<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "https://jats.nlm.nih.gov/nlm-dtd/publishing/3.0/journalpublishing3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article" dtd-version="3.0" xml:lang="en">
<front>
<journal-meta>
<journal-id journal-id-type="publisher">CP</journal-id>
<journal-title-group>
<journal-title>Climate of the Past</journal-title>
<abbrev-journal-title abbrev-type="publisher">CP</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Clim. Past</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1814-9332</issn>
<publisher><publisher-name>Copernicus Publications</publisher-name>
<publisher-loc>Göttingen, Germany</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.5194/cp-3-109-2007</article-id>
<title-group>
<article-title>Predicting Pleistocene climate from vegetation in North America</article-title>
</title-group>
<contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Loehle</surname>
<given-names>C.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
</contrib-group><aff id="aff1">
<label>1</label>
<addr-line>National Council for Air and Stream Improvement, Inc., 552 S Washington St., #224, Naperville, IL 60540, USA</addr-line>
</aff>
<pub-date pub-type="epub">
<day>12</day>
<month>02</month>
<year>2007</year>
</pub-date>
<volume>3</volume>
<issue>1</issue>
<fpage>109</fpage>
<lpage>118</lpage>
<permissions>
<copyright-statement>Copyright: &#x000a9; 2007 C. Loehle</copyright-statement>
<copyright-year>2007</copyright-year>
<license license-type="open-access">
<license-p>This work is licensed under the Creative Commons Attribution-NonCommercial-ShareAlike 2.5 Generic License. To view a copy of this licence, visit <ext-link ext-link-type="uri"  xlink:href="https://creativecommons.org/licenses/by-nc-sa/2.5/">https://creativecommons.org/licenses/by-nc-sa/2.5/</ext-link></license-p>
</license>
</permissions>
<self-uri xlink:href="https://cp.copernicus.org/articles/3/109/2007/cp-3-109-2007.html">This article is available from https://cp.copernicus.org/articles/3/109/2007/cp-3-109-2007.html</self-uri>
<self-uri xlink:href="https://cp.copernicus.org/articles/3/109/2007/cp-3-109-2007.pdf">The full text article is available as a PDF file from https://cp.copernicus.org/articles/3/109/2007/cp-3-109-2007.pdf</self-uri>
<abstract>
<p>Climates at the Last Glacial Maximum have been inferred from fossil pollen
assemblages, but these inferred climates are colder for eastern North
America than those produced by climate simulations. It has been suggested
that low CO&lt;sub&gt;2&lt;/sub&gt; levels could account for this discrepancy. In this study
biogeographic evidence is used to test the CO&lt;sub&gt;2&lt;/sub&gt; effect model. The
recolonization of glaciated zones in eastern North America following the
last ice age produced distinct biogeographic patterns. It has been assumed
that a wide zone south of the ice was tundra or boreal parkland
(Boreal-Parkland Zone or BPZ), which would have been recolonized from
southern refugia as the ice melted, but the patterns in this zone differ
from those in the glaciated zone, which creates a major biogeographic
anomaly. In the glacial zone, there are few endemics but in the BPZ there
are many across multiple taxa. In the glacial zone, there are the expected
gradients of genetic diversity with distance from the ice-free zone, but no
evidence of this is found in the BPZ. Many races and related species exist
in the BPZ which would have merged or hybridized if confined to the same
refugia. Evidence for distinct southern refugia for most temperate species
is lacking. Extinctions of temperate flora were rare. The interpretation of
spruce as a boreal climate indicator may be mistaken over much of the region
if the spruce was actually an extinct temperate species. All of these
anomalies call into question the concept that climates in the zone south of
the ice were extremely cold or that temperate species had to migrate far to
