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Biology of Earthworms Home Springer
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Biology of, Earthworm s, C A Edwards, Rotlwmsud Experimental Station. SPRINGER SCIENCE BUSINESS MEDIA B V, C A Edwards and J R Lofty 1972. Originally published by Chapman and Hall Ltd in 1972. Softcover reprint ofthe hardcover 1st edition 1972. SB N 412 11060 1, ISBN 978 0 412 11060 3 ISBN 978 1 4899 6912 5 eBook. DOI 10 1007 978 1 4899 6912 5, All rights reserved No part of this publication. may be produced stored in a retrieval system or, transmitted in any form or by any means electronic.
mechanical photocopying recording or otherwise, without the prior written permission of the publisher. Foreword pagex, Preface Xlll, 1 Morphology 1, 1 1 Segmentation external 1. 1 2 Chaetotaxy 4, 1 3 Genital and other apertures 7. 1 4 The clitellum and associated structures 9, 1 5 Pigmentation 13. 1 6 The body wall 13, 1 7 The coelom 17, 1 8 The alimentary canal 19.
1 9 The vascular system 21, 1 10 The respiratory system 23. 1 11 The excretory system 24, 1 12 The nervous system 28. 1 13 The reproductive system 33, 2 Taxonomy 37, 2 1 Systematic affinities and descent 37. 2 2 Families genera and species 39, 2 2 1 Moniligastridae 40. 2 2 2 Megascolecidae 41, a Megascolecidae sensu strictu 42.
b Ocnerodrilidae 44, c Acanthodrilidae 45, d Octochaetidae 46. Vl BIOLOGY OF EARTHWORMS, 2 2 3 Eudrilidae 47, 2 2 4 Glossoscolecidae 49. 2 2 5 Lumbricidae 51, 2 3 Geographical distribution 53. 3 Biology 56, 3 1 Life cycles 56, 3 2 Reproduction 62. 3 2 1 Spermatogenesis 62, 3 2 2 Oogenesis 63, 3 2 3 Copulation and fertilization 63.
3 3 Quiescence diapause and aestivation 67, 3 4 Growth 69. 3 5 Behaviour 71, 4 Physiology 78, 4 1 Respiration 78. 4 2 Digestion 80, 4 3 Excretion 81, 4 4 Circulation 84. 4 5 Nervous system 85, 4 5 1 General 85, 4 5 2 Light reactions 87. 4 5 3 Chemoreception 88, 4 5 4 Thigmotactic reactions 89.
4 5 5 Responses to electrical stimuli 90, 4 6 Water relationships 90. 4 7 Locomotion and peristalsis 92, 4 8 Regeneration 94. 4 9 Transplantation 96, 5 Ecology 98, 5 1 Estimation of populations 98. 5 1 1 Handsorting 98, 5 1 2 Soil washing 99, CONTENTS Vll. 5 1 3 Electrical methods 99, 5 1 4 Chemical methods 100.
5 1 5 Heat extraction 101, 5 1 6 Comparisons of methods 101. 5 1 7 Number and size of samples 102, 5 2 Size of populations 102. 5 3 Population structure 104, 5 4 Population distributions 106. 5 4 1 Horizontal distributions 106, 5 4 2 Vertical distributions 107. 5 5 Seasonal populations and activity 110, 5 6 Burrowing and casting 117.
5 7 Species associations 122, 5 8 Predators and parasites 123. 5 9 Effect of environmental factors 125, 5 9 1 pH 125. 5 9 2 Moisture 127, 5 9 3 Temperature 129, 5 9 4 Aeration and Carbon dioxide 133. 5 9 5 Organic matter 133, 5 9 6 Soil type 136, 5 9 7 Food supply 138. 6 The role of earthworms in organic matter, cycles 141.
6 1 Fragmentation and breakdown 141, 6 2 Consumption turnover and humification 145. 6 3 Nitrogen mineralization 147, 6 4 Effects on the C N ratio 150. 6 5 Effect on available mineral nutrients 152, 7 Earthworms and micro organisms 155. 7 1 Effect of earthworms on number of micro organisms 155. 7 2 Effects of pH earthworms on dispersal of micro. organisms 161, 7 3 Stimulation of microbial decomposition 161. Vlll BIOLOGY OF EARTHWORMS, 8 Earthworms and soil fertility 163.
