Animal Welfare Series Editor Clive Phillips School of Veterinary Science University of Queensland Gatton, QLD Australia Advisory Editors Marieke Cassia Gartner Atlanta, GA, USA Karen F. Mancera Mexico City, Mexico Fiona C. Rioja-Lang Mérida, Yucatán, Mexico More information about this series at http://www.springer.com/series/5675 Claudio Carere • Jennifer Mather Editors The Welfare of Invertebrate Animals Editors Claudio Carere Jennifer Mather Department of Ecological and Biological Department of Psychology Sciences, Ichthyogenic Experimental University of Lethbridge Marine Centre (CISMAR) Lethbridge, Alberta, Canada University of Tuscia Tarquinia, Viterbo, Italy ISSN 1572-7408 Animal Welfare ISBN 978-3-030-13946-9 ISBN 978-3-030-13947-6 (eBook) https://doi.org/10.1007/978-3-030-13947-6 © Springer Nature Switzerland AG 2019 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Editor Fiona C. Rioja-Lang deceased in 2019 at the time of publication My Vote for the “Invertebrates” At the time when I was a young student and quite some time beyond, most biologists were convinced: if you can’t even know what your best friend is thinking and feeling, how can you ever know what is going on in an animal? It was considered unscientific for such dogmatic reasons to seriously ask a question regarding the mental life, the thinking and feeling of an animal. You could not hope to find answers but would rather get lost, if you dared to delve into this black hole. Among widespread beliefs and attitudes were the following: (1) that we can never say anything about feelings or thoughts of animals; (2) that man is too distinct from animals to be compared with them; (3) that man is the centre of the world and human society has to take care of humanity only; and (4) that animals are dull and “bêtes machines”, as Descartes said, just things, which automatically follow their instincts in a preprogrammed way. I am of course exaggerating. Darwin had firmly embedded man in the process of evolution and placed us on a continuum with animals a long time ago, and there have always been keepers of cats, dogs and horses and maybe even birds who would insist that their pets do think and feel and have emotions and personality. Obviously, it is easier to assign such capabilities to animals the closer these are to us humans phylogenetically. But invertebrates like crickets and fruit flies? For most listeners, it was at best an entertaining joke when I talked about the individuality of spider males, impressed by the differences of their courtship behaviour. Remember that Jane Goodall even met with difficulties when giving her chimpanzees in Gombe human names and personality in the 1960s. Although much of the attitude prevailing at my time as a biology student still exists outside our field of expertise, big changes of our view of the world have occurred since. Many of us (though not enough yet) now better understand that humankind has responsibilities beyond itself or, put in another way, that we have to take care of the entire planet with all its diversity of life and environmental com- plexities. Nature is not just a machine endlessly producing for us only, or a big buffet table at which we can endlessly serve ourselves without paying. And wilderness is not just luxury. We have instead begun to better understand that all living beings are vii viii My Vote for the “Invertebrates” interconnected on this planet and depend on each other, that sustainability is a big issue and that nature cannot be treated as if it existed to serve our human needs and pleasures only. I hopefully assume that there is more awareness of biophilia now, following the psychoanalyst Erich Fromm and even more so the biologist Edward O Wilson: the appreciation of all living beings and a feeling of deep-seated connect- edness. Some unpleasant changes critically endangering our own human welfare like overpopulation, food and energy shortage and climate change have helped to promote this point of view. A few societies only, mostly small and of little global influence, have kept the old traditions of their balanced view of nature. Unfortu- nately, they are unable to stop the dramatic impact of industrialization and the unlimited endeavour for our human well-being and materialistic success. I am not an expert on questions of animal welfare but instead an experimental physiologist doing basic research. As is true for many biologists, my own biophilia implies a sense for the deep value of nature and an interest in its increasingly profound understanding, including an understanding of our human place in it. Apart from my professional interest in science, it is this background and wishful thinking which underlies my interest in animal welfare and the invertebrates. I read through this book on the welfare of invertebrate animals with great interest and much appreciate that the topic is given so much attention. Apart from the presentation of practical cases like that of butterfly houses, bee keeping, spiders in captivity and captive cephalopods, experts also address questions of philosophy and ethics. I was pleased to find so many honest “might”, “could”, “probably” and subjunctives. No wonder we are still far from a clear understanding of nociception and pain, the integration of sensory information in the central nervous systems of most invertebrates and the selection pressures and fitness gains associated with much of invertebrate behaviour. Feelings and thoughts of a snail or an earthworm or a cockroach are still obscure issues. And of course, one cannot expect any overarching general recipe to provide welfare for such a large and diverse group of animals as are the invertebrates. From a zoological point of view, the “animals without backbones” are much too heterogeneous to be put into one phylum. Their diversity could not be greater, considering their Bauplan, their habitat, nervous and sensory systems and behaviour. A sessile sponge filter feeding in the sea, an earthworm digging through the soil, a spider building mechanical masterpieces of aery silken webs, a cricket singing love songs or a dragonfly manoeuvring masterly through the air: the differences could not be greater. For the zoologist, the so-called invertebrates (not a serious terminology in taxonomy) comprise as many as 26 of a total of 27 phyla, the only additional one being the chordates which include us and all other vertebrates. “Invertebrates” make up for at least 95% of all animal species so far described. A few numbers illustrate their dominance impressively: insects c. 1,000,000 described species (and probably some 10 million existing) as opposed to vertebrates with only c. 66,000 species; even the arachnids are good for more than 100,000 species, with the spiders alone representing almost 50,000 described species. We are living in an age of insects and even more so of arthropods. Seen from this and an evolutionary point of view, it My Vote for the “Invertebrates” ix is frustrating that some people still talk about “insects (or even bugs) and animals”. Aren’t insects like all other “invertebrates” worth being called animals as well? When we talk about the welfare of the “invertebrates”, we should also gratefully appreciate how important they have become in research, allowing us to reveal many fundamental secrets of life at all levels of organization, from molecular to organis- mal. The fruit fly Drosophila, the nematode Caenorhabditis, the honeybee Apis, the coelenterate Nematostella and others have become outstandingly important model organisms. Even spiders, disliked by many for no good reason, have made their way into experimental research asking serious questions of cognition and personality. Neurobiology owes the understanding of action potentials to the giant fibres of the squid Loligo, that of lateral inhibition to the horseshoe crab Limulus and that of learning and memory to a large extent to the sea hare Aplysia. And who had thought only some thirty years ago that the fruit fly and bees and jumping spiders might play a significant role in cognition research? Cognitive abilities of sometimes startling complexity now reach down far in the phylogenetic tree. Man is not so distinct from the rest of the animal kingdom anymore, including the “invertebrates”. Latest research has brought to light rather surprising deep homologies conserved through- out the metazoans. There are many correspondences of neural circuits in arthropods and chordates (vertebrates), and there is evidence for the conservation of neural ground patterns and genetic mechanisms underlying brain development.