the south. An alternate hypothesis is that low CO&lt;sub&gt;2&lt;/sub&gt; levels gave an
advantage to pine and spruce, which are the dominant trees in the BPZ, and
to herbaceous species over trees, which also fits the observed pattern. Thus
climate reconstruction from pollen data is probably biased and needs to
incorporate CO&lt;sub&gt;2&lt;/sub&gt; effects. Most temperate species could have survived
across their current ranges at lower abundance by retreating to moist
microsites. These would be microrefugia not easily detected by pollen
records, especially if most species became rare. These results mean that
climate reconstructions based on terrestrial plant indicators will not be
valid for periods with markedly different CO&lt;sub&gt;2&lt;/sub&gt; levels.</p>
</abstract>
<counts><page-count count="10"/></counts>
</article-meta>
</front>
<body/>
<back>
<ref-list>
<title>References</title>
<ref id="ref1">
<label>1</label><mixed-citation publication-type="other" xlink:type="simple"> Austin, J. D., Lougheed, S. C., Neidrauer, L., Chek, A. A., and Boag, P. T.: Cryptic lineages in a small frog: The post-glacial history of the spring peeper, \textitPseudacris crucifer (Anura: Hylidae), Molecular Phylogenetics and Evolution, 25, 316&amp;ndash;329, 2002. </mixed-citation>
</ref>
<ref id="ref2">
<label>2</label><mixed-citation publication-type="other" xlink:type="simple"> Austin, J. D., Lougheed, S. C., and Boag, P. T.: Discordant temporal and geographic patterns in maternal lineages of eastern North American frogs, \textitRana castebeiana (Ranidae) and \textitPseudacris crucifer (Hylidae), Molecular Phylogenetics and Evolution, 32, 799&amp;ndash;816, 2004. </mixed-citation>
</ref>
<ref id="ref3">
<label>3</label><mixed-citation publication-type="other" xlink:type="simple"> Baskin, J. M. and Baskin, C. C.: Distribution and geographical/evolutionary relationships of cedar glade endemics in southeastern United States, ASB Bulletin, 33, 138&amp;ndash;154, 1986. </mixed-citation>
</ref>
<ref id="ref4">
<label>4</label><mixed-citation publication-type="other" xlink:type="simple"> Baskin, J. M. and Baskin, C. C.: Cedar glade endemics in Tennessee, and a review of their autecology, J. Tennessee Academy Sci., 64, 63&amp;ndash;74, 1989. </mixed-citation>
</ref>
<ref id="ref5">
<label>5</label><mixed-citation publication-type="other" xlink:type="simple"> Beerling, D. J.: Ecophysiological responses of woody plants to past CO&lt;sub&gt;2&lt;/sub&gt; concentrations, Tree Physiology, 16, 389&amp;ndash;396, 1996. </mixed-citation>
</ref>
<ref id="ref6">
<label>6</label><mixed-citation publication-type="other" xlink:type="simple"> Beerling, D. J. and Woodward, F. I..: Ecophysiological responses of plants to global environmental change since the Last Glacial Maximum, New Phytologist, 125, 641&amp;ndash;648, 1993. </mixed-citation>
</ref>
<ref id="ref7">
<label>7</label><mixed-citation publication-type="other" xlink:type="simple"> Bennett, K. D.: The spread of \textitFagus grandifolia across eastern North America during the last 18 000 years, J. Biogeogr., 12, 147&amp;ndash;164, 1985. </mixed-citation>
</ref>
<ref id="ref8">
<label>8</label><mixed-citation publication-type="other" xlink:type="simple"> Bennett, K. D. and Willis, K. J.: Effect of global atmospheric carbon dioxide on glacial-interglacial vegetation change, Global Ecol. Biogeogr., 9, 355&amp;ndash;361, 2000. </mixed-citation>
</ref>
<ref id="ref9">
<label>9</label><mixed-citation publication-type="other" xlink:type="simple"> Black, R. F.: Periglacial features indicative of permafrost: Ice and soil wedges, Quaternary Res., 6, 3&amp;ndash;26, 1976. </mixed-citation>
</ref>
<ref id="ref10">
<label>10</label><mixed-citation publication-type="other" xlink:type="simple"> Bradley, R. S.: Paleoclimatology: Reconstructing climates of the Quaternary, Second edition, Elsevier Academic Press, Amsterdam, 1999. </mixed-citation>
</ref>
<ref id="ref11">
