The evolution and frequency of female color morphs in holarctic odonata: Why are male-like females typically the minority?

International Journal of Odonatology

Volumn 8, Issue 2, 2005, Pages 183-212

  Fincke, Ola M ^a   Jödicke, Reinhard ^b   Paulson, Dennis R ^c   Schultz, Thomas D ^d  

^a UNIVERSITY OF OKLAHOMA   (United States)

^b University of Puget Sound ^*  (Germany)

^c UNIVERSITY OF PUGET SOUND   (United States)

^d DENISON UNIVERSITY   (United States)

Author keywords

Ancestral color; Andromorph frequencies; Color polymorphism; Dragonfly; Holarctic; Odonata

Indexed keywords

EID: 34147177461     PISSN: 13887890     EISSN: 21596719     Source Type: Journal    
DOI: 10.1080/13887890.2005.9748252     Document Type: Article

References (134)

1. Acorn, J., 2004. Damselflies of Alberta:flying neon toothpicks in the grass. Winnipeg:The University of Alberta Press.
2. Adamovic, Z. R., and Vijatov, S. T., 1996. Morphometric examination of Calopteryx balcanica Fudakowski, 1930 and C. splendens ancilla Selys, 1853(Zygoptera:Calopterygidae). Odonatologica, 25:109–118.
3. Andrés [Abad], J. A., and Cordero [Rivera], A., 1999. The inheritance of female colour morphs in the damselfly Ceriagrion tenellum (Odonata, Coenagrionidae). Heredity, 82:328–335.
4. Andrés [Abad], J. A, Sánchez-Guillén, R. A., and Cordero Rivera, A., 2002. Evolution of female colour polymorphism in damselflies:testing the hypotheses. Animal Behaviour, 63:677–685.
5. Askew, R. R., 1988. The dragonflies of Europe. Martins:Harley.
6. Belyshev, B. F., 1955. [Unknown forms and stages of dragonflies from Siberia. In Russian]. Zametki po Faune Sibiri, Tomsk State University, 18:27–29.
7. Beutler, H., 1987. Untersuchungen zur Populationsstruktur und-dynamik mitteleuropäischer Libellen (Odonata). Dissertation, Humboldt-Universität Berlin.
8. Bick, G. H., and Bick, J. C., 1965. Color variation and significance of color in reproduction in the damselfly Argia apicalis (Say) (Zygoptera:Coenagrionidae). Canadian Entomologist, 97:32–41.
9. Bilek, A., 1964. Beobachtungen über Odonaten in Südfrankreich mit besonderer Berücksichtigung der Färbungsstadien von Anax parthenope Selys. Nachrichtenblatt Bayerischer Entomologen, 13:59–64.
10. Borror, D. J., 1942. A revision of the libelluline genus Erythrodiplax (Odonata). Columbus:Ohio State University.
11. Brooks, S., and Lewington, R., 1999. Field guide to the dragonflies and damselflies of Great Britain and Ireland, revised edition. Rotherwick:British Wildlife Publishing.
12. Cannings, R. A., 1982. Notes on the biology of Aeshna sitchensis Hagen (Anisoptera:Aeshnidae). Odonatologica, 11:219–223.
13. Carini, A., 1995. Compartamento riproductive e compartamento individuale di Ceriagrion tenellum (de Villers, 1789), con particolare riferimento all competizione sessuale (Insecta, Odonata). MSc Thesis Università die Roma “La Sapienza”.
14. Chippendale, P. T., Varshal, D., Whitmore, D. H., and Robinson, J. V., 1999. Polygenetic relationships of North American damselflies of the genus Ischnura (Odonata:Zygoptera:Coenagrionidae) based on sequences of three mitochondrial genes. Molecular Phylogenetics and Evolution, 11:110–121.
