Cited by 87Animal Behavior and ReproductionPrimate Behavior and EcologyWildlife Ecology and Conservation

Evolution of ungulate mating systems: Integrating social and environmental factors

Open access
Bowyer, R. Terry, McCullough, Dale R., Rachlow, Janet L., Ciuti, Simone, Whiting, Jericho C.·2020·Ecology and Evolution

Abstract

Ungulates exhibit diverse mating systems that range from monogamous pair territories to highly polygynous leks. We review mating systems and behaviors across ungulates and offer a new approach synthesizing how interacting factors may shape those mating systems. Variability exists in mating systems among and within species of ungulates and likely is affected by predation risk, availability of resources (food and mates), habitat structure, and sociality. Ungulate mating systems may be labile as a consequence of the varying strength of those interacting factors. In addition, degree of polygyny and sexual dimorphism in size are associated with the evolution of mating systems. Neither male–male combat nor paternal care, however, can completely explain differences in sexual size dimorphism for ungulates, a necessary component in understanding the development of some mating systems. Whatever the evolutionary pathway, sexual segregation limits paternal care allowing more intense male–male competition. Selection of habitat structure, because it modifies risk of predation, is a major determinant of sociality for ungulates. Likewise, ruggedness and steepness of terrain limit the types of mating systems that can occur because of limitations in group size and cohesiveness, as well as the ability of males to herd even small groups of females effectively. The quality and defensibility of resources affect mating systems, as does the defensibility of females. Population density of females also may be a critical determinant of the types of mating systems that develop. Size of groups likewise constrains the types of mating tactics that males can employ. Our aim was to use those relationships to create a broad conceptual model that predicts how various environmental and social factors interact to structure mating systems in ungulates. This model provides a useful framework for future tests of the roles of both ecological and social conditions in influencing the social systems of ungulates.

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Integrity signals

Registry

Crossref · 10.1002/ece3.6246

Cites

1 of 1 references verified0 citation errors · 0 not checkable

Statistics

Claims the paper makes

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

The self-check needs the paper’s full text.

