Abstract
Appearance of Old world vipers of genus Vipera serves various purposes including crypsis and aposematism. Recent research showed that the zigzag pattern represents strong signal to predators to avoid vipers as a prey. It is also possible that vipers function within ecological community as Batesian model for numerous mimics, including other reptiles, birds, and invertebrates. It is then showed that Batesian models need to have prominent features to sustain the mimicry system. The main modulation of this system is presented here as iconicity. Iconicity is treated as quantitative variable resulting from open dynamic process with multiple inputs, also including iconicity of previous states of system. Batesian mimicry is based on mimics adopting the iconicity of the model. It is an example of ecological facilitation, and as such, it is part of niche construction. Since Batesian mimicry is based on semiotic processes, it is a special case of ecological facilitation, namely semiotic facilitation.
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References
Altieri, A. H., van Wesenbeeck, B. K., Bertness, M. D., & Silliman, B. R. (2010). Facilitation cascade drives positive relationship between native biodiversity and invasion success. Ecology, 91, 1269–1275.
Andrén, C., & Nilson, G. (1981). Reproductive success and risk of predation in normal and melanistic colour morphs of the adder, Vipera berus. Biological Journal of the Linnean Society, 15, 235–246.
Aubret, F., & Mangin, A. (2014). The snake hiss: Potential acoustic mimicry in a viper–colubrid complex. Biological Journal of the Linnean Society, 113, 1107–1114.
Barnett, J. B., & Cuthill, I. C. (2014). Distance-dependent defensive coloration. Current Biology, 24, R1157–R1158.
Bates, H. W. (1862). XXXII. Contributions to an insect Fauna of the Amazon Valley. Lepidoptera: Heliconidæ. Transactions of the Linnean Society of London, 23, 495–566.
Bertness, M. D., & Callaway, R. (1994). Positive interactions in communities. Trends in Ecology & Evolution, 9, 191–193.
Bond, A. B. (2007). The evolution of color polymorphism: Crypticity, searching images, and apostatic selection. Annual Review of Ecology, Evolution, and Systematics, 38, 489–514.
Brodie, E. D. (1992). Correlational selection for color pattern and antipredator behavior in the garter snake Thamnophis ordinoides. Evolution, 46, 1284–1298.
Brooker, R. W., Maestre, F. T., Callaway, R. M., Lortie, C. L., Cavieres, L. A., Kunstler, G., Liancourt, P., Tielbörger, K., Travis, J. M. J., & Anthelme, F. (2008). Facilitation in plant communities: The past, the present, and the future. Journal of Ecology, 96, 18–34.
Brown, J. L. (1965). Flicker and intermittent stimulation. Vision and Visual Perception, 1, 251–320.
Bruno, J. F., Stachowicz, J. J., & Bertness, M. D. (2003). Inclusion of facilitation into ecological theory. Trends in Ecology & Evolution, 18, 119–125.
Caro, T. (2017). Wallace on coloration: Contemporary perspective and unresolved insights. Trends in Ecology & Evolution, 32, 23–30.
Clarke, B. (1962). Balanced polymorphism and the diversity of sympatric species. In D. Nichols (Ed.), Taxonomy and geography (pp. 47–70). Oxford: Systematics Association.
Cott, H. B. (1940). Adaptive coloration in animals. London: Methuen.
Emeneau, M. B. (1948). Taboos on animal names. Language, 24, 56–63.
Endler, J. A. (1978). A predator’s view of animal color patterns. Evolutionary Biology, 11, 319–364.
Endler, J. A. (1981). An overview of the relationships between mimicry and crypsis. Biological Journal of the Linnean Society, 16, 25–31.
Endler, J. A. (1990). On the measurement and classification of colour in studies of animal colour patterns. Biological Journal of the Linnean Society, 41, 315–352.
Endler, J. A., & Mappes, J. (2004). Predator mixes and the conspicuousness of aposematic signals. American Naturalist, 163, 532–547.
Fernández, E. (2014). The role of semiosis in evolution - from biosemiotics to technosemiotics. Gatherings biosemiotics 14, Middlesex University London, 13 pp.