15. Conrad, K. F., and Pritchard, G., 1989. Female dimorphism and physiological colour change in the damselfly Argia vivida Hagen (Odonata:Coenagrionidae). Canadian Journal of Zoology, 67:298–304.
16. Corbet, P. S., 1957. The life-history of the Emperor dragonfly Anax imperator Leach (Odonata:Aeshnidae). Journal of Animal Ecology, 26:1–69.
17. Corbet, P. S., 1999. Dragonflies:behavior and ecology of Odonata. Ithaca:Cornell University Press.
18. Cordero Rivera, A., 1987. Estructura de población en Ischnura graellsi Rambur, 1842(Zygop. Coenagrionidae). Boletín de la Asociación Española de Entomología, 11:269–286.
19. Cordero [Rivera], A., 1990. The inheritance of female polymorphism in the damselfly Ischnura graellsii (Rambur) (Odonata:Coenagrionidae). Heredity:, 64:341–346.
20. Cordero [Rivera], A., 1992. Morphological variability, female polymorphism and heritability of body length in Ischnura graellsii (Rambur) (Zygoptera:Coenagrionidae). Odonatologica:, 21:409–419.
21. Cordero Rivera, A., and Egdigo Peréz, F. J., 1998. Mating frequency, population density and female polychromatism in the damselfly Ischnura graellsii:an analysis of four natural populations. Etología, 6:61–67.
22. Cordero Rivera, A., and Andrés Abad, J. A., 1999. Lifetime mating success, survivorship and synchronized reproduction in the damselfly Ischnura pumilio (Odonata:Coenagrionidae). International Journal of Odonatology, 2:105–114.
23. Córdoba-Aguilar, A., 1993. Population structure in Ischnura denticollis (Burmeister) (Zygoptera:Coenagrionidae). Odonatologica, 22:455–464.
24. d'Aguilar, J., and Dommanget, J.-L., 1998. Guide des libellules d'Europe et d'Afrique du Nord. 2nd edition. Delachaux & Niestlé, Lausanne & Paris.
25. de Marchi, G., 1990. Precopulatory reproductive isolation and wing colour dimorphism in Calopteryx splendens females in southern Italy (Zygoptera:Calopterygidae). Odonatologica, 19:243–250.
26. Dingemanse, N. J., 2000. Kleurpolymorfisme bij vrouwen van de Koraaljuffer (Ceriagrion tenellum). Brachytron, 4:18–24.
27. Duda, R. O., and Hart, P., 1973. Pattern classification and scene analysis. New York:John Wiley.
28. Dumont, H. J., 1977. A review of the dragonfly fauna of Turkey and adjacent Mediterranean islands (Insecta Odonata). Bulletin et Annales de la Société Royal Entomologique de Belgique, 113:119–171.
29. Dumont, H. J., 1991. Fauna Palaestina. Insecta V - Odonata of the Levant Jerusalem:Israel Academy of Sciences and Humanities.
30. Dumont, H. J., Mertens, J., and Miho, A., 1993. A contribution to the knowledge of the Odonata of Albania. Opuscula Zoologica Fluminensia, 113:1–10.
31. Dumont, H. J., Haritonov, A. Yu., Kosterin, O. E., Malikova, E. I., and Popova, O., 2005. A review of the Odonata of Kamchatka Peninsula, Russia. Odonatologica, 34:131–153.
32. Dunkle, S. W., 1989. Dragonflies of Florida, Bermuda and the Bahamas. Gainesville:Scientific Publishers.
33. Dunkle, S. W., 1990. Damselflies of Florida, Bermuda and the Bahamas. Gainesville:Scientific Publishers.
34. Dunkle, S. W., 2000. Dragonflies through binoculars:a field guide to dragonflies of North America. Oxford:Oxford University Press.
35. Endler, J. A., 1992. Signals, signal conditions, and the direction of evolution. American Naturalist, 139:s125–s153.