Cites

Flagged first, then by citations

Research works printed but not linked: 204

  1. Airst, J., & Lingle, S. (2019). Courtship strategies of white‐tailed deer and mule deer when living in sympatry. Behaviour, 156(3–4), 307– 330.
  2. Alexander, R. D. (1974). The evolution of social behavior. Annual Review of Ecology and Systematics, 5(1), 325– 383.
  3. Altmann, M. (1959). Group dynamics in Wyoming moose during the rutting season. Journal of Mammalogy, 40(3), 420– 424.
  4. Apollonio, M. (1989). Lekking in fallow deer: Just a matter of density? Ethology, Ecology and Evolution, 1(3), 291– 294.
  5. Apollonio, M., Brivio, F., Rossi, I., Bassano, B., & Grignolio, S. (2013). Consequences of snowy winters on male mating strategies and reproduction in a mountain ungulate. Behavioural Processes, 98, 44– 50. 10.1016/j.beproc.2013.05.001 23669063 · DOI
  6. Apollonio, M., De Cena, F., Bongi, P., & Ciuti, S. (2014). Female preference and predation risk models can explain the maintenance of a fallow deer (Dama dama) lek and its 'handy' location. PLoS ONE, 9(3), e89852 10.1371/journal.pone.0089852 24599036 PMC3943860 · DOI
  7. Apollonio, M., Festa‐Bianchet, M., & Mari, F. (1989). Correlates of copulatory success in a fallow deer lek. Behavioral Ecology and Sociobiology, 25(2), 89– 97. 10.1007/bf00302925 · DOI
  8. Apollonio, M., Festa‐Bianchet, M., Mari, F., Mattioli, S., & Sarno, B. (1992). To lek or not to lek: Mating strategies of male fallow deer. Behavioral Ecology, 3(1), 25– 31. 10.1093/beheco/3.1.25 · DOI
  9. Apollonio, M., Festa‐Bianchet, M., Mari, F., & Riva, M. (1990). Site‐specific asymmetries in male copulatory success in a fallow deer lek. Animal Behaviour, 39(2), 205– 212. 10.1016/s0003-3472(05)80864-4 · DOI
  10. Atwood, T. C., Gese, E. M., & Kunkel, K. E. (2009). Spatial partitioning of predation risk in a multiple predator‐multiple prey system. Journal of Wildlife Management, 73(6), 876– 884.
  11. Balmford, A., Albon, S., & Blakeman, S. (1992). Correlates of male mating success and female choice in a lek‐breeding antelope. Behavioral Ecology, 3(2), 112– 123. 10.1093/beheco/3.2.112 · DOI
  12. Balmford, A., Bartos, L., Brotherton, P., Herrmann, H., Lancingerova, J., Mika, J., & Zeeb, U. (1993). When to stop lekking–density‐related variation in the rutting behavior of sika deer. Journal of Zoology, 231(4), 652– 656. 10.1111/j.1469-7998.1993.tb01946.x · DOI
  13. Balmford, A., Deutsch, J. C., Nefdt, R. J. C., & Clutton‐Brock, T. (1993). Testing hotspot models of lek evolution: Data from three species of ungulates. Behavioral Ecology and Sociobiology, 33(1), 57– 65. 10.1007/bf00164347 · DOI
  14. Barboza, P. S., & Bowyer, R. T. (2000). Sexual segregation in dimorphic deer: A new gastrocentric hypothesis. Journal of Mammalogy, 81(2), 473– 489. 10.1644/1545-1542(2000)081<0473:SSIDDA>2.0.CO;2 · DOI · Paper page
  15. Barten, N. L., Bowyer, R. T., & Jenkins, K. J. (2001). Habitat use by female caribou: Tradeoffs associated with parturition. Journal of Wildlife Management, 5(1), 77– 92.
  16. Baskin, L., & Danell, K. (2003). Ecology of ungulates: A handbook of species in eastern Europe and northern and central Asia (434 pp.). Berlin, Germany: Springer.
  17. Beehler, B. M., & Foster, M. S. (1988). Hotshots, hotspots, and female preference in the organization of lek mating systems. The American Naturalist, 131(2), 203– 219. 10.1086/284786 · DOI
  18. Belovsky, G. E. (1981). Food plant selection by a generalist herbivore: The moose. Ecology, 62(4), 1020– 1030. 10.2307/1937001 · DOI
  19. Berger, J. (1978). Group size, foraging, and antipredator ploys: An analysis of bighorn sheep decisions. Behavioral Ecology and Sociobiology, 4(1), 91– 99. 10.1007/bf00302563 · DOI
  20. Berger, J. (1979). Predator harassment as a defensive strategy in ungulates. American Midland Naturalist, 102(1), 197– 199. 10.2307/2425087 · DOI
  21. Berger, J. (1986). Wild horses of the Great Basin: Social competition and population size (326 pp.). Chicago, IL: The University of Chicago Press.
  22. Berger, J. (1989). Female reproductive potential and its apparent evaluation by male mammals. Journal of Mammalogy, 70(2), 347– 358. 10.2307/1381518 · DOI
  23. Berger, J. (1991). Pregnancy incentives, predation constraints, and habitat shifts: Experimental and field evidence for wild bighorn sheep. Animal Behaviour, 41(1), 61– 77. 10.1016/s0003-3472(05)80503-2 · DOI
  24. Berger, J., & Cunningham, C. (1994). Bison: Mating and conservation in small populations (330 pp.). New York, NY: Columbia University Press.
  25. Bergerud, A. T. (1974). Rutting behaviour of Newfoundland caribou In Geist V., & Walther F. R. (Eds.), The behaviour of ungulates and its relation to management (pp. 395– 435). Morges, Switzerland: IUCN.
  26. Bertram, B. C. R. (1978). Living in groups: Predators and prey In Krebs J. R., & Davies N. B. (Eds.), Behavioural ecology: An evolutional approach (pp. 64– 96). Sunderland, MA: Sinauer Associates.
  27. Bleich, V. C. (1999). Mountain sheep and coyotes: Patterns of predator evasion in a mountain ungulate. Journal of Mammalogy, 80(1), 283– 289. 10.2307/1383228 · DOI
  28. Body, G., Weladji, R. D., Holand, Ø., & Nieminen, M. (2014). Highly competitive reindeer males control female behavior during the rut. PLoS ONE, 9(4), e95618. 24759701 PMC3997419
  29. Bowling, A. T., & Touchberry, R. W. (1990). Parentage of Great Basin feral horses. Journal of Wildlife Management, 54(3), 424– 429. 10.2307/3809652 · DOI
  30. Bowyer, R. T. (1981). Activity, movement, and distribution of Roosevelt elk during rut. Journal of Mammalogy, 62(3), 574– 582. 10.2307/1380404 · DOI
  31. Bowyer, R. T. (1987). Coyote group size relative to predation on mule deer. Mammalia, 51(4), 515– 526. 10.1515/mamm.1987.51.4.515 · DOI
  32. Bowyer, R. T. (2004). Sexual segregation in ruminants: Definitions, hypotheses, and implications for conservation and management. Journal of Mammalogy, 85(6), 1039– 1052. 10.1644/bbl-002.1 · DOI
  33. Bowyer, R. T., Bleich, V. C., Manteca, X., Whiting, J. C., & Stewart, K. M. (2007). Sociality, mate choice, and timing of mating in American bison (Bison bison): Effects of large males. Ethology, 113(11), 1048– 1060. 10.1111/j.1439-0310.2007.01411.x · DOI
