List of animals displaying homosexual behavior explained

For these animals, there is documented evidence of homosexual behavior of one or more of the following kinds: sex, courtship, affection, pair bonding, or parenting, as noted in researcher and author Bruce Bagemihl's 1999 book Biological Exuberance: Animal Homosexuality and Natural Diversity.

Bagemihl writes that the presence of same-sex sexual behavior was not "officially" observed on a large scale until the 1990s due to observer bias caused by social attitudes towards nonheterosexual people, making the homosexual theme taboo.^{[1] [2]} Bagemihl devotes three chapters, "Two Hundred Years at Looking at Homosexual Wildlife", "Explaining (Away) Animal Homosexuality" and "Not For Breeding Only" in his 1999 book Biological Exuberance to the "documentation of systematic prejudices" where he notes "the present ignorance of biology lies precisely in its single-minded attempt to find reproductive (or other) 'explanations' for homosexuality, transgender, and non-procreative and alternative heterosexualities."^[3] Petter Bøckman, academic adviser for the Against Nature? exhibit, stated "[M]any researchers have described homosexuality as something altogether different from sex. They must realize that animals can have sex with who they will, when they will and without consideration to a researcher's ethical principles." Homosexual behavior is found amongst social birds and mammals, particularly the sea mammals and the primates.^{[2] [4]} In 1986, it was even discovered amongst insects when butterfly scientist W.J. Tennent observed four male Mazarine blues competing for the attention of another male in Morocco.^[5]

Sexual behavior takes many different forms, even within the same species and the motivations for and implications of their behaviors have yet to be fully understood. Bagemihl's research shows that homosexual behavior, not necessarily sex, has been documented in about five hundred species as of 1999, ranging from primates to gut worms.^{[1] [6]} Homosexuality in animals is seen as controversial by social conservatives because it asserts the naturalness of homosexuality in humans, while others counter that it has no implications and is nonsensical to equate natural animal behaviors to morality.^{[7] [8]} Sexual preference and motivation is always inferred from behavior. Thus homosexual behavior has been given a number of terms over the years. The correct usage of the term homosexual is that an animal exhibits homosexual behavior, however this article conforms to the usage by modern research,^{[9] [10] [11] [12]} applying the term homosexuality to all sexual behavior (copulation, genital stimulation, mating games and sexual display behavior) between animals of the same sex.

In October 2023, biologists reported studies of mammals (over 1,500 different species) that found same-sex behavior (not necessarily related to human orientation) may help improve social stability by reducing conflict within the groups studied.^{[13] [14]}

__TOC__

Mammals

See main article: List of mammals displaying homosexual behavior.

Selected mammals from the full list

• Baboon
• Bison^[15]
• Bonobo^[16]
• Brown bear^[17]
• Brown Rat^[18]
  • Cavy Caribou^[19]
• Cat (domestic)^[20]
• Cattle (domestic)^[21]
• Chimpanzee^{[22] [23] [24] [25]}
• Common dolphin^[26]
• Common marmoset^[27]
• Dog^[28]
• Dolphin^[29]
• Elephant^[30]
• Fox^[31]
• Giraffe^{[2] [32] [33]}
• Goat^[15]
• Horse (domestic)^[34]
• Human^{[35] [36] [37]}
• Humpback whale^[38]
• Koala^[39]
• Lion
• Orca
• Wolf

Birds

See main article: List of birds displaying homosexual behavior. Homosexual behaviour is demonstrated by 120 known species of birds. ^[40] While an uptick in research on bird homosexuality – and animal homosexuality in general – has been coming out in recent years, it's common for some authors to labour in articulating the view any root cause or function of bird homosexuality is poorly understood.^[41]

Hypotheses contrived in an attempt to explain the behaviour – homosexuality in birds – typically diverge from one another, further exacerbating an apparent perception in the concerned scientific communities knowledge's quest to realise bird homosexuality remains elusive as an objective. Some authors posit the behaviours are a result of any bird demonstrably homosexual being less inclined to rear young, while other authors posit the timing of emergence from the egg is a factor.^[42]

