Springer. Adaptations of Desert Organisms. Edited by J. 1. Cloudsley-Thompson

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1 Adaptations of Desert Organisms Edited by J. 1. Cloudsley-Thompson Springer Berlin Heidelberg New York Barcelona Budapest Hong Kong London Milan Paris Tokyo

2 Volumes already published Ecophysiology of the Camelidae and Desert Ruminants By R. T. Wilson (1989) Ecophysiology of Desert Arthropods and Reptiles By J. L. Cloudsley-Thompson (1991) Plant Nutrients in Desert Environments By A. Day and K. Ludeke (1993) Seed Germination in Desert Plants By Y. Gutterman (1993) Behavioural Adaptations of Desert Animals By G. Costa (1995) Invertebrates in Hot and Cold Arid Environments By L. S0mme (1995) In preparation Energetics of Desert Invertebrates By H. Heatwole (1995) Ecophysiology of Desert Birds By G.L. Maclean (1996)

3 Lauritz S0mme Invertebrates in Hot and Cold Arid Environments With 77 Figures Springer

4 Professor Dr. LAURITZ S0MME University of Oslo Department of Biology P.O. Box 1050 Blindern N-0316 Oslo Norway Cover illustration: Photograph by /.L. Cloudsley-Thompson ISBN-13: e-isbn-13: DOl: / Die Deutsche Bibliothek - CIP-Einheitsaufnahme Semme, Lauritz: Invertebrates in hot and cold arid environments / Lauritz Semme. - Berlin; Heidelberg; New York: Springer,1995 (Adaptations of desert organisms) ISBN -13' CIP data applied for This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically of translation, reprinting reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. o Springer-Verlag Berlin Heidelberg 1995 Sof'tcover reprint of the hardcover 1st edition 1995 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relavant protective laws and regulations and therefore free for general use. Typesetting: K. Darms, GOttingen SPIN / Printed on acid-free paper

5 Preface The idea to write a book on the adaptation of invertebrates from both hot and cold arid environments in the series Adaptations of Desert Organisms originated from Professor J.L. Cloudsley-Thompson. My thanks are due to him for his encouragement during the preparation of the present volume, and for his many suggestions to improve the content. I am also most grateful to Professor K.E. Zacharias sen, University of Trondheim, Dr. H. Raml0v, University of Copenhagen and Dr. M. Holmstrup, University of Aarhus for valuable comments to parts of the manuscript. The scanning electron microscope photographs of tardigrades have been reproduced by the courtesy of Drs. R.M. Kristensen and H. Raml0v, University of Copenhagen, and the photographs of freezing in nematodes by the courtesy of Dr. David Wharton, University of Otago. Traditionally, the biology of plants and animals from the hot temperate and subtropical deserts is treated separately from those of cold polar deserts. Within the series Adaptations of Desert Organisms, edited by Professor J.L. Cloudsley-Thompson, most books are concerned with hot deserts, while particular attention is given to polar deserts in the present volume. To point out differences and similarities, an attempt is made to compare invertebrates from such different regions, in particular with regard to adaptations to their hostile environments. Questions are raised if species from hot and cold deserts exhibit similarities in their tolerance of drought, and how species from both regions are adapted to low temperatures. To provide a background to these problems, the climate and fauna of hot and cold deserts are described in the first two chapters. In the Arctic and the Antarctic, precipitation is low and comparable to that of many deserts in warmer regions. In winter, snow and ice predominate, and liquid water is unavailable to terrestrial organisms. Clearly, invertebrates inhabiting these regions must be able to survive both aridity and low temperatures. Similarly, species from hot temperate deserts are not only adapted to dry conditions but to cold as well. Winter severity increases with latitude, altitude and continentality. The fauna of insects, spiders and myriapods of hot deserts is fundamentally different from that of polar deserts, suggesting differences in historical development and specificity of adaptation. On the other hand, similarities are seen in the fauna of mites and springtails, comprising important elements

