Mating tactics and mating system of an aquatic-mating pinniped: the harbor seal, Phoca vitulina
- 490 Downloads
- 26 Citations
Abstract
Our best understanding of marine mammal mating systems comes from land-mating pinnipeds. Logistical problems of observing behavior at sea have limited our ability to make inferences about species with aquatic-mating systems, which comprise over half the pinnipeds. The mating systems of these species likely involve different mating tactics than land-mating species. We used several methods in combination (e.g., animal-borne cameras, radio telemetry, time-depth recorders, and DNA paternity assessment) to provide a comprehensive study of the aquatic-mating tactics of harbor seal males. Males decreased time offshore (26.0 vs 14.8%) and increased time near shore (33.8 vs 43.7%) between premating and mating periods, respectively. Concomitantly, males reduced foraging effort and increased activities associated with competition for females (e.g., visual/vocal displays and threats). As females come into estrus near the end of lactation and spend more time at sea, males reduced their near-shore ranges (4.2 vs 1.0 km2), which were clustered within 1–1.5 km of the beach where females attended their pups. Body mass of males was not a major factor affecting their reproductive behavior. From a small number of paternity assignments to study males, it appears that females select males. These combined results are more consistent with a lek-type mating system than with the territorial or female defense systems characteristic of land-mating pinnipeds.
Keywords
Harbor seals Mating system Lek Radio telemetry Time-depth recorders PaternityNotes
Acknowledgements
We thank the following people for assistance in the field: Brian Beck, Dave Coltman, Sarah Ellis, Sara Iverson, Jim McMillan, and Monica Muelbert. Gerry Forbes of the Department of Environment, Canada provided logistical assistance on Sable Island. Dave Coltman conducted the DNA paternity analysis and kindly provided the data. Sara Iverson, Martha Leonard, Chris Wemmer, Sophie van Parijs, and an anonymous reviewer provided helpful feedback on drafts of the manuscript. The project was supported by the Smithsonian Institution, Friends of the National Zoo, Department of Fisheries and Oceans of Canada, Natural Sciences and Engineering Research Council of Canada, and National Geographic Society.
References
- Ambs SM, Boness DJ, Bowen WD, Perry EA (1999) Proximate factors associated with high levels of extraconsort fertilization in polygynous grey seals. Anim Behav 58:527–535PubMedCrossRefGoogle Scholar
- Andersson M (1994) Sexual selection. Princeton University Press, Princeton, New JerseyGoogle Scholar
- Apollonio M, Festa-Bianchet M, Mari F, Mattioli S, Sarno B (1992) To lek or not to lek: mating strategies of male fallow deer. Behav Ecol Sociobiol 3:25–31Google Scholar
- Balmford A, Bartos L, Brotherton P, Herrmann H, Lancingerova J, Mika J, Zeeb U (1993a) When to stop lekking: density-related variation in the rutting behaviour of sika deer. J Zool 231:652–656Google Scholar
- Balmford AP, Deutsch JC, Nefdt RJC, Clutton-Brock T (1993b) Hotspot models of lek evolution: testing the predictions in three ungulate species. Behav Ecol Sociobiol 33:57–65CrossRefGoogle Scholar
- Bartholomew GA (1970) A model for the evolution of pinniped polygyny. Evolution 24:546–559CrossRefGoogle Scholar
- Bartsh SS, Johnston SD, Siniff DB (1992) Territorial behavior and breeding frequency of male Weddell seals (Leptonychotes weddelli) in relation to age, size and concentrations of serum testosterone and cortisol. Can J Zool 70:680–692Google Scholar
- Beehler BM, Fosters M (1988) Hotshots, hotspots and female preference in the organization of lek mating systems. Am Nat 131:203–219CrossRefGoogle Scholar