  34. Bowyer, R. T., Bleich, V. C., Stewart, K. M., Whiting, J. C., & Monteith, K. L. (2014). Density dependence in ungulates: A review of causes, and concepts with some clarifications. California Fish and Game, 100(3), 550– 572.
  35. Bowyer, R. T., & Kie, J. G. (2006). Effects of scale on interpreting life‐history characteristics of ungulates and carnivores. Diversity and Distributions, 12(3), 244– 257. 10.1111/j.1366-9516.2006.00247.x · DOI
  36. Bowyer, R. T., & Kitchen, D. W. (1987). Sex and age class differences in vocalization of Roosevelt elk during rut. American Midland Naturalist, 118(2), 225– 235. 10.2307/2425779 · DOI
  37. Bowyer, R. T., Rachlow, J. L., Stewart, K. M., & Van Ballenberghe, V. (2011). Vocalizations by Alaskan moose: Female incitation of male aggression. Behavioral Ecology and Sociobiology, 65(12), 2251– 2260. 10.1007/s00265-011-1234-y · DOI
  38. Bowyer, R. T., Stewart, K. M., Kie, J. G., & Gasaway, W. C. (2001). Fluctuating asymmetry in antlers of Alaskan moose: Size matters. Journal of Mammalogy, 82(3), 814– 824.
  39. Boyce, M. W. (1989). The Jackson elk herd: Intensive wildlife management in North America (306 pp.). New York, NY: Cambridge University Press.
  40. Boyd, L., Scorolli, A., Nowzari, H., & Bouskila, A. (2016). Social organization of wild equids In Ransom J. I., & Kaczensky P. (Eds.), Wild equids: Ecology, management, conservation (pp. 7– 22). Baltimore, MD: Johns Hopkins University Press.
  41. Bradbury, J. W. (1981). The evolution of leks In Alexander R. D., & Tinkle D. W. (Eds.), Natural selection and social behavior (pp. 138– 169). New York, NY: Chiron Press.
  42. Bradbury, J. W., & Gibson, R. M. (1983). Leks and mate choice In Bateson P. (Ed.), Mate choice (pp. 109– 138). Cambridge, UK: Cambridge University Press.
  43. Bradbury, J. W., Gibson, R., & Tsai, I. M. (1986). Hotspots and the dispersion of leks. Animal Behaviour, 34(6), 1694– 1709. 10.1016/s0003-3472(86)80257-3 · DOI
  44. Briefer, E. F., Farrell, M. E., Hayden, T. J., & McElligott, A. G. (2013). Fallow deer polyandry is related to fertilization insurance. Behavioral Ecology and Sociobiology, 67(4), 657– 665. 10.1007/s00265-013-1485-x · DOI
  45. Bro‐Jørgensen, J. (2002). Overt female mate competition and preference for central males in a lekking antelope. Proceedings of the National Academy of Sciences of the United States of America, 99(14), 9290– 9293. 10.1073/pnas.142125899 12089329 PMC123133 · DOI
  46. Bro‐Jørgensen, J. (2007). The intensity of sexual selection predicts weapon size in male bovids. Evolution, 61(6), 1316– 1326. 10.1111/j.1558-5646.2007.00111.x 17542842 · DOI
  47. Bro‐Jørgensen, J. (2011a). Intra‐ and intersexual conflicts and cooperation in the evolution of mating strategies: Lessons learnt from ungulates. Evolutionary Biology, 38(1), 28– 41.
  48. Bro‐Jørgensen, J. (2011b). Queuing in space and time reduces the lek paradox on an antelope lek. Evolutionary Ecology, 25(6), 1385– 1395. 10.1007/s10682-011-9523-5 · DOI
  49. Brotherton, P. N. M., & Manser, M. B. (1997). Female dispersion and the evolution of monogamy in the dik‐dik. Animal Behaviour, 54, 1413– 1424. 9794769
  50. Buechner, H. K., & Roth, H. D. (1974). Lek system in Uganda kob antelope. American Zoologist, 14(1), 145– 162.
  51. Burt, W. H. (1943). Territoriality and home range concepts as applied to mammals. Journal of Mammalogy, 24(3), 346– 352.
  52. Byers, J. A., & Kitchen, D. W. (1988). Mating system shift in a pronghorn population. Behavioral Ecology and Sociobiology, 22(5), 355– 360.
  53. Byers, J. A., Moodie, J. D., & Hall, N. (1994). Pronghorn females choose vigorous mates. Animal Behaviour, 47(1), 33– 43. 10.1006/anbe.1994.1005 · DOI
  54. Carling, M. D., Wiseman, P. A., & Byers, J. A. (2003). Microsatellite analysis reveals multiple paternity in a population of wild pronghorn antelopes (Antilocapra americana). Journal of Mammalogy, 84(4), 1237– 1243. 10.1644/brb-116 · DOI
  55. Caro, T. M. (1986). The functions of stotting: A review of the hypotheses. Animal Behaviour, 34(3), 649– 662. 10.1016/s0003-3472(86)80051-3 · DOI
  56. Caro, T. M. (2005). Antipredator defenses in birds and mammals (592 pp.). Chicago, IL: The University of Chicago Press.
  57. Caro, T. M., & Bateson, P. (1986). Organization and ontogeny of alternative tactics. Animal Behaviour, 34(5), 1483– 1499. 10.1016/s0003-3472(86)80219-6 · DOI
  58. Carranza, J. (2000). Environmental effects on the evolution of mating systems in endotherms In Apollonio M., Festa‐Bianchet M., & Mainardi D. (Eds.), Vertebrate mating systems (pp. 106– 139). River Edge, NJ: World Scientific Publishing Co.
  59. Carranza, J., Alvarez, F., & Redondo, T. (1990). Territoriality as a mating strategy in red deer. Animal Behaviour, 40(1), 79– 88. 10.1016/s0003-3472(05)80667-0 · DOI
  60. Carranza, J., & Valencia, J. (1999). Red deer females collect on male clumps at mating areas. Behavioral Ecology, 10(5), 525– 532. 10.1093/beheco/10.5.525 · DOI
  61. Cassini, M., Borgnia, M., Arzamendia, Y., Benítez, V., & Vilá, B. (2009). Sociality, foraging and habitat use by vicuña In Gordon I. J. (Ed.), The vicuña: The theory and practice on community‐based wildlife management (pp. 35– 48). New York, NY: Springer.
  62. Caughley, G., & Krebs, C. J. (1983). Are big mammals simply little mammals writ large. Oecologia, 59(1), 7– 17. 10.1007/bf00388066 25024141 · DOI
  63. Ciuti, S., & Apollonio, M. (2016). Reproductive timing in a lekking mammal: Male fallow deer getting ready for female estrus. Behavioral Ecology, 27(5), 1522– 1532. 10.1093/beheco/arw076 · DOI
  64. Ciuti, S., De Cena, F., Bongi, P., & Apollonio, M. (2011). Benefits of a risky life for fallow deer bucks (Dama dama) aspiring to patrol a lek territory. Behaviour, 148(4), 435– 460. 10.1163/000579511x563981 · DOI
  65. Clutton‐Brock, T. H. (1989). Mammalian mating systems. Proceedings of the Royal Society Series B‐Biological Sciences, 236(1285), 339– 372. 10.1098/rspb.1989.0027 2567517 · DOI
  66. Clutton‐Brock, T. H. (2016). Mammal societies (760 pp.). West Sussex, UK: John Wiley & Sons.