In 1977, The New York Times also reported on a colony of lesbian seagulls "off [the] coast of California."^[43]

Selected birds from the full list

• Barn owl^[44]
• Cassowary
• Black swan
• Chicken^[45]
• Common gull^[46]
• Emu^[47]
• House sparrow^[48]
• Kestrel^[44]
• King penguin^[49]
• Mallard^[50]
• Ostrich^[47]
• Raven^[51]
• Rock dove^[52]

Fish

• Amazon molly^[53]
• Anglerfish
• Blackstripe topminnow^[54]
• Bluegill sunfish^[54]
• Char
• Grayling
• European bitterling^[55]
• Green swordtail^[55]
• Guiana leaffish^[56]
• Houting whitefish
• Jewel cichlid^[57]
• Least darter (Microperca punctulata)^[55]
• Mouthbreeding fish sp.^[54]
• Salmon spp.^[58]
• Southern platyfish^[55]
• Ten-spined stickleback^[55]
• Three-spined stickleback^[55]

Reptiles

The all-female Whiptail lizard species Aspidoscelis neomexicanus (center), which reproduces via parthenogenesis, is shown flanked by two sexual species having males, A. inornatus (left) and A. tigris (right). Research has shown that simulated mating behavior increases fertility for Aspidoscelis neomexicanus. One female lies on top of another, playing the role of the male, the lizard that was on bottom has larger eggs, in most cases. The lizards switch off this role each mating season.^[59]

• Anole sp.^[60]
• Bearded dragon
• Blue-tailed day gecko (Phelsuma cepediana)^[61]
• Broad-headed skink^[55]
• Checkered whiptail lizard^[62]
• Chihuahuan spotted whiptail lizard^[62]
• Common ameiva^[62]
• Common garter snake^[55]
• Cuban green anole^[60]
• Desert grassland whiptail lizard^[62]
• Desert tortoise^[63]
• Eastern fence lizard^[62]
• Five-lined skink^[55]
• Gold dust day gecko (Phelsuma laticauda)^[61]
• Gopher (pine) snake^[54]
• Green anole^[60]
• Inagua curlytail lizard^[62]
• Jamaican giant anole^[60]
• Laredo striped whiptail lizard^[62]
• Largehead anole^[60]
• Mourning gecko^[64]
• Plateau striped whiptail lizard^[62]
• Red diamond rattlesnake^[55]
• Red-tailed skink^[55]
• Seychelles giant tortoise^[65]
• Side-blotched lizard^[62]
• Speckled rattlesnake^[55]
• Spectacled Caiman^[66]
• Water moccasin^[55]
• Western rattlesnake (Crotalus viridis)^[55]
• Western banded gecko^[64]
• Whiptail lizard spp.^[62]
• Wood turtle^[60]
• Blue-tongued skink

Amphibians

See also: Gay frogs (internet meme).

• Appalachian woodland salamander^[67]
• Black-spotted frog^[68]
• Mountain dusky salamander^[67]
• Tengger desert toad^[60]

Insects

Male homosexuality has been inferred in several species of dragonflies. A survey of damsel and dragonflies reveals characteristic cloacal pincher mating damage in 20–80 percent of the males, indicating a fairly high occurrence of sexual coupling between males.^{[69] [70]}