36. Fincke, O. M., 1982. Lifetime mating success in a natural population of the damselfly, Enallagma hageni (Walsh) (Odonata:Coenagrionidae). Behavioral Ecology and Socio-biology, 10:293–302.
37. Fincke, O. M., 1986. Lifetime reproductive success and the opportunity for selection in a nonterritorial damselfly (Odonata:Coenagrionidae). Evolution, 40:791–803.
38. Fincke, O. M., 1987. Female monogamy in Ischnura verticalis Say (Zygoptera, Coenagrionidae). Odonatologica, 16:129–143.
39. Fincke, O. M., 1994. Female dimorphism in damselflies:failure to reject the null hypothesis. Animal Behaviour, 47:1249–1266.
40. Fincke, O. M., 1997. Conflict resolution in the Odonata:implications for understanding female mating patterns and female choice. Biological Journal of the Linnean Society, 60:201–220.
41. Fincke, O. M., 2004. Polymorphic signals of harassed female odonates and the males that learn them support a novel frequency-dependent model. Animal Behaviour, 67:833–845.
42. Forbes, M. R., 1994. Tests of hypothesis for female-limited polymorphism in Enallagma boreale Selys. Animal Behaviour, 47:724–726.
43. Forbes, M., and Teather, K. L., 1994. Variation in attraction of male Enallagma ebrium (Hagen) to two female morphs (Zygoptera:Coenagrionidae). Odonatologica, 23:103–114.
44. Forbes, M. R., Richardson, J. M.L., and Baker, R. L., 1995. Frequency of female morphs is related to an index of male density in the damselfly, Nehalennia irene (Hagen). Ecoscience, 2:28–33.
45. Fudakowski, J., 1930. Über die Formen von Calopteryx splendens Harr. aus Dalmatien und Herzegovina (Odonata). Annales Musei Zoologici Polonici, 9:57–63. pl. X.
46. Fudakowski, J., 1932. Neue Beiträge zur Odonaten-Fauna Polens. Fragmenta Faunistica Musei Zoologici Polonici, 1:389–401.
47. Garrison, R. W., 1978. A mark-recapture study of imaginal Enallagma cyathigerum (Charpentier) and Argia vivida Hagen (Zygoptera:Coenagrionidae). Odonatologica, 7:223–236.
48. Garrison, R. W., and Hafernik, J. E., 1981. The distribution of Ischnura gemina (Kennedy) and a description of the andromorph female (Zygoptera:Coenagrionidae). Odonatologica, 10:85–91.
49. Gorb, S. N., 1998. Visual cues in mate recognition by males of the damselfly Coenagrion puella (L.) (Odonata:Coenagrionidae). Journal of Insect Behavior, 11:73–92.
50. Gorb, S. N., 1999. Visual cues in mate recognition in the damselfly Ischnura elegans Vander Linden (Zygoptera:Coenagrionidae). International Journal of Odonatology, 2:83–93.
51. Gross, M. R., 1996. Alternative reproductive strategies and tactics:diversity within sexes. Trends in Ecology and Evolution, 11:92–98.
52. Hammond, C. O., and Merritt, R., 1985. The dragonflies of Great Britain and Ireland (second edition). Martins:Harley Books.
53. Haritonov, A. Yu., 1988. “ [Dragonflies of the genus Ischnura Charp. (Insecta, Odonata) in the fauna of the USSR. In Russian] ”. In “Taksonomiya zhivotnykh Sibiri”. Novye i Maloizvestnye Vidy Fauny Sibiri 20. Nauka, Novosibirsk Edited by:Zolotarenko, G. S., 32–46.
54. Hilton, D. F.J., 1987. A terminology for females with color patterns that mimic males. Entomological News, 98:221–223.
55. Hinnekint, B. O.N., 1987. Population dynamics of Ischnura e. elegans (Vander Linden) (Insecta:Odonata) with special reference to morphological colour changes, female polymorphism, multiannual cycles and their influence on behaviour. Hydrobiologia, 146:3–31.
