Age, Size, and Spatiotemporal Variation in Ovulation Patterns of a Seasonal Breeder, the Norwegian Moose (Alces alces)

In seasonal environments, timing of reproduction is an important fitness component. However, in ungulates, our understanding of this biological process is limited. Here we analyze how age and body mass affect spatiotemporal variation in timing of ovulation of 6,178 Norwegian moose. We introduced a parametric statistical model to obtain inferences about the seasonal timing of ovulation peak, the degree of synchrony among individuals, and the proportion of individuals that ovulate. These components showed much more spatiotemporal variation than previously reported. Young (primiparous) and old (≥11.5 years of age) females ovulated later than prime‐aged (2.5–10.5 years of age) females. In all age classes, ovulation was delayed with decreasing body mass. Ovulation rates were lower and more variable among primiparous females than among older females. Young females required higher body mass than older females did to ovulate. The body‐mass‐to‐ovulation relationship varied with age, showed large regional variation, and differed among years within region. These results suggest that (1) environmental and population characteristics contribute to shape seasonal variation in the breeding pattern and (2) large regional variation exists in the size‐dependent age at maturity in moose. Hence, the life‐history trade‐off between reproduction and body growth should differ regionally in moose.

[1]  William N. Venables,et al.  Modern Applied Statistics with S , 2010 .

[2]  Joseph Hilbe,et al.  Data Analysis Using Regression and Multilevel/Hierarchical Models , 2009 .

[3]  B. Sæther,et al.  Lack of compensatory body growth in a high performance moose Alces alces population , 2008, Oecologia.

[4]  J. Gaillard,et al.  Selective harvesting and habitat loss produce long-term life history changes in a mouflon population. , 2007, Ecological applications : a publication of the Ecological Society of America.

[5]  B. Sæther,et al.  Annual variation in maternal age and calving date generate cohort effects in moose (Alces alces) body mass , 2007, Oecologia.

[6]  J. Gaillard,et al.  Indicators of ecological change: new tools for managing populations of large herbivores , 2007 .

[7]  K. Høgda,et al.  Population characteristics predict responses in moose body mass to temporal variation in the environment. , 2006, The Journal of animal ecology.

[8]  Kjell Arild Høgda,et al.  Environmental phenology and geographical gradients in moose body mass , 2006, Oecologia.

[9]  K. Høgda,et al.  The length of growing season and adult sex ratio affect sexual size dimorphism in moose. , 2006, Ecology.

[10]  J. Gaillard,et al.  Male age structure influences females’ mass change during rut in a polygynous ungulate: the reindeer (Rangifer tarandus) , 2006, Behavioral Ecology and Sociobiology.

[11]  A. Mysterud,et al.  Adaptive adjustment of offspring sex ratio and maternal reproductive effort in an iteroparous mammal , 2006, Proceedings of the Royal Society B: Biological Sciences.

[12]  Erlend B. Nilsen,et al.  Patterns of hunting mortality in Norwegian moose (Alces alces) populations , 2006, European Journal of Wildlife Research.

[13]  O. Jones,et al.  Different hunting strategies select for different weights in red deer , 2005, Biology Letters.

[14]  J. Gaillard,et al.  Reproductive output of female mouflon (Ovis gmelini musimon×Ovis sp.): a comparative analysis , 2005 .

[15]  S. Bertouille,et al.  Fertility of red deer in relation to area, age, body mass, and mandible length , 2002, Zeitschrift für Jagdwissenschaft.

[16]  J. Testa.  POPULATION DYNAMICS AND LIFE HISTORY TRADE‐OFFS OF MOOSE (ALCES ALCES) IN SOUTH‐CENTRAL ALASKA , 2004 .

[17]  Atle Mysterud,et al.  Timing and Synchrony of Ovulation in Red Deer Constrained by Short Northern Summers , 2004, The American Naturalist.

[18]  John G. Cook,et al.  EFFECTS OF SUMMER‐AUTUMN NUTRITION AND PARTURITION DATE ON REPRODUCTION AND SURVIVAL OF ELK , 2004, Wildlife Monographs.

[19]  H. Sand.  Life history patterns in female moose (Alces alces): the relationship between age, body size, fecundity and environmental conditions , 1996, Oecologia.

[20]  D. Coltman,et al.  Undesirable evolutionary consequences of trophy hunting , 2003, Nature.

