Behavioral changes associated with a population density decline in the facultatively social red fox

Behavioral Ecology

Volumn 20, Issue 2, 2009, Pages 385-395

  Iossa, Graziella ^a,b   Soulsbury, Carl D ^a   Baker, Philip J ^a   Edwards, Keith J ^a   Harris, Stephen ^a  

^a UNIVERSITY OF BRISTOL   (United Kingdom)

^b UNIVERSITY OF READING   (United Kingdom)

Author keywords

Carnivores; Dispersal; Dominance attainment; Extrapair fertilizations; Multiple paternity; Sociality

Indexed keywords

BEHAVIORAL RESPONSE; CANID; CARNIVORE; DISPERSAL; DOMINANCE; EXTRA-PAIR COPULATION; FITNESS; MULTIPLE BROODING; PATERNITY; PHILOPATRY; POPULATION DECLINE; POPULATION DENSITY; POPULATION STRUCTURE; REPRODUCTIVE STRATEGY; SEX RATIO;

BRISTOL [ENGLAND]; ENGLAND; EURASIA; EUROPE; UNITED KINGDOM; WESTERN EUROPE;

VULPES; VULPES VULPES;

EID: 64949083043     PISSN: 10452249     EISSN: 14657279     Source Type: Journal    
DOI: 10.1093/beheco/arn149     Document Type: Article

References (138)