Forsman, A., & Ås, S. (1987). Maintenance of colour polymorphism in adder, Vipera berus, populations: A test of a popular hypothesis. Oikos, 50, 13–16.
Gans, C. (1961). Mimicry in procryptically colored snakes of the genus Dasypeltis. Evolution, 15, 72–91.
Gans, C., & Maderson, P. F. A. (1973). Sound producing mechanisms in recent reptiles: Review and comment. American Zoologist, 13, 1195–1203.
Greene, H. W., & McDiarmid, R. W. (1981). Coral snake mimicry: Does it occur? Science, 213, 1207–1212.
Greene, H. W., & McDiarmid, R. W. (2005). Wallace and Savage: heroes, theories and venomous snake mimicry. In B. I. Crother, C. Guyer, M. H. Wake, & M. A. Donnelly (Eds.), Ecology and evolution in the tropics: a herpetological perspective (pp. 190–208). Chicago: University of Chicago Press.
Guilford, T. (1988). The evolution of conspicuous coloration. American Naturalist, 131, S7–S21.
Guimaraes, M., & Sawaya, R. J. (2011). Pretending to be venomous: Is a snake’s head shape a trustworthy signal to a predator? Journal of Tropical Ecology, 27, 437–439.
Guimaraes, M., & Sawaya, R. J. (2012). The snake head-shape signal: A reply to Valkonen & Mappes. Journal of Tropical Ecology, 28, 125–126.
Ham, A. D., Ihalainen, E., Lindström, L., & Mappes, J. (2006). Does colour matter? The importance of colour in avoidance learning, memorability and generalisation. Behavioral Ecology and Sociobiology, 60, 482–491.
Hanlon, R. T., Forsythe, J. W., & Joneschild, D. E. (1999). Crypsis, conspicuousness, mimicry and polyphenism as antipredator defences of foraging octopuses on indo-Pacific coral reefs, with a method of quantifying crypsis from video tapes. Biological Journal of the Linnean Society, 66, 1–22.
Hebets, E. A., & Papaj, D. R. (2005). Complex signal function: Developing a framework of testable hypotheses. Behavioral Ecology and Sociobiology, 57, 197–214.
Hossie, T. J., & Sherratt, T. N. (2013). Defensive posture and eyespots deter avian predators from attacking caterpillar models. Animal Behaviour, 86, 383–389.
Hunsinger, E., Root-Gutteridge, H., Cusano, D. A., & Parks, S. E. (2017). A description of defensive hiss types in the flat horned hissing cockroach (Aeluropoda insignis). Bioacoustics, 27, 261–271. https://doi.org/10.1080/09524622.2017.1327371.
Ioannou, C. C., & Krause, J. (2009). Interactions between background matching and motion during visual detection can explain why cryptic animals keep still. Biology Letters, 5, 191–193.
IUCN (2018). The IUCN red list of threatened species. Version 2017-3. <www.iucnredlist.org> downloaded on 31st January 2018.
Jackson, J. F., Ingram, W., III, & Campbell, H. W. (1976). The dorsal pigmentation pattern of snakes as an antipredator strategy: A multivariate approach. American Naturalist, 110, 1029–1053.
Kazemi, B., Gamberale-Stille, G., Tullberg, B. S., & Leimar, O. (2014). Stimulus salience as an explanation for imperfect mimicry. Current Biology, 24, 965–969.
Kikuchi, D. W., Mappes, J., Sherratt, T. N., & Valkonen, J. K. (2016). Selection for multicomponent mimicry: Equal feature salience and variation in preferred traits. Behavioral Ecology, 27, 1515–1521.
Kikvidze, Z., & Callaway, R. M. (2009). Ecological facilitation may drive major evolutionary transitions. Bioscience, 59, 399–404.
Kirchner, W. H., & Röschard, J. (1999). Hissing in bumblebees: An interspecific defence signal. Insectes Sociaux, 46, 239–243.
Kleisner, K. (2010). Re-semblance and re-evolution: Paramorphism and semiotic co-option may explain the re-evolution of similar phenotypes. Sign Systems Studies, 38, 378–392.
