; ovipositor, 4.50. Measurements (width i n mm.): head, 3.00; pronotum, 2.55; p r o t i b i a , 0.36; profemur, 0.95; metatibia, 0.40; metafemur, 0.70. Description of paratypes : Length Measurements of Para types (Range and Mean) Males Females Pronotum 2.98-3.00 (2.99) 2.92-3.00 (2.93) Pro t i b i a 2.85-3.00 (2.92) 2.90-3.00 (2.95) Profemur 3.50-3.58 (3.54) 3.40-3.50 (3.49) 238 Metatibia 4.65-4.80 (4.72) 4.75-4.95 (4.82) Metafemur 4.70 (4.70) 4.50-4.70 (4.61) Ovipositor 4.35-4.45 (4.43) General comments: The shape of the supra-anal plate separates G . paullnai from a l l other species except G. r o t h l and G. lava cola. G r y l l o b l a t t a paullnai d i f f e r s from G. r o t h i i n the number of antennal segments, shape and d e t a i l s of r i g h t gonocoxa, primary copulatory s c l e r i t e and secondary accessory s c l e r i t e s ; G. paulinal d i f f e r s from G. lavacola i n the d e t a i l s of the supra-anal plate, r i g h t gonocoxa and s c l e r i t e s of the g e n i t a l i a . G r y l l o b l a t t a paullnai i s r e s t r i c t e d to South Ice Cave or the hypolithion l n the immediate v i c i n i t y . The species occupies the cave except during the winter when the temperatures and humidities f a l l below the l e t h a l l i m i t s for the insect. This species i s named after the cat t l e r u s t l e r renegade Indian Chief, Paulina, of the Walapi Tribe of the Snake Indian nation. The renegade t i t l e was declared by white men when he and his band would not be forced onto a reservation with t h e i r natural enemy, the Warm Springs Tribe. Chief Paulina was k i l l e d i n ambush on A p r i l 25, 1867, for stealing a few cat t l e to feed his starving band. Since the type l o c a l i t y , South Ice Cave, occurs on the southeast slope of Paulina Mountain, i t seems f i t t i n g to recognize t h i s brave i n d i v i d u a l once again by naming t h i s new species a f t e r him. 239 G r y l l o b l a t t a campodeiformls athapaska n . 3 8 p . Type l o c a l i t y : Mt. St. Paul, 4925 feet, Summit Lake, Stone Mountain P r o v i n c i a l Park, B r i t i s h Columbia. Date: August 26, 1962. C o l l e c t o r : R. E. Leech. Habitat: east slope of hypolithion. Holotype: adult male, unaccessioned, Canadian National C o l l e c t i o n . A l l o t y p e : adult female, same l o c a l i t y data except collected at 5400 feet, unaccessioned, Canadian National C o l l e c t i o n . Paratypes: none. Description of male holotype: Size as i n G. c. campodeiformis; body pubescence pale and l e s s conspicuous than l n type o f species; 29 antennal segments l e f t , 28 r i g h t (broken); i n d i v i d u a l antennal segments longer than i n G. c. campodeiformis from Jasper National Park. Pronotal length shorter, than i n Jasper population o f G. c. campodelformia ? being nearly square and with l a t e r a l margins less convergent than i n topotypic material ( F i g . 28-9); postero-lateral angle of pronotum approximately acute; leg r a t i o s : (length divided by width): p r o t i b i a , 6.09; profemur, 2.73; metatibia, 10.00; metafemur, 5.45; legs shorter i n proportion than In G. _c. campodeiformis. Supra-anal plate (Fig. 28-1) s l i g h t l y asymmetrical to 10th abdominal segment; tergum shape as i n type (Fig.28-1); r i g h t gonocoxa (Figs. 28-2,3) d i f f e r i n g i n d e t a i l .. from that of G. c. campodeiformis, anterior margin a smooth elipse rather than obtuse angulate aa i n Jasper G._c. campodeiformis specimens; stylus length twice 240 Figure 28. G r y l l o b l a t t a campodeiformls athapa ska n. ssp. 241 width, shorter and e l i p s o i d rather than tapering as In G. c. campodeiformis. P r i n c i p a l copulatory s c l e r i t e (Figs. 28-6,7) d i f - ~ ferin g i n d e t a i l s from G. c. campodelform1a with apex of dorsal process pyramid-shaped rather than beak-like, d o r s o - l a t e r a l lobe l e s s alatus than normal f o r topotype material, ventro-lateral lobe short and equal to dorso-l a t e r a l lobe length, less d i l a t e d and more alatus than i n G. c. campodeiformis; secondary accessory s c l e r i t e s d i f f e r i n g as i n Figures 28-4,5. Color: not certain for specimen has faded In preserving. Measurements (length i n mm.): pronotum, 2.59; p r o t i b i a , 2.50; profemur, 2.73; metatibia, 4.00; metafemur, 3.71; cercus, 4.50. Measurements (width i n mm.): head, 2.71; pronotum, 2.60; p r o t i b i a , 0.41; profemur, 1.00; metatibia, 0.40; metafemur, 0.68. Description of female a l l o t y p e : Size equal to topotyplc Jasper material; 29 antennal segments each; leg r a t i o s (length divided by width): pro-t i b i a , 5.50; profemur, 3.50; metatibia, 9.15; metafemur, 5.07; p r o t i b i a , metatibia and metafemur much shorter i n proportion than Jasper female G. c. campodeiformls; basal tarsomere subequal i n length to next three t a r s i whereas tarsomere equal to next three i n Jasper G. c. campodeiformls. Pronotum robust and almost square i n shape, l a t e r a l margins almost straight rather than converging as i n G. c. campodeiformis. 