Hostname: page-component-77c89778f8-swr86 Total loading time: 0 Render date: 2024-07-19T15:27:44.414Z Has data issue: false hasContentIssue false

Late Pleistocene paleoecology of arctic ground squirrel (Urocitellus parryii) caches and nests from Interior Alaska's mammoth steppe ecosystem, USA

Published online by Cambridge University Press:  20 January 2017

Benjamin V. Gaglioti*
Department of Biology and Wildlife, University of Alaska Fairbanks, 902 N. Koyukuk Dr., Fairbanks, Alaska 99775, USA Alaska Stable Isotope Facility, Water and Environmental Research Center, University of Alaska Fairbanks, 306 Tanana Dr., Fairbanks, Alaska 99775, USA
Brian M. Barnes
Department of Biology and Wildlife, University of Alaska Fairbanks, 902 N. Koyukuk Dr., Fairbanks, Alaska 99775, USA Institute of Arctic Biology, University of Alaska, Fairbanks, 902 N. Koyukuk Dr., Fairbanks, Alaska 99775, USA
Grant D. Zazula
Yukon Palaeontology Program, Department of Tourism & Culture, Government of Yukon, Box 2703 L2-A, Whitehorse, Yukon Y1A 2C6 Canada
Alwynne B. Beaudoin
Royal Alberta Museum, 12845-102nd Avenue, Edmonton, Alberta T5N 0M6 Canada
Matthew J. Wooller
Alaska Stable Isotope Facility, Water and Environmental Research Center, University of Alaska Fairbanks, 306 Tanana Dr., Fairbanks, Alaska 99775, USA Institute of Marine Science, School of Fisheries and Ocean Sciences, University of Alaska Fairbanks, 905 N. Koyukuk Dr., Fairbanks, Alaska 99775, USA Alaska Quaternary Center, 373 Reichardt Building, 900 Yukon Dr., University of Alaska Fairbanks, Fairbanks, AK 99775, USA
Corresponding author. E-mail (B. V. Gaglioti).


Botanical analyses of fossil and modern arctic ground squirrel (Urocitellus parryii) caches and nests have been used to reconstruct the past vegetation from some parts of Beringia, but such archives are understudied in Alaska. Five modern and four fossil samples from arctic ground squirrel caches and nests provide information on late Pleistocene vegetation in Eastern Beringia. Modern arctic ground squirrel caches from Alaska's arctic tundra were dominated by willow and grass leaves and grass seeds and bearberries, which were widespread in the local vegetation as confirmed by vegetation surveys. Late Pleistocene caches from Interior Alaska were primarily composed of steppe and dry tundra graminoid and herb seeds. Graminoid cuticle analysis of fossil leaves identified Calamagrostis canadensis, Koeleria sp. and Carex albonigra as being common in the fossil samples. Stable carbon isotopes analysis of these graminoid specimens indicated that plants using the C3 photosynthetic pathways were present and functioning with medium to high water-use efficiency. Fossil plant taxa and environments from ground squirrel caches in Alaska are similar to other macrofossil assemblages from the Yukon Territory, which supports the existence of a widespread mammoth steppe ecosystem type in Eastern Beringia that persisted throughout much of the late Pleistocene.

Research Article
University of Washington

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)


