Author | Frank Evers Beddard |
---|---|
Illustrator | Peter Smit and others |
Country | United Kingdom |
Subject | Camouflage |
Genre | Natural history |
Publisher | Swan Sonnenschein |
Publication date | 1892 |
Animal Coloration, or in full Animal Coloration: An Account of the Principal Facts and Theories Relating to the Colours and Markings of Animals, is a book by the English zoologist Frank Evers Beddard, published by Swan Sonnenschein in 1892. It formed part of the ongoing debate amongst zoologists about the relevance of Charles Darwin's theory of natural selection to the observed appearance, structure, and behaviour of animals, and vice versa.
Beddard states in the book that it contains little that is new, intending instead to give a clear overview of the subject. The main topics covered are camouflage, then called 'protective coloration'; mimicry; and sexual selection. Arguments for and against these aspects of animal coloration are intensively discussed in the book.
The book was reviewed in 1892 by the major journals including The Auk, Nature, and Science. The scientist reviewers Joel Asaph Allen, Edward Bagnall Poulton and Robert Wilson Shufeldt took up different positions on the book and accordingly praised or criticized Beddard's work.
Modern evaluation of the book is from a variety of perspectives, including the history of Darwinism, the history of the Thayer debate on the purpose of camouflage, the mechanisms of camouflage, sexual selection, and mimicry. Beddard is seen as having covered a wide swath of modern biology with both theory and experiment.
Beddard (1858–1925) was an English zoologist specializing in Annelid worms, but writing much more widely on topics including mammals and zoogeography. He also contributed articles on earthworms, leeches and nematode worms to the 1911 Encyclopædia Britannica . His decision to write an accessible book on animal coloration falls into this pattern. Beddard wrote Animal Coloration at a time when scientists' confidence in Charles Darwin's theory of evolution by natural selection [1] was at a low ebb. Beddard's book [P 1] was part of an ongoing debate among zoologists about how far natural selection affected animals, and how far other forces – such as the direct action of light – might be the causes of observed features such as the colours of animals. Edward Bagnall Poulton's far more strongly pro-Darwinian book The Colours of Animals had appeared just two years earlier in 1890. [2]
Beddard explains in his preface that the book grew from his 1890 Davis Lectures given for the public at London Zoo. The book "contains hardly anything novel, but professes to give some account of the principal phenomena of coloration exhibited by animals." [P 2] He also notes that since Poulton's recent book "deal[s] with colour almost entirely from the point of view of natural selection, I have attempted to lay some stress upon other aspects of the question." [P 2] Similarly, because Poulton treated insects in some detail, Beddard chooses to give more attention to other groups, though "it is impossible not to devote a good deal of space to insects". [P 3] The examples are mainly from Beddard's own observation of "animals that may be usually seen in the Zoological Society's Gardens", [P 3] though he also introduces and quotes the work of other scientists, including Henry Walter Bates and Alfred Russel Wallace.
The book has four colour plates by Peter Smit, who both drew and prepared the chromolithographic plates. Plate 1 is stated in the List of Illustrations "To face page 108", but as bound in the first edition it is used as a Frontispiece, facing the title page.
There are also 36 woodcuts (in black and white) in the text, though one of these, "Eolis and Dendronotus" is intentionally repeated as figures 10 and 19 to accompany the text in two places. The woodcuts vary from small line drawings on a simple white background (as in the diagrammatic figure 28 of Psyche helix , and figure 34 of the winter moth) to page-width illustrations like figure 2 which shows ermines in winter pelage, in a realistic depiction with a detailed snowy scene in the background. The woodcuts are certainly by a number of different artists; many are unsigned, but figures 5 and 26 are signed "E.A. Brockhaus X.A" lower right (X=cut, A=Artist), while figure 29 is signed "GM" lower left, and figures 35 and 36 are signed "ES" lower left. Figure 2 bears a monograph "FR", lower left, and figure 7, of the penguin Aptenodytes patagonica is stated to be "from Brehm" ( Brehms Tierleben ).
Animal Coloration has a simple structure of six chapters in its 288 pages.