8 1 Effects of earthworms on soil structure 163, 8 1 1 Breakdown of soil particles 163. 8 1 2 Turnover of soil 164, 8 1 3 Formation of aggregates 165. 8 1 4 Aeration porosity and drainage 167, 8 2 Earthworms as indicators of soil type 168. 8 3 Effect of earthworms on crop yield 168, 8 4 Soil amelioration by earthworms 171. 9 Effects of agriculture on earthworm popula, 9 1 Effects of cultivations 174.
9 2 Effects of cropping 176, 9 3 The effects of fertilizers 178. 9 4 Mortality from pesticides 181, 9 5 Uptake of pesticides into earthworms 184. 9 6 Radioisotopes and earthworms 188, 10 Earthworms as pests and benefactors 190. 10 1 As pests of crops 190, 10 2 As pests of grassland and turf 191. 10 3 Control of earthworms 191, 10 4 Transmission of diseases 191.
10 5 Adverse effects on soil 196, 10 6 Earthworms as benefactors 197. 11 Simple experiments and field studies with, earthworms 198. 11 1 Cultures 198, 11 2 Preservation 200, 11 3 Dissection 200. 11 4 Field experiments with earthworms 200, 11 4 1 Distributions 200. 11 4 2 Effects of pH 201, CONTENTS ix, 11 4 3 Production of casts 201.
11 4 4 Incorporation of plant organic matter 201, 11 4 5 Palatability of leaves 203. 11 4 6 Burrowing experiments 204, 11 4 7 Life history studies 206. 11 4 8 Vertical and horizontal distribution of earth. 11 4 9 Effects of insecticides on earthworms 209, 11 4 10 Physiological experiments 209. 11 4 11 Behavioural experiments 210, A simplified key to common genera of terrestrial earth. References 219, Systematic 253, Author 261, General 268.
by Dr J E Satchell, Merlewood Research Station, Grange over Sands Lancashire. As the classroom type of the Annelida the earthworm is familiar to. every student of biology Dissection manuals neatly package its. anatomical features and numerous texts admirably relate earthworm. form and function Beyond these frontiers student and teacher alike. are met by a bewildering mass of publications ranging in content. from the military value of earthworms as survival food to the. species stored by moles The volume of this literature is quite. exceptional for an invertebrate group which is important neither as. pest nor food It stems from the inherent zoological interest of the. earthworm as a terrestrial form retaining many of the charac. teristics of its aquatic ancestors from its convenience as an experi. mental animal for behaviourists and physiologists and from its. effects on soil fertility, This last aspect had already been perceived two centuries ago by. Gilbert White of Selbome who in 1770 wrote Earthworms. though in appearance a small and despicable link in the chain of. Nature yet if lost would make a lamentable chasm worms seem. to be the great promoters of vegetation which would proceed but. lamely without them In more recent times such ideas have. been critically examined and extended at Rothamsted Experimental. Station in a programme of earthworm research pioneered by Sir. John Russell and continued by a number of workers during the last. twenty five years It is particularly appropriate that the task of. surveying the earthworm literature and summarising it at readable. length should have been undertaken by the current representatives. of this Rothamsted tradition, An important part of this book which will be particularly valuable. to soil zoologists is the bibliography of between five and six. BIOLOGY OF EARTHWORMS Xl, hundred literature references An analysis of their frequency dis. tribution in time throws an interesting light on the development of. earthworm studies Divided into twenty year periods they fall as. follows 1870 1889 6 1890 1909 12 1910 1929 68 1930 1949 106. 1950 1969 361 Allowing for some selection in favour of more. recent studies it seems that whereas around the turn of the century. earthworm papers were published on average about one every other. year in the last two decades one was published on average about. every three weeks If publication continues at this exponential rate. a revision of this book in twenty years time will involve the formid. able task of reviewing another 640 papers, In the face of this alarming publication explosion it is interesting.