<label>11</label><mixed-citation publication-type="other" xlink:type="simple"> Brant, S. V. and Ortí, G.: Phylogeography of the northern short-tailed shrew, \textitBlarina brevicauda (Insectivora: Soricidae): Past fragmentation and postglacial recolonization, Molecular Ecol., 12, 1435&amp;ndash;1449, 2003. </mixed-citation>
</ref>
<ref id="ref12">
<label>12</label><mixed-citation publication-type="other" xlink:type="simple"> Braun, E. L.: \textitDeciduous forests of eastern North America, Hafner Press, London, 1950. </mixed-citation>
</ref>
<ref id="ref13">
<label>13</label><mixed-citation publication-type="other" xlink:type="simple"> Braun, E. L.: Plant distribution in relation to the glacial boundary, Ohio J. Sci., 51, 139&amp;ndash;146, 1951. </mixed-citation>
</ref>
<ref id="ref14">
<label>14</label><mixed-citation publication-type="other" xlink:type="simple"> Brown, J. H. and Gibson, A. C.: Biogeography, C. V. Mosby Co., St. Louis, MO, 1983. </mixed-citation>
</ref>
<ref id="ref15">
<label>15</label><mixed-citation publication-type="other" xlink:type="simple"> Bryant, V.M., Jr.: A 16,000 year pollen record of vegetation change in central Texas, Palynology, 1, 143-156, 1977. </mixed-citation>
</ref>
<ref id="ref16">
<label>16</label><mixed-citation publication-type="other" xlink:type="simple"> Bryant, V. M. and Holloway, R. G.: A late Quaternary paleoenvironmental record of Texas: An overview of the pollen evidence, 39-70 in V.M. Bryant and R.G. Holloway (eds.), Pollen records of late Quaternary North American sediments, American Association of Stratigraphic Palynologists Foundation, Dallas, 1985. </mixed-citation>
</ref>
<ref id="ref17">
<label>17</label><mixed-citation publication-type="other" xlink:type="simple"> Burns, G. W.: Wisconsin age forests in western Ohio: II, Vegetation and burial conditions, Ohio J. Sci., 58, 220&amp;ndash;230, 1958. </mixed-citation>
</ref>
<ref id="ref18">
<label>18</label><mixed-citation publication-type="other" xlink:type="simple"> Burns, R. M. and Honkala, B. H.: Silvics of North America, Vol. 1, Conifers, USDA Handbook 654, 1990. </mixed-citation>
</ref>
<ref id="ref19">
<label>19</label><mixed-citation publication-type="other" xlink:type="simple"> Cain, S. A.: Foundations of plant geography, Harper &amp; Row, New York, 1944. </mixed-citation>
</ref>
<ref id="ref20">
<label>20</label><mixed-citation publication-type="other" xlink:type="simple"> Cain, M. L., Damman, H., and Muir, A.: Seed dispersal and the Holocene migration of woodland herbs, Ecol. Monogr., 68, 325&amp;ndash;347, 1998. </mixed-citation>
</ref>
<ref id="ref21">
<label>21</label><mixed-citation publication-type="other" xlink:type="simple"> Camp, W. H.: A biogeographic and paragenetic analysis of the American beech (\textitFagus), 166&amp;ndash;169 in Yearbook of the American Philosophical Society, 1950. </mixed-citation>
</ref>
<ref id="ref22">
<label>22</label><mixed-citation publication-type="other" xlink:type="simple"> Church, S. A., Kraus, J. M., Mitchell, J. C., Church, D. R., and Taylor, D. R.: Evidence for multiple Pleistocene refugia in the postglacial expansion of the eastern tiger salamander, \textitAmbystoma tigrinum tigrinum, Evolution, 57, 372&amp;ndash;383, 2003. </mixed-citation>
</ref>
<ref id="ref23">
<label>23</label><mixed-citation publication-type="other" xlink:type="simple"> Clayton, L., Attig, J. W., and Mickelson, D. M.: Effects of late Pleistocene permafrost on the landscape of Wisconsin, USA, Boreas, 30, 173&amp;ndash;188, 2001. </mixed-citation>
</ref>
<ref id="ref24">
<label>24</label><mixed-citation publication-type="other" xlink:type="simple"> Cole, D. R. and Monger, H. C.: Influence of atmospheric CO&lt;sub&gt;2&lt;/sub&gt; on the decline of C&lt;sub&gt;4&lt;/sub&gt; plants during the last deglaciation, Nature, 368, 533&amp;ndash;536, 1994. </mixed-citation>
</ref>
<ref id="ref25">
<label>25</label><mixed-citation publication-type="other" xlink:type="simple"> Cooper, A. W. and Mercer, E. P.: Morphological variation in \textitFagus grandifolia Ehrh. in North Carolina, Journal of Elisha Mitchell Scientific Society, 93, 136&amp;ndash;149, 1977. </mixed-citation>