Comparative Rarity of Green tree-frequenting Animals an Argument in favour of Selection
It is not a little surprising to find how few green animals there are ... it is precisely because the sandy colour of desert animals and the transparency of pelagic organisms is so universal, that some general environmental cause appears to be necessary for the explanation of the facts; on the other hand, the picking and choosing among arboreal animals savours distinctly of natural selection. [P 6]
The American zoologist and ornithologist Joel Asaph Allen reviewed Animal Coloration in The Auk in 1893. [3] Allen notes Beddard's remark that the book contains hardly anything novel, so that it is mainly a review of previous theories, but welcomes it as a review of the state of knowledge together with Beddard's critical commentary. Allen notes that Beddard could have gone further in criticising Weismann and Poulton on colour changes, but is "glad to see [that Beddard] is willing to grant that the influence of an animal's surroundings may exercise a direct influence upon its coloration without the intervention of the agency of 'natural selection.'" [3]
Allen praises Beddard's "commendable conservatism" in his discussion of camouflage, which he compares to the "credulous spirit" of other authors. Reviewing the chapter on warning coloration, Allen remarks that the great horned owl is known to prey on the skunk, showing that even such a disagreeably pungent animal can be subject to predation. [3]
On mimicry, Allen is critical of Bates's theory, arguing that edible mimics (such as flies) are often not protected by resembling distasteful models (such as wasps). Allen notes that Beddard deals with many special cases "as of .. spiders mimicking ants, etc." and finds the arguments against any selective advantage from Batesian mimicry, and so against natural selection, somewhat conclusive. Finally, reviewing the chapter on sexual selection, Allen writes (knowing that Wallace largely rejected sexual selection) [3]
Mr. Beddard appears to wholly reject Mr. Darwin's much admired theory of 'Sexual Selection,' and quotes at length Mr. Wallace's reductio ad absurdum, which, as illustrating the view of an ultra natural selectionist, may well be here transcribed... [3]
Allen then makes some remarks, praising Beddard for the "fine vein of irony" that he uses of
alleged instances of sexual selection, as in respect to the mating and 'love dances' of spiders. But on the whole his criticisms are suggestive rather than aggressive." and concludes with "we are glad to welcome so healthy an antidote to this mild phase of scientific lunacy as Mr. Beddard's book on 'Animal Coloration.' [3]
The zoologist Edward Bagnall Poulton, whose work is referred to throughout Beddard's book, reviewed Animal Coloration in Nature in 1892. Poulton is critical of Beddard and other authors, defending Darwin's theory of natural selection as "the most generally accepted explanation of organic evolution" and insisting that in "case after case" the Darwinian explanation turns out to be correct. [4]
The white supremacist scientist Robert Wilson Shufeldt reviewed Animal Coloration in Science in 1892, praising it as a concise and useful summary of the subject. He admires Macmillan Publishers' handling of the book with its attractive wood-cuts and coloured lithographic plates. He is pleased to find many Americans in the index. He quotes Beddard's distinction between colour and coloration. He considers that the book brings readers fully up to date and even adds a few new ideas. He recommends the book to all working American naturalists. [5]
The anonymous reviewer in Popular Science Monthly in December 1892 writes that Beddard has "made a book interesting to both the zoologist and the general reader." [6] On protective coloration, "he raises the question whether as a matter of fact animals are concealed from their foes by their protective resemblances, and shows that there is much evidence on the negative side", and further that such colours are sometimes produced "more simply and directly than by the operation of natural selection." On warning colours, the reviewer notes that Beddard gives "much weight" to Eisig's theory that "the usual bright pigments" in caterpillars (accidentally) cause inedibility, "instead of being produced to advertise it" and that Beddard cautions against assuming that "the sight or taste of animals were the same as that of man". [6]
Beddard's Animal Coloration is cited and discussed both by historians of science, and by practising scientists from a number of different fields. For example, the book illuminates the progress of Darwinism, camouflage research, sexual selection, mimicry and the debate on the purpose of animal coloration triggered by Abbott Thayer. These areas are described in turn below.