242 Ovipositor: shorter and more robust than i n G. c. campodeiformis. with ventral valves markedly curved over d i s t a l 1/3; ce r c i short with 6 cereal segments and equal to length of dorsal blades of ovipositor; dorsal blades of ovipositor equal to 5 segments i n the Jasper G. c. campodeiformls. Color: head, thorax and abdomen dark tan suede; venter and legs medium straw. Measurements (length i n mm.) : pronotum, 2.72; pro-t i b i a , 2.20; profemur, 2.80; metatibia, 3.00; metafemur, 3 . 6 0 ; ovipositor, 3.00; c e r c i , 4.40. Measurements (width i n mm.): head, 2.80; pronotum, 2.45; p r o t i b i a , 0.40; profemur, 0.80; metatibia, 0.40; metafemur, 0.71. General comments : Gr y l l o b l a t t a campodeiformls athapaska can be d i s t i n -guished from G. c. campodeiformls by the shorter, more square, pronotum and by the absolute measurements and proportional r a t i o s of the legs. Males of G. c. athapaska d i f f e r i n es s e n t i a l d e t a i l s of the p r i n c i p a l copulatory s c l e r i t e and the secondary accessory s c l e r i t e s . Female G. c. athapaska are distinguished by the shortness of the cerci and ovipositor blades. Mt. St. Paul, the type l o c a l i t y , i s over 450 miles north of the l a s t known Rocky Mountain Cordilleran popula-tion of G. c. campodeiformls (from Whistler Mt., Jasper National Park). The two subspecies are i s o l a t e d from each 243 other by the broad, low Peace River plateau. Survey c o l l e c t i n g between the two populations by R. E. Leech and E. E. MacDonald and my own survey t r i p s indicated that suitable hypolithion habitats over the plateau were extremely r a r e , and no a d d i t i o n a l populations of G r y l l o b l a t t a have been discovered. I believe that G. c. athapa ska i s a l a t e Pleistocene r e l i c t that has survived i n the refugium along the Nahan and L i a r d ranges. The subspecies i s named af t e r the native language of the Indian nation that inhabited northern B r i t i s h Columbia and the Yukon and Northwest T e r r i t o r i e s . G r y l l o b l a t t a campodeiformis nahanni n. ssp. Type l o c a l i t y ; Mt. McDame, 5400 f e e t , Cassiar Mountain Range, Cassiar, B r i t i s h Columbia. Date:. September 17, 1969. C o l l e c t o r : J . W. Kamp. Habitat: hypolithion near snow bank:. Holotype: adult male, un-accessioned, Canadian National C o l l e c t i o n . A l l o t y p e : adult female, Limestone Peak, 6000 f e e t , Cassiar Mountain Range, collected September 16, 1969, hy J . W. Kamp i n h y p o l i t h i o n ' i n snowstorm, unaccessioned, Spencer Entomological Museum, University of B r i t i s h Columbia. Paratypes: none. Description of male holotype: Size larger than G. c. athapaska; pubescence conspicu-ous; major setae prominent; 24 antennal segments l e f t (broken), 244 29 r i g h t ; t h i r d antennal segment 1.5 time the length of second segment. Pronotal length about equal to head width and longer than i n G. c. athapaska ; l a t e r a l margins of pronotum n o t i -ceably convergent, approaching G. c. campodeiformis speci-mens from Jasper National Park; pronotum wider l n propor-ti o n to length than i n G. c. campodeiformis and not square as i n G. c. athapaska ; leg r a t i o s (length divided by width): p r o t i b i a , 5.33; profemur, 2.75; metatibia, 10.62; meta-femur, 6.55; metatibia and femur longer than i n G. c. athapaska ; metatibia longer than i n c. campodeiformis from Jasper National Park. Supra-anal plate s l i g h t l y asymmetrical to 10th ab-dominal tergum (Fig. 29-1); d i f f e r s from other subspecies i n d e t a i l s of l a t e r a l and posterior margins (Fig. 28-1);, r i g h t gonocoxa (Figs. 29-2,3) with clasper process more obliquely expanded than i n other supspecies.. • -•-P r i n c i p a l copulatory s c l e r i t e (Figs. 29-6,7) d i f f e r i n g i n d e t a i l s from G. c. campodeiformis and G. _c. athapaska. apex of dorsal cap long and recurving, do r s o - l a t e r a l lobe as i n Jasper G. c. campodeiformis. but with greater l a t e r a l expansion of lobe; secondary accessory s c l e r i t e 1 ( F i g . 29-5) wider and more rounded than i n other forms; accessory s c l e r i t e 2 with constricted neck (Fig. 29-4); basal s c l e r i t e portion of r i g h t phallomere ( F i g . 29-8) more as In G. c. athapaska. . Color: head and thorax medium tan; abdomen grayish-tan; 245 Figure 29. G r y l l o b l a t t a campodeiformls nahanni n.ssp. 246 venter and legs medium straw. .., Measurements (length i n mm.): pronotum, 2.80; p r o t i b i a , 2.40; profemur, 2.70; metatibia, 4 . 