1. Addessi E, Visalberghi E. 2001. Social facilitation of eating novel food in tufted capuchin monkeys (Cebus apella): input provided by group members and responses affected in the observer. Anim Cogn. 4:297-303.
2. Adret P. 1997. Discrimination of video-images by zebra finches (Taeniopygia guttata): direct evidence from song performance. J Comp Psychol. 111:115-125.
3. Beauchamp G. 2003. Group-size effects on vigilance: a search for mechanisms. Behav Processes. 63:111-121.
4. Beauchamp G, Livoreil B. 1997. The effect of group size on vigilance and feeding rate in spice finches (Lonchura punctulata). Can J Zool. 75:1526-1531.
5. Beauchamp G, Ruxton GD. 2003. Changes in vigilance with group size under scramble competition. Am Nat. 161:672-675.
6. Bednekoff PA. 2003. Testing explanations of the group size effect on vigilance: let's be direct. Behav Processes. 63:135-136.
7. Bolyard KJ, Rowland WJ. 1996. Context-dependent response to red coloration in stickleback. Anim Behav. 52:923-927.
8. Clark CW, Mangel M. 1986. The evolutionary advantages of group foraging. J Theor Biol. 30:45-75.
9. Clark DL, Macedonia JM, Rosenthal GG. 1997. Testing video playback to lizards in the field. Copeia. 2:421-423.
10. Dittrich WH, Lea SEG. 1993. Motion as a natural category for pigeons: generalisation and a feature-positive effect. J Exp Anal Behav. 59:115-129.
11. Elgar MA. 1989. Predator vigilance and group size in mammals and birds: a critical review of the empirical evidence. Biol Rev. 64:13-33.
12. Elgar MA, Catterall CP. 1981. Flocking and predator surveillance in house sparrows: test of an hypothesis. Anim Behav. 29:868-872.
13. Evans CS, Macedonia JM, Marler P. 1993. Effects of apparent size and speed on the response of chickens, Gallus gallus, to computer-generated simulations of aerial predators. Anim Behav. 46:1-11.
14. Gauvin S, Giraldeau LA. 2004. Nutmeg mannikins (Lonchura punctulata) reduce their feeding rates in response to simulated competition. Oecologia. 139:150-156.
15. Giraldeau LA, Hogan JA, Clinchy MJ. 1990. The payoffs to producing and scrounging: what happens when patches are divisible? Ethology. 85:132-146.
16. Grand TC, Dill LM. 1999. The effect of group size on the foraging behaviour of juvenile coho salmon: reduction of predation risk or increased competition? Anim Behav. 58:443-451.
17. Lima SL. 1990. The influence of models on the interpretation of vigilance. In: Bekoff M, Jamieson D, editors. Interpretation and explanation in the study of animal behavior: vol. 2. Explanation, evolution and adaptation. Boulder (CO): Westview Press. p. 246-267.
18. Lima SL. 1995. Back to the basics of anti-predatory vigilance: the group-size effect. Anim Behav. 49:11-20.
19. Lima SL, Dill LM. 1990. Behavioral decisions made under the risk of predation: a review and prospectus. Can J Zool. 68:619-640.
20. Lima SL, Zollner PA, Bednekoff PA. 1999. Predation, scramble competition, and the vigilance group size effect in dark-eyed juncos (Junco hyemalis). Behav Ecol Sociobiol. 46:110-116.
21. McDonald CG, Reimchen TE, Hawryshyn CW. 1995. Nuptial colour loss and signal masking in Gasterosteus: an analysis using video imaging. Behaviour. 132:963-977.
22. McGregor PK. 2000. Playback experiments: design and analysis. Acta Ethol. 3:3-8.
23. McNamara JM, Houston AI. 1992. Evolutionarily stable levels of vigilance as a function of group size. Anim Behav. 43:641-658.
24. McQuoid LM, Galef BG. 1993. Social stimuli influencing feeding behaviour of Burmese fowl: a video analysis. Anim Behav. 46:13-22.
25. Ord TJ, Peters RA, Evans CS, Taylor AJ. 2002. Digital video playback and visual communication in lizards. Anim Behav. 63:879-890.
26. Plimpton EH, Swartz KB, Rosenblum LA. 1981. Responses of juvenile bonnet macaques to social stimuli presented through color videotapes. Dev Psychobiol. 14:109-115.
27. Pulliam HR. 1973. On the advantages of flocking. J Theor Biol. 38:419-429.
28. Randler C. 2005. Coots Fulica atra reduce their vigilance under increased competition. Behav Processes. 68:173-178.
29. Roberts G. 1996. Why individual vigilance declines as group size increases? Anim Behav. 51:1077-1086.
30. Rosenthal GG, Evans CS, Miller WL. 1996. Female preference for dynamic traits in the green swordtail Xiphophorus helleri. Anim Behav. 51:811-820.
31. Valone TJ. 1993. Patch estimation: a cost of group foraging. Oikos. 68:258-266.