[21]  L. Kruuk,et al.  ESTIMATING SELECTION ON NEONATAL TRAITS IN RED DEER USING ELASTICITY PATH ANALYSIS , 2003, Evolution; international journal of organic evolution.

[22]  Climate–vegetation dynamics in the fast lane , 2003 .

[23]  E. Post,et al.  Synchrony between caribou calving and plant phenology in depredated and non-depredated populations , 2003 .

[24]  J. Liao,et al.  Adjusted Coefficients of Determination for Logistic Regression , 2003 .

[25]  B. Sæther,et al.  Effects of Altering Sex Ratio Structure on the Demography of an Isolated Moose Population , 2003 .

[26]  E. Milner‐Gulland,et al.  Conservation: Reproductive collapse in saiga antelope harems , 2003, Nature.

[27]  E. Solberg,et al.  Status of moose populations and challenges to moose management in Fennoscandia. , 2003 .

[28]  A. Mysterud,et al.  The effect of sex ratio and male age structure on reindeer calving , 2003 .

[29]  B. Johnson,et al.  Effects of Male Age and Female Nutritional Condition on Elk Reproduction , 2002 .

[30]  Eric R. Ziegel,et al.  Generalized Linear Models , 2002, Technometrics.

[31]  B. Sæther,et al.  Biased adult sex ratio can affect fecundity in primiparous moose Alces alces , 2002, Wildlife Biology.

[32]  S. Palumbi,et al.  Humans as the world's greatest evolutionary force. , 2001, Science.

[33]  J. Gaillard,et al.  Phenotypic quality and senescence affect different components of reproductive output in roe deer , 2001 .

[34]  J. P. Ball,et al.  AGE-RELATED REPRODUCTIVE EFFORT AND SENESCENCE IN FREE-RANGING MOOSE, ALCES ALCES , 2001 .

[35]  B T Grenfell,et al.  Age, sex, density, winter weather, and population crashes in Soay sheep. , 2001, Science.

[36]  David R. Anderson,et al.  Kullback-Leibler information as a basis for strong inference in ecological studies , 2001 .

[37]  S. Côté,et al.  Birthdate, mass and survival in mountain goat kids: effects of maternal characteristics and forage quality , 2001, Oecologia.

[38]  Göran Ericsson,et al.  Age-specific moose (Alces alces) mortality in apredator-free environment: Evidence for senescence in females , 2001 .

[39]  J. Gaillard,et al.  Temporal Variation in Fitness Components and Population Dynamics of Large Herbivores , 2000 .

[40]  J. Hoef,et al.  Life-history consequences of maternal condition in Alaskan moose , 2000 .

[41]  David R. Anderson,et al.  Model selection and inference : a practical information-theoretic approach , 2000 .

[42]  B. Sæther,et al.  Age-specific harvest mortality in a Norwegian moose Alces alces population , 2000, Wildlife Biology.

[43]  J. Lindström,et al.  Early development and fitness in birds and mammals. , 1999, Trends in ecology & evolution.

[44]  P. E. Komers,et al.  Timing of Estrus in Fallow Deer Is Adjusted to the Age of Available Mates , 1999, The American Naturalist.

[45]  A. Loison,et al.  Body mass and winter mortality in red deer calves: disentangling sex and climate effects , 1999 .

[46]  J. Kie,et al.  Timing and Synchrony of Parturition in Alaskan Moose: Long-Term Versus Proximal Effects of Climate , 1998 .

[47]  G. Adams,et al.  Body condition and adjustments to reproductive effort in female moose (Alces alces) , 1998 .

[48]  T. Clutton‐Brock,et al.  Cohort variation in male survival and lifetime breeding success in red deer. , 1998, The Journal of animal ecology.

[49]  Marcel E. Visser,et al.  Warmer springs lead to mistimed reproduction in great tits (Parus major) , 1998, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[50]  F. Pérez-Barbería,et al.  Reproductive parameters, kidney fat index, and grazing activity relationships between the sexes in Cantabrian chamois Rupicapra pyrenaica parva , 1998 .

[51]  J. Gaillard,et al.  Mass‐ and Density‐Dependent Reproductive Success and Reproductive Costs in a Capital Breeder , 1998, The American Naturalist.

