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Müllerian and Batesian mimicry rings of white-variegated aposematic spiny and thorny plants: A hypothesis

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image of Israel Journal of Plant Sciences

Twenty-one wild spiny or thorny plant species growing in Israel have been found so far that are conspicuous because of white stripes and spots found on their leaves. Twenty of these species occupy open habitats, and only one is a climber (Smilax aspera) that is found in both shady and open habitats. I propose that these spiny, thorny, or prickly conspicuous plants form a defensive Müllerian mimicry ring. The genus Launaea (Asteraceae) includes several species that are both white variegated and spiny or thorny (a defended Müllerian mimicry ring), and four non-thorny but variegated plants (a Batesian mimicry ring). I propose that these four species that form a non-defended Batesian mimicry ring enjoy the indirect protection of both their co-generic spiny and thorny species and also of defended plants from other taxa. The long history of the considerable impact of grazing in this arid region seems to have selected for this character.

Affiliations: 1: Department of Science Education—Biology, Faculty of Science and Science Education, University of Haifa-Oranim


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1. Allen, J. A., Knill, R. 1991. Do grazers leave mottled leaves in the shade? Trends Ecol. Evol. 6: 109-110.
2. Archetti, M. 2000. The origin of autumn colours by coevolution. J. Theor. Biol. 205: 625-630.
3. Archetti, M. 2009. Classification of hypotheses for the evolution of autumn colours. Oikos 118: 328-333.
4. Archetti, M., Döring, T. F., Hagen, S. B., Hughes, N. M., Leather, S. R., Lee, D. W., Lev-Yadun, S., Manetas, Y., Ougham, H. J., Schaberg, P. G., Thomas, H. 2009. Unravelling the evolution of autumn colours: an interdisciplinary approach. Trends Ecol. Evol. 24: 166-173.
5. Atsatt, P. R., O'Dowd, D. J. 1976. Plant defense guilds. Science 193: 24-29.
6. Brady, J., Shereni, W. 1988. Landing responses of the tsetse fly Glossina morsitans morsitans Westwood and the stable fly Stomoxys calcitrans (L.) (Diptera: Glossinidae & Muscidae) to black-and-white patterns: a laboratory study. Bull. Entomol. Res. 78: 301-311.
7. Bremer, K. 1994. Asteraceae. Cladistics & classification. Timber Press, Portland, OR.
8. Cahn, M. G., Harper, J. L. 1976. The biology of the leaf mark polymorphism in Trifolium repens L. 2. Evidence for the selection of leaf marks by rumen fistulated sheep. Heredity 37: 327-333.
9. Campitelli, B. E., Steglik, I., Stinchcombe, J. R. 2008. Leaf variegation is associated with reduced herbivore damage in Hydrophyllum virginianum.Botany 86: 306-313.
10. Cook, A. D., Atsatt, P. R., Simon, C. A. 1971. Doves and dove weed: multiple defenses against avian predation. BioScience 21: 277-281.
11. Cott, H. B. 1940. Adaptive coloration in animals. Methuen, London.
12. Danin, A. 2004. Distribution atlas of plants in the Flora Palaestina area. The Israel Academy of Sciences and Humanities, Jerusalem.
13. Doku, C., Brady, J. 1989. Landing site preferences of Glossina morsitans morsitans Westwood (Diptera: Glossinidae) in the laboratory: avoidance of horizontal features? Bull. Entomol. Res. 79: 521-528.
14. Eisner, T. 1964. Catnip: its raison d'ětre. Science 146: 1318-1320.
15. Eisner, T., Grant, R. P. 1981. Toxicity, odor aversion, and "olfactory aposematism". Science 213: 476.
16. Endler, J. A., Mappes, J. 2004. Predator mixes and the conspicuousness of aposematic signals. Am. Nat. 163: 532-547.
17. Feinbrun-Dothan, N. 1978. Flora Palaestina. Vol. III. The Israel Academy of Sciences and Humanities, Jerusalem.
18. Feinbrun-Dothan, N. 1986. Flora Palaestina. Vol. IV. The Israel Academy of Sciences and Humanities, Jerusalem.
19. Feinbrun-Dothan, N., Danin, A. 1991. Analytical flora of Israel. Cana Publishing House Ltd., Jerusalem (in Hebrew).
20. Gamberale-Stille, G., Guilford, T. 2003. Contrast versus colour in aposematic signals. Anim. Behav. 65: 1021-1026.
21. Gibson, G. 1992. Do tsetse flies ‘see’ zebras? A field study of the visual response of tsetse to striped targets. Physiol. Entomol. 17: 141-147.