Ager, T.A. Late Quaternary Environmental History of the Tanana Valley, Alaska Ohio State University Institute for Polar Studies, Report 54. (1975). Columbus, Google Scholar
Armbruster, W.S., Rae, D., and Edwards, M.E. Topographic complexity and biotic response to high-latitude climate change: variance is as important as the mean. Ørbaek, J.B. et al. Arctic-Alpine Ecosystems and People in a Changing Environment. (2007). Springer, Verlag. pp. 105–121 Google Scholar
Barnosky, A.D., Koch, P.L., Feranec, R.S., Wing, S.L., and Shabel, A.B. Assessing the causes of Late Pleistocene extinctions on the continents. Science 306, (2004). 7075.Google Scholar
Berggren, G. Atlas of Seeds and Small Fruits of Northwest European Plant Species with Morphological Descriptions. (1969). Swedish Natural Science Research Council, Stockholm.Google Scholar
Bigelow, N.H., Zazula, G.D., and Atkinson, D.E. Plant Macrofossil Records / Arctic North America. Elias, S.A. Encyclopedia of Quaternary Science. (2006). Elsevier, Amsterdam. pp. 2434–2450 Google Scholar
Birks, H.J.B. Past and Present Vegetation on the Isle of Skye. (1973). University Press, London.Google Scholar
Buck, C.L., and Barnes, B.M. Annual cycle of body composition and hibernation in free-living arctic ground squirrels. Journal of Mammalogy 80, (1999). 430442.Google Scholar
Chapin, F.S. III, and McKendrick, J.D. Seasonal movement of nutrients in plants of differing growth form in an Alaskan tundra ecosystem: implications for herbivory. Journal of Ecology 68, (1980). 189209.Google Scholar
Cody, W.J. Flora of the Yukon Territory. (2000). National Research Council (NRC) Press, Ottawa.Google Scholar
Cowling, S.A. Simulated effects of low atmospheric CO2 on structure and composition of North American vegetation at the Last Glacial Maximum. Global Ecology and Biogeography 8, (1999). 8193.Google Scholar
Cowling, S.A., and Sykes, M.T. Physiological significance of low atmospheric CO2 for plant-climate interactions. Quaternary Research 52, (1999). 237242.Google Scholar
Cwynar, L.C., and Ritchie, J.C. Arctic steppe tundra — a Yukon perspective. Science 208, (1980). 13751377.Google Scholar
Dallwitz, M.J. A general system for coding taxonomic descriptions. Taxon 29, (1980). 4146.CrossRefGoogle Scholar
Dial, K.P., and Czaplewski, N.J. Do woodrat middens accurately represent the animal's environments and diets?. Betancourt, J.L., Van Devender, T.R., and Martin, P.S. Packrat Middens. (1990). University of Arizona Press, Phoenix. 4358.Google Scholar
Edwards, M.E., and Armbruster, W.S. A tundra-steppe transition on Kathul Mountain, Alaska, USA. Arctic and Alpine Research 21, (1989). 296304.Google Scholar
Elias, S.A. Mutual climatic range reconstructions of seasonal temperatures based on Late Pleistocene fossil beetle assemblages in Eastern Beringia. Quaternary Science Reviews 20, (2001). 7791.Google Scholar
Elias, S.A., Short, S.K., and Birks, H.H. Late Wisconsin environments of the Bering land bridge. Palaeogeography, Palaeoclimatology, Palaeoecology 196, (1997). 293308.CrossRefGoogle Scholar
Farquhar, G.D., Ehleringer, J.R., and Hubick, K.T. Carbon isotope discrimination and photosynthesis. Annual Review of Plant Physiology and Plant Molecular Biology 40, (1989). 503537.CrossRefGoogle Scholar
Friedli, H., Lotscher, H., Oeschger, H., Siegenthaler, U., and Stauffer, B. Ice Core Records of the 13C/12C ratio of atmospheric CO2 in the past two centuries. Nature 324, (1986). 237238.Google Scholar
Gaglioti, B.V., Severin, K., and Wooller, M.J. Developing graminoid cuticle analysis for application to Beringian palaeoecology. Review of Palaeobotany and Palynology 162, (2010). 95110.CrossRefGoogle Scholar
Gillis, E.A., Morrison, S.F., Zazula, G.D., and Hik, D.S. Evidence for selective caching by arctic ground squirrels living in alpine meadows in the Yukon. Arctic 58, 8 (2005). 354360.Google Scholar