• Alfalfa weevil^[71]
• Australian parasitic wasp sp.^[71]
• Bean weevil sp.^[71]
• Bedbug and other bug spp.^{[72] [73]}
• Blister beetle spp.^[74]
• Blowfly
• Broadwinged damselfly sp.^[75]
• Cabbage (small) white (butterfly)
• Checkerspot butterfly^[76]
• Club-tailed dragonfly spp.^[77]
• Cockroach spp.^[78]
• Codling moth^[79]
• Common skimmer dragonfly spp.^[77]
• Creeping water bug sp.^[80]
• Cutworm^[81]
• Digger bee^[82]
• Dragonfly spp.^[77]
• Eastern giant ichneumon wasp^[71]
• Eucalyptus longhorned borer
• Field cricket sp.^[83]
• Flour beetle^[84]
• Fruit fly spp.^{[85] [86]}
• Glasswing butterfly^[76]
• Hypoponera opacior ant^[87]
• Grape borer^[80]
• Green lacewing^[88]
• Hen flea^[88]
• House fly^[89]
• Ichneumon wasp sp.^[71]
• Japanese scarab beetle^[90]
• Larch bud moth^[79]
• Large milkweed bug^[73]
• Large white^[73]
• Leek moth^[91]
• Long-legged fly spp.
• Mazarine blue^[73]
• Mexican white (butterfly)^[73]
• Midge sp.^[92]
• Migratory locust^[93]
• Monarch butterfly^[76]
• Narrow-winged damselfly spp.^[75]
• Parsnip leaf miner^[92]
• Peach moth^[94]
• Pomace fly^[92]
• Queen butterfly^[76]
• Red ant sp.^[92]
• Red flour beetle^[73]
• Rice moth^[95]
• Reindeer warble fly (Hypoderma tarandi)^[92]
• Rose chafer (Macrodactylus subspinosus)^[92]
• Rove beetle spp.^[73]
• Scarab beetle (melolonthine)^[96]
• Screwworm fly^[92]
• Southeastern blueberry bee^[82]
• Southern green stink bug^[73]
• Southern masked chafer^[92]
• Southern one-year canegrub^[92]
• Spreadwinged damselfly spp.^[75]
• Spruce budworm moth^{[97] [98]}
• Stable fly sp.^[92]
• Stag beetle spp.^[73]
• Tsetse fly^[92]
• Tropical tasar silkmoth^[99]
• Vine moth^[100]
• Water boatman bug^[73]
• Water strider spp.^[73]

Other invertebrates

• Blood-fluke^[101]
• Box crab^[102]
• Harvestman sp.^[103]
• Hawaiian orb-weaver (spider)^[103]
• Incirrate octopus spp.^[102]
• Jumping spiders^[103]
• Mite sp.^[92]
• Spiny-headed worm^[104]

See also

• Against Nature?, an exhibit at the University of Oslo's Natural History Museum that took place until 19 August 2007.
• Anthropomorphism
• Behavioral ecology is the study of the ecological and evolutionary basis for animal behavior
• Biodiversity
• Bioethics
• Biology and sexual orientation
• Ethology is the scientific study of animal behavior, and a branch of zoology; cognitive ethology fuses cognitive science and classical ethology to observe animals under more-or-less natural conditions
• Evolutionary biology
  • Innate bisexuality
• Sexual selection