6 VI Preface in soil ecosystems. Nematodes are abundant in both hot deserts and polar regions. Like tardigrades, they are able to survive almost complete loss of water. The following chapters discuss an array of adaptational strategies required for life in hot and cold deserts. In arthropods, several physiological mechanisms of water conservation and cold hardiness have been developed. Of equal importance to insects and other terrestrial arthropods is the ability to avoid the most stressful situations through their behaviour. Microc1imatic niches, ameliorating extreme moisture and temperature fluctuations, are of paramount importance. The unpredictable moisture conditions of hot deserts favour opportunistic activity and development when water is available. Dormancy may be an important strategy to survive adverse periods. In polar areas, seasonal changes are regular, but the summer is short and cool; the winter is long and cold, retarding the phenology of most species. Recently, several authors have suggested that the tolerance of terrestrial arthropods to cold and drought may, in fact, be overlapping adaptations. Through their behaviour and physiology, many species are protected against both cold and drought. In tardigrades, nematodes and earthworms, tolerance to dehydration enhances resistance to cold. Our present knowledge on the adaptations of invertebrates to hot and cold arid environments is far from complete. To understand the role of the organisms in their environment, more physiological and ecological studies are required. In many taxa, behavioural and phenological adaptations are poorly understood. Surprisingly, even information on the biodiversity of many hot and cold desert regions is still incomplete. It is hoped that the presentation of the different problems, as described in this book, may stimulate to further research on the variety of desert invertebrates and their biology. The present volume covers a broad spectrum of topics, and relies heavily on the use of illustrations from other publications. I am most grateful for permission received from the different authors personally, and for the courtesy of the publishers in permitting reproduction of figures from their publications. Figures are reprinted with the kind permission of the following publishers and organizations: Academic Press Ltd., London (Figs. 2.1, 2.5, 5.5, 6.1); Academic Press Inc., Orlando (Figs. 1.5, 9.10, 9.12); American Physiological Society, Bethesda (Fig. 4.4); Annual Reviews Inc. (Annual Review of Physiology), Palo Alto (Fig. 5.6); Australian Museum, Sidney (Fig. 2.3); Blackwell Science Ltd., Oxford (Figs. 7.2, 11.4, Table 9.3); Canadian Museum of Nature, Ottawa (Fig. 2.10); Cambridge University Press, Cambridge (Figs. 1.1,2.7, 7.3); Company of Biologists Ltd., Cambridge (Figs. 4.3,6.3,6.4); Cryo-Letters, Cambridge (Fig. 9.6); Czech Academy of Science, Ceske Budejovice (Fig. 3.7); Elsevier Science Ltd., Kidlington (Figs. 4.2, 4.6, 8.1,8.2,8.3, 9.11); Entomological Society of Canada, Ottawa (Figs. 1.2, 1.3); Gauthier-Villars Publishers, Montrouge (Figs. 3.1, 4.5); Japan Plant Protection Association, Tokyo (Fig. 8.4); John Wiley & Sons Inc., New York

7 Preface VII (Figs. 5.2, 5.4, 7.7); National Research Council of Canada, Ottawa (Figs. 7.1, 11.2); Norwegian Institue of Nature Research, Trondheim (Fig. 4.1); Munksgaard International Booksellers and Publishers, Copenhagen (Fig. 2.4, 9.1, 9.5); Redacci6n de Vieraea, La Laguna (Fig. 3.3); Regents of the University of Colorado, Boulder (Figs. 1.9, 6.6, 6.7); Society of Nematologists, Washington D.C. (Fig. 5.1); Springer-Verlag, Heidelberg (Figs. 1.8,2.6,3.2, 6.5,7.5,7.6,9.2,9.13, ILl, 11.3); University of Arizona Press, Tucson (Fig. 2.2). Oslo, Spring 1995 Lauritz S0mme

8 Contents 1 The Climate of Arid Environments j 1.1 Subtropical and Temperate Deserts Definitions The Distribution of Deserts Microclimate Polar Deserts The Arctic..., The Antarctic Microclimate The Limitation of Water in Antarctic Terrestrial Ecosystems Latitudinal Variations Microclimatic Variations Conclusions The Fauna of Hot and Cold Arid Deserts The Diversity of Hot Desert Invertebrates Introduction..., Soil Fauna: Microarthropods and Nematodes Ants and Termites Herbivores Detritivorous Macroarthropods Necrophagous and Coprophagous Arthropods Predaceous Arthropods..., The Fauna of Polar Regions..., Introduction Nematodes Tardigrades Rotifers Other Invertebrates Terrestrial Arthropods..., Comparison of Hot and Cold Arid EnVironments... 51

