Climate‐driven ecological stability as a globally shared cause of Late Quaternary megafaunal extinctions: the Plaids and Stripes Hypothesis

Controversy persists about why so many large‐bodied mammal species went extinct around the end of the last ice age. Resolving this is important for understanding extinction processes in general, for assessing the ecological roles of humans, and for conserving remaining megafaunal species, many of which are endangered today. Here we explore an integrative hypothesis that asserts that an underlying cause of Late Quaternary megafaunal extinctions was a fundamental shift in the spatio‐temporal fabric of ecosystems worldwide. This shift was triggered by the loss of the millennial‐scale climate fluctuations that were characteristic of the ice age but ceased approximately 11700 years ago on most continents. Under ice‐age conditions, which prevailed for much of the preceding 2.6 Ma, these radical and rapid climate changes prevented many ecosystems from fully equilibrating with their contemporary climates. Instead of today's ‘striped’ world in which species' ranges have equilibrated with gradients of temperature, moisture, and seasonality, the ice‐age world was a disequilibrial ‘plaid’ in which species' ranges shifted rapidly and repeatedly over time and space, rarely catching up with contemporary climate. In the transient ecosystems that resulted, certain physiological, anatomical, and ecological attributes shared by megafaunal species pre‐adapted them for success. These traits included greater metabolic and locomotory efficiency, increased resistance to starvation, longer life spans, greater sensory ranges, and the ability to be nomadic or migratory. When the plaid world of the ice age ended, many of the advantages of being large were either lost or became disadvantages. For instance in a striped world, the low population densities and slow reproductive rates associated with large body size reduced the resiliency of megafaunal species to population bottlenecks. As the ice age ended, the downsides of being large in striped environments lowered the extinction thresholds of megafauna worldwide, which then increased the vulnerability of individual species to a variety of proximate threats they had previously tolerated, such as human predation, competition with other species, and habitat loss. For many megafaunal species, the plaid‐to‐stripes transition may have been near the base of a hierarchy of extinction causes whose relative importances varied geographically, temporally, and taxonomically.

[1]  J. Estes Serendipity , 2019 .

[2]  Michael C. Westaway,et al.  Big data little help in megafauna mysteries , 2018, Nature.

[3]  M. Bar-Matthews,et al.  The earliest modern humans outside Africa , 2018, Science.

[4]  John A. Kamm,et al.  Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans , 2018, Nature.

[5]  J. Weaver,et al.  Impact-related microspherules in Late Pleistocene Alaskan and Yukon “muck” deposits signify recurrent episodes of catastrophic emplacement , 2017, Scientific Reports.

[6]  K. Douka,et al.  Human Colonization of Asia in the Late Pleistocene , 2017, Current Anthropology.

[7]  B. Kirk,et al.  Serendipity: An Ecologist's Quest to Understand Nature , 2017, Journal of Mammalogy.

[8]  W. Deng,et al.  The common variants implicated in microstructural abnormality of first episode and drug-naïve patients with schizophrenia , 2017, Scientific Reports.

[9]  J. Southon,et al.  A case of early Wisconsinan ;over-chill;: New radiocarbon evidence for early extirpation of western camel (Camelops hesternus) in eastern Beringia , 2017 .

[10]  J. Dortch,et al.  Large mammal species richness and late Quaternary precipitation change in south‐western Australia , 2017 .

[11]  J. Wilmshurst,et al.  Island extinctions: processes, patterns, and potential for ecosystem restoration , 2017, Environmental Conservation.

[12]  R. Bovendorp,et al.  Defaunation and biomass collapse of mammals in the largest Atlantic forest remnant , 2017 .

[13]  G. Zazula,et al.  Megafaunal isotopes reveal role of increased moisture on rangeland during late Pleistocene extinctions , 2017, Nature Ecology &Evolution.

[14]  Martin Kircher,et al.  The evolutionary and phylogeographic history of woolly mammoths: a comprehensive mitogenomic analysis , 2017, Scientific Reports.

[15]  M. Meyer,et al.  Fossil and genomic evidence constrains the timing of bison arrival in North America , 2017, Proceedings of the National Academy of Sciences.

[16]  P. D. Polly,et al.  Merging paleobiology with conservation biology to guide the future of terrestrial ecosystems , 2017, Science.

[17]  G. Pazour,et al.  Ror2 signaling regulates Golgi structure and transport through IFT20 for tumor invasiveness , 2017, Scientific Reports.

[18]  K. Roucoux,et al.  Vegetation responses to abrupt climatic changes during the Last Interglacial Complex (Marine Isotope Stage 5) at Tenaghi Philippon, NE Greece , 2016 .

[19]  Michael K. Gagan,et al.  Disentangling the impacts of climate and human colonization on the flora and fauna of the Australian arid zone over the past 100 ka using stable isotopes in avian eggshell , 2016 .

[20]  A. Markova,et al.  Mammal faunas in Europe at the end of the Early – Beginning of the Middle Pleistocene , 2016 .

[21]  Menna E. Jones,et al.  A Nose for Death: Integrating Trophic and Informational Networks for Conservation and Management , 2016, Front. Ecol. Evol..