25 ; metafemur, 3.43; c e r c i , 4.80. Measurements (width i n mm.): head, 2.81; pronotum, 2.48; p r o t i b i a , 0.45; profemur, 0.98; metatibia, 0.40; metafemur, 0.60. Description of female a l l o t y p e : Larger than females of Jasper G. c. campodeiformls; 29 antennal segments l e f t and r i g h t . Pronotum length about equal to head; pronotum wider i n proportion than i n G. c. campodeiformlst shape being between square appearance of G. _c. athapaska and rectangular shape of G. c. campodeiformls; l a t e r a l margins of pronotum s l i g h t l y convergent, but les s so than i n G. c. campodeiformls material from Jasper; pronotum not as straight as i n G. c. athapa ska; basal tarsomere of metatarsus subequal i n length to next three tarsomeres, as i n G. _c. athapaska ; legs longer than i n other subspecies; leg r a t i o s (length divided by width): p r o t i b i a , 6.08; profemur, 3.06; metatibia, 11.07; metafemur, 5.33. Ovipositor: short; dorsal valves about as i n G. c. at ha pa ska ; c e r c i longer than i n other subspecies. Color: head, thorax and abdomen golden brown; legs and ventral region pale straw. Measurements (length i n mm.): pronotum, 3.05; pro-t i b i a , 2.80; profemur, 3.11; metatibia, 4.43; metafemur, 247 4.00; o v i p o s i t o r , 3.00; c e r c i , 5.00. Measurements (width i n mm.): head, 3.18; pronotum, 2.70; p r o t i b i a , 0.45; pro-femur, 0.88; metatibia, 0.40; metafemur, 0.75. General comments: Gr y l l o b l a t t a campodeiformis nahanni may be distinguished from G. c. campodeiformis by the length of the ovipositor and d i f f e r s from the other subspecies by the longer cerci and legs i n both sexes. Males can be distinguished from males of G. c. campodeiformis and G. c. athapaska by the d e t a i l s of the, p r i n c i p a l copulatory s c l e r i t e and secondary accessory s c l e r i t e s . In some features G. c. na ha nn.1 i s intermediate between Jasper material of G. c. campodeiformis and Mt. S t . Paul material of G. c. athapaska. The Cassiar region of Cassiar Mountain Range, the type l o c a l i t y for G. c. nahanni. i s approximately 200 miles west-northwest of the type l o c a l i t y of G. c. athapa ska. The two l o c a l i t i e s are separated by the intervening L i a r d P l a i n . For re l a t i o n s h i p s between G. c. athapaska and G. c. na ha nr\iand t h e i r possible l a t e Pleistocene o r i g i n see the section on d i s t r i b u t i o n . G r y l l o b l a t t a campodeiformis nahanni i s named a f t e r the NahanniIndian Tribe which s t i l l inhabits the region. These natives were a great source of information to me regarding the habitat of t h i s insect and reported seeing a strange "bug" on Limestone Peak. 248 G r y l l o b l a t t a s c u l l e n i cryocola n. sap. Type l o c a l i t y : Edison Ice Cave, 5200 f e e t , (T. 19S; R. 9E.; Sec. 14, S.E. \\ U. S. Forest Service map); Deschutes National Forest, Deschutes Co., Oregon. Date: November 15, 1970. Co l l e c t o r : J . W. Kamp. Habitat: dark zone on cave i c e . Holotype: adult male, unaccessioned, U. S. National Museum. Allotype: adult female, same l o c a l i t y data, collected July 20, 1962, by J . W. Kamp, unaccessioned, U. S. National Museum. Paratypes: 1 adult male and 5 adult females, J . W. Kamp c o l l e c t i o n . Description of male holotype: Size medium for males of genus, smaller than G. s. s c u l l e n i ; pubescence and major setae as i n G. s. s c u l l e n i ; antennae as long as i n G. s. s c u l l e n i . longer than t o t a l head-body length; t h i r d antennal segment longer than i n G. s. s c u l l e n i ; t h i r d antennal segment equal to combined lengths of segments 4 and 5 ( c f . subequal i n G. s. s c u l l e n i ) : 40 antennal segments l e f t and r i g h t . Pronotal length 1.2 times head width and proportion-ately longer than i n G. s. s c u l l e n i ; legs long; leg r a t i o s (length divided by width): p r o t i b i a , 8.72; profemur, 4.50; metatibia, 13.80; metafemur, 6.66; legs longer and wider than i n G. s. s c u l l e n i . Supra-anal plate ( F i g . 30-1) symmetrical to 10th abdominal tergum, plate d i f f e r i n g from that of G. s. s c u l l e n i : plate moderately curvate around postero-lateral Figure 30. G r y l l o b l a t t a s c u l l e n i cryocola n. ssp. 250 margins, convex rather than concave as i n G. a. s c u l l e n l ; cereal length as i n G. a. s c u l l e n i ; r i g h t gonocoxa (Figs. 30-2-3) with s l i g h t l y arcuate ventral margin, antero-dorsal margins abruptly recurving to meet clasper; i n l a t e r a l oblique view clasper narrower than i n G. a. s c u l l e n l . P r i n c i p a l copulatory s c l e r i t e (Figs. 30-6,7) d i f f e r i n g i n d e t a i l s of apex and d o r s o - l a t e r a l , ventr'o-lateral lobea; secondary accessory s c l e r i t e l ( F i g . 