9 X Contents 3 Tolerance to Arid Conditions in Terrestrial Arthropods Microarthropods Alpine Species Polar Species Coleoptera Alpine Species Polar Species Tropical and Temperate Lowland Species Other Terrestrial Arthropods From Lowland Deserts Alpine Millipedes Conclusions The Physiology of Tolerance to Desiccation Introduction Evaporative Water Losses Beetles Scorpions The Cuticula of Terrestrial Arthropods Structure Restriction of Water Loss Cuticular Lipids Desert Arthropods...; Evaporative Cooling Water Absorption The Hindgut of Insects The Cryptonephridial System Water Absorption in Other Terrestrial Arthropods Osmoregulation Occurrence in Different Taxa Desert Tenebrionids Alpine Beetles Conclusions Anhydrobiosis in Terrestrial Invertebrates Introduction Nematodes Tolerance to Desiccation Rates of Dehydration Nematodes from Hot Deserts Antarctic Nematodes Accumulation of Trehalose Tardigrades Tun Formation

10 Contents Ecological Implications Accumulation of Trehalose Recovery from Anhydrobiosis Rotifers Terrestrial Arthropods Collembola Chironomid Midges Biochemical Aspects of Anhydrobiosis Conclusions XI Behavioural Adaptations Arthropods from Hot Deserts Introduction Burrowing Circadian and Seasonal Rhythms Thermoregulation Polar and Alpine Terrestrial Arthropods Introduction Mating and Oviposition Microarthropods Behavioural Thermoregulation Avoidance of Heat and Cold by Alpine Tropical Arthropods Phenological Adaptations Life Cycles Hot Desert Invertebrates Polar Species Alpine Species Conclusions Diapause and Quiescence Introduction Principle Features of Diapause Dormancy in Tropical Insects Dormancy in Polar Species Conclusions Water Balance During Overwintering Introduction Choice of Overwintering Sites Environmental Factors Cues for Overwintering Anaerobiosis

11 XII ILl Contents Moisture Content of Overwintering Sites Lack of Fluid Water Water Vapour Pressure in Frozen Habitats Longevity and Saturation Deficit Inoculative Freezing Contact with Ice and Inoculation Protection Against Inoculative Freezing The Effect of Dehydration on Cold Hardiness Conclusions Cold Hardiness in Terrestrial Arthropods Introduction Concepts of Cold Hardiness Definition of Important Concepts Freeze Susceptible Species The Importance of Supercooling Supercooling Capacity in Microarthropods Removal of Ice-Nucleating Agents Seasonal Changes Gut Content and Ice-Nucleating Agents Low Molecular Weight Cryoprotectant Substances Thermal Hysteresis Proteins Cryoprotectants in Alpine and Polar Microarthropods Freeze Tolerant Species Why is Freezing Injurious? The Occurence of Freeze Tolerance in Insects Polar and alpine insects Variations in the Degree of Freeze Tolerance Ice Nucleation in Freeze Tolerant Species Cryoprotectant Substances Freeze Tolerance Versus Freeze Avoidance Hot Desert Arthropods at Low Temperatures Preadaptation to Cold Cold Hardiness of Different Taxa Scorpions Centipedes Spiders Insects Conclusions Cold Tolerance in "Lower" Invertebrates Nematodes Strategies of Cold Hardiness

12 Contents Antarctic Nematodes Freeze Tolerance During Anhydrobiosis Freeze Tolerance or Freeze Avoidance? Tardigrades Freezing in Liquid Gases Freezing at Environmental Temperatures Antarctic Tardigrades Rotifers Lumbricid Earthworms Effect of Dehydration Long Term Survival A Protective Dehydration Mechanism Conclusions Summary and Concluding Remarks Deserts The Fauna Behaviour Life Cycles Terrestrial Arthropods and Aridity Cold Hardiness Terrestrial Arthropods Tardigrades and Nematodes Lumbricid Earthworms Combined Tolerance of Cold and Drought Anhydrobiosis XIII References Subject Index