[22]  Lee A. Newsom,et al.  Timing and causes of mid-Holocene mammoth extinction on St. Paul Island, Alaska , 2016, Proceedings of the National Academy of Sciences.

[23]  M. Rummukainen,et al.  Vegetation–climate feedbacks modulate rainfall patterns in Africa underfuture climate change , 2016 .

[24]  Montgomery Slatkin,et al.  Excess of genomic defects in a woolly mammoth on Wrangel island , 2016, PLoS genetics.

[25]  S. K. Lyons,et al.  The changing role of mammal life histories in Late Quaternary extinction vulnerability on continents and islands , 2016, Biology Letters.

[26]  L. Halsey Terrestrial movement energetics: current knowledge and its application to the optimising animal , 2016, Journal of Experimental Biology.

[27]  Zenobia Jacobs,et al.  A late Quaternary vertebrate deposit in Kudjal Yolgah Cave, south‐western Australia: refining regional late Pleistocene extinctions , 2016 .

[28]  G. B. Golding,et al.  Mammuthus Population Dynamics in Late Pleistocene North America: Divergence, Phylogeography, and Introgression , 2016, Front. Ecol. Evol..

[29]  Jiyoul Lee,et al.  Pulse-modulated multilevel data storage in an organic ferroelectric resistive memory diode , 2016, Scientific Reports.

[30]  M. Wooller,et al.  High-resolution records detect human-caused changes to the boreal forest wildfire regime in interior Alaska , 2016 .

[31]  Erle C. Ellis,et al.  Late Holocene climate: Natural or anthropogenic? , 2016 .

[32]  T. L. Rasmussen,et al.  North Atlantic warming during Dansgaard-Oeschger events synchronous with Antarctic warming and out-of-phase with Greenland climate , 2016, Scientific Reports.

[33]  Christopher N. Johnson,et al.  Geographic variation in the ecological effects of extinction of Australia's Pleistocene megafauna , 2016 .

[34]  A. Timmermann,et al.  Millennial to orbital-scale variations of drought intensity in the Eastern Mediterranean , 2016 .

[35]  Jianping Huang,et al.  Global semi-arid climate change over last 60 years , 2016, Climate Dynamics.

[36]  Damien A. Fordham,et al.  Climate change not to blame for late Quaternary megafauna extinctions in Australia , 2016, Nature Communications.

[37]  J. Terborgh,et al.  Megafauna and ecosystem function from the Pleistocene to the Anthropocene , 2016, Proceedings of the National Academy of Sciences.

[38]  D. Harper,et al.  Onset of main Phanerozoic marine radiation sparked by emerging Mid Ordovician icehouse , 2016, Scientific Reports.

[39]  Alejandro Ordonez,et al.  The influence of paleoclimate on present-day patterns in biodiversity and ecosystems , 2015 .

[40]  S. Archibald,et al.  A continent-wide assessment of the form and intensity of large mammal herbivory in Africa , 2015, Science.

[41]  B. Shapiro,et al.  Life and extinction of megafauna in the ice-age Arctic , 2015, Proceedings of the National Academy of Sciences.

[42]  D. Doak,et al.  Sea otters, kelp forests, and the extinction of Steller’s sea cow , 2015, Proceedings of the National Academy of Sciences.

[43]  Todd A. Surovell,et al.  Test of Martin’s overkill hypothesis using radiocarbon dates on extinct megafauna , 2015, Proceedings of the National Academy of Sciences.

[44]  D. J. Meltzer,et al.  Pleistocene Overkill and North American Mammalian Extinctions , 2015 .

[45]  J. Svenning,et al.  Historic and prehistoric human‐driven extinctions have reshaped global mammal diversity patterns , 2015 .

[46]  R. Röthlisberger,et al.  Millennial changes in North American wildfire and soil activity over the last glacial cycle , 2015 .

[47]  Konrad A Hughen,et al.  Abrupt warming events drove Late Pleistocene Holarctic megafaunal turnover , 2015, Science.

[48]  A. Berger,et al.  Interglacial analogues of the Holocene and its natural near future , 2015 .

[49]  F. Chapin,et al.  Vegetation succession in deglaciated landscapes: implications for sediment and landscape stability , 2015 .

[50]  Carson A. Baughman,et al.  Soil surface organic layers in Arctic Alaska: Spatial distribution, rates of formation, and microclimatic effects , 2015 .

[51]  A. Stuart Late Quaternary megafaunal extinctions on the continents: a short review , 2015 .

[52]  Nathan J B Kraft,et al.  Linking environmental filtering and disequilibrium to biogeography with a community climate framework. , 2015, Ecology.

[53]  B. Jones,et al.  Hydrological transformation coincided with megafaunal extinction in central Australia , 2015 .

[54]  M. Raymo,et al.  An 800-kyr record of global surface ocean δ18O and implications for ice volume-temperature coupling , 2014 .

[55]  C. Buizert,et al.  Consistently dated records from the Greenland GRIP, GISP2 and NGRIP ice cores for the past 104 ka reveal regional millennial-scale δ18O gradients with possible Heinrich event imprint , 2014 .