56. Holland, B., and Rice, W. R., 1999. Experimental removal of sexual selection reversed intersexual antagonistic coevolution and removes a reproductive load. Proceedings of the National Academy of Science, 96:5083–5088.
57. Jödicke, R., 1986. Libellenbeobachtungen in Mooren Norwegens. Libellula, 5 (1/2;):41–42.
58. Jödicke, R., 1995. Frühjahrsaspekte der Odonatenfauna in Marokko südlich des Hohen Atlas. Opuscula Zoologica Fluminensia, 134:1–10.
59. Jödicke, R., 1996. Die Odonatenfauna der Provinz Tarragona (Catalunya, Spanien). Advances in Odonatology, Supplement, 1:77–111.
60. Jödicke, R., 1997. Die Binsenjungfern und Winterlibellen Europas. Lestidae Magdeburg:Westarp.
61. Jödicke, R., 1999. Libellenbeobachtungen in Podlasie, Nordost-Polen. Libellula, 18:31–48.
62. Jödicke, R., 2001. Die Libellen der Dianaseen (Insecta:Odonata). Drosera, 2001:117–125.
63. Jödicke, R., 2005. Bemerkungen zu Coenagrion intermedium (Odonata:Coenagrionidae). Libellula Supplement, 6:15–24.
64. Jödicke, R., Arlt, J., Kunz, B., Lopau, W., and Seidenbusch, R., 2000. The Odonata of Tunisia. International Journal of Odonatology, 3:41–71.
65. Johnson, C., 1964. The inheritance of the female dimorphism in the damselfly, Ischnura damula. Genetics, 49:513–519.
66. Johnson, C., 1966. Genetics of female dimorphism in Ischnura demorsa. Heredity, 21:453–459.
67. Johnson, C., 1975. Polymorphism and natural selection in ischnuran damselflies. Evolutionary Theory, 1:81–90.
68. Kotarac, M., 1996. A note on the existence of androchrome females in Crocothemis erythraea (Brullé) (Anisoptera:Libellulidae). Notulae Odonatologicae, 4:123–124.
69. Krüner, U., 1986. Die Späte Adonislibelle (Ceriagrion tenellum [de Villers]) im südwestlichen niederrheinischen Tiefland (Nordrhein-Westfalen). Libellula, 5 (3/4;):85–94.
70. Krüner, U., 1989. Die Verbreitung, Biologie und Ökologie der Späten Adonislibelle Ceriagion tenellum (Odonata, Coenagrionidae) in Nordwesteuropa, insbesondere im Gebiet zwischen Maas und Rhein. Verhandlungen Westdeutscher Entomologen Tag, 1988:133–140.
71. Krüner, U., 2003. Die Häufigkeitsverteilung der Weibchenfarben von Ceriagrion tenellum an drei Gewässern im Naturpark Schwalm-Nette (Odonata:Coenagrionidae). Libellula, 22:133–140.
72. Lacroix, J. L., 1915. Notes névroptérologiques. II. Excursions en Charente-Inférieure. I. Insecta, 5:106–118.
73. Lacroix, J. L., 1919. Notes névroptérologiques. VIII. Excursions en Charente-Inférieure. III. Boletin de la Sociedad Ibérica de Ciencas Naturales, 1919:98–102.
74. Lajeunesse, M. J., and Forbes, M. R., 2003. A comparison of structural size and condition of two female morphs of the damselfly Nehalennia irene (Hagen) (Zygoptera:Coenagrionidae). Odonatologica, 32:281–287.
75. Lam, E., 2004. Damselflies of the Northeast:a guide to the species for eastern Canada and the northeastern United States. Forest Hills, NY:Biodiversity Books.
76. Langenbach, A., 1993. Verhaltensökologie von Ischnura pumilio (Charpentier) unter besonderer Berücksichtigung des Farbwechsels der Weibchen (Insecta:Odonata). Diplomarbeit, Technische Universität Braunschweig.