  67. Clutton‐Brock, T. H., Deutsch, J. C., & Nefdt, R. J. C. (1993). The evolution of ungulate leks. Animal Behaviour, 46(6), 1121– 1138. 10.1006/anbe.1993.1302 · DOI
  68. Clutton‐Brock, T. H., Green, D., Hiraiwa‐Hasegawa, M., & Albon, S. D. (1988). Passing the buck: Resource defense, lek breeding and mate choice in fallow deer. Behavioral Ecology and Sociobiology, 23(5), 281– 296. 10.1007/bf00300575 · DOI
  69. Clutton‐Brock, T. H., Guinness, F. E., & Albon, S. D. (1982). Red deer: Behavior and ecology of two sexes (400 pp.). Chicago, IL: University of Chicago Press.
  70. Clutton‐Brock, T. H., & Huchard, E. (2013). Social competition and its consequences in female mammals. Journal of Zoology, 289(3), 151– 171.
  71. Clutton‐Brock, T. H., & McAuliffe, K. (2009). Female mate choice in mammals. Quarterly Review of Biology, 84(1), 3– 27. 19326786
  72. Clutton‐Brock, T. H., Price, O. F., & Maccoll, A. D. C. (1992). Mate retention, harassment, and the evolution of ungulate leks. Behavioral Ecology, 3(3), 234– 242. 10.1093/beheco/3.3.234 · DOI
  73. Coltman, D. W., Festa‐Bianchet, M., Jorgenson, J. T., & Strobeck, C. (2002). Age‐dependent sexual selection in bighorn rams. Proceedings of the Royal Society of London Series B: Biological Sciences, 269(1487), 165– 172. 11798432 PMC1690873
  74. Corlatti, L., Caroli, M., Pietrocini, V., & Lovari, S. (2013). Rutting behaviour of territorial and nonterritorial male chamois: Is there a home advantage? Behavioural Processes, 92, 118– 124. 10.1016/j.beproc.2012.11.008 23182914 · DOI
  75. Debeffe, L., Focardi, S., Bonenfant, C., Hewston, A. J. M., Morellet, N., Vampé, C., … Cagnacci, F. (2014). A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic. Oecologia, 176(2), 431– 443. 25027183
  76. Dehn, M. M. (1990). Vigilance for predators: Detection and dilution effects. Behavioral Ecology and Sociobiology, 26(5), 337– 342.
  77. Deutsch, J. C. (1994). Lekking by default: Female habitat preferences and male strategies in Uganda kob. Journal of Animal Ecology, 63(1), 101– 115. 10.2307/5587 · DOI
  78. DeYoung, R. W., Demarais, S., Gonzales, R. A., Honeycutt, R. L., & Gee, K. L. (2002). Multiple paternity in white‐tailed deer (Odocoileus virginianus) revealed by DNA microsatellites. Journal of Mammalogy, 83(3), 884– 892. 10.1644/1545-1542(2002)083<0884:mpiwtd>2.0.co;2 · DOI
  79. Endo, A. (2009). Variation in mating behavior of sika deer: Mating behavior of sika deer on Nozaki Island In McCullough D. R., Takatsuki S., & Kaji K. (Eds.), Sika deer: Biology and management of native and introduced populations (pp. 285– 296). New York, NY: Springer.
  80. Endo, A., & Doi, T. (2002). Multiple copulations and post‐copulatory guarding in a free‐living population of sika deer (Cervus nippon). Ethology, 108(8), 739– 747. 10.1046/j.1439-0310.2002.00803.x · DOI
  81. Espmark, Y. (1964). Rutting behaviour in reindeer (Rangifer tarandus L.). Animal Behaviour, 12(1), 159– 160. 10.1016/0003-3472(64)90117-4 · DOI
  82. Estes, R. (1974). Social organization of the African Bovidae In Walther F. R., & Geist V. (Eds.), The behaviour of ungulates and its relation to management (pp. 166– 205). Morges, Switzerland: IUCN.
  83. Fairbanks, W. S. (1994). Dominance, age and aggression among female pronghorn, Antilocapra americana (Family, Antilocapridae). Ethology, 97(4), 278– 293.
  84. Feh, C. (1999). Alliances and reproductive success in Camargue stallions. Animal Behaviour, 57(3), 705– 713. 10.1006/anbe.1998.1009 10196062 · DOI
  85. Feist, J. D., & McCullough, D. R. (1976). Behavior patterns and communication in feral horses. Zeitschrift für Tierpsychologie, 41(4), 337– 371. 983427
  86. Feldhamer, G. A., Drickamer, L. C., Vessey, S. H., Merritt, J. F., & Krajewski, C. (2015). Mammalogy: Adaptation, diversity, ecology (p. 747). Baltimore, MD: Johns Hopkins University Press.
  87. Festa‐Bianchet, M., & Côté, S. D. (2008). Mountain goats: Ecology, behavior, and conservation of an alpine ungulate (290 pp.). Washington, DC: Island Press.
  88. Franklin, W. L. (1983). Contrasting sociologies of South American's wild camelids: The vicuña and guanaco In Eisenberg J. F., & Kleiman D. G. (Eds.), Special publication of the American Society of Mammalogists 7 (pp. 573– 629). Stillwater, OK.
  89. Fryxell, J. M. (1987). Lek breeding and territorial aggression in white‐eared kob. Ethology, 75(3), 211– 220.
  90. Geist, V. (1966). The evolution of horn‐like organs. Behaviour, 27(3), 175– 214.
  91. Geist, V. (1968). On the interrelation of external appearance, social behavior and social structure of mountain sheep. Zeitschrift für Tierpsychologie, 25, 199– 215. 5684151
  92. Geist, V. (1971). Mountain sheep: A study in behavior and evolution (383 pp.). Chicago, IL: The University of Chicago Press.
  93. Geist, V. (1974). On the relationship of social evolution and ecology in ungulates. Annual Review of Ecology and Systematics, 8(1), 193– 207.
  94. Gosling, L. M. (1986a). Biased sex ratios in stressed mammals. The American Naturalist, 127(6), 893– 896. 10.1086/284532 · DOI
  95. Gosling, L. M. (1986b). The evolution of the mating strategies in male antelopes In Rubenstein D. I., & Wrangham R. W. (Eds.), Ecological aspects of social evolution: Birds and mammals (pp. 244– 281). Princeton, NJ: Princeton University Press.
  96. Gosling, L. M. (1991). The alternative mating strategies of male topi, Damaliscus lunatus. Applied Animal Behaviour Science, 29(1–4), 107– 119. 10.1016/0168-1591(91)90240-x · DOI
  97. Gosling, L. M., & Petrie, M. (1990). Lekking in topi: A consequence of satellite behaviour by small males at hotspots. Animal Behaviour, 40(2), 272– 287. 10.1016/s0003-3472(05)80922-4 · DOI
  98. Grignolio, S., Rossi, I., Bassano, B., & Apollonio, M. (2007). Predation risk as a factor affecting sexual segregation in alpine ibex. Journal of Mammalogy, 88(6), 1488– 1497.
  99. Gross, M. R. (1996). Alternative reproductive strategies and tactics: Diversity within sexes. Trends in Ecology and Evolution, 11(2), 92– 98. 10.1016/0169-5347(96)81050-0 21237769 · DOI
  100. Grovenburg, T. W., Jenks, J. A., Jacques, C. N., Klaver, R. W., & Swanson, C. C. (2009). Aggressive defensive behavior by free‐ranging white‐tailed deer. Journal of Mammalogy, 90(5), 1218– 1223. 10.1644/08-mamm-a-360.1 · DOI