[56]  H. Fischer,et al.  A stratigraphic framework for abrupt climatic changes during the Last Glacial period based on three synchronized Greenland ice-core records: refining and extending the INTIMATE event stratigraphy , 2014 .

[57]  George L. W. Perry,et al.  A high-precision chronology for the rapid extinction of New Zealand moa (Aves, Dinornithiformes) , 2014 .

[58]  H. Gregory McDonald,et al.  American mastodon extirpation in the Arctic and Subarctic predates human colonization and terminal Pleistocene climate change , 2014, Proceedings of the National Academy of Sciences.

[59]  Charlotte L. Oskam,et al.  An extremely low-density human population exterminated New Zealand moa , 2014, Nature Communications.

[60]  J. Wilmshurst,et al.  A High-Resolution Chronology of Rapid Forest Transitions following Polynesian Arrival in New Zealand , 2014, PloS one.

[61]  M. Maslin,et al.  East African climate pulses and early human evolution , 2014 .

[62]  C. Ramsey,et al.  Sea-level variability over five glacial cycles , 2014, Nature Communications.

[63]  M. Wooller,et al.  Radiocarbon age‐offsets in an arctic lake reveal the long‐term response of permafrost carbon to climate change , 2014 .

[64]  C. Sandom,et al.  Global late Quaternary megafauna extinctions linked to humans, not climate change , 2014, Proceedings of the Royal Society B: Biological Sciences.

[65]  J. Alroy A simple Bayesian method of inferring extinction , 2014, Paleobiology.

[66]  Thomas Guillerme,et al.  Ecology and mode-of-life explain lifespan variation in birds and mammals , 2014, Proceedings of the Royal Society B: Biological Sciences.

[67]  C. McCain,et al.  Body size and activity times mediate mammalian responses to climate change , 2014, Global change biology.

[68]  M. Leuenberger,et al.  Temperature reconstruction from 10 to 120 kyr b2k from the NGRIP ice core , 2014 .

[69]  M. Griffiths,et al.  Rapid interhemispheric climate links via the Australasian monsoon during the last deglaciation , 2013, Nature Communications.

[70]  H. Lischke,et al.  Vegetation responses to rapid warming and to minor climatic fluctuations during the Late-Glacial Interstadial (GI-1) at Gerzensee (Switzerland) , 2013 .

[71]  J. Erlandson,et al.  Archeology, deep history, and the human transformation of island ecosystems , 2013 .

[72]  P. deMenocal,et al.  Abrupt Shifts in Horn of Africa Hydroclimate Since the Last Glacial Maximum , 2013, Science.

[73]  S. Elias The problem of conifer species migration lag in the Pacific Northwest region since the last glaciation , 2013 .

[74]  C. Tovar,et al.  Determining the response of African biota to climate change: using the past to model the future , 2013, Philosophical Transactions of the Royal Society B: Biological Sciences.

[75]  Heike Lischke,et al.  Tree Migration-Rates: Narrowing the Gap between Inferred Post-Glacial Rates and Projected Rates , 2013, PloS one.

[76]  Henry T. Wright,et al.  Stone tools and foraging in northern Madagascar challenge Holocene extinction models , 2013, Proceedings of the National Academy of Sciences.

[77]  W. Humphreys,et al.  North Atlantic forcing of millennial-scale Indo-Australian monsoon dynamics during the Last Glacial period , 2013 .

[78]  N. Owen‐Smith Contrasts in the large herbivore faunas of the southern continents in the late Pleistocene and the ecological implications for human origins , 2013 .

[79]  J. Svenning,et al.  Disequilibrium vegetation dynamics under future climate change. , 2013, American journal of botany.

[80]  B. Gaglioti,et al.  Ice-age megafauna in Arctic Alaska: extinction, invasion, survival , 2013 .

[81]  J. Faith Ungulate diversity and precipitation history since the Last Glacial Maximum in the Western Cape, South Africa , 2013 .

[82]  D. Grayson,et al.  Climate change frames debate over the extinction of megafauna in Sahul (Pleistocene Australia-New Guinea) , 2013, Proceedings of the National Academy of Sciences.

[83]  J. Long,et al.  Quantifying the periodicity of Heinrich and Dansgaard–Oeschger events during Marine Oxygen Isotope Stage 3 , 2013, Quaternary Research.

[84]  J. Singarayer,et al.  Millennial Climatic Fluctuations Are Key to the Structure of Last Glacial Ecosystems , 2013, PloS one.

[85]  T. Blackburn,et al.  Magnitude and variation of prehistoric bird extinctions in the Pacific , 2013, Proceedings of the National Academy of Sciences.

[86]  L. Werdelin,et al.  Temporal Change in Functional Richness and Evenness in the Eastern African Plio-Pleistocene Carnivoran Guild , 2013, PloS one.

[87]  A. Timmermann,et al.  Links between tropical rainfall and North Atlantic climate during the last glacial period , 2013 .

[88]  K. Roucoux,et al.  Environmental variability during the Last Interglacial: a new high‐resolution pollen record from Tenaghi Philippon, Greece , 2013 .

[89]  Kenji Kawamura,et al.  Eemian interglacial reconstructed from a Greenland folded ice core , 2013, Nature.