30-5) much longer than i n G. a. s c u l l e n l ; accessory s c l e r i t e 2 with apex long and not recurved as i n G. s. s c u l l e n l ; d i s t a l portion of s c l e r i t i z e d region of r i g h t phallomere (Fig.30-8) gently rounded,while pointed i n G. s. s c u l l e n l . Description of female a l l o t y p e ; Size as large as i n G. s. s c u l l e n l ; Al antennal segments r i g h t and l e f t . Pronotum proportionately wider than i n topotype G. a, s c u l l e n l material; p o s t e r o - l a t e r a l angle of pronotum more obtuse than i n G. a. s c u l l e n l and somewhat acute; l a t e r a l margin of pronotum straight for anterior 3/4 of length, then converging noticeably; legs longer than l n G. s. s c u l l e n l ; leg r a t i o s (length divided by width): p r o t i b i a , 9.28; profemur, 4 . 5 0 ; metatibia, 13.89; meta-femur, 8.00; legs proportionately longer except metatibia and with greater proportional length than l n G. a. s c u l l e n l (mean leg r a t i o s of 6 topotype G. s. s c u l l e n i : p r o t i b i a , 7.84; profemur, 3.83; metatibia, 14.15; metafemur, 6.21). Ovipositor: length of dorsal valves about equal to 251 those of G. a. s c u l l e n i ; ventral valves of ovipositor more, acute i n curve than i n G. s.. s c u l l e n i ; cerci longer than i n G. s. s c u l l e n i . Color: head and dorsal thorax-abdomen very pale straw; ventral surface and legs dry yellow. Measurements (length i n mm.): pronotum, 3.70; pro-t i b i a , 4.18; profemur, 4.50; metatibia, 6.67; metafemur, 6.00; c e r c i , 9.25; ovipositor, 3.70. Measurements (width i n mm.): head, 3.35; pronotum, 2.85; p r o t i b i a , 0.45; pro-femur, 1.00; metatibia, 0.48; metafemur, 0.75. Description of paratypes: Thirty-nine to 41 antennal segments. Ratios of Leg Measurements of 5 Paratype Females Range Mean Pr o t i b i a 8.33-9.22 8.63 Profemur 3.97-4.59 4.33 Metatibia 13.30-14.78 13.91 Metafemur 7.46-8.00 7.81 General comments: Gr y l l o b l a t t a s c u l l e n i cryocola females are very d i f f i c u l t to d i s t i n g u i s h from those of G. s. s c u l l e n i . G r y l l o b l a t t a s c u l l e n i cryocola d i f f e r s i n the greater length of the p r o t i b i a , profemur and metafemur. The leg r a t i o s are greater than i n G. a. s c u l l e n i . G r y l l o b l a t t a  s c u l l e n i cryocola Is much l i g h t e r i n color than i s G. s. s c u l l e n i and has longer c e r c i . Males of G. s. cryocola may be distinguished by the d e t a i l s of the g e n i t a l s c l e r i t e s . G r y l l o b l a t t a s c u l l e n i cryocola i s r e s t r i c t e d to Edison Ice Cave and the hypolithion i n the immediate v i c i n i t y . I t occurs sympatrically with G. r o t h i . G r y l l o b l a t t a s c u l l e n i  cryocola was not found to be sympatric with G. r o t h i In other l o c a l i t i e s and i t was not found i n the higher ele-vations of the Three Si s t e r s Mountains. G r y l l o b l a t t a hoodalles n. sp. Type l o c a l i t y : Mt. Hood, 5900 feet, junction of Phlox Point and Tlmberline roads, Oregon. Date: June 18, 1970. Col l e c t o r : J . W. Kamp. Habitat: hypolithion. Holotype: adult male, unaccessioned, U. S. National Museum. A l l o -type: adult female, same data except collected June 18, 1970. Paratypes: 1 adult male and A adult females, J . W. Kamp c o l l e c t i o n . Description of male holotype: Small for genus; smaller than G. chlrugica and more resembling size of G. r o t h i ; fine body pubescence conspic-uous; 30 antennal segments l e f t , 2A r i g h t ; legs short and stout, more so than i n topotypic G. r o t h i . Pronotum shorter than i n G. r o t h i ; leg r a t i o s (length divided by width): p r o t i b i a , 5.OA; profemur, 2.72; meta-t i b i a , 9.52; metafemur, 3.96. Supra-anal plate ( F i g . 31-1) i s asymmetrical with base of l e f t cercus posterior to r i g h t , as i n G. r o t h i ; plate shorter than i n G. r o t h i , with l e f t a p i c a l lobe less 253 Figure 31. G r y l l o b l a t t a hoodalles n. sp. 254 developed; l e f t gonocoxa as broad as long; r i g h t gonocoxa with clasper process ( F i g . 31-3) more angulate to dorsal margin than i n G. r o t h l ; i n l a t e r a l oblique view ( F i g . 