32. Vásquez RA, Kacelnik A 2000. Foraging rate versus sociality in the starling Sturnus vulgaris. Proc R Soc Lond B Biol Sci. 267:157-164.
33. Baker PJ, Funk SM, Bruford MW, Harris S. 2004. Polygynandry in a red fox population: implications for the evolution of group living in canids? Behav Ecol. 15:766-778.
34. Baker PJ, Funk SM, Harris S, White PCL. 2000. Flexible spatial organization of urban foxes, Vulpes vulpes, before and during an outbreak of sarcoptic mange. Anim Behav. 59:127-146.
35. Baker PJ, Harris S. 2004. Red foxes: the behavioural ecology of red foxes in urban Bristol. In: Macdonald DW, Sillero-Zubiri C, editors. Biology and conservation of wild canids. Oxford: Oxford University Press. p. 207-216.
36. Baker PJ, Harris S, Robertson CPJ, Saunders G, White PCL. 2001. Differences in the capture rate of cage-trapped red foxes Vulpes vulpes and an evaluation of rabies control measures in Britain. J Appl Ecol. 38:823-835.
37. Baker P, Newman T, Harris S. 2001. Bristol's foxes-40 years of change. Br Wildl. 12:411-417.
38. Baker PJ, Robertson CPJ, Funk SM, Harris S. 1998. Potential fitness benefits of group living in the red fox, Vulpes vulpes. Anim Behav. 56:1411-1424.
39. Bekoff M, Diamond J, Mitton JB. 1981. Life-history patterns and sociality in canids: body size, reproduction, and behavior. Oecologia. 50:386-390.
40. Bellemain E, Swenson JE, Taberlet P. 2006. Mating strategies in relation to sexually selected infanticide in a non-social carnivore: the brown bear. Ethology. 112:238-246.
41. Cant MA. 1998. A model for the evolution of reproductive skew without reproductive suppression. Anim Behav. 55:163-169.
42. Carpenter PJ, Pope LC, Greig C, Dawson DA, Rogers LM, Erven K, Wilson GJ, Delahay RJ, Cheeseman CL, Burke T. 2005. Mating system of the Eurasian badger, Meles meles, in a high density population. Mol Ecol. 14:273-284.
43. Cavallini P. 1996. Variation in the social system of the red fox. Ethol Ecol Evol. 8:323-342.
44. Chakraborty R, Meagher TR, Smouse PE. 1988. Parentage analysis with genetic markers in natural populations 1: The expected proportion of offspring with unambiguous paternity. Genetics. 118:527-536.
45. Clutton-Brock TH, Gaynor D, Kansky R, MacColl ADC, McIlrath G, Chadwick P, Brotherton PNM, O'Riain JM, Manser M, Skinner JD. 1998. Costs of cooperative behaviour in suricates (Suricata suricatta). Proc R Soc Lond B Biol Sci. 265:185-190.
46. Clutton-Brock TH, Hodge SJ, Spong G, Russell AF, Jordan NR, Bennett NC, Sharpe LL, Manser MB. 2006. Intrasexual competition and sexual selection in cooperative mammals. Nature. 444:1065-1068.
47. Courchamp F, Grenfell BT, Clutton-Brock TH. 2000. Impact of natural enemies on obligately cooperative breeders. Oikos. 91:311-322.
48. Courchamp F, Macdonald DW. 2001. Crucial importance of pack size in the African wild dog Lycaon pictus. Anim Conserv. 4:169-174.
49. Craighead L, Paetkau D, Reynolds HV, Vyse ER, Strobeck C. 1995. Microsatellite analysis of paternity and reproduction in Arctic grizzly bears. J Hered. 86:255-261.
50. Creel S, Macdonald D. 1995. Sociality, group size, and reproductive suppression among carnivores. In: Slater PJB, Rosenblatt JS, Snowdon CT, Milinski M, editors. Advances in the study of behavior. Vol. 24. San Diego (CA): Academic Press. p. 203-257.
51. Creel S, Sands JL. 2003. Is social stress a consequence of subordination or a cost of dominance? In: de Waal FBM, Tyack PL, editors. Animal social complexity: intelligence, culture, and individualized societies. Cambridge (MA): Harvard University Press. p. 153-169.
52. Creel SR, Creel NM. 1991. Energetics, reproductive suppression and obligate communal breeding in carnivores. Behav Ecol Sociobiol. 28:263-270.
53. Dierkes P, Heg D, Taborsky M, Skubic E, Achmann R. 2005. Genetic relatedness in groups is sex-specific and declines with age of helpers in a cooperatively breeding cichlid. Ecol Lett. 8:968-975.
54. Dugdale HL, Macdonald DW, Pope LC, Burke T. 2007. Polygynandry, extra-group paternity and multiple-paternity litters in European badger (Meles meles) social groups. Mol Ecol. 16:5294-5306.
55. Dunn PO, Cockburn A, Mulder RA. 1995. Fairy-wren helpers often care for young to which they are unrelated. Proc R Soc Lond B Biol Sci. 259:339-343.