[90]  F. Chapin,et al.  Mammoth steppe: a high-productivity phenomenon , 2012 .

[91]  A. Berger,et al.  Modelling the climatic diversity of the warm interglacials , 2012 .

[92]  V. Brovkin,et al.  Can we predict the duration of an interglacial , 2012 .

[93]  M. Huston Precipitation, soils, NPP, and biodiversity: resurrection of Albrecht's curve , 2012 .

[94]  Jacquelyn L. Gill,et al.  Climatic and megaherbivory controls on late-glacial vegetation dynamics: a new, high-resolution, multi-proxy record from Silver Lake, Ohio , 2012 .

[95]  C. Bradshaw,et al.  Robust estimates of extinction time in the geological record , 2012 .

[96]  S. Burns,et al.  High-Latitude Forcing of the South American Summer Monsoon During the Last Glacial , 2012, Science.

[97]  R. Edwards,et al.  800,000 Years of Abrupt Climate Variability , 2011, Science.

[98]  Mark A. Wilson,et al.  Sea-level oscillations during the last interglacial highstand recorded by Bahamas corals , 2011 .

[99]  J. Calabrese,et al.  How landscape dynamics link individual‐ to population‐level movement patterns: a multispecies comparison of ungulate relocation data , 2011 .

[100]  V. Pitulko,et al.  Last straw versus Blitzkrieg overkill: Climate-driven changes in the Arctic Siberian mammoth population and the Late Pleistocene extinction problem , 2011 .

[101]  J. Kutzbach,et al.  The role of GCM resolution in simulating glacial inception , 2011 .

[102]  Kelly K. Caylor,et al.  Climatological determinants of woody cover in Africa , 2011, Proceedings of the National Academy of Sciences.

[103]  N. Drake,et al.  Ancient watercourses and biogeography of the Sahara explain the peopling of the desert , 2010, Proceedings of the National Academy of Sciences.

[104]  E. Wolff,et al.  Interglacial and glacial variability from the last 800 ka in marine, ice and terrestrial archives , 2010 .

[105]  E. Wolff,et al.  Millennial-scale variability during the last glacial: The ice core record , 2010 .

[106]  W. Ripple,et al.  Linking Top-Down Forces to the Pleistocene Megafaunal Extinctions , 2010 .

[107]  J. M. Fryxell,et al.  Foraging theory upscaled: the behavioural ecology of herbivore movement , 2010, Philosophical Transactions of the Royal Society B: Biological Sciences.

[108]  W. J. Sanders,et al.  Cenozoic Mammals of Africa , 2010 .

[109]  E. Milner‐Gulland,et al.  Tracking greenery across a latitudinal gradient in central Asia – the migration of the saiga antelope , 2010 .

[110]  J. Jouzel,et al.  Millennial and sub-millennial scale climatic variations recorded in polar ice cores over the last glacial period , 2010 .

[111]  R. Bradshaw,et al.  Factors influencing the Holocene history of Fagus , 2010 .

[112]  A. Barnosky,et al.  Timing of Quaternary megafaunal extinction in South America in relation to human arrival and climate change , 2010 .

[113]  Y. Kuzmin Extinction of the woolly mammoth (Mammuthus primigenius) and woolly rhinoceros (Coelodonta antiquitatis) in Eurasia: Review of chronological and environmental issues , 2010 .

[114]  J. Stewart,et al.  Refugia revisited: individualistic responses of species in space and time , 2010, Proceedings of the Royal Society B: Biological Sciences.

[115]  B. Noon,et al.  Pitfalls and challenges of estimating population growth rate from empirical data: consequences for allometric scaling relations , 2010 .

[116]  L. Stejneger,et al.  Results of a Biological Survey of the San Francisco Mountain Region and Desert of the Little Colorado, Arizona , 2010 .

[117]  D. Gavin The coastal‐disjunct mesic flora in the inland Pacific Northwest of USA and Canada: refugia, dispersal and disequilibrium , 2009 .

[118]  M. Pärtel,et al.  Extinction debt: a challenge for biodiversity conservation. , 2009, Trends in ecology & evolution.

[119]  S. Turvey,et al.  Probabilistic methods for determining extinction chronologies , 2009 .

[120]  A. Lézine,et al.  Plant migration and plant communities at the time of the “green Sahara” , 2009 .

[121]  M. Hofreiter,et al.  Ecological Change, Range Fluctuations and Population Dynamics during the Pleistocene , 2009, Current Biology.

[122]  A. Dobson,et al.  Food-web structure and ecosystem services: insights from the Serengeti , 2009, Philosophical Transactions of the Royal Society B: Biological Sciences.

[123]  S. Turvey,et al.  Holocene deforestation: a history of human–environmental interactions, climate change, and extinction , 2009 .

[124]  P. Tzedakis The MIS 11 - MIS 1 analogy, southern European vegetation, atmospheric methane and the , 2009 .

[125]  Kaustuv Roy,et al.  The use of sighting records to infer species extinctions: an evaluation of different methods. , 2009, Ecology.

[126]  J. Berger,et al.  Global decline in aggregated migrations of large terrestrial mammals , 2009 .