31-2) clasper more expanded than i n G. r o t h i . with posterior arm of gonocoxa narrower; stylus borne on basal 1/3» length 2 times width. -P r i n c i p a l copulatory s c l e r i t e (Figs. 31-6,7) d i f f e r i n g i n d e t a i l s from G. r o t h l ; apex of dorsal process of t h i s s c l e r i t e shorter and more curved than i n topotypic G, r o t h l material, d o r s o - l a t e r a l lobe 1.5 times that of G. r o t h l p straighter i n p r o f i l e and more l a t e r a l l y expanded, i n oblique view (Fig.31-7) ve n t r o - l a t e r a l lobe with acute ventral point lacking as i n G. r o t h l ; secondary accessory s c l e r i t e 1 (Fig.31-5) short, globular with basal constric-t i o n d i f f e r i n g from the longer tapering s c l e r i t e of G. r o t h l ; accessory s c l e r i t e 2 d i f f e r s i n d e t a i l s of head-like portion and apex of beak; dorsal s c l e r i t i z e d portion of ri g h t phallomere ( F i g . 31-8) longer and thinner than i n G. r o t h l . Color: darker than G. r o t h l ; general body medium brown ventral abdomen, legs and cerci l i g h t e r buff. Measurements (length i n mm.): pronotum, 2.75; p r o t i b i a 2.52; profemur, 3.00; metatibia, 4.00; metafemur, 3.77. Measurements (width i n mm.): head, 2.78; pronotum, 2*40; p r o t i b i a , 0.50; profemur, 1.10; metatibia, 0.42; metafemur, 0.95. 255 Description of female allotype ; Thorax and abdomen short compared to G. r o t h i ; thorax 1/5 longer i n G. r o t h i ; 22 antennal segments l e f t (broken), 30 segments r i g h t ; t h i r d antennal segment less than 2 times length of second and shorter than In G. r o t h i ; antennae short, 3/4 length of topotypic G. r o t h i ; segments of antennae short, terminal segments smaller than i n any other form and les s than g- length of terminal segments of G. r o t h i . Head and pronotum small compared to those of G. r o t h i ; posterior margins of pronotum obtuse rather than d i -emarginate as i n G. r o t h i ; legs short and robust; leg r a t i o s (length divided by width): p r o t i b i a , 5.34; profemur, 2.95; metatibia, 10.00; metafemur, 4.44. Ovipositor: length of dorsal blade about equal to that of G. r o t h i ; cerci shorter than i n G. r o t h i ; dorsal valve of ovipositor reaching middle of 8th cereal segment. Color: general body color about as i n G. r o t h i ; ab-dominal terga grayish-brown, darker than i n G. r o t h i . Measurements (length i n mm.): pronotum, 3.00; pro-t i b i a , 2.51; profemur, 2.95; metatibia, 4.00; metafemur, 3.87; ovipositor, 3.62. Measurements (width i n mm.): head, 3.00; pronotum, 2.50; p r o t i b i a , 0.47; profemur, 1.00; metatibia, 0.40; metafemur, 0.87. Description of paratypes: Agree with type material i n leg r a t i o s and d e t a i l s of g e n i t a l i a . General comments: Gr y l l o b l a t t a hoodalles may be distinguished from G. 256 r o t h i by the short robust legs and the general d e t a i l s i n the d e s c r i p t i o n . The type l o c a l i t y , near Timberline Lodge, i s i n an unsorted debris from an eruption thought to be less than 1000 years old (see discussion of d i s t r i b u t i o n ) . G r y l l o -b l a t t a hoodalles i s geographically i s o l a t e d from G. r o t h l by the low intervening elevations of the Cascade Range. It seems to be r e s t r i c t e d to the i s o l a t e d stratovolcano (Mt. Hood), for no populations have been found between the type l o c a l i t y and the Three Sisters Mountains. The species name re f e r s to Mt. Hood i n The Dalles region of the Columbia River. G r y l l o b l a t t a lava cola n. sp. Type l o c a l i t y : Belknap Crater Lava F i e l d s , 5384 feet, (T. 15S; R. 8E., U. S. Geological Survey Topographic Map, Three S i s t e r s Quadrangle), McKenzie Pass, Mt. Washington Wilderness Area, Cascade Mountains, Oregon. Date: June 18, 1970. C o l l e c t o r : J . W. Kamp. Habitat: collected on snow f i e l d at night. Holotype: adult male, unaccessioned, U. S. National Museum. Allotype: adult female, same data except collected July 18, 1971, by J . M. Taylor, unaccessioned, U. S. National Museum. Paratypea: 4 adult males and 12 adult females, J . W. Kamp c o l l e c t i o n . Description of male holotypei Size large for genua, as large as G. a. s c u l l e n l ; body 257 and legs densely clothed with fine pubescence; major setae les s conspicuous than i n G. s. s c u l l e n i ; antennae longer than i n G. r o t h i ; antenna about 14.5 mm.; 34 antennal segments l e f t , 33 r i g h t ; t h i r d segment 1.2 times as long as second. Head width equal to pronotal length; pronotum larger . than i n G. s.. s c u l l e n i ; pronotum (Fig. 32-9) with l a t e r a l margins moderately converging, more than i n G. s. s c u l l e n i ; dorsum of pronotum lacking rows of setae t y p i c a l of G. s. s c u l l e n i ; legs longer than i n G. r o t h i ; leg r a t i o s (length divided by width): p r o t i b i a , 5.72; profemur, 3.34; meta-t i b i a , 11.11; metafemur, 4 .80. Supra-anal plate ( F i g . 