56. East ML, Burke T, Wilhelm K, Greig C, Hofer H. 2003. Sexual conflicts in spotted hyenas: male and female mating tactics and their reproductive outcome with respect to age, social status and tenure. Proc R Soc Lond B Biol Sci. 270:1247-1254.
57. Emlen ST. 1982. The evolution of helping 2: The role of behavioral conflict. Am Nat. 119:40-53.
58. Emlen ST. 1984. Cooperative breeding in birds and mammals. In: Krebs JR, Davies NB, editors. Behavioural ecology: an evolutionary approach. 2nd ed. Oxford: Blackwell. p. 305-339.
59. Emlen ST, Oring LW 1977. Ecology, sexual selection, and evolution of mating systems. Science. 197:215-223.
60. Engh AL, Funk SM, Van Horn RC, Scribner KT, Bruford MW, Libants S, Szykman M, Smale L, Holekamp KE. 2002. Reproductive skew among males in a female-dominated mammalian society. Behav Ecol. 13:193-200.
61. Gilbert DA, Packer C, Pusey AE, Stephens JC, O'Brien SJ. 1991. Analytical DNA fingerprinting in lions: parentage, genetic diversity, and kinship. J Hered. 82:378-386.
62. Girman DJ, Mills MGL, Geffen E, Wayne RK. 1997. A molecular genetic analysis of social structure, dispersal, and interpack relationships of the African wild dog (Lycaon pictus). Behav Ecol Sociobiol. 40:187-198.
63. Gittleman JL. 1989. Carnivore group living: comparative trends. In: Gittleman JL, editor. Carnivore behaviour, ecology and evolution. Vol. 1. London: Chapman & Hall. p. 183-207.
64. Gottelli D, Wang J, Bashir S, Durrant SM. 2007. Genetic analysis reveals promiscuity among female cheetahs. Proc R Soc Lond B Biol Sci. 274:1993-2001.
65. Griffin AS, Pemberton JM, Brotherton PNM, McIlrath G, Gaynor D, Kansky R, O'Riain J, Clutton-Brock TH. 2003. A genetic analysis of breeding success in the cooperative meerkat (Suricata suricatta). Behav Ecol. 14:472-480.
66. Guo SW, Thompson EA. 1992. Performing the exact test of Hardy-Weinberg proportion for multiple alleles. Biometrics. 48:361-372.
67. Hardy OJ, Vekemans X. 2002. SPAGe Di: a versatile computer program to analyse spatial genetic structure at the individual or population levels. Mol Ecol Notes. 2:618-620.
68. Harris S. 1978. Age determination of the red fox (Vulpes vulpes)-an evaluation of technique efficiency as applied to a sample of suburban foxes. J Zool. 184:91-117.
69. Harris S, Cresswell WJ, Forde PG, Trewhella WJ, Woollard T, Wray S. 1990. Home-range analysis using radiotracking data-a review of problems and techniques particularly as applied to the study of mammals. Mammal Rev. 20:97-123.
70. Harris S, Trewhella WJ. 1988. An analysis of some of the factors affecting dispersal in an urban fox (Vulpes vulpes) population. J Appl Ecol. 25:409-422.
71. Hartley FGL, Follett BK, Harris S, Hirst D, McNeilly AS. 1994. The endocrinology of gestation failure in foxes (Vulpes vulpes). J Reprod Fertil. 100:341-346.
72. Hauber ME, Lacey EA. 2005. Bateman's principle in cooperatively breeding vertebrates: the effects of non-breeding alloparents on variability in female and male reproductive success. Integr Comp Biol. 45:903-914.
73. Hedmark E, Persson J, Segerström P, Landa A, Ellegren H. 2007. Paternity and mating system in wolverines Gulo gulo. Wildl Biol. 13(Suppl 2): 13-30.
74. Higashi M, Yamamura N. 1993. What determines animal group size? Insider-outsider conflict and its resolution. Am Nat. 142:553-563.
75. Hoffman JI, Amos W 2005. Microsatellite genotyping errors: detection approaches, common sources and consequences for paternal exclusion. Mol Ecol. 14:599-612.
76. Holland OJ, Gleeson DM. 2005. Genetic characterization of blastocysts and the identification of an instance of multiple paternity in the stoat (Mustela erminea). Conserv Genet. 6:855-858.
77. Iossa G, Soulsbury CD, Baker PJ, Harris S. 2008. Body mass, territory size and life history tactics in a socially monogamous canid. J Mammal. 89:1481-90.
78. Isvaran K, Clutton-Brock TH. 2007. Ecological correlates of extra-group paternity in mammals. Proc R Soc Lond B Biol Sci. 274:219-224.
79. Janečka JE, Blankenship TL, Hirth DH, Tewes ME, Kilpatrick CW, Grassman LI. 2006. Kinship and social structure of bobcats (Lynx rufus) inferred from microsatellite and radio-telemetry data. J Zool. 269:494-501.
80. Johnson DDP, Kays R, Blackwell PG, Macdonald DW 2002. Does the resource dispersion hypothesis explain group living? Trends Ecol Evol. 17:563-570.
81. Johnson DDP, Macdonald DW, Dickman AJ. 2000. An analysis and review of models of the socio-biology of the Mustelidae. Mammal Rev. 30:171-196.