Bibliography

• News: Gay Penguins Resist 'Aversion Therapy' . 365Gay.com . 11 February 2005 . https://web.archive.org/web/20070929090349/http://www.365gay.com/newscon05/02/021105penguins.htm . 29 September 2007 . 16 June 2010.
• Bagemihl, Bruce (1999). Biological Exuberance: Animal Homosexuality and Natural Diversity. St. Martin's Press
• Caramagno, Thomas C (2002). Irreconcilable Differences? Intellectual Stalemate in the Gay Rights Debate; Praeger/Greenwood, .
• Cooper, J.B. "An Exploratory Study on African Lions" in Comparative Psychology Monographs 17:1-48.
• Cziko, Gary (2000) The Things We Do: Using the Lessons of Bernard and Darwin to Understand the What, How, and Why of Our Behavior; MIT Press, .
• de Waal, Frans B. M. (2001) The Ape and The Sushi Master: Cultural Reflections by a Primatologist; Basic Books (chapter Bonobos and Fig Leaves).
• Dunkle, S.W. (1991), "Head damage from mating attempts in dragonflies (Odonata:Anisoptera)". Entomological News 102, pp. 37–41. Retrieved on 16 June 2010.
• Eaton, R. L. (1974). "The Biology and Social Behavior of Reproduction in the Lion" in Eaton, ed. The World's Cats, vol. II; pp. 3–58; Seattle.
• Forger, Nancy G., Laurence G. Frank, S. Marc Breedlove, Stephen E. Glickman (6 December 1998). "Sexual Dimorphism of Perineal Muscles and Motoneurons in Spotted Hyenas"; The Journal of Comparative Neurology, Volume 375, Issue 2, Pages 333 - 343. Retrieved 11 September 2007.
• Web site: Gay Animals: Alternate Lifestyles in the Wild . Live Science . 19 June 2018 . 20 September 2011.
• Goudarzi, Sara (16 November 2006). "Gay Animals Out of the Closet?: First-ever Museum Display Shows 51 Species Exhibiting Homosexuality". MSNBC. Retrieved on 12 September 2007.
• News: Harrold. Max. February 16, 1999. Creature Comforts. The Advocate. March 10, 2018. 61–62. 779. In his news book, Biological Exuberance: Animal Homosexuality and Natural Diversity ... author Bruce Bagemihl portrays an animal kingdom that embraces a whole spectrum of sexual orientations ... [and] paints a complex mosaic that resembles humanity ... At 751 pages and with photos and documentation of homosexual behaviour in more than 450 species of mammals, birds, repties, and insects, Biological Exuberance brings the dusty facts to light as Bagemihl deconstructs the all-heterosexual Noah's Ark we've been sold..
• Holekamp, Kay E. (2003). Research: Spotted Hyena - Introduction and Overview. Michigan State University, Department of Zoology]. Retrieved 16 June 2010.
• Kick, Russ (2001). You Are Being Lied to: The Disinformation Guide to Media Distortion, Historical Whitewashes and Cultural Myths. The Disinformation Company, . Retrieved on 18 November 2007.
• Web site: The Science of Sex . 19 September 2007 . 16 June 2010 . https://web.archive.org/web/20071108213627/http://www.nerve.com/Regulars/ScienceOfSex/09-19-00/09-19-00.asp . 2007-11-08 .
