SERIES ENTOMOLOGICA EDITOR E. SCHIMITSCHEK GOTTINGEN VOLUMEN 4 SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. 196 8 ECOLOGY AND BIOGEOGRAPHY OF HIGH ALTITUDE INSECTS BY Pro£ Dr. M. S. MANI, M. A., D. S c., Of the School of Entomology, St. John's College, Agra SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. 196 8 ISBN 978-90-481-8511-5 ISBN 978-94-017-1339-9 (eBook) DOI 10.1007/978-94-017-1339-9 Copyright Springer Science+Business Media Dordrecht 1968 Originally published by Dr. W.Junk N.V. Publishers. The Hague in 1968 Softcover reprint ofthe hardcover Ist edition 1968 Printed by Belinfante N. V. Dedicated to my daughter Prema in affectionate token of her discovery of the Coccinellid Mass Assemblage on the Himalaya a discovery that stimulated interest in high altitude entomology CONTENTS Page PREFACE. • . xiii CHAPTER 1. INTRODUCTION 1 I. The concept of high altitude 2. Mountains as insect habitat . 2 3. Mountain insects . 4 4. The forest-line on mountains: the threshold of high altitude biota . 4 5. The biome above the forest 7 6. Hypsobiont insects . 8 CHAPTER II. THE HIGH ALTITUDE ENVIRONMENT. 9 I. The concept of high altitude environment 9 2. Reduced atmospheric pressure . 10 3. Atmospheric temperature. 16 a. Temperature lapse rate. b. Shade and sun temperatures. c. Diurnal variations of atmospheric temperature. d. Annual variations of atmo­ spheric temperature. e. Ecological importance of atmospheric cold at high elevations. 4. Atmospheric precipitation . .. 23 5. Snow-cover . .. 25 a. The winter snow. b. Snowline. c. Ecological importance of snow­ cover. 6. Wind. 30 7. Light . 31 8. Microclimate. 32 a. Microclimatic conditions on rock. b. Microclimatic conditions under stones. c. Soil temperature. d. Soil moisture. 9. Slope aspect and massiveness of the mountains 41 10. Gravity . 43 II. Biotic factors. 44 a. Absence of trees: Open habitats. b. Nutritional conditions. c. Crowd- ing and isolation. 12. Interdependence of the lowland and high altitude environments . .. 48 CHAPTER III. ECOLOGICAL SPECIALIZATIONS OF HIGH ALTITUDE INSECTS . 51 I. The concept of high altitude specializations. 51 2. High altitude melanism . 52 3. Reduction and loss of wings . 54 4. Some other structural peculiarities 58 5. Cold stenothermy. 61 6. Hygrophily and terricoly. 63 7. Other habits . 65 8. Hibernation and life-cycle 71 CHAPTER IV. ECOLOGICAL INTERRELATIONS AT HIGH ALTI- TUDES AND HIGH ALTITUDE INSECT COMMUNITIES. .. 75 I. Interrelations of the high altitude, the montane forest and the low- land biota . .. 75 VIII Contents Page 2. Interrelations within the high altitude biota. 78 3. High altitude insect communities 82 4. Rock communities . 84 S. Hypolithic communities . 8S 6. Soil communities. 88 7. Cavernicole communities. 89 8. Snow communities . 90 9. Aquatic communities . 92 a. Communities of stagnant waters. b. Communities of torrential streams. c. Communities of thermal and chemical springs. CHAPTER V. SOME TYPICAL HIGH ALTITUDE INSECTS AND OTHER ARTHROPODA .. 99 A.Insects. .. 99 Order I. Ephemerida 99 Order 2. Plecoptera . 100 Order 3. Orthoptera 102 Order 4. Dermaptera 104 Order S. Grylloblattodea. lOS Order 6. Heteroptera . ros Order 7. Coleoptera... ro6 Family Carabidae. Family Paussidae. Family Amphizoidae. Family Dytiscidae. Families Silphidae and Scydmaenidae. Families Staphy­ linidae and Pselaphidae. Family Histeridae. Family Hydrophilidae. Family Byrrhidae. Family Dryopidae. Family Tenebrionidae. Family Chrysomelidae. Family Curculionidae. Other families. Order 8. Hymenoptera II4 Order 9. Trichoptera . lIS Order ro. Lepidoptera. II6 Family Pamassiidae. Families Nymphalidae and Satyridae. Family Pieridae. Order II. Diptera. Nematocera. Family Tipulidae. Family Bibionidae. Family Blepha­ roceridae. Family Simuliidae. Family Deuterophlebiidae. Family Chironomidae. Family Psychodidae. Family Culicidae. Family Sciariidae. Brachycera-Orthorrhapha. Brachycera-Cydorrhapha. Aschiza. Hypocera. Schizophora. Order 12. Thysanura . 123 Order 13. Collembola ..... 123 B. Other Arthropoda ....... 12S Crustacea. Arachnida. Scorpionida. Chelonethida. Araneida. Acarina. Chilopoda-Diplopoda-Symphyla Complex. CHAPTER VI. DISTRIBUTION OF HIGH ALTITUDE INSECTS. 130 I. Factors governing the distribution of high altitude insects 130 2. Vertical distribution. ..... 132 3. Horizontal distribution . 137 4. The component elements of the high altitude faunas, their origin and evolution. .•......• 139 Contents IX 'Page CHAPTER VII. THE MOUNTAINS OF EQUATORIAL EAST AFRICA 143 I. The East African mountains and the Rift Valley system 143 2. Mt. Kilimanjaro . 144 i. Ecological conditions on Mt. Kilimanjaro . 146 ii. Biotic zones on Mt. Kilimanjaro. 148 iii. Early studies on insects from Kilimanjaro . 150 iv. Ecological characteristics of high altitude insects from Kilimanjaro. 