</ref>
<ref id="ref26">
<label>26</label><mixed-citation publication-type="other" xlink:type="simple"> Cowling, S. A .: Simulated effects of low atmospheric CO&lt;sub&gt;2&lt;/sub&gt; on structure and composition of North American vegetation at the Last Glacial Maximum, Global Ecol. Biogeogr., 8, 81&amp;ndash;93, 1999. </mixed-citation>
</ref>
<ref id="ref27">
<label>27</label><mixed-citation publication-type="other" xlink:type="simple"> Cowling, S. A. and Sykes, M. T.: Physiological significance of low atmospheric CO&lt;sub&gt;2&lt;/sub&gt; for plant-climate interactions, Quaternary Res., 52, 237&amp;ndash;242, 1999. </mixed-citation>
</ref>
<ref id="ref28">
<label>28</label><mixed-citation publication-type="other" xlink:type="simple"> Crowley, T. J.: CLIMAP SSTs re-visited, Clim. Dyn., 16, 241&amp;ndash;255, 2000. </mixed-citation>
</ref>
<ref id="ref29">
<label>29</label><mixed-citation publication-type="other" xlink:type="simple"> Davis, M. B.: Quaternary history of deciduous forests of eastern North America and Europe, Annals of the Missouri Botanical Garden, 70, 550&amp;ndash;563, 1983. </mixed-citation>
</ref>
<ref id="ref30">
<label>30</label><mixed-citation publication-type="other" xlink:type="simple"> Davis, M. B.: Holocene vegetational history of the eastern United States, 166&amp;ndash;181, in: Late-Quaternary environments of the United States, edited by: Wright Jr., H. E., Vol. 2, The Holocene, Longman, London, 1984. </mixed-citation>
</ref>
<ref id="ref31">
<label>31</label><mixed-citation publication-type="other" xlink:type="simple"> Davis, M. B. and Shaw, R. G.: Range shifts and adaptive responses to quaternary climate change, Science, 292, 673&amp;ndash;679, 2001. </mixed-citation>
</ref>
<ref id="ref32">
<label>32</label><mixed-citation publication-type="other" xlink:type="simple"> Deevey Jr., E. S.: Biogeography of the Pleistocene, Geological Soc. Am. Bull., 60, 1315&amp;ndash;1416, 1949. </mixed-citation>
</ref>
<ref id="ref33">
<label>33</label><mixed-citation publication-type="other" xlink:type="simple"> Delcourt, H. R. and Delcourt, P. A.: Ice age haven for hardwoods, Natural History, 84, 22&amp;ndash;28, 1984. </mixed-citation>
</ref>
<ref id="ref34">
<label>34</label><mixed-citation publication-type="other" xlink:type="simple"> Delcourt, P. A. and Delcourt, H. R.: Paleoclimates, paleovegetation, and paleofloras during the late Quaternary, 71&amp;ndash;94 in Flora of North America Editorial Committee (ed.), Flora of North America, Oxford University Press, New York, 1993. </mixed-citation>
</ref>
<ref id="ref35">
<label>35</label><mixed-citation publication-type="other" xlink:type="simple"> Denny, C. S.: Pleistocene frost action near the border of the Wisconsin drift in Pennsylvania, Ohio J. Sci., 51, 116&amp;ndash;125, 1951. </mixed-citation>
</ref>
<ref id="ref36">
<label>36</label><mixed-citation publication-type="other" xlink:type="simple"> Drake, B. G., Gonzàlez-Meler, M. A., and Long, S. P.: More efficient plants: A consequence of rising atmospheric CO&lt;sub&gt;2&lt;/sub&gt;?, Ann. Rev. Plant Physiol. Plant Molecular Biol., 48, 609&amp;ndash;639, 1997. </mixed-citation>
</ref>
<ref id="ref37">
<label>37</label><mixed-citation publication-type="other" xlink:type="simple"> Elenga, H., Peyron, O., Bonnefille, R., Jolly, D., Cheddadi, R., Guiot, J., Andrieu, V., Bottema, S., Buchet, G., de Beaulieu, J.-L., Hamilton, A.C., Maley, J., Marchant, R., Perez-Obiol, R., Reille, M., Riollet, G., Scott, L., Straka, H., Taylor, D., Van Campo, E., Vincens, A., Laarif, F., and Jonson, H.: Pollen-based biome reconstruction for southern Europe and Africa 18 000 yr bp, J. Biogeogr., 27, 621&amp;ndash;634, 2000. </mixed-citation>
</ref>
<ref id="ref38">
<label>38</label><mixed-citation publication-type="other" xlink:type="simple"> Estill, J. C. and Cruzan, M. B.: Phytogeography of rare plant species endemic to the southeastern United States, Castanea, 66, 3&amp;ndash;23, 2001. </mixed-citation>