77. Manolis;, T., 2003. Dragonflies and damselflies of California. Berkeley:University of California Press.
78. May, M. L., 1976. Physiological color changes in New World damselflies. Odonatologica, 7:27–47.
79. McKee, D., Harvey, I. F., Thompson, D. J., and Sherratt, T. N., 2005. Frequency of female colour morphs in populations of four coenagrionid damselflies (Zygoptera:Coenagrionidae). Odonatologica, 34:37–49.
80. Miller, M. N., and Fincke, O. M., 1999. Cues for mate recognition and the effect of prior experience on mate recognition in Enallagma damselflies. Journal of Insect Behavior, 12:801–814.
81. Miller, M. N., and Fincke, O. M., 2004. Mistakes in sexual recognition among sympatric zygoptera vary with time of day and color morphism (Odonata:Coenagrionidae). International Journal of Odonatology, 7:471–491.
82. Naraoka, H., 2003. Changes of the body color of Nehalennia speciosa (Coenagrionidae:Odonata). Gekkan-Mushi, 388:38–40. [In Japanese; English title].
83. Nikula, B., Loose, J. L., and Burne, M. R., 2003. A field guide to the dragonflies and damselflies of Massachusetts. Massachusetts Division of Fisheries & Wildlife Natural Heritage & Endangered Species Program, Westborough.
84. Parr, M. J., 1965. A population study of a colony of imaginal Ischnura elegans (Van der Linden) (Odonata:Coenagrionidae) at Date, Pembrokeshire. Field Studies, 2:237–282.
85. Parr, M. J., 1969. Population studies of some zygopteran dragonflies (Odonata). Ph.D. Thesis University of Salford.
86. Parr, M. J., 1999. The terminology of female polymorphs of Ischnura (Zygoptera:Coenagrionidae). International Journal of Odonatology, 2:95–99.
87. Parr, M. J., and Palmer, M., 1971. The sex ratios, mating frequencies and mating expectancies of three coenagriids in northern England. Entomologica Scandinavica, 2:191–204.
88. Paulson, D. R., 1974. Reproductive isolation in damselflies. Systematic Zoology, 23:40–49.
89. Paulson, D. R., 1998. Possible morphological and behavioral male mimicry in a libellulid dragonfly, Erythrodiplax umbrata (L.) (Anisoptera:Libellulidae). Odonatologica, 27:249–252.
90. Paulson, D. R., 1999. Ischnura perparva Mclachlan (Zygoptera:Coenagrionidae) has an andromorphic female, and another suggestion to modify the terminology of female color polymorphism in Odonata. International Journal of Odonatology, 2:101–103.
91. Perl, F., 1998. Verhaltensökologische Untersuchungen an der Späten Adonislibelle (Ceriagrion tenellum) im Landkreis Celle. Staatsexamensarbeit, Universität Hannover.
92. Peters, G., 1985. Die Libellenfauna der westlichen und nördlichen Mongolei und einige Phänomene ihrer intrakontinentalen Isolation. Mitteilungen des Zoologischen Museums Berlin, 61:11–42.
93. Peters, G., 1987. Die Edellibellen Europas. Die Neue Brehm-Bücherei 585. Ziemsen, Wittenberg.
94. Petzold, F., 2002. Beobachtungen zum Verhalten von Aeshna crenata und A. grandis an einem Gewässer in Westsibirien (Odonata:Aeshnidae). Libellula, 21:79–100.
95. Prum, R. O., Cole, J. A., and Torres, R. H., 2004. Blue integumentary structural colors in dragonflies (Odonata) are not produced by incoherent Tyndall scattering. Journal of Experimental Biology, 207:3999–4009.
96. Rehn, A. C., 2003. Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology, 28:181–239.