150 v. Distributional characters of the high altitude insects from Kili- manjaro. 153 vi. Vertical distribution ofinsects on Mt. Kilimanjaro 154 3. Mt. Meru, Mt. Kenya, Mt. Aberdare and Mt. Elgon. 157 4. The Ruwenzori Range . 162 i. General ecological conditions and biotic zonation on the Ruwenzori Range. ii. High altitude insects from the Ruwenzori Range. 5. The Ethiopian Highlands. 167 i. Biotic zones on the Abyssinian highlands. ii. Characteristics of the high altitude insect life of the Ethiopian Highlands. CHAPTER VIII. OTHER TROPICAL MOUNTAINS. 176 I. New Guinea mountains . 177 2. The Andes. 180 3. Some ecological peculiarities of tropical mountains. 183 4. Altitudinal zonation of life on the Andes. 187 5. Some peculiarities of the insect life at high altitudes on the Andes of the tropics. 191 6. Boreal faunal elements on tropical mountains. 194 CHAPTER IX. THE HIMALAYA. 196 I. The Himalayan System . 197 2. The ranges of the Himalaya 201 3. Geographical and biogeographical divisions of the Himalaya 204 4. Geology and glaciology . 206 5. Ecological characters and biotic zones . 208 6. The insect life of the Himalaya. 211 Ephemerida. Plecoptera. Orthoptera. Dermaptera. Heteroptera. Coleoptera. Carabidae. Dytiscidae. Hydrophilidae. Staphylinidae. Tenebrionidae. Chrysomelidae. Curculionidae. Hymenoptera. Tri­ choptera. Lepidoptera. Papilionidae. Parnassiidae. Nymphalidae. Satyridae. Pieridae. Lycaenidae. Diptera. Thysanura. Collembola. 7. The Hindu Kush Range. 226 8. Biogeographical characters. 227 CHAPTER X. THE ALAI-PAMIRS AND THE TIEN SHAN. 229 I. The Alai-Pamirs Region. 229 2. Subdivisions of the Pamirs. 231 3. Ecological characters of the Pamirs Region. 234 4. Vegetation and biotic zones of the Pamirs Region. 240 i. The Pamir proper . 242 ii. The Western Valleys. 242 x Contents Page iii. The Alai vegetation . 243 iv. Altitudinal biotic zonation. 243 5. Insect life of the Pamirs Region. 245 Orthoptera. Dermaptera. Heteroptera. Coleoptera. Carabidae. Staphylinidae. Tenebrionidae. Chrysomelidae. Curculionidae. Hy­ menoptera. Formicidae. Apidae. Bombidae. Lepidoptera. Diptera. 6. The Tien Shan Mountains . 7. General ecological and vegetational characters of the Tien Shan 8. Insect life of the Tien Shan. 9. Biogeographical characters of the Pamirs-Tien-Shan-Complex. i. Steppecole elements . ii. Montane elements . iii. Boreal and boreo-alpine elements. iv. Endemic elements . v. Faunal subdivisions . vi. Faunistic affinities. CHAPTER XI. THE ALPS AND THE CARPATHIANS. 275 I. The elevated areas of Europe. 275 2. The Alps ................... 279 3. The ecological characters and biotic zonation of the Alps . 284 4. The insect life of the West Alps. 288 Orthoptera. Heteroptera. Coleoptera. Lepidoptera. Diptera. Collem­ bola. 5. The insect life of the East Alps . 292 Orthoptera. Dermaptera. Coleoptera. Hymenoptera. Lepidoptera. Diptera. Collembola. 6. Other Arthropoda. Araneida. Acarina. Diplopoda and Chilopoda. 7. Communities and ecological succession on the East Alps 299 i. Some typical communities. 299 ii. Some peculiarities of ecological succession. 30 1 8. The Carpathians . 302 9. The ecological characters and biotic zonation of the Carpathians. 30 4 IO. Characteristics of high altitude insects from the Carpathians . 306 II. Biogeographical characters of the Alps and the Carpathians . 3IO CHAPTER XII. THE BOREO-ALPINE INSECTS. 31 3 I. What are boreo-alpine insects? . 313 2. Some typical boreo-alpine insects . 314 Orthoptera. Heteroptera. Coleoptera. Hymenoptera. Lepidoptera. 3. Some typical boreo-alpine Arachnida . 324 4. The Boreo-alpine insects in the north area 324 5. The boreo-alpine insects in the south area. 327 6. The origin of boreo-alpine insects. 330 CHAPTER XIII. OTHER MOUNTAINS OF THE NORTH TEMPERATE REGIONS OF THE OLD WORLD 336 I. The Pyrenees. 336 2. The Spanish Sierra Nevada and other Spanish mountains. 340 Contents XI Page 3. The Adas Mountains . 344 4. The French Central Plateau, the Jura Mountains, Vosges, the Sudeten, etc. 344 5. The Apennines. 346 6. The Balkan Mountains. 348 7. The Caucasus. 349 8. The Armenian Highlands. 354 9. The Ural Mountains. 355 10. The Altai and other mountains of the Angar-Beringea . 356 II. General ecological characters of the Altai. 357 12. The insect life of the Altai . 358 13. Other mountains of Angar-Beringea. 360 i. The Sikhote-Alin mountains. 361 ii. The mountains of Kamchatka . 361 iii. The Japanese mountains. 362 14. Biogeographical affinities of the European and Angaran mountains. 363 CHAPTER XIV. THE MOUNTAINS OF THE NORTH TEMPERATE REGIONS OF THE NEW WORLD. 365 I. The Appalachian Highlands. 365 2. Ecological characters of the White Mountains. .' . 366 3. Altitudinal biotic zones on the North American mountains. 368 4. Characters of the high