Kleisner, K. (2011). Perceive, co-opt, modify, and live! Organism as a Centre of experience. Biosemiotics, 4, 223–241.
Kleisner, K. (2015). Semantic organs: The concept and its theoretical ramifications. Biosemiotics, 8, 367–379.
Kleisner, K., & Markoš, A. (2005). Semetic rings: Towards the new concept of mimetic resemblances. Theory in Biosciences, 123, 209–222.
Komárek, S. (2003). Mimicry, aposematism and related phenomena: Mimetism in nature and the history of its study (167 pp). Lincom Europa: München.
Krams, I., Vrublevska, J., Koosa, K., Krama, T., Mierauskas, P., Rantala, M. J., & Tilgar, V. (2014). Hissing calls improve survival in incubating female great tits (Parus major). Acta Ethologica, 17, 83–88.
Kroon, C. (1975). A possible Müllerian mimetic complex among snakes. Copeia, 1975, 425–428.
Lindell, L. E., & Forsman, A. (1996). Sexual dichromatism in snakes: Support for the flicker-fusion hypothesis. Canadian Journal of Zoology, 74, 2254–2256.
Lindström, L., Alatalo, R. V., & Mappes, J. (1997). Imperfect Batesian mimicry—the effects of the frequency and the distastefulness of the model. Proceedings of the Royal Society B: Biological Sciences, 264, 149–153.
Mallet, J., & Joron, M. (1999). Evolution of diversity in warning color and mimicry: Polymorphisms, shifting balance, and speciation. Annual Review of Ecology and Systematics, 30, 201–233.
Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica, 2007, 223–248.
Maran, T. (2011). Becoming a sign: The mimic’s activity in biological mimicry. Biosemiotics, 4, 243–257.
Maran, T., & Kleisner, K. (2010). Towards an evolutionary biosemiotics: Semiotic selection and semiotic co-option. Biosemiotics, 3, 189–200.
Merriam-Webster (2018). “Adder.” <www.merriam-webster.com/dictionary/adder> Accessed on 31st January 2018.
Moser, A., Graber, C., & Freyvogel, T. A. (1984). Observations sur l’ethologie et l’evolution d’une population de Vipera aspis (L.) au nord du Jura Suisse. Amphibia-Reptilia, 5, 373–393.
Neumeyer, R. (1987). Density and seasonal movements of the adder (Vipera berus L. 1758) in a subalpine environment. Amphibia-Reptilia, 8, 259–275.
Niskanen, M., & Mappes, J. (2005). Significance of the dorsal zigzag pattern of Vipera latastei gaditana against avian predators. Journal of Animal Ecology, 74, 1091–1101.
Odling-Smee, F. J., Laland, K. N., & Feldman, M. W. (1996). Niche construction. American Naturalist, 147, 641–648.
Öhman, A., & Mineka, S. (2003). The malicious serpent: Snakes as a prototypical stimulus for an evolved module of fear. Current Directions in Psychological Science, 12, 5–9.
Pfennig, D. W. (2016). Evolutionary biology: To mimicry and back again. Nature, 534, 184–185.
Pfennig, D. W., & Mullen, S. P. (2010). Mimics without models: causes and consequences of allopatry in Batesian mimicry complexes. Proceedings of the Royal Society B: Biological Sciences, 277, 2577–2585.
Pfennig, D. W., Harcombe, W. R., & Pfennig, K. S. (2001). Frequency-dependent Batesian mimicry. Nature, 410, 323.
Pfennig, D. W., Harper, G. R., Brumo, A. F., Harcombe, W. R., & Pfennig, K. S. (2007). Population differences in predation on Batesian mimics in allopatry with their model: Selection against mimics is strongest when they are common. Behavioral Ecology and Sociobiology, 61, 505–511.
Pough, F. H. (1976). Multiple cryptic effects of crossbanded and ringed patterns of snakes. Copeia, 1976, 834–836.
Pough, F. H. (1988). Mimicry of vertebrates: Are the rules different? American Naturalist, 131, S67–S102.