The historian Robinson M. Yost explains that Darwinism went into eclipse during the 1890s. At that time, most zoologists felt that natural selection could not be the main cause of biological adaptation, and sought alternative explanations. As a result, many zoologists rejected both Batesian mimicry and Müllerian mimicry. [7] Beddard, writes Yost, explained some problems in the theory of mimicry including that, given how many insect species there are, resemblances between species could arise by chance, and that mimicry was sometimes either useless or actually harmful. In Yost's view, Beddard wanted more evidence that natural selection really was responsible. Yost cites the staunch Darwinist Poulton's hostile review of 1892, which asserts the pre-eminence of Darwin's theory. [4] But, writes Yost, Beddard was not alone in being wary of natural selection. [7]
The zoologist Martin Stevens and colleagues, in 2006, write that "almost all early discussions of camouflage were of the background-matching type", citing Wallace, Poulton, and Beddard, "until the pioneering work of Thayer (1909) and Cott (1940)", which added disruptive coloration. [8] Cott however both makes use of Beddard as an authority (for the fact that the Hudson's Bay lemming turns white in winter whereas the Scandinavian lemming does not, [9] and for his experiments on the effectiveness of prey coloration on predators [10] ) and is critical of him for the "extreme and illogical" opinion held by Beddard and other authors that keeping perfectly still is vital to camouflage. [11] Cott pointed out on that subject that a cryptic colour scheme makes an animal harder to track and to recognize, even while it is moving. [11]
The ornithologist Geoffrey Edward Hill, writing in 2002, notes that both Poulton and Beddard discuss sexual selection, and both agreed that "sexual selection by female choice is a likely explanation for the bright coloration of at least some species of birds". In contrast, Hill observes, Cott's detailed 1940 book does not mention it at all; like other zoologists including Wallace and Huxley, Cott preferred explanations "firmly rooted in natural selection". [12]
The American evolutionary zoologists Jane Van Zandt Brower and Lincoln Pierson Brower followed up the experiments described in the book (pp. 153–159). Beddard, they write, observed the results of feeding the drone fly Eristalis tenax , a harmless but intimidating Batesian mimic of honeybees, to various predators. A chameleon, a green lizard, and a sand skink eagerly consumed the flies, whereas a thrush and a great spotted woodpecker did not. However, they — like Cott before them, they note — were unable to replicate Beddard's claim that toads would eat insects of any kind, including stinging bees and wasps. They describe their own experimental investigations of bees and their drone fly mimics, like Beddard using toads as the predators, concluding that the Batesian mimicry of the honeybee by the drone fly was "highly effective". [13]
The historian of science Sharon Kingsland, in a 1978 paper on Abbott Thayer and the protective coloration debate, uses Beddard repeatedly to illuminate the different strands of the argument. She quotes Beddard (p. 94) on how difficult the question of animal coloration seemed in the 1890s. Thayer — an artist, not a scientist — had dived head-first into the debate. One of the protagonists, notes Kingsland, was Allen, who had reviewed Beddard's book, and who believed that the environment directly influenced animal coloration — Kingsland cites Beddard p. 54 here —, so natural selection seemed to him an unlikely factor, and he pointed out that blending inheritance would dilute the effect of selection. Furthermore, argues Kingsland, again citing Beddard (p. 148), another major protagonist, Alfred Russel Wallace, was emphasizing the problem of conspicuous markings, which could be selected for as warning coloration. [14]
Wallace went so far as to argue, notes Kingsland, that bright colours in sexual dimorphism "resulted from a surplus of vital energy", citing Beddard p. 263 ff. Thayer, on the other hand, had exactly one explanation for everything: natural selection for protective coloration, in particular camouflage by countershading, which radically departed from earlier explanations such as Allen's environmental influences (colours might be affected by light) or Beddard's suggestion that dolphins might have dark backs and light bellies as camouflage when seen from above and from below (Kingsland cites Beddard, p. 115). [14]
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the battledress of a modern soldier, and the leaf-mimic katydid's wings. A third approach, motion dazzle, confuses the observer with a conspicuous pattern, making the object visible but momentarily harder to locate, as well as making general aiming easier. The majority of camouflage methods aim for crypsis, often through a general resemblance to the background, high contrast disruptive coloration, eliminating shadow, and countershading. In the open ocean, where there is no background, the principal methods of camouflage are transparency, silvering, and countershading, while the ability to produce light is among other things used for counter-illumination on the undersides of cephalopods such as squid. Some animals, such as chameleons and octopuses, are capable of actively changing their skin pattern and colours, whether for camouflage or for signalling. It is possible that some plants use camouflage to evade being eaten by herbivores.
Henry Walter Bates was an English naturalist and explorer who gave the first scientific account of mimicry in animals. He was most famous for his expedition to the rainforests of the Amazon with Alfred Russel Wallace, starting in 1848. Wallace returned in 1852, but lost his collection on the return voyage when his ship caught fire. When Bates arrived home in 1859 after a full eleven years, he had sent back over 14,712 species of which 8,000 were new to science. Bates wrote up his findings in his best-known work, The Naturalist on the River Amazons.
Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on butterflies in the rainforests of Brazil.
Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration, sounds, odours, or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.
Sir Edward Bagnall Poulton, FRS HFRSE FLS was a British evolutionary biologist, a lifelong advocate of natural selection through a period in which many scientists such as Reginald Punnett doubted its importance. He invented the term sympatric for evolution of species in the same place, and in his book The Colours of Animals (1890) was the first to recognise frequency-dependent selection. Poulton is also remembered for his pioneering work on animal coloration. He is credited with inventing the term aposematism for warning coloration, as well as for his experiments on 'protective coloration' (camouflage). Poulton became Hope Professor of Zoology at the University of Oxford in 1893.