- Boness DJ (1991) Determinants of mating systems in the Otariidae (Pinnipedia). In: Renouf D (ed) Behaviour of pinnipeds. Chapman & Hall, London, pp 1–44Google Scholar
- Boness DJ, Bowen WD, Francis JM (1993) Implications of DNA fingerprinting for mating systems and reproductive strategies of pinnipeds. Symp Zool Soc Lond 66:61–93Google Scholar
- Boness DJ, Bowen WD, Oftedal OT (1994) Evidence of a maternal foraging cycle resembling that of otariid seals in a small phocid, the harbor seal. Behav Ecol Sociobiol 34:95–104CrossRefGoogle Scholar
- Boness DJ, Clapham PJ, Mesnick SL (2002) Life history and reproductive strategies. In: Hoelzel AR (ed) Marine mammal biology. Blackwell Science, Oxford, pp 278–324Google Scholar
- Bowen WD, Iverson SJ, Boness DJ, Oftedal OT (2001) Foraging effort, food intake and lactation performance depend on maternal mass in a small phocid. Funct Ecol 15:325–334CrossRefGoogle Scholar
- Bowen WD, Tully D, Boness DJ, Bulheier BM, Marshall GJ (2002) Prey-dependent foraging tactics and prey profitability in a marine mammal. Mar Ecol Prog Ser 244:235–245Google Scholar
- Bowen WD, Ellis SL, Iverson SJ, Boness DJ (2003) Maternal and newborn life-history traits during periods of contrasting population trends: implications for explaining the decline of harbour seals (Phoca vitulina), on Sable Island. J Zool 261:155–163CrossRefGoogle Scholar
- Bradbury JW (1981) The evolution of leks. In: Alexander RD, Tinkle D (eds) Natural selection and social behavior. Chiron Press, New York, pp 138–169Google Scholar
- Clutton-Brock TH (1989) Mammalian mating systems. Proc R Soc Lond B236:339–372CrossRefGoogle Scholar
- Clutton-Brock TH, Green D, Hiraiwa-Hasegawa M, Albon SD (1988) Passing the buck: resource defence, lek breeding and mate choice in fallow deer. Behav Ecol Sociobiol 23:281–296CrossRefGoogle Scholar
- Coltman DW, Bowen WD, Boness DJ, Iverson SJ (1997) Balancing foraging and reproduction in the male harbour seal, an aquatically mating pinniped. Anim Behav 54:663–678PubMedCrossRefGoogle Scholar
- Coltman DW, Bowen WD, Iverson SJ, Boness DJ (1998a) The energetics of male reproduction in an aquatically mating pinniped: the harbour seal. Physiol Zool 71:387–399PubMedGoogle Scholar
- Coltman DW, Bowen WD, Wright JM (1998b) Male mating success in an aquatically mating pinniped, the harbor seal (Phoca vitulina), assessed by microsatellite DNA markers. Mol Ecol 7:627–638PubMedCrossRefGoogle Scholar
- Coltman DW, Bowen WD, Wright JM (1999) A multivariate analysis of phenotype and paternity in male harbor seals, Phoca vitulina, at Sable Island, Nova Scotia. Behav Ecol 10:169–177CrossRefGoogle Scholar
- Davies NB (1991) Mating systems. In: Krebs JR, Davies NB (eds) Behavioural ecology. Blackwell, London, pp 263–299Google Scholar
- Emlen ST, Oring LW (1977) Ecology, sexual selection and the evolution of mating systems. Science 197:215–223PubMedGoogle Scholar
- Gjerde I, Wegge P, Rolstad J (2000) Lost hotspots and passive female preference: the dynamic process of lek formation in capercaillie Tetrao urogallus. Wildl Biol 6:291–298Google Scholar
- Goldsworthy SD, Boness DJ, Fleischer RC (1999) Mate choice among sympatric fur seals: female preference for conphenotypic. Behav Ecol Sociobiol 45:253–267CrossRefGoogle Scholar
- Hanggi EB, Schusterman RJ (1994) Underwater acoustic displays and individual variation in male harbour seals, Phoca vitulina. Anim Behav 48:1275–1283CrossRefGoogle Scholar
- Hayes SA, Costa DP, Harvey JT, Le Boeuf BJ (2004) Aquatic mating strategies of the male Pacific harbor seal (Phoca vitulina Richardsii): are males defending the hotspot? Mar Mamm Sci 20:639–656CrossRefGoogle Scholar
- Hoelzel AR (1999) Alpha-male paternity in elephant seals. Behav Ecol Sociobiol 46:298–306CrossRefGoogle Scholar
- Höglund J, Alatalo RV (1995) Leks. Princeton University Press, Princeton, New JerseyGoogle Scholar