[52]  B. Sæther,et al.  Ecological Correlates of Regional Variation in Life History of the Moose Alces Alces , 1998 .

[53]  J. Linnell,et al.  Timing and synchrony of birth in a hider species, the roe deer Capreolus capreolus , 1998 .

[54]  B. Dale,et al.  Timing and Synchrony of Parturition in Alaskan Caribou , 1998 .

[55]  Costs of reproduction in female moose (Alces alces) as measured by means of phenotypic correlations , 1998 .

[56]  B. Sæther.  Environmental stochasticity and population dynamics of large herbivores: a search for mechanisms. , 1997, Trends in ecology & evolution.

[57]  R. Andersen,et al.  Variation in maternal investment in a small cervid; the effects of cohort, sex, litter size and time of birth in roe deer (Capreolus capreolus) fawns , 1997, Oecologia.

[58]  T. Clutton‐Brock,et al.  Climate, plant phenology and variation in age of first reproduction in a temperate herbivore , 1996 .

[59]  A. Hewison.  Variation in the fecundity of roe deer in Britain: effects of age and body weight , 1996 .

[60]  Å. Pehrson,et al.  Predicting body mass from chest circumference in moose Alces alces , 1996, Wildlife Biology.

[61]  Jean-Michel Gaillard,et al.  Body mass of roe deer fawns during winter in 2 contrasting populations , 1996 .

[62]  M. Festa‐Bianchet,et al.  Life History Consequences of Variation in Age of Primiparity in Bighorn Ewes , 1995 .

[63]  S. Engen,et al.  Retrospective Studies of Red Deer Reproduction Using Regressing Luteal Structures , 1994 .

[64]  J. Hoef,et al.  Predicting Parturition Rate of Caribou from Autumn Body Mass , 1994 .

[65]  S. Engen,et al.  Optimal allocation of resources to growth and reproduction , 1994 .

[66]  R. Bon,et al.  Mating and Lambing Periods as Related to Age of Female Mouflon , 1993 .

[67]  B. Sæther,et al.  Ecological correlates of individual variation in age at maturity in female moose (Alces alces): the effects of environmental variability , 1993 .

[68]  C. C. Schwartz,et al.  Reproductive characteristics of Alaskan moose , 1993 .

[69]  S. Albon,et al.  Plant phenology and the benefits of migration in a temperate ungulate , 1992 .

[70]  R. Langvatn.  Analysis of ovaries in studies of reproduction in red deer (Cervus elaphus, L.): Application and limitations , 1992 .

[71]  Jean-Michel Gaillard,et al.  Effects of age and body weight on the proportion of females breeding in a population of roe deer (Capreolus capreolus) , 1992 .

[72]  P. A. Jewell,et al.  Early development and population fluctuations in Soay sheep , 1992 .

[73]  T. Clutton‐Brock,et al.  Red Deer: Behavior and Ecology of Two Sexes , 1992 .

[74]  J. Berger.  Facilitation of Reproductive Synchrony by Gestation Adjustment in Gregarious Mammals: A New Hypothesis , 1992 .

[75]  J. Rachlow,et al.  Interannual Variation in Timing and Synchrony of Parturition in Dall's Sheep , 1991 .

[76]  R. Bowyer.  Timing of Parturition and Lactation in Southern Mule Deer , 1991 .

[77]  Warren B. Ballard,et al.  Population dynamics of moose in South-central Alaska , 1991 .

[78]  C. M. Lessells,et al.  The Evolution of Life Histories , 1994 .

[79]  T. Skogland.  Natural selection of wild reindeer life history traits by food limitation and predation , 1989 .

[80]  K. Risenhoover,et al.  Growth rates and birthing period of bighorn sheep in low-elevation environments in Colorado , 1988 .

[81]  M. Festa‐Bianchet.  Birthdate and survival in bighorn lambs (Ovis canadensis) , 1988 .

[82]  M. Festa‐Bianchet.  Age-Specific Reproduction of Bighorn Ewes in Alberta, Canada , 1988 .

[83]  A. Rutberg.  Adaptive Hypotheses of Birth Synchrony in Ruminants: An Interspecific Test , 1987, The American Naturalist.

[84]  L. J. Verme,et al.  Relationship of Photoperiod to Puberty in Doe Fawn White-Tailed Deer , 1987 .