22. Gittleman, J. L., Harvey, P. H. 1980. Why are distasteful prey not cryptic? Nature 286: 149-150.
23. Givnish, T. J. 1990. Leaf mottling: relation to growth form and leaf phenology and possible role as camouflage. Funct. Ecol. 4: 463-474.
24. Guilford, T., Nicol, C., Rothschild, M., Moore, B. P. 1987. The biological roles of pyrazines: evidence for a warning odour function. Biol. J. Linn. Soc. 31: 113-128.
25. Halpern, M., Raats, D., Lev-Yadun, S. 2007a. Plant biological warfare: thorns inject pathogenic bacteria into herbivores. Environ. Microbiol. 9: 584-592.
26. Halpern, M., Raats, D., Lev-Yadun, S. 2007b. The potential anti-herbivory role of microorganisms on plant thorns. Plant Signal. Behav. 2: 503-504.
27. Hamilton, W. D., Brown, S. P. 2001. Autumn tree colours as a handicap signal. Proc. R. Soc. London B 268: 1489-1493.
28. Hara, N. 1957. Study of the variegated leaves, with special reference to those caused by air spaces. Jpn. J. Bot. 16: 86-101.
29. Harborne, J. B. 1982. Introduction to ecological biochemistry. Academic Press, London.
30. Harper, J. L. 1977. Population biology of plants. Academic Press, London.
31. Harvey, P. H., Paxton, R. J. 1981. The evolution of aposematic coloration. Oikos 37: 391-396.
32. Hill, M. E. 2006. The effect of aposematic coloration on the food preference of Aphelocoma coerulescens, the Florida scrub jay. Bios 77: 97-106.
33. Hinton, H. E. 1973. Natural deception. In: Gregory, R. L., Gombrich, E. H., eds. Illusion in nature and art. Duckworth, London, pp. 97-159.
34. Inbar, M., Lev-Yadun, S. 2005. Conspicuous and aposematic spines in the animal kingdom. Naturwissenschaften 92: 170-172.
35. Kaye, H., Mackintosh, N. J., Rothschild, M., Moore, B. P. 1989. Odour of pyrazine potentiates an association between environmental cues and unpalatable taste. Anim. Behav. 37: 563-568.
36. Knight, R. S., Siegfried, W. R. 1983. Inter-relationships between type, size and color of fruits and dispersal in southern African trees. Oecologia 56: 405-412.
37. Komárek, S. 1998. Mimicry, aposematism and related phenomena in animals and plants: bibliography 1800-1990. Vesmir, Prague.
38. Konoplyova, A., Petropoulou, Y., Yiotis, C., Psaras, G. K., Manetas, Y. 2008. The fine structure and photosynthetic cost of structural leaf variegation. Flora 203: 653-662.
39. Launchbaugh, K. L., Provenza, F. D. 1993. Can plants practice mimicry to avoid grazing by mammalian herbivores? Oikos 66: 501-504.
40. Lee, D. 2007. Nature's palette. The science of plant color. University of Chicago Press, Chicago.
41. Lee, D. W., Brammeler, S., Smith, A. P. 1987. The selective advantages of anthocyanins in developing leaves of mango and cacao. Biotropica 19: 40-49.
42. Levin, D. A. 1973. The role of trichomes in plant defense. Q. Rev. Biol. 48: 3-15.
43. Lev-Yadun, S. 2001. Aposematic (warning) coloration associated with thorns in higher plants. J. Theor. Biol. 210: 385-388.
44. Lev-Yadun, S. 2003a. Why do some thorny plants resemble green zebras? J. Theor. Biol. 244: 483-489.
45. Lev-Yadun, S. 2003b. Weapon (thorn) automimicry and mimicry of aposematic colorful thorns in plants. J. Theor. Biol. 244: 183-188.
46. Lev-Yadun, S. 2006a. Defensive coloration in plants: a review of current ideas about anti-herbivore coloration strategies. In: Teixeira da Silva, J. A., ed. Floriculture, ornamental and plant biotechnology: advances and topical issues. Vol. IV. Global Science Books, London, pp. 292-299.
47. Lev-Yadun, S. 2006b. Defensive functions of white coloration in coastal and dune plants. Isr. J. Plant Sci. 54: 317-325.
48. Lev-Yadun, S. 2009. Aposematic (warning) coloration in plants. In: Baluska, F., ed. Plant-environment interactions. From sensory plant biology to active behavior. Vol. II. Springer Verlag, Berlin, pp. 167-202.