Rabosky, A. R. D., Cox, C. L., Rabosky, D. L., Holmes, I. A., Feldman, A., & McGuire, J. A. (2016). Coral snakes predict the evolution of mimicry across New World snakes. Nature Communications, 7, 11484.
Ripple, W. J., Estes, J. A., Beschta, R. L., Wilmers, C. C., Ritchie, E. G., Hebblewhite, M., Berger, J., Elmhagen, B., Letnic, M., Nelson, M. P., Schmitz, O. J., Smith, D. W., Wallach, A. D., & Wirsing, A. J. (2014). Status and ecological effects of the world’s largest carnivores. Science, 343, 1241484.
Rix, H., & Kümmel, M. (2001). Lexikon der indogermanischen Verben, Die Wurzeln und ihre Primärstammbildungen (822 pp). Dr. Ludwig Reichert Verlag: Wiesbaden.
Rodríguez-Robles, J. A., & De Jesús-Escobar, J. M. (2000). Molecular systematics of New World gopher, bull, and pinesnakes (Pituophis: Colubridae), a transcontinental species complex. Molecular Phylogenetics and Evolution, 14, 35–50.
Rowe, M. P., Coss, R. G., & Owings, D. H. (1986). Rattlesnake rattles and burrowing owl hisses: A case of acoustic Batesian mimicry. Ethology, 72, 53–71.
Ruxton, G. D., Franks, D. W., Balogh, A. C. V., & Leimar, O. (2008). Evolutionary implications of the form of predator generalization for aposematic signals and mimicry in prey. Evolution, 62, 2913–2921.
Santos, J. C., Baquero, M., Barrio-Amorós, C., Coloma, L. A., & Erdtmann, L. K. (2014). Aposematism increases acoustic diversification and speciation in poison frogs. Proceedings of the Royal Society B: Biological Sciences, 281, 20141761.
Santos, X., Azor, J. S., Cortés, S., Rodríguez, E., Larios, J., & Pleguezuelos, J. M. (2017). Ecological significance of dorsal polymorphism in a Batesian mimic snake. Current Zoology. https://doi.org/10.1093/cz/zox058.
Sherbrooke, W. C., & Westphal, M. F. (2006). Responses of greater roadrunners during attacks on sympatric venomous and nonvenomous snakes. Southwest Naturalist, 51, 41–47.
Sherratt, T. N., & Beatty, C. D. (2003). The evolution of warning signals as reliable indicators of prey defense. American Naturalist, 162, 377–389.
Shiffrar, M., & Lorenceau, J. (1996). Increased motion linking across edges with decreased luminance contrast, edge width and duration. Vision Research, 36, 2061–2067.
Shine, R., & Madsen, T. (1994). Sexual dichromatism in snakes of the genus Vipera: A review and a new evolutionary hypothesis. Journal of Herpetology, 28, 114–117.
Smal-Stocki, R. (1950). Taboos on animal names in Ukrainian. Language, 26, 489–493.
Smith, R. H. (1974). Is the slow worm a Batesian mimic? Nature, 247, 571–572.
Souchet, J., & Aubret, F. (2016). Revisiting the fear of snakes in children: The role of aposematic signalling. Scientific Reports, 6, 37619.
Speybroeck, J., Beukema, W., Bok, B., & Van Der Voort, J. (2016). Field guide to the amphibians and reptiles of Britain and Europe (432 pp). London: Bloomsbury Publishing.
Stachowicz, J. J. (2001). Mutualism, facilitation, and the structure of ecological communities: Positive interactions play a critical, but underappreciated, role in ecological communities by reducing physical or biotic stresses in existing habitats and by creating new habitats on. AIBS Bulletin, 51, 235–246.
Štěrbová, L. (2017). Etymologie vybraných názvů zvířat v bulharštině (59 pp). Brno: Masaryk University.
Stevens, M. (2007). Predator perception and the interrelation between different forms of protective coloration. Proceedings of the Royal Society B: Biological Sciences, 274, 1457–1464.