  101. Guernsey, N. C., Lohse, K. A., & Bowyer, R. T. (2015). Rates of decomposition and nutrient release of herbivore inputs driven by microhabitat characteristics. Ecological Research, 30, 951– 961. 10.1007/s11284-015-1296-9 · DOI
  102. Gunn, A. (1992). Differences in the sex and age composition of two muskox populations and implications for male breeding strategies. Rangifer, 12(1), 17– 19.
  103. Hass, C. C., & Jenni, D. A. (1991). Structure and ontogeny of dominance relationships among bighorn rams. Canadian Journal of Zoology, 69(2), 471– 476.
  104. Hirth, D. H., & McCullough, D. R. (1977). Evolution of alarm signals in ungulates with special reference to white‐tailed deer. The American Naturalist, 111(977), 31– 42. 10.1086/283136 · DOI
  105. Hogg, J. T. (1984). Mating in bighorn sheep: Multiple creative male strategies. Science, 225(4661), 526– 529. 6539948
  106. Hogg, J. T. (1988). Copulatory tactics in relation to sperm competition in Rocky Mountain bighorn sheep. Behavioral Ecology and Sociobiology, 22(1), 49– 59.
  107. Hogg, J. T. (2000). Mating systems and conservation at large spatial scales In Apollonio M., Festa‐Bianchet M., & Mainardi D. (Eds.), Vertebrate mating systems (pp. 214– 252). River Edge, NJ: World Scientific Publishing Co.
  108. Hogg, J. T., & Forbes, S. H. (1997). Mating in bighorn sheep: Frequent male reproduction via a high‐risk “unconventional” tactic. Behavioral Ecology and Sociobiology, 41(1), 33– 48.
  109. Höglund, J., & Alatalo, R. V. (1995). Leks (264 pp.). Princeton, NJ: Princeton University Press.
  110. Ihl, C., & Bowyer, R. T. (2011). Leadership in mixed‐sex groups of muskoxen during the snow‐free season. Journal of Mammalogy, 92(4), 819– 827. 10.1644/10-mamm-a-109.1 · DOI
  111. Isvaran, K. (2005). Variation in male mating behaviour within ungulate populations: Patterns and processes. Current Science, 89(7), 1192– 1199.
  112. Isvaran, K., & Jhala, Y. V. (2000). Variation in lekking costs in blackbuck (Antilope cervicapra): Relationship to lek‐territory location and female mating patterns. Behaviour, 137(5), 547– 563.
  113. Isvaran, K., & Ponkshe, A. (2013). How general is a female mating preference for clustered males in lekking species? A meta‐analysis Animal Behaviour, 86(2), 417– 425. 10.1016/j.anbehav.2013.05.036 · DOI
  114. Janis, C. M. (1982). Evolution of horns in ungulates: Ecology and paleoecology. Biological Reviews of the Cambridge Philosophical Society, 57(2), 261– 318.
  115. Jarman, P. J. (1974). Social organisation of antelope in relation to their ecology. Behaviour, 48(1–4), 215– 267. 10.1163/156853974x00345 · DOI
  116. Jarman, P. J. (1983). Mating system and sexual dimorphism in large, terrestrial mammalian herbivores. Biology Review, 58(4), 485– 520.
  117. Jhala, Y. V., & Isvaran, K. (2016). Behavioural ecology of a grassland antelope, the blackbuck Antilope cervicapra: linking habitat, ecology and behaviour In Ahrestani F. S., & Sankaran M. (Eds.) The ecology of large herbivores in South and Southeast Asia. Ecological Studies (vol. 225, pp. 151– 176). New York, NY: Springer.
  118. Kie, J. G., Bowyer, R. T., Nicholson, M. C., Boroski, B. B., & Loft, E. R. (2002). Landscape heterogeneity at differing scales: Effects on spatial distribution of mule deer. Ecology, 83(2), 530– 544. 10.2307/2680033 · DOI
  119. Kie, J. G., Johnson, B. K., Noyes, J. H., Williams, C. L., Dick, B. L., Rhodes, O. E., … Bowyer, R. T. (2013). Reproduction in North American elk Cervus elaphus: Paternity of calves sired by males of mixed age classes. Wildlife Biology, 19(3), 302– 310.
  120. Klingel, H. (1974). A comparison of the social behaviour of the Equidae In Geist V., & Walther F. R. (Eds.), The behaviour of ungulates and its relation to management (pp. 124– 132). Morges, Switzerland: IUCN.
  121. Klingel, H. (1978). Sociobiology of zebras and antelopes. Recherche, 9(86), 112– 120.
  122. Koford, C. B. (1957). The vicuna and the puna. Ecological Monographs, 27(2), 153– 219. 10.2307/1948574 · DOI
  123. Kohl, M. T., Ruth, T. K., Metz, D. T., Smith, D. W., White, P. J., & MacNulty, D. R. (2019). Do prey select for vacant hunting domains to minimize a multi‐predator threat? Ecology Letters, 2019, 10.1111/ele13319 31373137 · DOI
  124. Kokko, H. (2007). Modelling for field biologists and other interesting people (242 pp.). Cambridge, UK: Cambridge University Press.
  125. Komers, P. E. (1996). Obligate monogamy without paternal care in Kirk's dikdik. Animal Behaviour, 51(1), 131– 140. 10.1006/anbe.1996.0011 · DOI
  126. Kruuk, H. (1972). The spotted hyena: A study of predation and social behavior (388 pp.). Chicago, IL: The University of Chicago Press.
  127. Kruuk, L. E. B., Slate, J., Pemberton, J. M., Brotherstone, S., Guinness, F., & Clutton‐Brock, T. (2002). Antler size in red deer: Heritability and selection but no evolution. Evolution, 56(8), 1683– 1695. 12353761
  128. Kucera, T. E. (1976). Social behavior and breeding system of the desert mule deer. Journal of Mammalogy, 40(3), 420– 424.
  129. Langbein, J., & Thirgood, S. J. (1989). Variation in mating systems of fallow deer (Dama dama) in relation to ecology. Ethology, 83(3), 195– 214.
  130. Lent, P. C. (1965). Rutting behaviour in barren‐ground caribou population. Animal Behaviour, 13(2–3), 259– 264. 10.1016/0003-3472(65)90044-8 5835842 · DOI
  131. Leuthold, W. (1977). African ungulates: A comparative review of their ethology and behavioral ecology (p. 310). New York, NY: Springer.
  132. Linklater, W. L., & Cameron, E. Z. (2000). Tests for cooperative behaviour between stallions. Animal Behaviour, 60(6), 731– 743. 10.1006/anbe.2000.1525 11124871 · DOI
  133. Linklater, W. L., Cameron, E. Z., Stafford, K. J., & Minot, E. O. (2013). Removal experiments indicate that subordinate stallions are not helpers. Behavioural Processes, 94, 1– 4. 10.1016/j.beproc.2013.02.005 23428706 · DOI
  134. Loison, A., Gaillard, J. M., Pelabon, C., & Yoccoz, N. G. (1999). What factors shape sexual size dimorphism in ungulates? Evolutionary Ecology Research, 1(5), 611– 633.