32-1) asymmetrical to 10th abdominal"tergum; l e f t lobe on posterior margin of plate well developed, larger than i n G. r o t h i . and r i g h t margin with lobe as long as l e f t lobe In G. r o t h i ; cerci about •§• antennal length, longer than i n G. r o t h i . and about 2/3 length of G. s. s c u l l e n i ; l e f t gonocoxa wider at base than long, heavily pubescent with scattered setae, lacking i n G. s. s c u l l e n i and G. r o t h i ; r i g h t gonocoxa (Figs. 32-2,3) has general shape of that of G. r o t h i , anterior margin more curved than i n G. r o t h i and much l i k e that found i n G. .s. s c u l l e n i ; stylus on gonocoxa borne l a t e r a l l y on basal t h i r d . P r i n c i p a l copulatory s c l e r i t e as i n Figures 32-6,7 d i f f e r i n g i n d e t a i l s from G. r o t h i ; dorso-lateral lobe of s c l e r i t e shorter than that of G. r o t h i and expanded l a t e r a l l y , dorsal processes and a p i c a l l i p approaching that of G. a. 258 Figure 3 2 . G r y l l o b l a t t a lava cola n. sp. 259 s c u l l e n i ; secondary accessory s c l e r i t e 1 (Fig. 32-5) shorter and wider than that of G. r o t h l found i n the Three S i s t e r s Mountains; second accessory s c l e r i t e (Fig. 32-4) with shorter beak-like apex than i n G. r o t h i . Color: medium brown body; legs and antennae medium straw. Measurements (length i n mm.); pronotum, 3.40; pro-t i b i a , 3.15; profemur, 3.75; metatibia, 5.00; metafemur, 5.05. Measurements (width i n mm.): head, 3.35; pronotum, 2.90; p r o t i b i a , 0.55; profemur, 1.30; metatibia, 0.45; metafemur, 1.05.. Description of female a l l o t y p e : Larger than G. r o t h l , and approaching G. s. s c u l l e n i i n s i z e ; major setae well developed, but less conspicuous than i n G. a. s c u l l e n i ; 34 antennal segments ri g h t and l e f t ; t h i r d segment 1.2 times length of second. Pronotum with noticeably converging l a t e r a l margins, pos t e r o - l a t e r a l margin obtuse; leg r a t i o s (length divided by width): p r o t i b i a , 7.44; profemur, 3.04; metatibia, 11.30; metafemur, 4.89. Ovipositor: dorsal valves longer than i n G. a. s c u l l e n l . without marked curvature to ventral blades; dorsal valves reaching to middle of s i x t h cereal segment; cerci shorter than i n G. .s. s c u l l e n i and each segment longer than i n G. r o t h i . Color: head-thorax medium brown; abdomen grayish-tan; legs l i g h t brown. . 2 6 0 Measurements (length In mm.): pronotum, 3.45; pro-t i b i a , 3.50; profemur, 3.65; metatibia, 5.10; metafemur, 4.75; ovipositor, 4.05. Measurements (width i n mm.): head, 3.30; pronotum, 2.90; p r o t i b i a , 0.47; profemur, 1.20; metatibia, 0.45; metafemur, 0.97. Description of paratypes: Agree in ra t i o s and measurements with holotype and alloty p e . Do not approximate either G. r o t h i or G. a. s c u l l e n i from the McKenzie Pass-Three Sisters region. General comments: Gr y l l o b l a t t a lava cola occurs sympatrically with both i l - s c u l l e n i and G. r o t h i i n the Belknap Crater Lava f i e l d s . G r y l l o b l a t t a lava cola seems to be limi t e d to the lavas and i s the most numerous species. G r y l l o b l a t t a o c c i d e n t a l l s . S i l v e s t e r ! , G r y l l o b l a t t a campodeiform!s occidentalls S i l v e s t e r ! , 1931 (Trans. Amer. Ent. Soc., 57:291-295) The type l o c a l i t y i s Mt. Baker, Washington, and i t was o r i g i n a l l y described from a nymphal male. The subsequent discovery of Gr y l l o b l a t t a scudderi and G_,_ skapitensls and additional material of G. _c. campodeiformls shows that G. c. occidentalls i s not a subspecies of G. campodeiformis. It i s closely related to G. scudderi, from Garibaldi P r o v i n c i a l Park, B r i t i s h Columbia, and to G. skagitensi s. from Glacier Peak, Washington. Based on the differences of male g e n i t a l i a figured by Gurney (1948), number of 261 antennal segments, and proportional and absolute measure-ments, I believe t h i s population warrants s p e c i f i c status as G. occidental!s. 1 2 3 4 5 7 8 9 10 11 12 13 APPENDIX II TABLE VI1 84 EXTERNAL CHARACTERS OF THE ORTHOPTEROIDS (Modified from G i l e s , 1963) 262 No. Character HEAD AND NECK O c e l l i absent Pleurostomal s u l c i angulate Subocular s u l c i Epistomal sulcus complete Antennal sockets near mandibular a r t i c u l a t i o n s Antennal sulcus and anterior t e n t o r i a l p i t confluent Tentorial body enti r e a n t e r i o r l y Tentorial body imperforate Tento r i a l body elongate Anterior t e n t o r i a l arms twisted Dorsal t e n t o r i a l arms stout Tentorial maculae near eyes Dorsal t e n t o r i a l arms a r i s e from body CO CD r) CD 0 c<S +•> •n ca -p H u CO •H © CD CO CD 1-1 CO CO CO •H CO -P XI 0 •d CD ft 0 60 •H •H * CO TJ CO rH rt r-1 •d +3 0 a r-1 -P r-1 •H co +> p u >> -P u CO CO c CD u CO U 0 X! r-1 CO Q EH <: PQ ss X X 0 0 0 0 0 0 X X X X X 0 X 0 X 0 0 X X 0 X 0 X X 0 X X X 0 X X X 0 0 0 X 0 0 x x o o o o o o X X X X X X X X X X 0 X 0 x> 0 0 0 X X X