[127]  A. Lotter,et al.  Modest summer temperature variability during DO cycles in western Europe - an answer to a long time paradox? , 2009 .

[128]  John M. Fryxell,et al.  Opposing Rainfall and Plant Nutritional Gradients Best Explain the Wildebeest Migration in the Serengeti , 2009, The American Naturalist.

[129]  M. Williams,et al.  Climatic patterns in equatorial and southern Africa from 30,000 to 10,000 years ago reconstructed from terrestrial and near-shore proxy data , 2008 .

[130]  J. Russell,et al.  Northern Hemisphere Controls on Tropical Southeast African Climate During the Past 60,000 Years , 2008, Science.

[131]  M. Kearney,et al.  Birth of a biome: insights into the assembly and maintenance of the Australian arid zone biota , 2008, Molecular ecology.

[132]  B. Jones,et al.  Alluvial evidence for major climate and flow regime changes during the middle and late Quaternary in eastern central Australia , 2008 .

[133]  F. Gugerli,et al.  Genetic consequences of glacial survival and postglacial colonization in Norway spruce: combined analysis of mitochondrial DNA and fossil pollen , 2008, Molecular ecology.

[134]  A. Lister,et al.  The impact of climate change on large mammal distribution and extinction: Evidence from the last glacial/interglacial transition , 2008 .

[135]  W. Newmark Isolation of African protected areas , 2008 .

[136]  Corey J A Bradshaw,et al.  Synergies among extinction drivers under global change. , 2008, Trends in ecology & evolution.

[137]  G. Possnert,et al.  Collection of radiocarbon dates on the mammoths (Mammuthus Primigenius) and other genera of Wrangel Island, northeast Siberia, Russia , 2008, Quaternary Research.

[138]  John E. Kutzbach,et al.  Climate model tests of the anthropogenic influence on greenhouse-induced climate change: the role of early human agriculture, industrialization, and vegetation feedbacks , 2008 .

[139]  Janet M. Wilmshurst,et al.  Dating the late prehistoric dispersal of Polynesians to New Zealand using the commensal Pacific rat , 2008, Proceedings of the National Academy of Sciences.

[140]  R. Whittaker,et al.  ORIGINAL ARTICLE: A general dynamic theory of oceanic island biogeography , 2008 .

[141]  G. Possnert,et al.  Rapid ecosystem response to abrupt climate changes during the last glacial period in western Europe, 40-16 ka , 2008 .

[142]  R. Edwards,et al.  Millennial‐scale precipitation changes in southern Brazil over the past 90,000 years , 2007 .

[143]  W. Ruddiman The early anthropogenic hypothesis: Challenges and responses , 2007 .

[144]  T. McMahon,et al.  Updated world map of the Köppen-Geiger climate classification , 2007 .

[145]  Angela Schwarm,et al.  A case of non-scaling in mammalian physiology? Body size, digestive capacity, food intake, and ingesta passage in mammalian herbivores. , 2007, Comparative biochemistry and physiology. Part A, Molecular & integrative physiology.

[146]  C. Buck,et al.  A calendar chronology for Pleistocene mammoth and horse extinction in North America based on Bayesian radiocarbon calibration , 2007 .

[147]  M. McCulloch,et al.  Global sea-level fluctuations during the Last Interglaciation (MIS 5e) , 2007 .

[148]  M. Hubbe,et al.  Early Holocene survival of megafauna in South America , 2007 .

[149]  T. Stocker,et al.  Four Climate Cycles of Recurring Deep and Surface Water Destabilizations on the Iberian Margin , 2007, Science.

[150]  Peter Kershaw,et al.  A complete pollen record of the last 230 ka from Lynch's Crater, north-eastern Australia , 2007 .

[151]  P. Moss,et al.  A Quaternary marine palynological record (oxygen isotope stages 1 to 13) for the humid tropics of northeastern Australia based on ODP site 820 , 2007 .

[152]  S. Burns,et al.  A high-resolution, absolute-dated deglacial speleothem record of Indian Ocean climate from Socotra Island, Yemen , 2007, Earth and Planetary Science Letters.

[153]  R. Hilborn,et al.  Long‐Term Ecosystem Dynamics in the Serengeti: Lessons for Conservation , 2007, Conservation biology : the journal of the Society for Conservation Biology.

[154]  H. Pontzer Effective limb length and the scaling of locomotor cost in terrestrial animals , 2007, Journal of Experimental Biology.

[155]  M. Loutre,et al.  Dansgaard–Oeschger climatic variability revealed by fire emissions in southwestern Iberia , 2007 .

[156]  A. Dyke Late Quaternary Vegetation History of Northern North America Based on Pollen, Macrofossil, and Faunal Remains , 2007 .

[157]  D. Froese,et al.  Arctic ground squirrels of the mammoth-steppe: paleoecology of Late Pleistocene middens (∼24000–29450 14C yr BP), Yukon Territory, Canada , 2007 .

[158]  Flemming Skov,et al.  Ice age legacies in the geographical distribution of tree species richness in europe , 2007 .

[159]  W. Peltier,et al.  Global glacial ice volume and Last Glacial Maximum duration from an extended Barbados sea level record , 2006 .