</ref>
<ref id="ref39">
<label>39</label><mixed-citation publication-type="other" xlink:type="simple"> Farquhar, G. D.: Carbon dioxide and vegetation, Science, 278, 1411, 1997. </mixed-citation>
</ref>
<ref id="ref40">
<label>40</label><mixed-citation publication-type="other" xlink:type="simple"> Farrera, I., Harrison, S. P., Prentice, I. C., Ramstein, G., Guiot, J., Bartlein, P. J., Bonnefille, R., Bush, M., Cramer, W., von Grafenstein, U., Holmgren K., Hooghiemstra, H., Hope, G., Jolly, D., Lauritzen, S.-E., Ono, Y., Pinot, S., Stute, M., and Yu, G.: Tropical climates at the Last Glacial Maximum: A new synthesis of terrestrial palaeoclimate data. I. Vegetation, lake-levels and geochemistry, Clim. Dyn., 15, 823&amp;ndash;856, 1999. </mixed-citation>
</ref>
<ref id="ref41">
<label>41</label><mixed-citation publication-type="other" xlink:type="simple"> Finkelstein, S. A., Gajewski, K., and Viau, A. E.: Improved resolution of pollen taxonomy allows better biogeographical interpretation of post-glacial forest development: Analyses from the North American Pollen Database, J. Ecol., 94, 415&amp;ndash;430, 2006. </mixed-citation>
</ref>
<ref id="ref42">
<label>42</label><mixed-citation publication-type="other" xlink:type="simple"> Froyd, C. A.: Fossil stomata reveal early pine presence in Scotland: Implications for postglacial colonization analyses, Ecology, 86, 579&amp;ndash;586, 2005. </mixed-citation>
</ref>
<ref id="ref43">
<label>43</label><mixed-citation publication-type="other" xlink:type="simple"> Gamache, I., Jaramillo-Correa, J. P., Payette, S., and Bousquet, J.: Diverging patterns of mitochondrial and nuclear DNA diversity in subarctic black spruce: Imprint of a founder effect associated with postglacial colonization, Molecular Ecol., 12, 891&amp;ndash;901, 2003. </mixed-citation>
</ref>
<ref id="ref44">
<label>44</label><mixed-citation publication-type="other" xlink:type="simple"> Ganopolski, A., Rahmstorf, S., Petoukhov, V., and Claussen, M.: Simulation of modern and glacial climates with a coupled global model of intermediate complexity, Nature, 391, 351&amp;ndash;356, 1998. </mixed-citation>
</ref>
<ref id="ref45">
<label>45</label><mixed-citation publication-type="other" xlink:type="simple"> Gilbert, C. R.: Zoogeography of the freshwater fish fauna of southern Georgia and peninsular Florida, Brimleyana, 13, 25&amp;ndash;54, 1987. </mixed-citation>
</ref>
<ref id="ref46">
<label>46</label><mixed-citation publication-type="other" xlink:type="simple"> Givens, C. R., and Givens, F. M.: Age and significance of fossil white spruce (\textitPicea glauca), Tunica Hills, Louisiana-Mississippi, Quaternary Res., 27, 283&amp;ndash;296, 1987. </mixed-citation>
</ref>
<ref id="ref47">
<label>47</label><mixed-citation publication-type="other" xlink:type="simple"> Gooding, A. M.: Illinoian and Wisconsin glaciations in the Whitewater basin, southeastern Indiana, and adjacent areas, J. Geol., 71, 665&amp;ndash;682, 1963. </mixed-citation>
</ref>
<ref id="ref48">
<label>48</label><mixed-citation publication-type="other" xlink:type="simple"> Greene, A. M., Seager, R. and Broecker, W. S.: Tropical snowline depression at the Last Glacial Maximum: Comparison with proxy records using a single-cell tropical climate model, J. Geophys. Res., 107(D8), https://doi.org/10.1029/2001JD000670, 2002. </mixed-citation>
</ref>
<ref id="ref49">
<label>49</label><mixed-citation publication-type="other" xlink:type="simple"> Grüger, E.: Late Quaternary vegetational development in south-central Illinois, Quaternary Res., 2, 217&amp;ndash;231, 1972a. </mixed-citation>
</ref>
<ref id="ref50">
<label>50</label><mixed-citation publication-type="other" xlink:type="simple"> Grüger, E.: Pollen and seed studies of Wisconsinan vegetation in Illinois, USA, Geological Soc. Am. Bull., 83, 2715&amp;ndash;2734, 1972b. </mixed-citation>
</ref>