to note how slowly concepts and attitudes change even amongst. contemporary ecologists Charles Darwin s famous book The. Formation of Vegetable Mould through the Action of Worms. published in 1881 provides a good example When Henry James. described the requirements for a successful mid Victorian genre. painting as that it shall embody some comfortable incident of. the daily life of our period suggestive more especially of its. familiar moralities he might well have had in mind Darwin s. analogies of earthworms as beneficent gardeners and industrious. ploughmen It is significant to find this anthropocentric approach. epitomised almost a century later in this present book in the chapter. title Earthworms as benefactors Indeed as the authors show some. of our ideas about earthworms can be traced back to Aristotle. Despite the voluminous literature of the intervening period. E W Russell summarizing our knowledge of the importance of. earthworms in agriculture wrote in 1950 They may play an im. portant role in the conversion of plant into humus but this has. not yet been rigorously proved Twenty two years later this is still. substantially true for we now know that earthworm activity is. primarily important through its effects on soil microflora and these. effects are extremely difficult to measure outside the artificial con. ditions of laboratory cultures Advances in this field await develop. ments in the techniques of microbiology, The prospective student of earthworm ecology should not be. inhibited by the weight of past research for much remains to be. done Important land use changes may be expected in Europe in the. Xll FOREWORD, coming decades with the restoration of former industrial sites and. the withdrawal of five million hectares from agricultural use under. the Mansholt plan It is in this marginal and restored land rather. than in the agricultural lowlands that earthworm research is likely. to be most relevant and fruitful for it is under the less intensively. managed land on base rich sites that soil processes generally become. dominated by earthworm activity Outside the temperate zone and. particularly in the tropics much progress has been made in recent. years in the basic taxonomy of indigenous earthworm species but. knowledge of their functional role as components of ecosystems is. at the best fragmentary Dr Edwards and Mr Lofty s review of the. current state of earthworm knowledge will prepare the way for. another generation of research in these diverse and important fields. Earthworms can be defined as terrestrial invertebrates that belong. to the Order Oligochaeta Class Chaetopoda Phylum Annelida. Their characteristic features are that they are externally segmented. with a corresponding internal segmentation and usually have setae. on all segments They are hermaphrodite and produce cocoons. containing eggs which hatch into immature worms that are similar. to the adults except in size and development of their genital organs. Earthworms range in size from a fraction of a em to exceptional. individuals of Megascolides australis which may measure 2 75 min. length and 3 em in diameter, Oligochaetes can be aquatic or terrestrial members of nine. families are aquatic and rather small Microdrili whereas worms in. the other five families are mostly terrestrial and much larger Mega. drili It is worms in the latter group that we normally call earth. worms There are about 1 800 species of earthworms distributed. all over the world The commonest worms in Europe Western. Asia and most of North America belong to the family Lumbricidae. the twenty eight species belonging to eight genera reported from. the British Isles to date all belong to this family Other regions. of the world are dominated by species from different families. thus members of the Glossoscolecidae are most numerous in. Central and South America and the megascolecid group is the. most important family in southern and eastern Asia and Australasia. Nevertheless some lumbricids that are found commonly in. North America India Ceylon New Zealand and South Africa. have probably been introduced from Europe because their dis. tribution often follows European colonization and they are. most numerous close to populated areas Where these species. XIV BIOLOGY OF EARTHWORMS, are introduced they often multiply rapidly and supplant the native. Earthworms make a large contribution to the total weight or. biomass of invertebrates in soil particularly in temperate regions. Aristotle first drew attention to their role in turning over the soil. and called them intestines of the earth However it was Charles. Darwin 1881 who concentrated attention on the role of earth. worms in breakdown of dead plant and animal material in soil and. forest litter and in maintenance of soil structure aeration and. fertility His famous book The Formation of Vegetable Mould through. the Action of Worms summarized the conclusions he had reached. during the preceding forty years of experimental work and led him. to express the opinion that earthworms have played a most important. part in the history of the world Other scientists contemporary. with Darwin Hensen 1877 Muller 1884 Urquhart 1887 also. believed that earthworms played a beneficial part in soil formation. and some stated that they were essential for soil fertility. Many of the conclusions and hypotheses reached by Darwin have. been repeated so many times in the literature that they are almost. taken as unassailable fact and have been little tested either in field or. laboratory One of the difficulties of interpreting Darwin s conclu. sions is that he did not identify the species of worms in. Vlll BIOLOGY OF EARTHWORMS 8 Earthworms and soil fertility 163 8 1 Effects of earthworms on soil structure 163 8 1 1 Breakdown of soil particles 163 8 1 2 Turnover of soil 164 8 1 3 Formation of aggregates 165 8 1 4 Aeration porosity and drainage 167 8 2 Earthworms as indicators of soil type 168

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