LARGE‐SCALE SPATIAL GRADIENTS IN HERBIVORE POPULATION DYNAMICS

Spatial gradients in density dependence and cyclicity are familiar features of the population dynamics of small mammals, particularly Fennoscandian rodents. The most well-documented of such gradients is a weakening of direct density dependence and an increase in the tendency of populations to cycle the farther north they occur, a phenomenon that has been attributed to gradients in predation and seasonality. Among large mammals, however, for which evidence of cyclicity is less clear, geographic gradients in population dynamics are limited to spatial variation in the strength of density independence. The population dynamics of caribou and muskoxen in Greenland, for example, display latitudinal gradients in the response of populations to large-scale climatic fluctuation. To my knowledge, the existence of spatial gradients in density dependence has not been explicitly investigated in large mammals. Here I present an analysis of the dynamics of 27 populations of caribou and reindeer in Greenland, Finland, and Russia, spanning 21 degrees of latitude (51.7°–72.7° N) and 215 degrees of longitude (56.4° W–159.5° E), to identify spatial gradients in density dependence and independence. Results of autoregressive time series analysis show a clear gradient in the strength of direct density dependence exhibited by these populations that declines from southern to northern latitudes. Although this pattern mirrors the latitudinal gradient evident in Fennoscandian rodent dynamics, an analysis of the dimensionality of these time series suggests that few, if any, of the populations are limited by predators. The existence of an inverse latitudinal gradient in the magnitude of the influence of large-scale climate on the dynamics of these populations suggests there may be a tension in the strength of density-dependent vs. density-independent limitation experienced by them.

[1]  C. Krebs,et al.  Are big mammals simply little mammals writ large? , 1983, Oecologia.

[2]  L. Birch,et al.  The intrinsic rate of natural increase of an insect population , 1948 .

[3]  N. Stenseth,et al.  Population dynamics of Norwegian red deer: density–dependence and climatic variation , 1998, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[4]  D. Lack The natural regulation of animal numbers , 1954 .

[5]  T. Royama,et al.  Analytical Population Dynamics , 1994, Population and Community Biology Series.

[6]  D. Klein The introduction, increase, and crash of reindeer on St. Matthew Island. , 1968 .

[7]  J. Davidson,et al.  The Influence of Rainfall, Evaporation and atmospheric Temperature on Fluctuations in the Size of a natural Population of Thrips imaginis (Thysanoptera). , 1948 .

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

[9]  A. J. Lotka Elements of Physical Biology. , 1925, Nature.

[10]  Jan Lindström,et al.  Climate and population density induce long‐term cohort variation in a northern ungulate , 2001 .

[11]  Laurence D. Mueller,et al.  Stability in Model Populations , 2000 .

[12]  E. Post,et al.  Pervasive influence of large-scale climate in the dynamics of a terrestrial vertebrate community , 2001, BMC Ecology.

[13]  N. Stenseth,et al.  Long-term responses in arctic ungulate dynamics to changes in climatic and trophic processes , 2002, Population Ecology.

[14]  N. Stenseth,et al.  Seasonal forcing on the dynamics ofClethrionomys rufocanus: Modeling geographic gradients in population dynamics , 1998, Researches on Population Ecology.

[15]  E. Ranta,et al.  Is the impact of environmental noise visible in the dynamics of age-structured populations? , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[16]  Eric Post Time Lags in Terrestrial and Marine Environments , 2005 .

[17]  N. Stenseth,et al.  A gradient from stable to cyclic populations of Clethrionomys rufocanus in Hokkaido, Japan , 1996, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[18]  E. Post,et al.  Vigilance and foraging behaviour of female caribou in relation to predation risk , 1997 .

[19]  F. Messier,et al.  THE SIGNIFICANCE OF LIMITING AND REGULATING FACTORS ON THE DEMOGRAPHY OF MOOSE AND WHITE-TAILED DEER , 1991 .

[20]  H. G. Andrewartha,et al.  The distribution and abundance of animals. , 1954 .

[21]  Ottar N. Bjørnstad,et al.  The impact of specialized enemies on the dimensionality of host dynamics , 2001, Nature.

[22]  I. Kojola,et al.  Body mass variation in semidomesticated reindeer , 1994 .