82. Kamler JF, Ballard WB, Lemons PR, Mote K. 2004. Variation in mating system and group structure in two populations of swift foxes, Vulpes velox. Anim Behav. 68:83-88.
83. Keane B, Creel SR, Waser PM. 1996. No evidence of inbreeding avoidance or inbreeding depression in a social carnivore. Behav Ecol. 7:480-489.
84. Keane B, Waser PM, Creel SR, Creel NM, Elliott LF, Minchella DJ. 1994. Subordinate reproduction in dwarf mongooses. Anim Behav. 47:65-75.
85. Kitchen AM, Gese EM, Karki SM, Schauster ER. 2005. Spatial ecology of swift fox social groups: from group formation to mate loss. J Mammal. 86:547-554.
86. Kleiman DG. 1977. Monogamy in mammals. Q Rev Biol. 52:39-69.
87. Kokko H, Johnstone RA, Clutton-Brock TH. 2001. The evolution of cooperative breeding through group augmentation. Proc R Soc Lond B Biol Sci. 268:187-196.
88. Komdeur J. 1992. Importance of habitat saturation and territory quality for evolution of cooperative breeding in the Seychelles warbler. Nature. 358:493-495.
89. Lindström E. 1986. Territory inheritance and the evolution of group-living in carnivores. Anim Behav. 34:1825-1835.
90. Lindström E. 1988. Reproductive effort in the red fox, Vulpes vulpes, and future supply of a fluctuating prey. Oikos. 52:115-119.
91. Lindström E. 1989. Food limitation and social regulation in a red fox population. Holarctic Ecol. 12:70-79.
92. Lloyd HG. 1980. The red fox. London: Batsford.
93. Lucia KE, Keane B, Hayes LD, Lin YK, Schaefer RL, Solomon NG. 2008. Philopatry in prairies voles: an evaluation of the habitat saturation hypothesis. Behav Ecol. 19:774-783.
94. Macdonald DW. 1979. 'Helpers' in fox society. Nature. 282:69-71.
95. Macdonald DW. 1980. Social factors affecting reproduction amongst red foxes (Vulpes vulpes L., 1758). In: Zimen E, editor. The red fox: symposium on behaviour and ecology. Hague (the Netherlands): Junk. p. 123-175.
96. Macdonald DW, Moehlman PD. 1982. Cooperation, altruism and restraint in the reproduction of carnivores. In: Bateson PPG, Klopfer PH, editors. Perspectives in ethology. Vol. 5. New York: Plenum Press. p. 433-467.
97. Magrath RD, Whittingham LA. 1997. Subordinate males are more likely to help if unrelated to the breeding female in cooperatively breeding white-browed scrubwrens. Behav Ecol Sociobiol. 41: 185-192.
98. Mantel N. 1967. The detection of disease clustering and a generalized regression approach. Cancer Res. 27:209-220.
99. Marshall TC, Slate J, Kruuk LEB, Pemberton JM. 1998. Statistical confidence for likelihood-based paternity inference in natural populations. Mol Ecol. 7:639-655.
100. Moehlman PD, Hofer H. 1997. Cooperative breeding, reproductive suppression, and body size in canids. In: Solomon N, French J, editors. Cooperative breeding in mammals. Cambridge: Cambridge University Press. p. 76-128.
101. Murphy KM. 1998. The ecology of the cougar (Puma concolor) in the Northern Yellowstone ecosystem: interactions with prey, bears, and humans [PhD thesis]. Moscow (ID): University of Idaho. p. 147.
102. Nielsen CLR, Nielsen CK. 2007. Multiple paternity and relatedness in southern Illinois raccoons (Procyon lotor). J Mammal. 88:441-447.
103. Olsen JB, Busack C, Britt J, Bentzen P. 2001. The aunt and uncle effect: an empirical evaluation of the confounding influence of full sibs of parents on pedigree reconstruction. J Hered. 92:243-247.
104. Onorato DP, Hellgren EC, Van Den Bussche RA, Skiles JR. 2004. Paternity and relatedness of American black bears recolonizing a desert montane island. Can J Zool. 82:1201-1210.
105. Queller DC, Goodnight KF. 1989. Estimating relatedness using genetic markers. Evolution. 43:258-275.
106. Ralls K, Pilgrim KL, White PJ, Paxinos EE, Schwartz MK, Fleischer RC. 2001. Kinship, social relationships, and den sharing in kit foxes. J Mammal. 82:858-866.
107. Randall DA, Pollinger JP, Wayne RK, Tallents LA, Johnson PJ, Macdonald DW. 2007. Inbreeding is reduced by female-biased dispersal and mating behavior in Ethiopian wolves. Behav Ecol. 18:579-589.
108. Ratnayeke S, Tuskan GA, Pelton MR. 2002. Genetic relatedness and female spatial organization in a solitary carnivore, the raccoon, Procyon lotor. Mol Ecol. 11:1115-1124.
109. Raymond M, Rousset F. 1995. GENEPOP (version 1.2): population genetics software for exact tests and ecumenicism. J Hered. 86: 248-249.