Darwinism: An Exposition of the Theory of Natural Selection with Some of Its Applications is an 1889 book on evolution by Alfred Russel Wallace, the co-discoverer of evolution by natural selection together with Charles Darwin. This was a book Wallace wrote as a defensive response to the scientific critics of natural selection. Of all Wallace's books, it is cited by scholarly publications the most.
Hymenopus coronatus is a mantis from the tropical forests of Southeast Asia. It is known by various common names, including walking flower mantis, orchid-blossom mantid and (pink) orchid mantis. It is one of several species known as flower mantids, a reference to their unique physical form and behaviour, which often involves moving with a “swaying” motion, as if being “blown” in the breeze. Several species have evolved to mimic orchid flowers as a hunting and camouflaging strategy, “hiding” themselves in plain view and preying upon pollinating insects that visit the blooms. They are known to grab their prey with blinding speed.
Hugh Bamford Cott was a British zoologist, an authority on both natural and military camouflage, and a scientific illustrator and photographer. Many of his field studies took place in Africa, where he was especially interested in the Nile crocodile, the evolution of pattern and colour in animals. During the Second World War, Cott worked as a camouflage expert for the British Army and helped to influence War Office policy on camouflage. His book Adaptive Coloration in Animals (1940), popular among serving soldiers, was the major textbook on camouflage in zoology of the twentieth century. After the war, he became a Fellow of Selwyn College, Cambridge. As a Fellow of the Zoological Society of London, he undertook expeditions to Africa and the Amazon to collect specimens, mainly reptiles and amphibians.
Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, and insects, both in predators and in prey.
Decorator crabs are crabs of several different species, belonging to the superfamily Majoidea, that use materials from their environment to hide from, or ward off, predators. They decorate themselves by sticking mostly sedentary animals and plants to their bodies as camouflage, or if the attached organisms are noxious, to ward off predators through aposematism.
Animal colouration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male has strong patterns, conspicuous colours and is iridescent, while the female is far less visible.
Disruptive coloration is a form of camouflage that works by breaking up the outlines of an animal, soldier or military vehicle with a strongly contrasting pattern. It is often combined with other methods of crypsis including background colour matching and countershading; special cases are coincident disruptive coloration and the disruptive eye mask seen in some fishes, amphibians, and reptiles. It appears paradoxical as a way of not being seen, since disruption of outlines depends on high contrast, so the patches of colour are themselves conspicuous.
Adaptive Coloration in Animals is a 500-page textbook about camouflage, warning coloration and mimicry by the Cambridge zoologist Hugh Cott, first published during the Second World War in 1940; the book sold widely and made him famous.
The Colours of Animals is a zoology book written in 1890 by Sir Edward Bagnall Poulton (1856–1943). It was the first substantial textbook to argue the case for Darwinian selection applying to all aspects of animal coloration. The book also pioneered the concept of frequency-dependent selection and introduced the term "aposematism".
Concealing-Coloration in the Animal Kingdom: An Exposition of the Laws of Disguise Through Color and Pattern; Being a Summary of Abbott H. Thayer’s Discoveries is a book published ostensibly by Gerald H. Thayer in 1909, and revised in 1918, but in fact a collaboration with and completion of his father Abbott Handerson Thayer's major work.
Snow camouflage is the use of a coloration or pattern for effective camouflage in winter, often combined with a different summer camouflage. Summer patterns are typically disruptively patterned combinations of shades of browns and greys, up to black, while winter patterns are dominated by white to match snowy landscapes.
Animal coloration provided important early evidence for evolution by natural selection, at a time when little direct evidence was available. Three major functions of coloration were discovered in the second half of the 19th century, and subsequently used as evidence of selection: camouflage ; mimicry, both Batesian and Müllerian; and aposematism.
Self-decoration camouflage is a method of camouflage in which animals or soldiers select materials, sometimes living, from the environment and attach these to themselves for concealment.
Dazzled and Deceived: Mimicry and Camouflage is a 2009 book on camouflage and mimicry, in nature and military usage, by the science writer and journalist Peter Forbes. It covers the history of these topics from the 19th century onwards, describing the discoveries of Henry Walter Bates, Alfred Russel Wallace and Fritz Müller, especially their studies of butterflies in the Amazon. The narrative also covers 20th-century military camouflage, begun by the painter Abbot Thayer who advocated disruptive coloration and countershading and continued in the First World War by the zoologist John Graham Kerr and the marine artist Norman Wilkinson, who developed dazzle camouflage. In the Second World War, the leading expert was Hugh Cott, who advised the British army on camouflage in the Western Desert.