[23]  I. Kojola,et al.  Reproduction and mortality of Finnish semi-domesticated reindeer in relation to density and management strategies , 1993 .

[24]  James W. Hurrell,et al.  The North Atlantic Oscillation: Climate Significance and Environmental Impact , 2003 .

[25]  Nils Chr. Stenseth,et al.  Population cycles in voles and lemmings : density dependence and phase dependence in a stochastic world , 1999 .

[26]  E. Post,et al.  Spatial synchrony of local populations has increased in association with the recent Northern Hemisphere climate trend. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[27]  R. May,et al.  Stability and Complexity in Model Ecosystems , 1976, IEEE Transactions on Systems, Man, and Cybernetics.

[28]  J. M. Elliott,et al.  Population regulation in contrasting populations of trout Salmo trutta in two lake district streams , 1987 .

[29]  D. Klein The roles of climate and insularity in establishment and persistence ofRangifer tarandus populations in the high Arctic , 1999 .

[30]  G. Kitagawa,et al.  Akaike Information Criterion Statistics , 1988 .

[31]  Robert M. May,et al.  Stability in Randomly Fluctuating Versus Deterministic Environments , 1973, The American Naturalist.

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

[33]  N. Stenseth,et al.  A geographic gradient in small rodent density fluctuations: a statistical modelling approach , 1995, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[34]  H. Steen Untangling the causes of disappearance from a local population of root voles, Microtus oeconomus: a test of the regional synchrony hypothesis , 1995 .

[35]  H. Tong,et al.  Common dynamic structure of canada lynx populations within three climatic regions , 1999, Science.

[36]  David A. Elston,et al.  Estimating the contributions of population density and climatic fluctuations to interannual variation in survival of Soay sheep , 1999 .

[37]  Anthony R. Ives,et al.  Predicting the response of populations to environmental change , 1995 .

[38]  R. Boonstra,et al.  Concurrent density dependence and independence in populations of arctic ground squirrels , 2000, Nature.

[39]  Ilkka Hanski,et al.  Microtine Rodent Dynamics in Northern Europe: Parameterized Models for the Predator‐Prey Interaction , 1995 .

[40]  A. Nicholson,et al.  Supplement: the Balance of Animal Populations , 1933 .

[41]  O. Hoegh‐Guldberg,et al.  Ecological responses to recent climate change , 2002, Nature.

[42]  R. McRoberts,et al.  Relationship of deer and moose populations to previous winters' snow , 1987 .

[43]  E. Post,et al.  Synchronization of animal population dynamics by large-scale climate , 2002, Nature.

[44]  S. Pennings,et al.  LINKING BIOGEOGRAPHY AND COMMUNITY ECOLOGY: LATITUDINAL VARIATION IN PLANT–HERBIVORE INTERACTION STRENGTH , 2005 .

[45]  Nils Chr. Stenseth,et al.  CLIMATIC VARIABILITY, PLANT PHENOLOGY, AND NORTHERN UNGULATES , 1999 .

[46]  R. Tibshirani,et al.  Generalized Additive Models , 1991 .

[47]  E. Post,et al.  Using large-scale climate indices in climate change ecology studies , 2004, Population Ecology.

[48]  E. Ranta,et al.  Self–organized dynamics in spatially structured populations , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[49]  A. Nicholson,et al.  The Balance of Animal Populations.—Part I. , 1935 .

[50]  Mercedes Pascual,et al.  ENSO and cholera: A nonstationary link related to climate change? , 2002, Proceedings of the National Academy of Sciences of the United States of America.

[51]  J. Hurrell Decadal Trends in the North Atlantic Oscillation: Regional Temperatures and Precipitation , 1995, Science.

[52]  H. Henttonen,et al.  Gradients in density variations of small rodents: the importance of latitude and snow cover , 1985, Oecologia.

[53]  A. Agrawal,et al.  Intraspecific variation in the strength of density dependence in aphid populations , 2004 .

[54]  N. Stenseth,et al.  The population dynamics of the voleClethrionomys rufocanus in Hokkaido, Japan , 1998, Researches on Population Ecology.

[55]  Ilkka Hanski,et al.  Specialist predators, generalist predators, and the microtine rodent cycle. , 1991 .