X X 0 X X X 0 X 0 X X 0 0 X X 0 0 0 X 0 0 0 0 0 0 0 0 X X 0 X 0 0 X & 263 14 Antennae short X X 0 0 X. X 0 0 15 Tip of labrum membranous X X O X O X X X 16 Mandibles with two a p i c a l teeth X X O 0 0 0 0 0 17 Membranous area basally on inner edge of mandibles X X X X X X O O 18 Lacinia with two a p i c a l teeth X 0 0 0 0 X X X : 19 Galea c y l i n d r i c a l X X X X 0 0 0 0 20 Maxillary palps not membranous d l s t a l l y X X 0 0 0 X 0 X 21 Labiostipes meslally divided X X 0 0 0 0 X X 22 Paraglossae c y l i n d r i c a l X X 0 0 0 0 0 0 23 Lab i a l palps s c l e r o t i z e d d i s t a l l y X X 0 0 0 X 0 X 24 Dorsal c e r v i c a l s c l e r i t e X 0 X X 0 X X X 25 Anterior ventral c e r v i c a l s c l e r i t e narrow X 0 0 0 0 0 X X 26 Posterior ventral c e r v i c a l s c l e r i t e wide X 0 0 0 X 0 0 0 27 Posterior l a t e r a l c e r v i c a l s c l e r i t e s X X O O X X X X 28 Posterior l a t e r a l c e r v i c a l not a r t i c u l a t e d with prothorax X 0 X X 0 0 0 0 264 THORAX PROTHORAX Pleural ResIon3 29 Generally well s c l e r o t i z e d X X 0 0 0 0 X X. 30 Cryptopleury absent X 0 0 0 0 X X X 31 Pleural apophysis fused to sternal apophysis X 0 0 0 X 0 X 0 32 Pleuro-coxal a r t i c u l a t i o n an-t e r i o r to trochantino-coxal X X 0 0 0 0 X X 33 Anterior edge of trochantin r o l l e d X X X 0 0 0 0 X 34 Accessory coxal plate X 0 X X X 0 X 0 35 Postpleural s c l e r i t e Sternum X 0 0 X 0 X X X 36 Presternum absent X X 0 0 0 X X X; 37 Sternellum X 0 X X X X X X) 38 Sternacostal suture absent X X 0 X 0 0 X 0 39 Separate s c l e r i t e s for p i t s X X 0 0 0 0 0 0 40 Separate spinasternite X X X X 0 0 X 0 41 Spina smaller than meso-thoracic spina X X X X X X 0 0 42 Coxae of each segment wide apart X X X 0 X X 0 0 43 Coxae wider than long X 0 X X X X 0 0 44 Three t a r s a l segments on a l l legs X 0 0 0 0 0 0 0 265 MESOTHORAX Tergum 45 Phragmata absent or very small X X 0 X 0 0 X X 46 Tegmina X 0 X X X X X X Pleural Regions 47 Pleural apophysis joined by muscle to sternal apophysis X X X 0 0 X X X 48. PIeuro-coxa1 a r t i c u l a t i o n an-t e r i o r to trochantino-coxal X X X X X 0 0 X 49 Anterior edge of trochantin r o l l e d X X X X X 0 0 X 50 Precoxale a separate s c l e r i t e X X 0 X 0 X X X 51 Accessory coxal plate X 0 0 X X X X 0 52 Peritremal s c l e r i t e absent x; 0 X X- 0 0 0 0 .53 Postpleural s c l e r i t e X 0 0 0 0 X X X Sternum 54 Sternellura absent X X 0 0 0 0 0 0 55 Sterna costal suture absent X X X X 0 ..0 0 0 56 Separate spinasternite X X 0 0 0 0 x: 0 Left 57 Mesothoracic and metathoracic coxae wider than long X X X X X X 0 0 META THORAX Tergum 58 Phragmata large X 0 X X X 0 X 266 Pleural Regions 59 Episternum and eplmeron nearly horizontal X X O O O O O O 60 Pleural ridge without apophysis X 0 0 0 0 0 0 0 61 Pleuro-coxal a r t i c u l a t i o n pos-t e r i o r to trochantino-coxal X 0 X X X X 0 0 62 Anterior edge of trochantin r o l l e d X X X X X X 0 X 63 Precoxale fused with basisternum X 0 0 0 0 0 0 0 64 Accessory coxa1 plate X 0 0 X 0 X X 0 Sternum 65 Two sternal p i t s X X 0 0 X 0 X 0 66 Roughly same length as front and middle legs X X O O O X X X GENERAL 67 Sterna overlap: pro-/meso-/ me ta sternum « X X 0 0 0 0 X 0 68 Intersegmental s c l e r i t e s small X X X X X X X X ABDOMEN GENERAL 69 Abdomen dorso-ventrally compressed X X O O O X X X GENERALIZED SEGMENT 2 67 Tergum 70 Antecosta forms d i s t i n c t ridge X X 0 0 . 0 X 0 0 Pleural Regions 71 E n t i r e l y membranous X X X X X O 0 0 72 Spiracles i n membrane X X X X 0 0 0 0 ' PREGENITAL SEGMENTS Segment 1. o* and 9 73 Tergum I has large phragmata X 0 X, 0 X X 0 0 74 Tergum I smaller than other terga X 0 X X 0 0 X X 75 Sternum I absent X O O O O O O O 76 Pleural regions membranous X X 0 0 0 0 X X 77 Spiracle I larg e l y surrounded by tergum I X 0 0 0 X 0 X X GENITAL SEGMENTS Segment 9. o* 78 Tergum IX approximately size of generalized tergum X X O 0 0 X 0 0 79 S t y l i absent from subgenital plate X X 0 X X X 0 0 Segment 7. $ 80 Sternum 7 i s long subgenital plate X 0 0 0 0 0 X X Segment 8, 9 81 Pleural region membranous X X X X X 0 0 0 268' POSTGENITAL SEGMENTS Segment 10. d and g 82 Tergum X large, well s c l e r o t l z e d X X X X O X O O Paraprocta 83 Well s c l e r o t l z e d , conspicuous plates X O O X X X X X Eplproct 84 Well s c l e r o t l z e d X O O X X X O O 269 TABLE VI11 80 EXTERNAL AND INTERNAL CHARACTERS OF THE ORTHOPTEROIDS (Modified from B l a c k i t h and Bl a c k i t h , 1968) CD CD CO CO CO CO <H cO •d "d >d U CD •H -H ri CO "d rH -d B -P O r-I «H CO -p -P >-, $-1 CO cO R U O Si r-» CO O <! At a) s HEAD AND NECK 1 Fewer than 30 antennal segments present X 0 0 0 X 0 0 0 2 Lat e r a l o c e l l i present i n apterous 0 0 0 X 0 0 X X 3 Frontal suture absent X 0 X 0 0 0 0 0 4 Post f r o n t a l suture