[85]  T. Clutton‐Brock,et al.  Early development and population dynamics in red deer. I: Density-dependent effects on juvenile survival , 1987 .

[86]  T. Clutton‐Brock,et al.  Early development and population dynamics in red deer. II: Density-independent effects and cohort variation , 1987 .

[87]  R. Sadleir.  Reproduction of female cervids , 1987 .

[88]  L. J. Verme,et al.  Induced early estrus in penned white-tailed deer does , 1987 .

[89]  S. Stearns,et al.  THE EVOLUTION OF PHENOTYPIC PLASTICITY IN LIFE‐HISTORY TRAITS: PREDICTIONS OF REACTION NORMS FOR AGE AND SIZE AT MATURITY , 1986, Evolution; international journal of organic evolution.

[90]  J. Bauer.  Factors determining the onset of sexual maturity in New Zealand chamois (Rupicapra rupicapra L.) , 1986 .

[91]  L. J. Verme.  Birth Weights of Fawns from Doe Fawn White-Tailed Deer , 1985 .

[92]  T. Skogland.  The effects of density-dependent resource limitations on the demography of wild reindeer , 1985 .

[93]  C. C. Schwartz,et al.  Moose twinning rates: a possible population condition assessment , 1985 .

[94]  B. Mitchell,et al.  FERTILITY AND BODY WEIGHT IN FEMALE RED DEER: A DENSITY-DEPENDENT RELATIONSHIP , 1983 .

[95]  A. Loudon.  Red Deer: Behaviour and Ecology of Two Sexes, T.H. Clutton-Brock, F.E. Guinness, S.D. Albon. Edinburgh University Press, Edinburgh (1982), xxii, + 378 Price £20.10 hardback, £9.75 paperback , 1983 .

[96]  B. Saether,et al.  Life history of the moose (Alces alces): fecundity rates in relation to age and carcass weight [Moose, Norway] , 1983 .

[97]  E. Reimers.  Reproduction in wild reindeer in Norway , 1983 .

[98]  R. W. Thompson,et al.  Temporal geographic variation in the lambing season of bighorn sheep , 1982 .

[99]  R. Lande,et al.  A Quantitative Genetic Theory of Life History Evolution , 1982 .

[100]  F. Bunnell.  The lambing period of mountain sheep: synthesis, hypotheses, and tests , 1982 .

[101]  Female breeding age and lamb survival in desert bighorn sheep (Ovis canadensis) , 1982 .

[102]  F. Bunnell.  Factors controlling lambing period of Dall's sheep. , 1980, Canadian journal of zoology.

[103]  R. Lande.  QUANTITATIVE GENETIC ANALYSIS OF MULTIVARIATE EVOLUTION, APPLIED TO BRAIN:BODY SIZE ALLOMETRY , 1979, Evolution; international journal of organic evolution.

[104]  H. Haagenrud.  Layers in secondary dentine of incisors as age criteria in moose (Alces alces). , 1978, Journal of mammalogy.

[105]  C. Pond,et al.  Morphological Aspects and the Ecological and Mechanical Consequences of Fat Deposition in Wild Vertebrates , 1978 .

[106]  R. L. Downing,et al.  Factors Affecting the Peak of White-Tailed Deer Fawning in Virginia , 1977 .

[107]  Jr . T. Charles Dauphini.  SYNCHRONOUS MATING IN CANADIAN BARREN-GROUND CARIBOU , 1974 .

[108]  D. Stevens,et al.  Reproduction of Shiras Moose in Montana , 1973 .

[109]  V. Geist.  Mountain sheep;: A study in behavior and evolution , 1972 .

[110]  G. Markgren.  Reproduction of Moose in Sweden , 1969 .

[111]  D. Simkin.  Reproduction and Productivity of Moose in Northwestern Ontario , 1965 .

[112]  L. J. Verme.  Reproduction Studies on Penned White-Tailed Deer , 1965 .

[113]  J. Peek.  Studies of Moose in the Gravelly and Snowcrest Mountains, Montana , 1962 .

[114]  D. Pimlott.  Reproduction and Productivity of Newfoundland Moose , 1959 .

[115]  R. Ritcey,et al.  REPRODUCTION IN A MOOSE POPULATION , 1958 .

[116]  G. A. Moore,et al.  Vertebrates of the United States , 1958 .