49. Lev-Yadun, S., Gould, K. S. 2007. What do red and yellow autumn leaves signal? Bot. Rev. 73: 279-289.
50. Lev-Yadun, S., Gould, K. S. 2008. Role of anthocyanins in plant defense. In: Gould, K. S., Davies, K. M., Winefield, C., eds. Life's colorful solutions: the biosynthesis, functions, and applications of anthocyanins. Springer-Verlag, Berlin, pp. 21-48.
51. Lev-Yadun, S., Halpern, M. 2008. External and internal spines in plants insert pathogenic microorganisms into herbivore's tissues for defense. In: Van Dijk, T., ed. Microbial ecology research trends. Nova Scientific Publishers, New York, pp. 155-168.
52. Lev-Yadun, S., Inbar, M. 2002. Defensive ant, aphid and caterpillar mimicry in plants. Biol. J. Linn. Soc. 77: 393-398.
53. Lev-Yadun, S., Ne'eman, G. 2004. When may green plants be aposematic? Biol. J. Linn. Soc. 81: 413-416.
54. Lev-Yadun, S., Ne'eman, G. 2006. Color changes in old aposematic thorns, spines, and prickles. Isr. J. Plant Sci. 54: 327-333.
55. Lindström, L., Alatalo, R. V., Mappes, J., Riipi, M., Vertainen, L. 1999. Can aposematic signals evolve by gradual change? Nature 397: 249-251.
56. Livneh, M., Heller, D., eds. 1982. Flowering plants A. In: Alon, A., ed. Plants and animals of the land of Israel. Vol. 10. Ministry of Defence/Publishing House, and Society for Protection of Nature, Israel, Tel Aviv (in Hebrew).
57. Livneh, M., Heller, D., eds. 1983. Flowering plants B. In: Alon, A., ed. Plants and animals of the land of Israel. Vol. 11. Ministry of Defence/Publishing House, and Society for Protection of Nature, Israel, Tel Aviv (in Hebrew).
58. Mallet, J., Gilbert, L. E. 1995. Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies. Biol. J. Linn. Soc. 55: 159-180.
59. Mallet, J., Joron, M. 1999. Evolution of diversity in warning color and mimicry: polymorphisms, shifting balance, and speciation. Annu. Rev. Ecol. Syst. 30: 201-233.
60. Marples, N. M., Kelly, D. J., Thomas, R. J. 2005. The evolution of warning coloration is not paradoxical. Evolution 59: 933-940.
61. Massei, G., Cotterill, J. V., Coats, J. C., Bryning, G., Cowan, D. P. 2007. Can Batesian mimicry help plants to deter herbivory? Pest Manag. Sci. 63: 559-563.
62. Midgley, J. J. 2004. Why are spines of African Acacia species white? Afr. J. Range For. Sci. 21: 211-212.
63. Midgley, J. J., Botha, M. A., Balfour, D. 2001. Patterns of thorn length, density, type and colour in African Acacias. Afr. J. Range Forage Sci. 18: 59-61.
64. Moore, B. P., Brown, W. V., Rothschild, M. 1990. Methylalkylpyrazines in aposematic insects, their hostplants and mimics. Chemoecology 1: 43-51.
65. Niemelä, P., Tuomi, J., Siren, S. 1984. Selective herbivory on mosaic leaves of variegated Acer pseudoplatanus.Experientia 40: 1433-1434.
66. Plitmann, U., Heyn, C., Danin, A., Shmida, A. 1983. Pictorial flora of Israel. Massada Ltd., Tel Aviv (in Hebrew).
67. Provenza, F. D., Kimball, B. A., Villalba, J. J. 2000. Roles of odor, taste, and toxicity in the food preferences of lambs: implications for mimicry in plants. Oikos 88: 424-432.
68. Ronel, M., Khateeb, S., Lev-Yadun, S. 2009. Protective spiny modules in thistles of the Asteraceae in Israel. J. Torrey Bot. Soc. 136: 46-56.
69. Rothschild, M. 1972. Some observations on the relationship between plants, toxic insects and birds. In: Harborne, J. B., ed. Phytochemical ecology. Academic Press, London, pp. 1-12.
70. Rothschild, M. 1973. Secondary plant substances and warning colouration in insects. In: van Emden, H. F., ed. Insect/plant relationships. Symposia of the Royal Entomological Society of London, Number six. Blackwell Scientific Publications, Oxford, pp. 59-83.
71. Rothschild, M. 1980. Remarks on carotenoids in the evolution of signals. In: Gilbert, L. E., Raven P. H., eds. Coevolution of animals and plants. University of Texas Press, Austin, pp. 20-51.