97. Reiter, C., 1993. Beobachtungen zur Autökologie von Ischnura pumilio, Charpentier, 1825, und Ischnura elegans, v.d. Linden, 1820, in der Umgebung von Freising (Insecta, Odonata). Diplomarbeit, Universität München.
98. Robertson, H. M., 1985. Female dimorphisms and mating behaviour in a damselfly, Ischnura ramburii:females mimicking males. Animal Behaviour, 33:805–809.
99. Robinson, J. V., and Jordan, W. H., 1996. Ontogenetic color change in Ischnura kellicotti Williamson females (Zygoptera:Coenagrionidae). Odonatologica, 25:83–85.
100. Robinson, J. V., and Allgeyer, R., 1996. Covariation in life-history traits, demographics, and behaviour in ischnuran damselflies; the evolution of monandry. Biological Journal of the Linnean Society, 58:85–98.
101. Rowe, L., Arnqvist, G., Sih, A., and Krupa, J. J., 1994. Sexual conflict and the evolutionary ecology of mating patterns:water striders as a model system. Trends in Ecology and Evolution, 9:289–293.
102. Samraoui, B., Weekers, P. H.H., and Dumont, H. J., 2002. The Enallagma of the western and central Palaearctic. Odonatologica, 31:371–381.
103. Sánchez-Guillén, R. A., Van Gossum, H., and Cordero Rivera, A., 2005. Hybridization and the inheritance of female colour polymorphism in two ischnurid damselflies (Odonata:Coenagrionidae). Biological Journal Linnean Society, 85:471–481.
104. Schiess, H., 1973. Beitrag zur Kenntnis der Biologie von Nehalennia speciosa (Charpentier, 1840) (Zygoptera:Coenagrionidae). Odonatologica, 2:33–37.
105. Schmidt, E., 1929. “ 7. Ordnung:Libellen, Odonata. ”. In “Die Tierwelt Mitteleuropas 4(1b)” Edited by:Brohmer, P., Ehrmann, P., and Ulmer, G., 1–66. Leipzig:Quelle & Meyer.
106. Schmidt, E., 1954. Ergebnisse der Österreichischen Iran-Expedition 1949/50. Die Libellen Irans. Sitzungsberichte der Österreichischen Akademie der Wissenschaften (Abt. I), 163:223–260.
107. Schmidt, E., 1964. Zur Genealogie der Libelle Agrion pulchellum (v.d. Lind. 1825) Selys-Hagen 1850(Odonata, Zygoptera). Schmidt, Bonn.
108. Schmidt, E., 1967. Versuch einer Analyse der Ischnura elegans-Gruppe (Zygoptera). Entomologiske Tidskrift, 88:188–225.
109. Seidenbusch, R., 1995. Dichromism in females of Ischnura pumilio (Charpentier), with special reference to homeochromic females. Journal of the British Dragonfly Society, 11:21–22.
110. de Selys Longchamps, E., 1854. Monographie des Caloptérygines. Mémoires de la Société d'Science d'Liege 9:i-xi, 1-291. [avec H.A. Hagen].
111. Sherratt, T. N., 2001. The evolution of female-limited polymorphisms in damselflies:a signal detection model. Ecology Letters, 4:22–29.
112. Sherratt, T. N., and Forbes, M. R., 2001. Sexual differences in coloration of coenagrionid damselflies (Odonata):a case of intraspecific aposematism? Animal Behaviour, 62:653–660.
113. Sirot, L. K., and Brockman, H. J., 2001. Costs of sexual interactions to females in Rambur's forktail damselfly, Ischnura ramburi (Zygoptera:Coenagrionidae). Animal Behaviour, 61:415–424.
114. Sirot, L. K., Brockman, H. J., Marinis, C., and Muschett, G., 2003. Maintenance of a female-limited polymorphism in Ischnura ramburi (Zygoptera:Coenagrionidae). Animal Behaviour, 66:763–775.