Goetcheus, V.G., and Birks, H.H. Full-glacial upland tundra vegetation preserved under tephra in the Beringia National Park, Seward Peninsula. Quaternary Science Reviews 20, (2001). 135147.CrossRefGoogle Scholar
Gower, J.C. A general coefficient of similarity and some of its properties. Biometrics 27, (1971). 657671.Google Scholar
Gubin, S.V.K. Fossil burrows of mammals in the loess-ice deposits of the Kolyma-Indigirka lowland. Doklady Biological Sciences 346, (1996). 2627.Google Scholar
Guthrie, R.D. Paleoecology of a Late Pleistocene large-mammal community from Interior Alaska. American Midland Naturalist 79, 2 (1968). 346363.Google Scholar
Guthrie, R.D. Mammals of the mammoth steppe as paleoenvironmental indicators. Hopkins, D.M., Schweger, C.E., and Young, S.B. Paleoecology of Beringia. (1982). Academic Press, New York. 307329.Google Scholar
Guthrie, R.D. Frozen Fauna of the Mammoth Steppe: The Story of Blue Babe. (1990). University of Chicago Press, Chicago.Google Scholar
Guthrie, R.D. Origin and causes of the mammoth steppe: a story of cloud cover, woolly mammal tooth pits, buckles, and inside-out Beringia. Quaternary Science Reviews 20, (2001). 549574.CrossRefGoogle Scholar
Helgen, K.M., Cole, F.R., Helgen, L.E., and Wilson, D.E. Generic revision in the Holarctic ground squirrel genus Spermophilus . Journal of Mammalogy 90, 2 (2009). 270305.Google Scholar
Hultén, E. Flora of Alaska. (1968). Stanford University Press, Stanford.Google Scholar
Körber-Grohne, U. Bestimmungsschüssel für subfossile Gramineae-Früchte. In Probleme der Küstendforschung im Südlichen Nordseegebiet. Band 18, (1991). 169234.Google Scholar
Krog, J. Storing of food items in the winter nest of the Alaskan ground squirrel, Citellus undulatas . Journal of Mammalogy 35, (1954). 586 Google Scholar
Kubien, D.S., and Sage, R.F. Low-temperature photosynthetic performance of a C4 grass and a co-occurring C3 grass native to high latitudes. Plant, Cell & Environment 27, (2004). 907916.Google Scholar
Laxton, N.F., Burn, C.R., and Smith, C.A.S. Productivity of loessal grasslands in the Kluane Lake region, Yukon Territory, and the Beringian “production paradox”. Arctic 49, (1996). 129140.Google Scholar
Lloyd, A.H., Armbruster, W.S., and Edwards, M.E. Ecology of a steppe-tundra gradient in Interior Alaska. Journal of Vegetation Science 5, (1994). 897912.Google Scholar
Martinson, D.G., Pisias, N.G., Hays, J.D., Imbrie, J., Moore, T.C. Jr., and Shackleton, N.J. Age dating and the orbital theory of the Ice Ages — development of a high-resolution 0 to 300,000-year chronostratigraphy. Quaternary Research 27, (1987). 129.CrossRefGoogle Scholar
Matthews, J.V. Wisconsin environment of Interior Alaska: pollen and macrofossil analysis of a 27 meter core from the Isabella Basin (Fairbanks, Alaska). Canadian Journal of Earth Sciences 11, (1974). 828841.CrossRefGoogle Scholar
Montgomery, F.H. Seeds and Fruits of Plants of Eastern Canada and Northeastern United States. (1977). University of Toronto Press, Toronto.CrossRefGoogle Scholar
O'Donnell, J.A., Harden, J.W., McGuire, D.A., Kanevskiy, M.Z., Jorgenson, M.T., and Xiaomei, X. The effect of fire and permafrost interactions on soil carbon accumulation in an upland black spruce ecosystem of Interior Alaska: implications for post-thaw carbon loss. Global Change Biology 17, (2011). 14611474.Google Scholar
Palmer, P.G. Grass cuticles: a new paleoecological tool for East African lake sediments. Canadian Journal of Botany 54, 15 (1976). 17251734.CrossRefGoogle Scholar
Palmer, P.G., Tucker, A.E., and Gerbeth-Jones, S. A Scanning Electron Microscope Survey of the Epidermis of East African Grasses. (1981). Smithsonian Institution Press, Washington.Google Scholar
Pitulko, V.V., Nikolsky, P.A., Girya, E.Y., Basilyan, A.E., Tumskoy, V.E., Koulakov, S.A., Astakhov, S.N., Pavlova, E.Y., and Anisimov, M.A. The Yana RHS site: humans in the Arctic before the Last Glacial Maximum. Science 303, (2004). 5256.Google Scholar
Stuiver, M., Reimer, P.J., and Reimer, R.W. CALIB 6.0. [WWW program and documentation;]. (2009). Google Scholar