  135. Lott, D. F. (1974). Sexual and aggressive behavior of adult male American bison (Bison bison) In Geist V., & Walther F. R. (Eds.), The behaviour of ungulates and its relation to management (pp. 382– 394). Morges, Switzerland: IUCN.
  136. Lott, D. F. (1991). Intraspecific variation in the social systems of wild vertebrates (256 pp.). Cambridge, UK: Cambridge University Press.
  137. Lovari, S., & Ale, S. B. (2001). Are there multiple mating strategies in blue sheep? Behavioural Processes, 53(1–2), 131– 135. 10.1016/s0376-6357(00)00134-0 11255000 · DOI
  138. Lukas, D., & Clutton‐Brock, T. H. (2013). The evolution of social monogamy in mammals. Science, 341(6145), 526– 530. 10.1126/science.1238677 23896459 · DOI · Paper page
  139. Lundrigan, B. (1996). Morphology of horns and fighting behavior in the family Bovidae. Journal of Mammalogy, 77(2), 462– 475.
  140. Maher, C. R., & Lott, D. F. (2000). A review of ecological determinants of territoriality within vertebrate species. American Midland Naturalist, 143(1), 1– 29. 10.1674/0003-0031(2000)143[0001:aroedo]2.0.co;2 · DOI
  141. Mainguy, J., Côté, S. D., Cardinal, E., & Houle, M. (2008). Mating tactics and mate choice in relation to age and social rank in male mountain goats. Journal of Mammalogy, 89(3), 626– 635. 10.1644/07-mamm-a-234r.1 · DOI
  142. Markussen, S. S., Herfindall, I., Loison, K. A., Solberg, E. J., Haanes, H., Røed, M. H., & Sæther, B.‐E. (2019). Determinations of age at first reproduction and life‐time breeding success revealed by full paternity assignment in a male ungulate. Oikos, 128(3), 328– 337.
  143. May, R. M. (2001). Stability and complexity in model ecosystems (304 pp.). Princeton, NJ: Princeton University Press.
  144. McCullough, D. R. (1979). The George Reserve deer herd: Population ecology of a K‐selected species (271 pp.). Ann Arbor, MI: The University of Michigan Press.
  145. McCullough, D. R., Pei, K. C. J., & Wang, Y. (2000). Home range, activity patterns, and habitat relations of Reeves' muntjacs in Taiwan. Journal of Wildlife Management, 64(2), 430– 441. 10.2307/3803241 · DOI
  146. McDonald, D. B. (2000). Demographic analyses of mating systems In Apollonio M., Festa‐Bianchet M., & Mainardi D. (Eds.), Vertebrate mating systems (pp. 80– 105). River Edge, NJ: World Scientific Publishing Co.
  147. McNab, B. K. (1963). Bioenergetics and determination of home range size. The American Naturalist, 97(894), 133– 140.
  148. McNaughton, S. J. (1979). Grazing as an optimization process: Grass‐ungulate relationships in the Serengeti. The American Naturalist, 113(5), 691– 703.
  149. Mech, L. D. (1970). The wolf: The ecology and behavior of an endangered species (384 pp.). Garden City, NY: The Natural History Press.
  150. Mekonnen, A., Bekele, A., & Balakrishnan, M. (2018). Population ecology of the giant forest hog, Hylochoerus meinertzhageni, in Chebera Churchura National Park, Ethiopia. African Journal of Ecology, 56(2), 272– 278.
  151. Molvar, E. M., & Bowyer, R. T. (1994). Costs and benefits of group living in a recently social ungulate: The Alaskan moose. Journal of Mammalogy, 75(3), 621– 630. 10.2307/1382509 · DOI
  152. Molvar, E. M., Van Ballenberghe, V., & Bowyer, R. T. (1993). Moose herbivory, browse quality, and nutrient cycling in an Alaskan treeline community. Oecologia, 94(4), 472– 479. 28313986
  153. Monteith, K. L., Long, R. A., Bleich, V. C., Heffelfinger, J. R., Krausman, P. R., & Bowyer, R. T. (2013). Effects of harvest, culture, and climate on trends in size of horn‐like structures in trophy ungulates. Wildlife Monographs, 183(1), 1– 28. 10.1002/wmon.1007 · DOI
  154. Monteith, K. L., Long, R. A., Stephenson, T. R., Bleich, V. C., Bowyer, R. T., & LaSharr, T. N. (2018). Horn size and nutrition in mountain sheep: Can ewe handle the truth? The Journal of Wildlife Management, 82(1), 67– 84.
  155. Monteith, K. L., Schmitz, L. E., Jenks, J. A., Delger, J. A., & Bowyer, R. T. (2009). Growth of male white‐tailed deer: Consequences of maternal effects. Journal of Mammalogy, 90(3), 651– 660.
  156. Morton, T. L., Haefner, J. W., Nugala, V., Decino, R. D., & Mendes, L. (1994). The selfish herd revisited: Do simple movement rules reduce relative predation risk? Journal of Theoretical Biology, 167(1), 73– 79. 10.1006/jtbi.1994.1051 · DOI
  157. Myers, P. (1978). Sexual dimorphism in size of vespertilionid bats. The American Naturalist, 112(986), 701– 711.
  158. Mysterud, A., Coulson, T., & Stenseth, N. C. (2002). The role of males in the dynamics of ungulate populations. Journal of Animal Ecology, 71(6), 907– 915.
  159. Mysterud, A., Langvatn, R., & Stenseth, N. C. (2004). Patterns of reproductive effort in male ungulates. Journal of Zoology, 264(2), 209– 215. 10.1017/s0952836904005618 · DOI
  160. Nefdt, R. J. C., & Thirgood, S. J. (1997). Lekking, resource defense, and harassment in two subspecies of lechwe antelope. Behavioral Ecology, 8(1), 1– 9. 10.1093/beheco/8.1.1 · DOI
  161. Newbolt, C. H., Acker, P. K., Newman, T. J., Hoffman, S. T., Ditckoff, S. S., & Steury, T. D. (2017). Factors influencing reproductive success in male white‐tailed deer. Journal of Wildlife Management, 81(2), 206– 217.
  162. Noyes, J. H., Johnson, B. K., Dick, B. L., & Kie, J. G. (2002). Effects of male age and female nutritional condition on elk reproduction. Journal of Wildlife Management, 66(4), 1301– 1307.
  163. Owen‐Smith, R. N. (1971). Territoriality in the white rhinoceros (Ceratotherium simum Burchell). Nature, 231(5301), 294– 296. 4930978
  164. Pelletier, F., & Festa‐Bianchet, M. (2006). Sexual selection and social rank in bighorn rams. Animal Behaviour, 71(3), 649– 655.
  165. Pelletier, F., Hogg, J. T., & Festa‐Bianchet, M. (2006). Male mating effort in a polygynous ungulate. Behavioral Ecology and Sociobiology, 60(5), 645– 654. 10.1007/s00265-006-0208-y · DOI
  166. Pemberton, J. M., Albon, S. D., Guinness, F. E., Clutton‐Brock, T. H., & Dover, G. A. (1992). Behavioral estimates of male mating success tested by DNA fingerprinting in a polygynous mammal. Behavioral Ecology, 3(1), 66– 75. 10.1093/beheco/3.1.66 · DOI