110. Reeve HK, Emlen ST, Keller L. 1998. Reproductive sharing in animal societies: reproductive incentives or incomplete control by dominant breeders? Behav Ecol. 9:267-278.
111. Rice WR. 1989. Analyzing tables of statistical tests. Evolution. 43: 223-225.
112. Roemer GW, Smith DA, Garcelon DK, Wayne RK. 2001. The behavioural ecology of the island fox (Urocyon littoralis). J Zool. 255:1-14.
113. Ross KG. 2001. Molecular ecology of social behaviour: analyses of breeding systems and genetic structure. Mol Ecol. 10:265-284.
114. Russell AF. 2004. Mammals: comparisons and contrasts. In: Koenig WD, Dickinson JL, editors. Ecology and evolution of cooperative breeding in birds. Cambridge: Cambridge University Press. p. 210-227.
115. Say L, Devillard S, Natoli E, Pontier D. 2002. The mating system of feral cats (Felis catus L.) in a sub-Antarctic environment. Polar Biol. 25:838-842.
116. Say L, Pontier D, Natoli E. 1999. High variation in multiple paternity of domestic cats (Felis catus L.) in relation to environmental conditions. Proc R Soc Lond B Biol Sci. 266:2071-2074.
117. Schenk A, Kovacs KM. 1995. Multiple mating between black bears revealed by DNA fingerprinting. Anim Behav. 50:1483-1490.
118. Silk JB. 2007. The adaptive value of sociality in mammalian groups. Philos Trans R Soc Lond B Biol Sci. 362:539-559.
119. Slobodchikoff CN. 1988. The ecology of social behavior. San Diego (CA): Academic Press.
120. Soulsbury CD, Baker PJ, Iossa G, Harris S. 2008. Fitness costs of dispersal in red foxes (Vulpes vulpes). Behav Ecol Sociobiol. 62: 1289-1298.
121. Soulsbury CD, Iossa G, Baker PJ, Cole NC, Funk SM, Harris S. 2007. The impact of sarcoptic mange Sarcoptes scabiei on the British fox Vulpes vulpes population. Mammal Rev. 37:278-296.
122. Soulsbury CD, Iossa G, Baker PJ, Harris S. 2008. Environmental variation at the onset of independent foraging affects full-grown body mass in the red fox. Proc R Soc Lond B Biol Sci. 275:2411-2418.
123. Soulsbury CD, Iossa G, Edwards KJ, Baker PJ, Harris S. 2007. Allelic dropout from a high-quality DNA source. Conserv Genet. 8: 733-738.
124. Spong G, Stone J, Creel S, Björklund M. 2002. Genetic structure of lions (Panthera leo L.) in the Selous Game Reserve: implications for the evolution of sociality. J Evol Biol. 15:945-953.
125. Stiver KA, Dierkes P, Taborsky M, Gibbs HL, Balshine S. 2005. Relatedness and helping in fish: examining the theoretical predictions. Proc R Soc Lond B Biol Sci. 272:1593-1599.
126. Støen O-G, Bellemain E, Sæbø S, Swenson JE. 2005. Kin-related spatial structure in brown bears Ursus arctos. Behav Ecol Sociobiol. 59: 191-197.
127. Van Horn RC, Altmann J, Alberts SC. 2008. Can't get there from here: inferring kinship from pairwise genetic relatedness. Anim Behav. 75:1173-1180.
128. Vehrencamp SL. 1983. Optimal degree of skew in cooperative societies. Am Zool. 23:327-335.
129. Verwey R, Begg C, Begg K, Matthee CA. 2004. A microsatellite perspective on the reproductive success of subordinate male honey badgers, Mellivora capensis. Afr Zool. 39:305-308.
130. von Schantz T. 1981. Female cooperation, male competition, and dispersal in the red fox Vulpes vulpes. Oikos. 37:63-68.
131. von Schantz T. 1984a. 'Non-breeders' in the red fox Vulpes vulpes: a case of resource surplus. Oikos. 42:59-65.
132. von Schantz T. 1984b. Spacing strategies, kin selection, and population regulation in altricial vertebrates. Oikos. 42:48-58.
133. Wagner AP, Creel S, Frank LG, Kalinowski ST. 2007. Patterns of relatedness and parentage in an asocial, polyandrous striped hyena population. Mol Ecol. 16:4356-4369.
134. Wang J. 2004. Sibship reconstruction from genetic data with typing errors. Genetics. 166:1963-1979.
135. Westneat DF, Stewart IRK. 2003. Extra-pair paternity in birds: causes, correlates, and conflicts. Annu Rev Ecol Evol Syst. 34:365-396.
136. Wolff JO, Sherman PW 2007. Rodent societies: an ecological and evolutionary perspective. Chicago: University of Chicago Press.
137. Yamaguchi N, Sarno RJ, Johnson WE, O'Brien SJ, Macdonald DW. 2004. Multiple paternity and reproductive tactics of free-ranging American minks, Mustela vison. J Mammal. 85:432-439.
138. Zabel CJ, Taggart SJ. 1989. Shift in red fox, Vulpes vulpes, mating system associated with El Niño in the Bering Sea. Anim Behav. 38:830-838.