Mimicry, Aposematism, and Related Phenomena
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Stanislav Komárek Mimicry, Aposematism, and related Phenomena Mimetism in Nature and the History of its Study Spilogale putorius in the aposematic display Lincom München Contents Introduction .................................................................................................. 7 Defining the scope of interest ....................................................................... 9 Surface, interiority, similarity, and kinship................................................. 10 The time-period up to the year 1800 .......................................................... 17 The time-period from 1800-1860 ............................................................... 23 The time-period between 1859 and 1900 ................................................... 28 Darwinism and sociomorphic modeling .......................................................................................... 37 Sexual dimorphism and selection, exaggerated structures.............................................................. 44 Wallace’s concept of adaptive coloration ....................................................................................... 50 Aposematism .................................................................................................................................. 55 Wallace’s concept of mimicry ........................................................................................................ 59 Müllerian mimicry ........................................................................................................................... 62 Mimicry rings .................................................................................................................................. 64 The time period from 1890 to 1953 (with a note on later r ............................ development trends up to the nineties of the 20th century)..................... 67 Teachings on mimicry and interpretations of the external appearance of organisms in England ... 67 Poulton, a classic of Darwinian interpretation ................................................................................ 70 Hingston and his original conception of animal coloration .............................................................. 81 Poulton’s coworkers and followers................................................................................................. 84 Research and interpretation of mimetic phenomena a and external appearance of organisms on the Continent ................................................................ 89 Eimer and the problem of animal color patterns ............................................................................. 90 Linking color patterns and Portmann’s concept ............................................................................. 99 Piepers, Wasmann, Heikertinger, and other Continental authors .................................................. 106 Mimicry in plants and fungi ...................................................................... 118 Summary ................................................................................................... 126 Glossary ................................................................................................... 128 Bibliography ............................................................................................ 129 List of Illustrations ................................................................................... 152 Name Index .............................................................................................. 155 Subject Index ............................................................................................ 161 About the author ....................................................................................... 167 5 6 Introduction The history of the research of mimetic phenomena in nature (and the interpretation of the external appearance of organisms generally) has not been studied in depth so far, even thou- gh it is an absolutely pivotal theme in biology, as was noted by Mayr (1982), for example. A certain hint of a historical overview of this research area, albeit not very extensive, none- the-less clear and deep, can by found in certain texts by Heikertinger (1921-27, 1954). Re- search of mimicry as exemplary instances of evolutionary theory was pursued by W. C. Kimler (1982) in his dissertation, which was partially published later in year 1983. This work is an example of Anglo-Saxon point of view, which disregardes almost all continental work on the subject and interprets only the most important Anglo-Saxon authors between the era of natu- ral theology, Bates (1862a), and Neo-Darwinian syntheses of the British population’s genetics in the style of reports on the linear progression of science (this method of research is appli- cable only when considering main-stream British and American research, which gives the impression of cumulation of knowledge). All other approaches, for example Hingston (1933), lead in a different direction, not to mention the continental, especially the German school’s approach to mimetic expressions in nature. At the same time Kimler’s work is not aware of the deeper aspects of mimetic expressions in nature, especially the general problem of forms. However large the extent of knowledge attained by Anglo-Saxon authors in the field of research and theoretical explanations of mimetic phenomena in nature, I consider it necessary to study Continental authors as well, and that for two reasons. First of all, Central European biology and the author himself, belong to this school and intellectual tradition, and secondly the postmodern era characteristically accentuates that, which is often considered marginal. The- se marginal themes have a tendency to become central, if only because central themes lost their potential for development due to intensive exploitation. The materials for this study were gathered from 1985 to 1995, first in Vienna and later in Prague, Leiden, and Amsterdam. The study is complemented by a bibliography (Komárek, 1998), and a database of works about mimetic phenomena, which contains around 5000 entries causing it to be the largest piece of work of this kind ever created. While the bibliography only encompasses the years 1800 to 1990 and concentrates only on cases of mimicry, aposematism, and certain important cases of crypsis, the study itself is wider because it encompasses a larger time period and a broader range of themes, especially concerning the history of the interpretation of the exterior appea- rance of living organisms. For that reason only quotations from the most important texts are cited in this book and in cases of need for a wider referential system the citations will point to the above mentioned Bibliography (in the text with a capital ‘B’). Only works concerning skin and wing patterns of animals, which are not included in the Bibliography, are referred to in full. The goal of this book is to point out the centrality of the problem of resemblance and mimicry in nature and the dependence on the understanding of this phenomena in the biolo- gical paradigm of the given era, the intellectual atmosphere, and the language in which the problem is interpreted. This work can also be interpreted to an extent as an „archeological“ study, because the swamp of forgetfulness, into which most written and later duly published works eventually sink can be more easily compared to studying deep layers of an archeological excavation than to the study of recent, easily accessible knowledge. The myth of linearly pro- gressing science, where nothing disappears beyond the horizon, is one of the most flagrant of unconscious modern hypocrisies. Only after concrete work in libraries, directly with historical texts, does the number of particularities and even complete alternative concepts, which have almost entirely disappeared beyond the horizon, become apparent. This research is similar to uncovering material archeologically or paleontologically, material which is difficult to find and also is to an extent fragmentary and which today does not address us directly, but in the best of cases through its „descendants“. Therefore, it is often strongly derivative making it difficult to trace back to its roots. This permanent masking of the past in science has the advantage that research starts with an almost entirely clean slate and doesn’t drag „deadwood“ from the past along with it (this phenomenon is generally acknowledged even outside of scientific histo- ry, where early Christianity discards „heathen prattle“, the Renaissance „scholastic trash“, Communism and Nazism „city-slicker deadwood“, etc.). It is always necessary to prepare a clean stage so the next generation has room to make themselves known, and this is only possible by moving the clutter of theses and particularities of the older era „off stage“, in other words into 7 the province of forgetfulness. This ends not only in their loss in memory but eventually even their true loss (this doom isn’t limited for example to Alexandrian science, but after another „reorganization of libraries“ may even afflict older layers of the modern period). Every social structure, including modern science, actively and even unconsciously camouflages and lays shadows on its own past. The past, incorporated into skinny textbooks or brochures, can only be viewed in this metamorphosed and simplified or even caricatured form. No historical study can compensate for studying original historical texts for the serious