• Web site: https://web.archive.org/web/20020602221325/http://www.colszoo.org/animalareas/aforest/bonobo.html . African Forest: Bonobo . Dave . Liggett . Columbus Zoo and Aquarium staff . Columbus Zoo and Aquarium . June 2, 2002 . ...frequent sex (including male-to-male and female-to-female) characterize bonobo society. . November 14, 2011.
• News-medical.net (23 October 2006). "1,500 Animal Species Practice Homosexuality" Retrieved on 10 September 2007.
• Poiani, Aldo (2010). Animal Homosexuality: A Biosocial Perspective. Cambridge University Press.
• Roselli, Charles E., Kay Larkin, John A. Resko, John N. Stellflug and Fred Stormshak (2004). "The Volume of a Sexually Dimorphic Nucleus in the Ovine Medial Preoptic Area/Anterior Hypothalamus Varies with Sexual Partner Preference". Endocrinology, Department of Physiology and Pharmacology, Oregon Health & Science University (C.E.R., K.L., J.A.R.), Portland, Oregon; Department of Animal Sciences, Oregon State University (F.S.), Corvallis, Oregon; and Agricultural Research Service, United States Sheep Experiment Station (J.N.S.), Dubois, Idaho, Vol. 145, No. 2. Retrieved on 10 September 2007.
• Roughgarden, Joan (2004). Evolutions Rainbow: Diversity, Gender and Sexuality in Nature and People; University of California Press, Berkeley, pages p. 13-183.
• Schaller, G. B. (1972). The Serengeti Lion; University of Chicago Press.
• Smith, Dinitia (7 February 2004). "Love That Dare Not Squeak Its Name" New York Times. Retrieved on 10 September 2007. Reprinted as "Central Park Zoo's Gay Penguins Ignite Debate", San Francisco Chronicle.
• Sommer, Volker & Paul L. Vasey (2006). Homosexual Behaviour in Animals, An Evolutionary Perspective. Cambridge University Press, Cambridge; ISBN 0521864461.
• Srivastav, Suvira (15–31 December 2001). "Lion, Without Lioness"
• Stein, Edward (1999) The Mismeasure of Desire: The Science, Theory, and Ethics of Sexual Orientation; Oxford University Press, US; .
• Tatarnic, Nikolai J., Gerasimos Cassis, Dieter F. Hochuli; 22 March 2006 "Traumatic insemination in the plant bug genus Coridromius Signoret (Heteroptera: Miridae)" Biology Letters Journal Volume 2, Number 1, pg 58-61: Royal Society Publishing; Retrieved 16 June 2010.
• Terry, Jennifer (2000) "'Unnatural Acts' In Nature: The Scientific Fascination with Queer Animals"; GLQ: A Journal of Lesbian and Gay Studies (6(2):151–193; OI:10.1215/10642684-6-2-151); Duke University Press.
• Utzeri, C. & C. Belfiore (1990): "Anomalous tandems in Odonata". Fragmenta Entomologica 22(2), pp. 271–288. Retrieved 11 September 2007.
• Vasey, Paul L. (1995), "Homosexual Behaviour in Primates: A Review of Evidence and Theory"; International Journal of Primatology 16: p 173-204.
• Wilson. Anna. 10.1086/345320. Sexing the Hyena: Intraspecies Readings of the Female Phallus. University of Chicago Press. 10.1086/345320. Signs. 28. 3. 755–790. 2003. 146640802 .
• Zimmer, Carl (2000); Parasite Rex: Inside the Bizarre World of Nature's Most Dangerous Creatures; Simon and Schuster, . Retrieved 18 November 2007.