[160]  A. Barnosky,et al.  Late Quaternary Extinctions: State of the Debate , 2006 .

[161]  S. Wroe,et al.  A review of the evidence for a human role in the extinction of Australian megafauna and an alternative interpretation , 2006 .

[162]  M. Loutre,et al.  Equatorial insolation: from precession harmonics to eccentricity frequencies , 2006 .

[163]  Kyle Joly,et al.  Simulating the influences of various fire regimes on caribou winter habitat. , 2006, Ecological applications : a publication of the Ecological Society of America.

[164]  Jingyun Fang,et al.  Climatic limits for the present distribution of beech (Fagus L.) species in the world , 2006 .

[165]  B. C. Hansen,et al.  Evidence for warm wet Heinrich events in Florida , 2006 .

[166]  R. D. Guthrie New carbon dates link climatic change with human colonization and Pleistocene extinctions , 2006, Nature.

[167]  D. Roberts,et al.  Significance of Sighting Rate in Inferring Extinction and Threat , 2006, Conservation biology : the journal of the Society for Conservation Biology.

[168]  T. Stocker,et al.  Isotope calibrated Greenland temperature record over Marine Isotope Stage 3 and its relation to CH4 , 2006 .

[169]  Alfred Russel Wallace,et al.  The geographical distribution of animals : with a study of the relations of living and extinct faunas as elucidating the past changes of the earth's surface / by Alfred Russel Wallace ; in two volumes ; with maps and illustrations. , 1876 .

[170]  P. Antoine,et al.  Lateglacial palaeoenvironments and palaeoclimates from Conty and Houdancourt, northern France, reconstructed from Beetle remains , 2005 .

[171]  W. Tinner,et al.  Rapid responses of high‐mountain vegetation to early Holocene environmental changes in the Swiss Alps , 2005 .

[172]  D. S. Glazier,et al.  Beyond the ‘3/4‐power law’: variation in the intra‐and interspecific scaling of metabolic rate in animals , 2005, Biological reviews of the Cambridge Philosophical Society.

[173]  Kate E. Jones,et al.  Multiple Causes of High Extinction Risk in Large Mammal Species , 2005, Science.

[174]  G. Milne,et al.  Modelling Holocene relative sea-level observations from the Caribbean and South America , 2005 .

[175]  Craig R. White,et al.  Allometric scaling of mammalian metabolism , 2005, Journal of Experimental Biology.

[176]  D. Bowman,et al.  One equation fits overkill: why allometry underpins both prehistoric and modern body size-biased extinctions , 2005, Population Ecology.

[177]  Guy Cowlishaw,et al.  How Far Do Animals Go? Determinants of Day Range in Mammals , 2004, The American Naturalist.

[178]  Beth Shapiro,et al.  Rise and Fall of the Beringian Steppe Bison , 2004, Science.

[179]  Walter Jetz,et al.  The Scaling of Animal Space Use , 2004, Science.

[180]  A. Lister,et al.  Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth , 2004, Nature.

[181]  A. Barnosky,et al.  Assessing the Causes of Late Pleistocene Extinctions on the Continents , 2004, Science.

[182]  S. Pääbo,et al.  Lack of phylogeography in European mammals before the last glaciation. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[183]  Henry T. Wright,et al.  A chronology for late prehistoric Madagascar. , 2004, Journal of human evolution.

[184]  James H. Brown,et al.  Toward a metabolic theory of ecology , 2004 .

[185]  J. Gaillard,et al.  What limits the Serengeti zebra population? , 2004, Oecologia.

[186]  A. Barnosky,et al.  Exceptional record of mid-Pleistocene vertebrates helps differentiate climatic from anthropogenic ecosystem perturbations. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[187]  R. D. Guthrie Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea island , 2004, Nature.

[188]  S. Hemming,et al.  Heinrich events: Massive late Pleistocene detritus layers of the North Atlantic and their global climate imprint , 2004 .

[189]  G. Coope Several million years of stability among insect species because of, or in spite of, Ice Age climatic instability? , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[190]  A. Lister The impact of Quaternary Ice Ages on mammalian evolution. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[191]  P. Bartlein,et al.  LATE‐QUATERNARY VEGETATION DYNAMICS IN NORTH AMERICA: SCALING FROM TAXA TO BIOMES , 2004 .

[192]  C. Bell,et al.  7. The Blancan, Irvingtonian, and Rancholabrean Mammal Ages , 2004 .

[193]  Richard Field,et al.  ENERGY, WATER, AND BROAD‐SCALE GEOGRAPHIC PATTERNS OF SPECIES RICHNESS , 2003 .

[194]  W. Ruddiman,et al.  The Anthropogenic Greenhouse Era Began Thousands of Years Ago , 2003 .

[195]  R. Negrini,et al.  Response of North American Great Basin Lakes to Dansgaard–Oeschger oscillations , 2003 .

[196]  F. Chapin,et al.  Non‐equilibrium succession dynamics indicate continued northern migration of lodgepole pine , 2003 .

[197]  J. Jouzel,et al.  Magnitude of isotope/temperature scaling for interpretation of central Antarctic ice cores , 2003 .