115. Staddon, J. E.R., and Gendron, R. P., 1983. Optimal detection of cryptic prey may lead to predator switching. American Naturalist, 122:843–848.
116. Sternberg, K., 1995. Experimentelle Erzeugung androchromer Weibchen durch Einwirkung hoher Temperaturen bei Arten der Libellen-Gattung Aeshna (Anisoptera:Aeshnidae). Entomologia Generalis, 20:37–42.
117. Sternberg, K., 1996. Colours, colour pattern and ‘cuticular windows’ as light traps - their thermoregulatoric and ecological significance in some Aeshna species (Odonata:Aeshnidae). Zoologischer Anzeiger, 235:77–88.
118. Sternberg, K., and Buchwald, R., eds. 1999. Die Libellen Baden-Württembergs. Band Vol. 1, Allgemeiner Teil, Kleinlibellen (Zygoptera). Ulmer, Stuttgart.
119. Sternberg, K., and Buchwald, R., eds. 2000. Die Libellen Baden-Württembergs. Band Vol. 2, Großlibellen (Anisoptera), Literatur. Ulmer, Stuttgart.
120. Stobbe, H., 1990. Bemerkungen zur Gattung Calopteryx in Griechenland. Naturkundlicher Rundbrief, 4:5–20.
121. Sugimura, M., Ishida, S., Kojima, K., Ishida, K., and Aoki, T., 2001. Dragonflies of the Japanese Archipelago in color. Sapporo:Hokkaido University Press.
122. Svensson, E. I., Abbott, J., and Härdling, R., 2005. Female polymorphism, frequency dependence, and rapid evolutionary dynamics in natural populations. American Naturalist, 165:567–576.
123. Tennessen, K. J., 1975. Reproductive behavior and isolation of two sympatric coenagrionid damselflies in Florida. Ph.D. Dissertation, University of Florida, Gainesville.
124. Thompson, D. J., 1989. Lifetime reproductive success in andromorph females of the damselfly Coenagrion puella (L.) (Zygoptera:Coenagrionidae). Odonatologica, 18:209–213.
125. Turgeon, J., Stoks, R., Thum, R. A., Brown, J. M., and McPeek, M. A., 2005. Simultaneous Quaternary radiations of three damselfly clades across the Holarctic. American Naturalist, 165:78–107.
126. Valle, K. J., 1927. Zur Kenntnis der Odonatenfauna Finnlands III. Acta Societatis pro Fauna et Flora Fennica, 56 (11):1–36.
127. Van Gossum, H., Stoks, R., Matthysen, E., Valck, F., and De Bruyn, L., 1999. Male choice for female colour morphs in Ischnura elegans (Odonata, Coenagrionidae):testing the hypotheses. Animal Behaviour, 57:1229–1232.
128. Van Gossum, H., Stoks, R., and De Bruyn, L., 2001a. Discriminative mate choice in relation with female maturation in Ischnura elegans (Odonata:Coenagrionidae). International Journal of Odonatology, 4:83–91.
129. Van Gossum, H., Stoks, R., and De Bruyn, L., 2001b. Reversible frequency-dependent switches in male mate choice. Proceedings of the Royal Society of London (B), 268:83–85.
130. Van Gossum, H., Stoks, R., and De Bruyn, L., 2001c. Frequency dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly, Ischnura elegans. Behavioral Ecology and Sociobiology, 51:69–75.
131. Van Gossum, H., Adriaens, T., Dumont, H., and Stoks, R., 2004. Sex- and morph-specific predation risk:Colour or behaviour dependency? European Journal of Entomology, 101:373–377.
132. Walker, E. M., 1953. The Odonata of Canada and Alaska. Vol. 1, Toronto:University of Toronto Press.
133. Walker, E. M., 1958. The Odonata of Canada and Alaska. Vol. 2, Toronto:University of Toronto Press.
134. Westfall, M. J., and May, M. L., 1996. Damselflies of North America. Gainesville:Scientific Publishers.