References

1. Bagemihl (1999)
2. News-medical.net (2006)
3. Bagemihl (1999) page 213
4. Web site: Same-Sex Behavior Among Animals Isn’t New. Science Is Finally Catching Up. . 2024-01-17 . National Wildlife Federation . en.
5. Web site: Same-Sex Behavior Among Animals Isn’t New. Science Is Finally Catching Up. . 2024-01-17 . National Wildlife Federation . en.
6. Harrold (1999)
7. Solimeo (2004)
8. Solimeo (2004b)
9. Bagemihl (1999) pages 122-166
10. Roughgarden (2004) pp.13-183
11. Vasey (1995) pages 173-204
12. Sommer & Vasey (2006)
13. News: Zimmer . Carl . Carl Zimmer . Same-Sex Behavior Evolved in Many Mammals to Reduce Conflict, Study Suggests - But the researchers cautioned that the work could not shed much light on sexual orientation in humans. + comment . 3 October 2023 . . live . https://ghostarchive.org/archive/sxXGh . 4 October 2023 . 4 October 2023 .
14. Gómez, José M. . et al. . The evolution of same-sex sexual behaviour in mammals . 3 October 2023 . . 14 . 5719 . 5719 . 10.1038/s41467-023-41290-x . free . 37788987 . 10547684 . 2023NatCo..14.5719G .
15. Bagemihl (1999) page 405
16. Hohmann. G.. Fruth. B.. July 2000. Use and function of genital contacts among female bonobos. Animal Behaviour. 60. 1. 107–120. 10.1006/anbe.2000.1451. 10924210. 39702173. 0003-3472. 2021-02-19. 2023-02-06. https://web.archive.org/web/20230206003707/https://www.sciencedirect.com/science/article/abs/pii/S0003347200914519?via%3Dihub. live.
17. Bagemihl (1999) page 441
18. Bagemihl (1999) page 469
19. Bagemihl (1999) pages 388,389
20. Bagemihl 1999) pages 81, 88
21. Bagemihl (1999) pages 81, 82, 89
22. de Waal (2001)
23. Liggett (1997–2006)
24. Imaginova (2007j)
25. Bagemihl (1999) pages 276–279
26. Bagemihl (1999) page 339
27. Bagemihl (1999) page 334
28. Bagemihl (1999) pages 310, 314
29. name="Biol.Exuberance: Dolphin
30. Bagemihl (1999) page 427
31. Bagemihl (1999) pages 218, 231, 317
32. Bagemihl (1999) page 391
33. Imaginova (2007d)
34. Bagemihl (1999) pages 81, 165, 205, 226, 231
35. Bagemihl (1999) page 432
36. 10.1007/BF01541598. Sell RL, Wells JA, Wypij D . The prevalence of homosexual behavior and attraction in the United States, the United Kingdom and France: results of national population-based samples . Archives of Sexual Behavior . 24 . 3 . 235–48 . June 1995 . 7611844. 12929812 .
37. Wellings, K., Field, J., Johnson, A., & Wadsworth, J. (1994). Sexual behavior in Britain: The national survey of sexual attitudes and lifestyles. London, UK: Penguin Books.
38. An observation of sexual behavior between two male humpback whales. Stephanie H. Stack. Lyle Krannichfeld. Brandi Romano. Marine Mammal Science. 2024. 10.1111/mms.13119. free.
39. Bagemihl (1999) page 455-457
40. MacFarlane . Geoff R. . Blomberg . Simon P. . Vasey . Paul L. . 2010-09-01 . Homosexual behaviour in birds: frequency of expression is related to parental care disparity between the sexes . Animal Behaviour . 80 . 3 . 375–390 . 10.1016/j.anbehav.2010.05.009 . 53148085 . 0003-3472.
41. Regaiolli . Barbara . July 2018 . Investigating parental care behaviour in same-sex pairing of zoo greater flamingo (Phoenicopterus roseus) . The Open Access Journal for Life and Environment. 6 . e5227 . 10.7717/peerj.5227 . 30042888 . 6054785 . free .
42. Web site: Same-sex sexual behavior in birds: expression is related to social mating system and state of development at hatching . 2023-12-04 . academic.oup.com.
43. Web site: November 23, 1977 . Extensive Homosexuality Is Found Among Seagulls Off Coast of California . 4 December 2023 . The New York Times.
44. Bagemihl (1999) pages 632-5
45. Bagemihl (1999) pages 83
46. Bagemihl (1999) pages 544-8
47. Bagemihl (1999) page 621-6
48. Bagemihl (1999) pages 602-5
49. 365 Gay.com (2005)
50. Bagemihl (1999) pages 491-5
51. Bagemihl (1999) pages 606-10
52. Bagemihl (1999) page 639
53. Bagemihl (1999), page 37
54. Bagemihl (1999), pages 658, 664
55. Bagemihl (1999), page 658