- Le Boeuf BJ (1991) Pinniped mating systems on land, ice and in the water: emphasis on the Phocidae. In: Renouf D (ed) Behaviour of Pinnipeds. Chapman & Hall, London, pp 45–65Google Scholar
- Lidgard DC, Boness DJ, Bowen WD, McMillan JI, Fleischer RC (2004) The rate of fertilization in male mating tactics of the polygynous grey seal. Mol Ecol 13:3543–3548PubMedCrossRefGoogle Scholar
- Lidgard DC, Boness DJ, Bowen WD, McMillan JI (2005) State-dependent male mating tactics in the grey seal: the importance of body size. Behav Ecol 16:541–549CrossRefGoogle Scholar
- Lott DF (1991) Intraspecific variation in the social systems of wild vertebrates. Cambridge University Press, CambridgeGoogle Scholar
- Marshall GJ (1998) Cittercam: an animal-borne imaging and data logging system. Mar Technol Soc J 32:11–17Google Scholar
- Nicholson TE (2000) Social structure and underwater behavior of harbor seals in Monterey Bay, California. Ph.D. thesis, San Francisco State University, San FranciscoGoogle Scholar
- Perry EA (1993) Aquatic territory defence by male harbour seals (Phoca vitulina) at Miquelon: relationship between active defence and male reproductive success. Ph.D. thesis, Memorial University of Newfoundland, St. JohnsGoogle Scholar
- Rubenstein DI (1986) Ecology and sociality in horses and zebras. In: Rubenstein DI, Wrangham RW (eds) Ecological aspects of social evolution. Princeton University Press, Princeton, pp 282–302Google Scholar
- Sjare B, Stirling I (1996) The breeding behavior of Atlantic walruses, Odobenus rosmarus rosmarus, in the Canadian High Arctic. Can J Zool 74:897–911Google Scholar
- Stirling I (1983) The evolution of mating systems in pinnipeds. In: Eisenberg JF, Kleiman DG (eds) Advances in the study of behavior. American Society of Mammalogists, Special Publication No. 7, pp 489–527Google Scholar
- Sullivan RM (1981) Aquatic displays and interactions in harbor seals (Phoca vitulina), with comments on mating systems. J Mammal 62:825–831CrossRefGoogle Scholar
- Van Parijs SM, Thompson PM, Tollit DJ, Mackay A (1997) Distribution and activity of male harbour seals during the mating season. Anim Behav 54:35–43CrossRefGoogle Scholar
- Van Parijs SM, Hastie GD, Thompson PM (1999) Geographical variation and spatial vocalization patterns of male harbour seals in the mating season. Anim Behav 58:1231–1239PubMedCrossRefGoogle Scholar
- Van Parijs SM, Hastie GD, Thompson PM (2000a) Individual and geographical variation in display behaviour of male harbour seals in Scotland. Anim Behav 59:559–568PubMedCrossRefGoogle Scholar
- Van Parijs SM, Janik VM, Thompson PM (2000b) Display-area size, tenure length, and site fidelity in the aquatically mating male harbour seal, Phoca vitulina. Can J Zool 78:2209–2217CrossRefGoogle Scholar
- Van Parijs SM, Kovacs KM, Lydersen C (2001) Spatial and temporal distribution of vocalising male bearded seals—implications for male strategies. Behaviour 138:905–922CrossRefGoogle Scholar
- Van Parijs SM, Lydersen C, Kovacs KM (2003) Vocalizations and movements suggest alternative mating tactics in male bearded seals. Anim Behav 65:273–283CrossRefGoogle Scholar
- Walker BG, Bowen WD (1993) Behavioural differences among adult male harbour seals during the breeding season may provide evidence of reproductive strategies. Can J Zool 71:1585–1591Google Scholar
- Widemo F, Owens IPF (1995) Lek size, male mating skew and the evolution of lekking. Nature (Lond) 373:148–151CrossRefGoogle Scholar
- Widemo F, Owens IPF (1999) Size and stability of vertebrate leks. Anim Behav 58:1217–1221PubMedCrossRefGoogle Scholar
- Wiley RH (1991) Lekking in birds and mammals: behavioral and evolutionary issues. Adv Stud Behav 20:201–291CrossRefGoogle Scholar
- Wilson EO (1975) Sociobiology. A new synthesis. Belknapp Press, Cambridge, MAGoogle Scholar