72. Rothschild, M. 1986. The red smell of danger. New Sci. 111 (September 4): 34-36.
73. Rothschild, M., Moore, B. 1987. Pyrazines as alerting signals in toxic plants and insects. In: Labeyrie, V., Fabres, G., Lachaise, D., eds. Insects—plants. Dr W. Junk Publishers, Dordrecht, pp. 97-101.
74. Rothschild, M., Moore, B. P., Brown, W. V. 1984. Pyrazines as warning odour components in the Monarch butterfly, Danaus plexippus, and in moths of the genera Zygaena and Amata (Lepidoptera). Biol. J. Linn. Soc. 23: 375-380.
75. Rowland, H. M., Ihalainen, E., Lindström, L., Mappes, J., Speed, M. 2007. Co-mimics have a mutualistic relationship despite unequal defences. Nature 448: 64-67.
76. Rubino, D. L., McCarthy, B. C. 2004. Presence of aposematic (warning) coloration in vascular plants of southeastern Ohio. J. Torrey Bot. Soc. 131: 252-256.
77. Ruxton, G. D. 2002. The possible fitness benefits of striped coat coloration for zebra. Mammal Rev. 32: 237-244.
78. Ruxton, G. D., Sherratt, T. N., Speed, M. P. 2004. Avoiding attack. The evolutionary ecology of crypsis, warning signals & mimicry. Oxford University Press, Oxford.
79. Scarchuk, J., Lent, J. M. 1965. The structure of mottled-leaf summer squash. J. Hered. 56: 167-168.
80. Schaefer, H. M., Schaefer, V. 2007. The evolution of visual fruit signals: Concepts and constraints. In: Dennis, A. J., Schupp, E. W., Green, R., Wescott, D. W., eds. Seed dispersal: theory and its application in a changing world. CAB International, Wallingford, UK, pp. 59-77.
81. Servedio, M. R. 2000. The effect of predator learning, forgetting, and recognition errors on the evolution of warning coloration. Evolution 54: 751-763.
82. Sherratt, T. N. 2007. Mimicry on the edge. Nature 448: 34-36.
83. Shifriss, O. 1981. Do Cucurbita plants with silvery leaves escape virus infection? Cucurbit Genetics Cooperative, Report 4: 42-43.
84. Shmida, A., Darom, D. 1985. Handbook of wildflowers of Israel. Mediterranean flora. Keter Publishing House Ltd., Jerusalem (in Hebrew).
85. Shmida, A., Darom, D. 1986. Handbook of wildflowers of Israel. Desert flora. Keter Publishing House Ltd., Jerusalem (in Hebrew).
86. Shmida, A., Darom, D. 1992. Handbook of trees and bushes of Israel. Keter Publishing House Ltd., Jerusalem (in Hebrew).
87. Smith, A. P. 1986. Ecology of leaf color polymorphism in a tropical forest species: habitat segregation and herbivory. Oecologia 69: 283-287.
88. Soltau, U., Dötterl, S., Liede-Schumann, S. 2009. Leaf variegation in Caladium steudneriifolium (Araceae): a case of mimicry? Evol. Ecol. 23: 503-512.
89. Speed, M. P., Ruxton, G. D. 2005. Warning displays in spiny animals: one (more) evolutionary route to aposematism. Evolution 59: 2499-2508.
90. Tsukaya, H., Okada, H., Mohamed, M. 2004. A novel feature of structural variegation in leaves of the tropical plant Schismatoglottis calyptrata.J. Plant Res. 117: 477-480.
91. Tuomi, J., Augner, M. 1993. Synergistic selection of unpalatability in plants. Evolution 47: 668-672.
92. Waage, J. K. 1981. How the zebra got its stripes—biting flies as selective agents in the evolution of zebra coloration. J. Entomol. Soc. S. Afr. 44: 351-358.
93. Wiens, D. 1978. Mimicry in plants. Evol. Biol. 11: 365-403.
94. Wiklund, C., Järvi, T. 1982. Survival of distasteful insects after being attacked by naive birds: a reappraisal of the theory of aposematic coloration evolving through individual selection. Evolution 36: 998-1002.
95. Woolfson, A., Rothschild, M. 1990. Speculating about pyrazines. Proc. R. Soc. London B 242: 113-119.
96. Zohary, M. 1966. Flora Palaestina. Vol. I. The Israel Academy of Sciences and Humanities, Jerusalem.
97. Zohary, M. 1972. Flora Palaestina. Vol. II. The Israel Academy of Sciences and Humanities, Jerusalem.

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