Sweet, S. S. (1985). Geographic variation, convergent crypsis and mimicry in gopher snakes (Pituophis melanoleucus) and western rattlesnakes (Crotalus viridis). Journal of Herpetology, 19, 55–67.
Titcomb, G. C., Kikuchi, D. W., & Pfennig, D. W. (2014). More than mimicry? Evaluating scope for flicker-fusion as a defensive strategy in coral snake mimics. Current Zoology, 60, 123–130.
Tullberg, B. S., Merilaita, S., & Wiklund, C. (2005). Aposematism and crypsis combined as a result of distance dependence: Functional versatility of the colour pattern in the swallowtail butterfly larva. Proceedings of the Royal Society B: Biological Sciences, 272, 1315–1321.
Valkonen, J. K., & Mappes, J. (2012). Comments on Guimarães & Sawaya. Pretending to be venomous: Is a snake’s head shape a trustworthy signal to a predator? Journal of Tropical Ecology, 28, 123–124.
Valkonen, J. K., & Mappes, J. (2014). Resembling a viper: Implications of mimicry for conservation of the endangered smooth snake. Conservation Biology, 28, 1568–1574.
Valkonen, J., Niskanen, M., Björklund, M., & Mappes, J. (2011a). Disruption or aposematism? Significance of dorsal zigzag pattern of European vipers. Evolutionary Ecology, 25, 1047–1063.
Valkonen, J. K., Nokelainen, O., & Mappes, J. (2011b). Antipredatory function of head shape for vipers and their mimics. PLoS One, 6, e22272.
Vallin, A., Jakobsson, S., Lind, J., & Wiklund, C. (2005). Prey survival by predator intimidation: An experimental study of peacock butterfly defence against blue tits. Proceedings of the Royal Society of London B: Biological Sciences, 272, 1203–1207.
Waldbauer, G. P., & Sternburg, J. G. (1987). Experimental field demonstration that two aposematic butterfly color patterns do not confer protection against birds in northern Michigan. American Midland Naturalist, 118, 145–152.
Wang, M.-Y., Vasas, V., Chittka, L., & Yen, S.-H. (2017). Sheep in wolf’s clothing: Multicomponent traits enhance the success of mimicry in spider-mimicking moths. Animal Behaviour, 127, 219–224.
Werner, Y. L. (1983). Behavioural triangulation of the head in three boigine snakes: Possible cases of mimicry. Israel Journal of Zoology, 32, 205–228.
Werner, Y. L., & Frankenberg, E. (1982). Head triangulation in two colubrine snakes: Probable behavioural reinforcement of Batesian mimicry. Israel Journal of Zoology, 31, 137–150.
Wertheimer, M. (1912). Experimentelle studien uber das sehen von bewegung. Zeitschrift fur Psychologie, 61, 161–265.
Wilmers, C. C., Crabtree, R. L., Smith, D. W., Murphy, K. M., & Getz, W. M. (2003). Trophic facilitation by introduced top predators: Grey wolf subsidies to scavengers in Yellowstone National Park. Journal of Animal Ecology, 72, 909–916.
Wolf, M., & Werner, Y. L. (1994). The striped colour pattern and striped/non-striped polymorphism in snakes (Reptilia: Ophidia). Biological Reviews, 69, 599–610.
Wüster, W., Allum, C. S. E., Bjargardóttir, I. B., Bailey, K. L., Dawson, K. J., Guenioui, J., Lewis, J., McGurk, J., Moore, A. G., Niskanen, M., & Pollard, C. P. (2004). Do aposematism and Batesian mimicry require bright colours? A test, using European viper markings. Proceedings of the Royal Society B: Biological Sciences, 271, 2495–2499.
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Work is part of the research goal VaV10/300/PM DKRVO 2017/05 of Czech Ministry of Culture. I would like to dedicate this work to Kristýna Eliášová.
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Brejcha, J. Viper as a Batesian Model – its Role in an Ecological Community. Biosemiotics 12, 25–38 (2019). https://doi.org/10.1007/s12304-019-09347-x
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DOI: https://doi.org/10.1007/s12304-019-09347-x