  167. Pérez‐Barbería, F. J., Gordon, I. J., & Pagel, M. (2002). The origins of sexual dimorphism in body size in ungulates. Evolution, 56(6), 1276– 1285. 12144026
  168. Pipia, A., Ciuti, S., Grignolio, S., Luchetti, S., Madau, R., & Apollonio, M. (2009). Effect of predation risk on grouping pattern and whistling behaviour in a wild mouflon Ovis aries population. Acta Theriologica, 54(1), 77– 86. 10.1007/bf03193139 · DOI
  169. Povilitis, A. J. (1983). Social organization and mating strategy of the huemul (Hippocamelus bisulcus). Journal of Mammalogy, 64(1), 156– 158.
  170. Pulliam, H. R., & Caraco, T. (1984). Living in groups: Is there an optimal group size? In Krebs J. R., & Davies N. B. (Eds.), Behavioural ecology: An evolutional approach (pp. 122– 147). Sunderland, MA: Sinauer Associates.
  171. Putman, R. (1988). The natural history of deer (224 pp.). Ithaca, NY: Cornell University Press.
  172. Putman, R., & Flueck, W. T. (2011). Intraspecific variation in biology and ecology of deer: Magnitude and causation. Animal Production Science, 51(4), 277– 291. 10.1071/an10168 · DOI
  173. Rachlow, J. L., Berkeley, E. V., & Berger, J. (1998). Correlates of male mating strategies in white rhinos (Ceratotherium simum). Journal of Mammalogy, 79(4), 1317– 1324. 10.2307/1383023 · DOI
  174. Ralls, K. (1977). Sexual dimorphism in mammals: Avian models and unanswered questions. The American Naturalist, 111(981), 917– 938. 10.1086/283223 · DOI
  175. Reyna‐Hurtago, R., Rojas‐Flores, E., & Tanner, G. W. (2009). Home range and habitat preferences of white‐lipped peccaries (Tayassu pecari) in Calakmul, Campeche, Mexico. Journal of Mammalogy, 90(5), 1199– 1209.
  176. Roberts, G. (1996). Why individual vigilance declines as group size increases. Animal Behaviour, 51(5), 1077– 1086. 10.1006/anbe.1996.0109 · DOI
  177. Rubenstein, D. I. (1978). On predation, competition, and the advantages of group living In Bateson P. P. G., & Klopfer P. H. (Eds.), Perspectives in ethology (pp. 205– 231). New York, NY: Plenum Press.
  178. Rubenstein, D. I. (1986). Ecology and sociality in horses and zebras In Rubenstein D. I., & Wrangham R. W. (Eds.), Ecological aspects of social evolution: Birds and mammals (pp. 282– 302). Princeton, NJ: Princeton University Press.
  179. Rughetti, M., & Festa‐Bianchet, M. (2011). Seasonal changes in sexual size dimorphism in northern chamois. Journal of Zoology, 284(4), 257– 264. 10.1111/j.1469-7998.2011.00800.x · DOI
  180. Schaller, G. B. (1972). The Serengeti lion: A study of predator‐prey relations (480 pp.). Chicago, IL: University of Chicago Press.
  181. Schaller, G. B. (1998). Wildlife of the Tibetan Steppe (383 pp.). Chicago, IL: University of Chicago Press.
  182. Schoener, T. W. (1968). Sizes of feeding territories among birds. Ecology, 49(1), 123– 141.
  183. Schroeder, C. A., Bowyer, R. T., Bleich, V. C., & Stephenson, T. R. (2010). Sexual segregation in Sierra Nevada bighorn sheep, Ovis canadensis sierrae: Ramifications for conservation. Arctic Antarctic and Alpine Research, 42(4), 476– 489. 10.1657/1938.4246.42.4.476 · DOI
  184. Schuster, S. M., & Wade, M. J. (2003). Mating systems and strategies. Monographs in Behavior and Ecology. Princeton, NJ: Princeton University Press.
  185. Sinclair, A. R. E. (1977). The African buffalo: A study of resource limitation of populations (p. 355). Chicago, IL: University of Chicago Press.
  186. Sowls, L. K. (1984). The Peccaries (p. 251). Tucson, AZ: University of Arizona Press.
  187. Spinage, C. A. (1982). A territorial antelope: The Uganda waterbuck (350 pp.). New York, NY: Academic Press.
  188. Stevens, E. F. (1990). Instability of harems of feral horses in relation to season and presence of subordinate stallions. Behaviour, 112(3–4), 149– 161.
  189. Stewart, K. M., Bowyer, R. T., & Weisberg, P. J. (2011). Spatial use of landscapes In Hewitt D. G. (Ed.), Biology and management of white‐tailed deer (pp. 181– 217). Boca Raton, FL: CRC Press.
  190. Stillman, R. A., Deutsch, J. C., Clutton‐Brock, T. H., & Sutherland, W. J. (1996). Black hole models of ungulate lek size and distribution. Animal Behaviour, 52(5), 891– 902. 10.1006/anbe.1996.0237 · DOI
  191. Stockley, P., & Bro‐Jørgensen, J. (2011). Female competition and its evolutionary consequences in mammals. Biological Reviews, 86(2), 341– 366. 20636474
  192. Thirgood, S. J., Robertson, A., Jarvis, A. M., Belbin, S. B., Robertson, D., Nefdt, R. J., & Kamwenshe, B. (1992). Mating system and the ecology of black lechwe (Kobus: Bovidae) in Zambia. Journal of Zoology London, 228(1), 155– 172.
  193. Trivers, R. L. (1972). Parental investment and sexual selection In Campbell B. (Ed.), Sexual selection and the descent of man 1871–1971 (pp. 136– 179). Chicago, IL: Aldine Publishing Company.
  194. Uccheddu, S., Body, G., Weladji, R. B., Holand, Ø., & Nieminen, M. (2015). Foraging competition in larger groups overrides harassment avoidance benefits in female reindeer (Rangifer tarandus). Oecologia, 179(3), 711– 718. 26188521
  195. Vampé, C., Gaillard, J.‐M., Kjellander, P., Mysterud, A., Magnien, P., Delorme, D., … Hewiston, A. J. M. (2007). Antlers size provides an honest signal of male quality in roe deer. The American Naturalist, 169(4), 481– 493. 17273980
  196. Weckerly, F. W. (1998). Sexual‐size dimorphism: Influence of mass and mating systems in the most dimorphic mammals. Journal of Mammalogy, 79(1), 33– 52. 10.2307/1382840 · DOI
  197. Weckerly, F. W. (2017). Population ecology of Roosevelt elk: Conservation and management in Redwood National and State Parks (190 pp.). Reno, NV: University of Nevada Press.
  198. Weladji, R. B., Body, G., Holand, Ø., Meng, X., & Nieminen, M. (2017). Temporal variation in the operational sex ratio and male mating behaviours in reindeer (Rangifer tarandus). Behavioural Processes, 140, 96– 103. 28438692
  199. White, K. S., Testa, J. W., & Berger, J. (2001). Behavioral and ecologic effects of differential predation pressure on moose in Alaska. Journal of Mammalogy, 82(2), 422– 429. 10.1644/1545-1542(2001)082<0422:baeeod>2.0.co;2 · DOI
  200. Whiting, J. C., Bowyer, R. T., & Flinders, J. T. (2008). Young bighorn (Ovis canadensis) males: Can they successfully woo females? Ethology, 114(1), 32– 41.