Vander Wall, S.B. Food Hoarding in Animals. (1990). University of Chicago Press, Chicago.Google Scholar
Vetter, M.A. Grasslands of the Aishihik-Sekulmun Lakes area, Yukon Territory, Canada. Arctic 53, (2000). 165173.CrossRefGoogle Scholar
Williams, J.W., and Jackson, S.T. Novel climates, no-analog communities, and ecological surprises. Frontiers in Ecology and the Environment 5, (2007). 475482.Google Scholar
Wooller, M.J., Swain, D.L., Ficken, K.J., Agnew, A.D.Q., Street-Perrott, F.A., and Eglinton, G. Late Quaternary vegetation changes around Lake Rutundu, Mount Kenya, East Africa: evidence from grass cuticles, pollen and stable carbon isotopes. Journal of Quaternary Science 18, (2003). 315.Google Scholar
Wooller, M.J., Zazula, G.D., Edwards, M.E., Froese, D.G., Boone, R.D., Parker, C., and Bennett, B. Stable carbon isotope compositions of Eastern Beringian grasses and sedges: investigating their potential as paleoenvironmental indicators. Arctic, Antarctic and Alpine Research 39, (2007). 318331.CrossRefGoogle Scholar
Wooller, M. J., Zazula, G. D., Blinnikov, M., Gaglioti, B. V., Bigelow, N. H., Sanborn, P., Kuzmina, S., and La Farge, C. (2011). Paleoecology of a Late Pleistocene (31,000 14C years) paleo-turf from interior Alaska. J. Quat. Sci. Corrected Proof. doi:10.1002/jqs.1497.Google Scholar
Young, S.B. The vegetation of the Bering Land Bridge. Hopkins, D.M., Schweger, C.E., and Young, S.B. Paleoecology of Beringia. (1982). Academic Press, New York. 179194.Google Scholar
Yurtsev, B.A. Relicts of xerophyte vegetation of Beringia in Northeastern Asia. Hopkins, D.M., Schweger, C.E., and Young, S.B. Paleoecology of Beringia. (1982). Academic Press, New York. 157174.Google Scholar
Yurtsev, B.A. The Pleistocene “Tundra-Steppe” and the productivity paradox: the landscape approach. Quaternary Science Reviews 20, (2001). 165174.Google Scholar
Zazula, G.D., Froese, D.G., Westgate, J.A., La Farge, C., and Mathewes, R.W. Paleoecology of Beringian “packrat” middens from central Yukon territory, Canada. Quaternary Research 63, (2005). 189198.CrossRefGoogle Scholar
Zazula, G.D., Froese, D.G., Elias, S.E., Kuzmina, S., Lafarge, C., Reyes, A.V., Sanborn, P.T., Schweger, C.E., Smith, C.A.S., and Mathewes, R.W. Vegetation buried under Dawson tephra (25,300 14C years BP) and locally diverse late Pleistocene paleoenvironments of Goldbottom Creek, Yukon, Canada. Palaeogeography, Palaeoclimatology, Palaeoecology 242, (2006). 253286.Google Scholar
Zazula, G.D., Mathewes, R.W., and Harestad, A.S. Cache selection by arctic ground squirrels inhabiting boreal-steppe meadows of Southwest Yukon Territory, Canada. Arctic, Antarctic and Alpine Research 38, (2006). 631638.Google Scholar
Zazula, G.D., Schweger, C.E., Beaudoin, A.B., and McCourt, G.H. Macrofossil and pollen evidence for full-glacial steppe within an ecological mosaic along the Bluefish River, Eastern Beringia. Quaternary International 142, (2006). 219.Google Scholar
Zazula, G.D., Froese, D.G., Elias, S.E., Kuzmina, S., and Mathewes, R.W. Arctic ground squirrels of the mammoth-steppe: paleoecology of Late Pleistocene middens (~ 24000-29450 14C yr BP), Yukon Territory, Canada. Quaternary Science Reviews 26, (2007). 9791003.Google Scholar
Zazula, G.D., Froese, D.G., Elias, S.A., Kuzmina, S., and Mathewes, R.W. Early Wisconsinan (MIS 4) Arctic ground squirrel middens and a squirrel-eye-view of the mammoth-steppe. Quaternary Science Reviews 30, (2010). 22202237.CrossRefGoogle Scholar
Zhang, Z., Meixun, Z., Huayu, L., and Falla, A.M. Lower temperature as the main cause of C4 plant declines during the glacial periods on the Chinese Loess Plateau. Earth and Planetary Science Letters 214, (2003). 467481.Google Scholar
Zimov, N.S., Zimov, S.A., Zimova, A.E., Zimova, G.M., Chuprynin, V.I., and Chapin, F.S. Carbon storage in permafrost and soils of the mammoth tundra-steppe biome: role in the global carbon budget. Geophysical Research Letters 36, (2009). L02502 CrossRefGoogle Scholar