[198]  D. J. Meltzer,et al.  A requiem for North American overkill , 2003 .

[199]  D. Steadman,et al.  The late Quaternary extinction and future resurrection of birds on Pacific islands , 2003 .

[200]  Shai Meiri,et al.  On the validity of Bergmann's rule , 2003, Journal of Biogeography.

[201]  A. Solow Estimation of stratigraphic ranges when fossil finds are not randomly distributed , 2003, Paleobiology.

[202]  C. Johnson,et al.  Determinants of loss of mammal species during the Late Quaternary ‘megafauna’ extinctions: life history and ecology, but not body size , 2002, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[203]  Victor Brovkin,et al.  Climate-vegetation interaction , 2002 .

[204]  Lenore Fahrig,et al.  DISPERSAL DISTANCE OF MAMMALS IS PROPORTIONAL TO HOME RANGE SIZE , 2002 .

[205]  S. Gould The Structure of Evolutionary Theory , 2002 .

[206]  R. L. Edwards,et al.  A High-Resolution Absolute-Dated Late Pleistocene Monsoon Record from Hulu Cave, China , 2001, Science.

[207]  M. Raupach,et al.  Balances of Water, Carbon, Nitrogen and Phosphorus in Australian Landscapes: (1) Project Description and Results , 2001 .

[208]  J. Diamond,et al.  Dinosaurs, dragons, and dwarfs: The evolution of maximal body size , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[209]  D. Schreve Differentiation of the British late Middle Pleistocene interglacials: the evidence from mammalian biostratigraphy , 2001 .

[210]  H Eugene Stanley,et al.  Scaling in the growth of geographically subdivided populations: invariant patterns from a continent-wide biological survey. , 2001, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[211]  N. Stenseth,et al.  Genetic structure of Siberian lemmings (Lemmus sibiricus) in a continuous habitat: large patches rather than isolation by distance , 2001, Heredity.

[212]  D. Grayson The Archaeological Record of Human Impacts on Animal Populations , 2001 .

[213]  John Alroy,et al.  New methods for quantifying macroevolutionary patterns and processes , 2000, Paleobiology.

[214]  Glenn D. Sutherland,et al.  Scaling of Natal Dispersal Distances in Terrestrial Birds and Mammals , 2000 .

[215]  R. Holdaway,et al.  Rapid extinction of the moas (Aves: Dinornithiformes): model, test, and implications. , 2000, Science.

[216]  H. H. Birks,et al.  Two terrestrial records of rapid climatic change during the glacial-Holocene transition (14,000- 9,000 calendar years B.P.) from Europe. , 2000, Proceedings of the National Academy of Sciences of the United States of America.

[217]  Ray Hilborn,et al.  Food regulates the Serengeti wildebeest: a 40‐year record , 1999 .

[218]  Jörg Keller,et al.  Rapid environmental changes in southern Europe during the last glacial period , 1999, Nature.

[219]  James H. Brown,et al.  The fourth dimension of life: fractal geometry and allometric scaling of organisms. , 1999, Science.

[220]  K. Lambeck,et al.  Refining the eustatic sea-level curve since the Last Glacial Maximum using far- and intermediate-field sites , 1998 .

[221]  J. W. Valentine,et al.  Scales of climatic variability and time averaging in Pleistocene biotas: implications for ecology and evolution. , 1996, Trends in ecology & evolution.

[222]  I. Atkinson Introductions of wildlife as a cause of species extinctions , 1996, Wildlife Biology.

[223]  Adrian M. Lister,et al.  Ice cores and mammoth extinction , 1995, Nature.

[224]  H. Fritz,et al.  On the carrying capacity for large ungulates of African savanna ecosystems , 1994, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[225]  Graeme Caughley,et al.  Directions in conservation biology , 1994 .

[226]  J. Jouzel,et al.  Evidence for general instability of past climate from a 250-kyr ice-core record , 1993, Nature.

[227]  D. Currie,et al.  Global patterns of animal abundance and species energy use , 1993 .

[228]  A. Solow Inferring Extinction from Sighting Data , 1993 .

[229]  Patrick J. Bartlein,et al.  VEGETATION AND CLIMATE CHANGE IN EASTERN NORTH AMERICA SINCE THE LAST GLACIAL MAXIMUM , 1991 .

[230]  A. Stuart MAMMALIAN EXTINCTIONS IN THE LATE PLEISTOCENE OF NORTHERN EURASIA AND NORTH AMERICA , 1991, Biological reviews of the Cambridge Philosophical Society.

[231]  S. McNaughton,et al.  Mineral nutrition and seasonal movements of African migratory ungulates , 1990, Nature.

[232]  N. Owen‐Smith,et al.  Body Size, Population Metabolism, and Habitat Specialization Among Large African Herbivores , 1989, The American Naturalist.

[233]  R. Owen-Smith,et al.  Megaherbivores: The Influence of Very Large Body Size on Ecology , 1990 .

[234]  John M. Fryxell,et al.  Why are Migratory Ungulates So Abundant? , 1988, The American Naturalist.

[235]  Norman A. Slade,et al.  Relating Body Size to the Rate of Home Range Use in Mammals , 1988 .