200 of 204 shown

Cited by

Sorted by citations

Authors
Bowyer, R. T., McCullough, D. R., Rachlow, J. L., Ciuti, S., & Whiting, J. C.
Affiliations
  • University of Alaska Fairbanks (US)
  • University of California, Berkeley (US)
  • University of Idaho (US)
  • University College Dublin (IE)
  • Brigham Young University - Idaho (US)
Corresponding author
Bowyer, R. Terry
Venue
Ecology and Evolution
Publisher
Wiley
Published
2020-04-15
Volume
10
Issue
11
Pages
5160-5178
ISSN
2045-7758 (ISSN-L)
OpenAlex
W3016297730
Cited by
87
References
1
Citations by year
2026: 15 · 2025: 16 · 2024: 15 · 2023: 10 · 2022: 16 · 2021: 11 · 2020: 4
FWCI
1.24
Type
journal article
Language
en
Open access
Locations
  • Ecology and Evolution (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
Topics
  • Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
  • Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
  • Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
Keywords
ungulates, mating systems, polygyny, sexual size dimorphism, predation risk, habitat structure, sociality, male-male competition, paternal care, lek mating, group size, resource availability, female density, mating system evolution, conceptual model

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

What counts as a claim ›How we measure confidence ›How we check this ›

Are you an author, or do you see a problem with this page? Request a review