absent 0 X X X X 0 X 0 5 Eplstomal suture absent 0 0 0 0 0 0 X 0 6 Subantennal suture absent X X X X 0 X. 0 X 7 Subocular suture absent 0 X X 0 0' 0 0 X ' 8 Genoepicranial suture absent 0 X 0 X X 0 X Xv 9 Keel on hypopharynx present 0 0 0 0 X 0 0 0 10 Mesal hook on torma absent 0 0 0 0 X 0 0 0 11 Superlinguae absent 0 0 X X X X X X 12 Salivary cup impactor present 0 0 0 0 X 0 0 0 13 Glossae not well developed X 0 0 0 X 0 X X 14 Dorsal s c l e r i t e s on cervix absent 0 X X 0 X X 0 0 No. Character CO CD •d CD o CO •P •d CO -p •H U CO •H Q> r-1 -P Xi o ft o to CO rH a rH -P >> -P CD u CD Q EH 15 Ventral s c l e r i t e s on cervix absent THORAX AND LEGS 16 Pleurosternal suture absent i n adult 17 Mesal prosternal process present 18 Trochantln i n metathorax absent 19 Dorsal t i b i a l spur or spurs present on hind legs 20 Ventral t i b i a l spur or spurs present 21 Hind t a r s i with fewer than 4 segments 22 Hind t a r s i with fewer than 5 segments 23 A r o l i a present 24 Brunner 1s organ present 25 Tympanum present on fore t i b i a ABDOMEN.AND GENITALIA 26 Dorsal sulcus well developed 27 Spiracles not i n te r g i t e s 28 Cerci unsegmented 29 Lat e r a l apodemes present on st e r n i t e s 270 0 0 0 0 X X O O 0 0 0 0 X 0 0 0 0 X 0 0 0 0 0 0 0 0 X 0 0 X 0 0 o o o x x o x o o x x x x o x x x o o x x o o o x o x x x o o o o o o o x x x o 0 0 0 0 X 0 0 0 o o x x o o o o 0 0 0 0 0 0 0 0 x x x x o x x x x o x x x x o o X 0 0 0 X 0 0 0 271 30 Tympanum present on f i r s t segment 0 0 0 0 X 0 0 0 31 Dorsal pouch present 0 0 0 0 X 0 0 0 32 Male g e n i t a l i a strongly asymmetric X X 0 0 0 0 X. X MUSCULAR CHARACTERS HEAD AND NECK 33 L a b i a l r e t r a c t o r absent 0 0 0 0 X 0 0 0 34 No muscles attached to ventral c e r v i c a l s c l e r i t e s of membrane 0 0 0 0 0 X 0 X THORAX AND LEGS 35 Crossed prosternals to f i r s t c e r v i c a l s c l e r i t e s present 0 X X X X 0 0 X 36 Tergopleurals i n meso or metathorax absent 0 0 X X X 0 0 0 37 La t e r a l intersegmentals i n prothorax absent 0 0 0 0 0 0 0 X 38 Lat e r a l intersegmentals i n meso or metathorax absent 0 0 X 0 X 0 0 0 39 Muscle joining meso and metapophyses present X X X X X O X X 40 Ventral transverse f u r c a l s absent X O X X X O X X Al Reductors of fore femora not well developed 0 0 0 0 O X 0 0 272 ABDOMEN AND GENITALIA 42 Two spiracular muscles present X 0 X X X 0 X X 43 Antagonistic muscle a r i s i n g from sternum X 0 X X X 0 0 0 44 Transverse eternals on anterior segments absent X X 0 0 0 X X X 45 Paradorsal present 0 0 0 0 X 0 0 0 46 Sternopleurals absent X 0 0 0 0 X X X 47 Tergopleurals absent X 0 X 0 X 0 X 0 48 No muscles joining s t e r n i t e or pl e u r a l s c l e r i t e to t e r g i t e cephalad X 0 X X X X 0 0 49 Tergosternals In 2 or more bands X X X 0 X 0 0 X 50 Internal l o n g i t u d i n a l tergals banded X 0 0 0 X X 0 X 51 Alary muscles not more than 10 0 0 0 0 X 0 0 0 52 Alary muscles not more than 11 X 0 0 X X X 0 0 NEURAL CHARACTERS 53 Posterior recurrent nerve single X X 0 0 0 X X X 54 Mandibular nerves close to c l r cumesophageal ''J connectives X X X X X X 0 0 273 55 Corpora a l l a t a large and separate from subesophageal ganglion O O X X X O X X 56 More than 1 abdominal ganglion fused to t h i r d t h oracic ganglion 0 0 0 X X 0 0 X 57 More than 2 abdominal ganglia fused to t h i r d thoracic ganglion 0 0 0 0 X 0 O X 58 Not more than h abdominal ganglia free 0 0 0 0 0 0 0 X 59 Not more than 5 abdominal ganglia free 0 0 0 X X 0 0 X 60 Not more than 6 abdominal ganglia free 0 O X X X X X X 61 Circumesophageal connectives pass through hole i n body of tentorium 0 0 0 0 0 0 X X INTESTINAL CHARACTERS 62 Intima of proventricuius without 6 l o n g i t u d i n a l folds 0 X 0 0 0 X 0 0 63 Neck of proventricuius not tubular 0 0 0 0 0 0 X X 64 Proventricuius globular i n part 0 X X X 0 0 0 0 0 0 0 0 X 0 0 0 0 X X X X 0 X X 0 0 0 0 X 0 X X 0 0 0 0 0 0 X X 274 65 V-shaped plates i n cardiac valve 66 Gastric caeca two or more 67 Gas t r i c caeca s i x or more 68. Gastric caeca eight or more OTHER CHARACTERS 69 Phytophagous insects of elongate form 0 0 0 0 0 X 0 0 70 Segmental vessels i n thorax present 0 0 0 0 0 0 X 0 71 Ovariole ligaments a r i s e i n thorax X X 0 0 X 0 0 X 72 Paired incurrent ostia absent i n thorax 0 0 0 0 0 X 0 0 73 Fewer than 3 incurrent os t i a i n thorax X X X X 0 :,X 0 0 74 Phagocytic organs present i n abdomen 0 0 X X 0 0 0 0 75 Segmental vessels present i n abdomen 0 0 0 0 0 0 X X 76 Ovariole ligaments a r i s e i n abdomen 0 0 X X 0 X 0 0 77 Male accessory glands few (less than 15 pai r s ) and not convoluted 0 X 0 0 0 X 0 0 78 Testis confined to terminal segments 0 0 0 0 0 0 O X 275 79 Ootheca formed O O O O X O X X 80 Subgenital plate i n female X O X X X X X X 

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