56. Bagemihl (1999), page 664
57. Bagemihl (1999), pages 658, 665
58. Bagemihl (1999), pages 232, 233, 244
59. LeVay, (19 September 2007)
60. Bagemihl (1999), page 657
61. Budzinski, R.-M. (1997) Homosexuelles Verhalten bei Geckos der Gattung Phelsuma. Sauria 19 (3): 33-34
62. Bagemihl (1999), page 657, 658
63. Bagemihl (1999), pages 232, 664
64. Bagemihl (1999), pages 663–664
65. Web site: Turns out this 186-year-old tortoise has a gay lover . 19 October 2017 . 2018-01-23 . 2018-01-24 . https://web.archive.org/web/20180124005744/https://nypost.com/2017/10/19/turns-out-this-186-year-old-tortoise-has-a-gay-lover/amp/ . live .
66. Antelo . R . Caiman crocodilus (spectacled caiman) homosexual behavior. . Herpetological Review . 2012 . 43 . 2 . 327–328 .
67. Bagemihl (1999), pages 657, 658
68. Bagemihl (1999), pages 243, 664
69. Dunkle (1991)
70. Young, L.C., et al. (2008)
71. Bagemihl (1999), page 666
72. Tatarnic1 et al., 22 March 2006
73. Bagemihl (1999) page 660
74. Bagemihl (1999) page 667
75. Bagemihl (1999) pages 704, 713
76. Bagemihl (1999) pages 150, 232, 236, 246
77. Bagemihl (1999) pages 33–34, 196, 217, 219, 232
78. Bagemihl (1999) page 668
79. 10.1007/BF01926659. Benz G. Role of sex pheromone, and its insignificance for heterosexual and homosexual behaviour of larch bud moth. Experientia. 29. 5 . 553–554 . May 1, 1973 . 39282068 .
80. Bagemihl (1999) page 666
81. Bagemihl (1999) page 595
82. Bagemihl (1999) pages 9, 649, 665
83. Bagemihl (1999) page 658
84. http://www.out.com/ Out magazine
85. 10.1016/j.anbehav.2010.08.017. Dukas, R . Causes and consequences of male–male courtship in fruit flies . Animal Behaviour . 80 . 5 . 913–919 . November 2010. 54253398 .
86. Bagemihl (1999) page 3
87. Kureck, I. M. . Neumann, A. . Foitzik, S. . 2011 . Wingless ant males adjust mate-guarding behaviour to the competitive situation in the nest . Animal Behaviour . 82 . 2 . 339–346 . 10.1016/j.anbehav.2011.05.008. 53164955 .
88. Bagemihl (1999) pages 666, 660
89. Bagemihl (1999) pages 661–2
90. Bagemihl (1999) page 661
91. Lecomte. C.. Thibout. E.. Pierre. D.. Auger. J.. 1998-04-01. Transfer, Perception, and Activity of Male Pheromone of Acrolepiopsis assectella with Special Reference to Conspecific Male Sexual Inhibition. Journal of Chemical Ecology. en. 24. 4. 655–671. 10.1023/a:1022390102631. 10001697. 0098-0331.
92. Bagemihl (1999), pages 661–2
93. Bagemihl (1999), page 659
94. 10.1007/BF01960660. Baker T. Variations in male oriental fruit moth courtship patterns due to male competition. Experientia. 39. 1. 112–114 . January 15, 1983 . 99995 .
95. 10.1007/BF00980201. Hall D, Cork A, Lester R, Nesbitt B, Zagatti P. Sex pheromones of rice moth, Corcyra cephalonica Stainton. Journal of Chemical Ecology. 13. 7. 1575–1589. July 1, 1987 . 24302328 . 1987JCEco..13.1575H . 11253920.
96. Bagemihl (1999), page 661
97. 10.1093/AESA/72.4.544. Palaniswamy P, Seabrook W, Ross R. Precopulatory behavior of males and perception of a potential male pheromone in spruce budworm. Annals of the Entomological Society of America. 72. 4. 544–551 . July 15, 1979 . 84678927 .
98. 10.4039/ENT107967-9. Sanders C. Factors affecting adult emergence and mating behavior of the eastern spruce budworm, Choristoneura fumiferana (Lepidoptera: Totricidae). Canadian Entomologist. 107. 9. 967–77 . September 1, 1975 . 86477332 .
99. Chaudhuri A, Sinha A. A male-male pairing in the tropical tasar silkmoth, Antheraea mylitta (Saturniidae). News of the Lepidopterists' Society. 39. 3. 39 . July 15, 1997.
100. 10.1111/j.1439-0418.1990.tb00062.x. Schmieder-Wenzel C, Schruft G. Courtship behaviour of the European Grape Berry Moth, Eupoecilia ambiguella Hb. (Lep., Tortricidae) in regard to pheromonal and tactile stimuli . Journal of Applied Entomology . 109 . 1–5 . 341–346 . January 12, 1990 . 86384073 .
101. Zimmer (2000)
102. Bagemihl (1999) page 657
103. Bagemihl (1999) pages 236, 704, 713
104. Bagemihl (1999), pages 668, 667