[236]  K. Briffa,et al.  Seasonal temperatures in Britain during the past 22,000 years, reconstructed using beetle remains , 1987, Nature.

[237]  N. Owen‐Smith,et al.  Pleistocene extinctions: the pivotal role of megaherbivores , 1987, Paleobiology.

[238]  Mark S. Boyce,et al.  Seasonality, Fasting Endurance, and Body Size in Mammals , 1985, The American Naturalist.

[239]  S. McNaughton,et al.  Ecology of a Grazing Ecosystem: The Serengeti , 1985 .

[240]  R. Peters The Ecological Implications of Body Size , 1983 .

[241]  D. Janzen The Pleistocene Hunters Had Help , 1983, The American Naturalist.

[242]  H. Birks,et al.  Soil Development on Recent End Moraines of the Klutlan Glacier, Yukon Territory, Canada , 1980, Quaternary Research.

[243]  M. Coe,et al.  Biomass and production of large African herbivores in relation to rainfall and primary production , 1976, Oecologia.

[244]  Tom Fenchel,et al.  Intrinsic rate of natural increase: The relationship with body size , 1974, Oecologia.

[245]  I. Hiscock Communities and Ecosystems , 1970, The Yale Journal of Biology and Medicine.

[246]  L. Wollman Man and the Vertebrates , 1966 .

[247]  N. Gillett,et al.  Interactive comment on “ Ice melt , sea level rise and superstorms : evidence from paleoclimate data , climate modeling , and modern observations that 2 ◦ C global warming is highly dangerous ” , 2015 .

[248]  A. Munn,et al.  Assessing the Jarman-Bell Principle: Scaling of intake, digestibility, retention time and gut fill with body mass in mammalian herbivores. , 2013, Comparative biochemistry and physiology. Part A, Molecular & integrative physiology.

[249]  L. Polishchuk The three-quarter power scaling of extinction risk in Late Pleistocene mammals, and a new theory of the size selectivity of extinction , 2010 .

[250]  Eelco J. Rohling,et al.  High rates of sea-level rise during the last interglacial period , 2008 .

[251]  R. Aarde,et al.  Megaparks for metapopulations: Addressing the causes of locally high elephant numbers in southern Africa , 2007 .

[252]  G. Zazula,et al.  Macrofossil and pollen evidence for full-glacial steppe within an ecological mosaic along the Bluefish River, eastern Beringia , 2006 .

[253]  S. K. Lyons,et al.  Of mice, mastodons and men: human-mediated extinctions on four continents , 2004 .

[254]  R. Dale Guthrie Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea island , 2004 .

[255]  R. H. Hart Where the Buffalo Roamed – Or Did They? , 2001 .

[256]  H. H. Birks,et al.  Full-glacial upland tundra vegetation preserved under tephra in the Beringia National Park, Seward Peninsula, Alaska , 2001 .

[257]  M. Williams Morphoclimatic maps at 18 ka, 9 ka, & 0 ka , 2001 .

[258]  R. D. Guthrie Origin and causes of the mammoth steppe: a story of cloud cover, woolly mammal tooth pits, buckles, and inside-out Beringia , 2001 .

[259]  Lo Forte,et al.  Ice Age , 2001, SIGGRAPH Video Review on Electronic Theater Program.

[260]  F. Gasse Hydrological changes in the African tropics since the Last Glacial Maximum , 2000 .

[261]  C. Torrence,et al.  A Practical Guide to Wavelet Analysis. , 1998 .

[262]  K. Arslanov,et al.  Radiocarbon Dating Evidence for Mammoths on Wrangel Island, Arctic Ocean, Until 2000 BC , 1995, Radiocarbon.

[263]  G. Coope,et al.  The Devensian (Weichselian) Lateglacial palaeoenvironmental record from Gransmoor, East Yorkshire, England: A contribution to the ‘North Atlantic seaboard programme’ of IGCP-253, ‘Termination of the Pleistocene’ , 1993 .

[264]  Keith Bennett,et al.  QUATERNARY REFUGIA OF NORTH EUROPEAN TREES , 1991 .

[265]  A. Lister Frozen fauna of the mammoth steppe: the story of blue babe , 1991 .

[266]  J. Maccracken,et al.  Browse regrowth and use by moose after fire in interior Alaska. , 1990 .

[267]  R. D. Guthrie Frozen Fauna of the Mammoth Steppe , 1990 .

[268]  M. B. Davis,et al.  Climatic Instability, Time, Lags, and Community Disequilibrium , 1984 .

[269]  Claudio M. Ghersa,et al.  Weed Ecology: Implications for Management , 1984 .

[270]  K. Schmidt-Nielsen,et al.  Scaling, why is animal size so important? , 1984 .

[271]  J. V. Matthews EAST BERINGIA DURING LATE WISCONSIN TIME: A REVIEW OF THE BIOTIC EVIDENCE , 1982 .

[272]  L. Cwynar,et al.  6 – THE LATE QUATERNARY VEGETATION OF THE NORTH YUKON , 1982 .

[273]  N. Heglund,et al.  Energetics and mechanics of terrestrial locomotion. , 1982, Annual review of physiology.