<ref id="ref51">
<label>51</label><mixed-citation publication-type="other" xlink:type="simple"> Guiot, J., Torre, F., Jolly, D., Peyron, O., Boreux, J. J., and Cheddadi, R.: Inverse vegetation modeling by Monte Carlo sampling to reconstruct palaeoclimates under changed precipitation seasonality and CO&lt;sub&gt;2&lt;/sub&gt; conditions: application to glacial climate in Mediterranean region. Ecol. Modell., 127, 119&amp;ndash;140, 2000. </mixed-citation>
</ref>
<ref id="ref52">
<label>52</label><mixed-citation publication-type="other" xlink:type="simple"> Harrison, S. P. and Prentice, C. I.: Climate and CO&lt;sub&gt;2&lt;/sub&gt; controls on global vegetation distribution at the last glacial maximum: Analysis based on palaeovegetation data, biome modeling and palaeoclimate simulations, Global Change Biol., 9, 983&amp;ndash;1004, 2003. </mixed-citation>
</ref>
<ref id="ref53">
<label>53</label><mixed-citation publication-type="other" xlink:type="simple"> Harvill Jr., A.M.: Phytogeography of the Virginias and the equilibrium concept of landscape, Castanea, 38, 266&amp;ndash;268, 1973. </mixed-citation>
</ref>
<ref id="ref54">
<label>54</label><mixed-citation publication-type="other" xlink:type="simple"> Hewitt, G. M.: Genetic consequences of climatic oscillations in the Quaternary, Philosophical Transactions of the Royal Society of London B, 359, 183&amp;ndash;195, 2004. </mixed-citation>
</ref>
<ref id="ref55">
<label>55</label><mixed-citation publication-type="other" xlink:type="simple"> Hultén, E.: Outline of the history of Arctic and boreal biota during the Quaternary Period, Bokforlags Aktiebolager Thule, Stockholm, 1937. </mixed-citation>
</ref>
<ref id="ref56">
<label>56</label><mixed-citation publication-type="other" xlink:type="simple"> Huntley, B., Alfano, M. J., Allen, J. R. M., Pollard, D., Tzedakis, P. C., de Beaulieu, J.-L., Grüger, E., and Watts, B.: European vegetation during Marine Oxygen Isotope Stage-3, Quaternary Res., 59, 195&amp;ndash;212, 2003. </mixed-citation>
</ref>
<ref id="ref57">
<label>57</label><mixed-citation publication-type="other" xlink:type="simple"> Jackson, S. T. and Givens, C. R.: Late Wisconsinan vegetation and environment of the Tunica Hills regions, Louisiana and Mississippi, Quaternary Res., 41, 316&amp;ndash;325, 1994. </mixed-citation>
</ref>
<ref id="ref58">
<label>58</label><mixed-citation publication-type="other" xlink:type="simple"> Jackson, S. T. and Weng, C.: Late Quaternary extinction of a tree species in eastern North America, PNAS, 96, 13 847&amp;ndash;13 852, 1999. </mixed-citation>
</ref>
<ref id="ref59">
<label>59</label><mixed-citation publication-type="other" xlink:type="simple"> Jacobson Jr., G. L., Webb III, T., and Grimm, E. C.: Patterns and rates of vegetation change during the deglaciation of eastern North America, 277&amp;ndash;288, in: The geology of North America, Ruddiman, W. F. and Wright Jr., H. E., Vol. K-3, North America and adjacent oceans during the last deglaciation, Geological Soc. Am., 1987. </mixed-citation>
</ref>
<ref id="ref60">
<label>60</label><mixed-citation publication-type="other" xlink:type="simple"> Jolly, D. and Haxeltine, A.: Effect of low glacial atmosphere CO&lt;sub&gt;2&lt;/sub&gt; on tropical African montane vegetation, Science, 276, 786&amp;ndash;788, 1997. </mixed-citation>
</ref>
<ref id="ref61">
<label>61</label><mixed-citation publication-type="other" xlink:type="simple"> Kageyama, M., Peyron, O., Pinot, S., Tarasov, P., Guiot, J., Joussaume, S., and Ramstein, G.: The Last Glacial Maximum climate over Europe and western Siberia: a PMIP comparison between models and data, Clim. Dyn., 17, 23&amp;ndash;43, 2001. </mixed-citation>
</ref>
<ref id="ref62">
<label>62</label><mixed-citation publication-type="other" xlink:type="simple"> Levin, D. A. and Wilson, A. C.: Rates of evolution in seed plants: Net increase in diversity of chromosome numbers and species numbers through time, Proceedings of the National Academy of Sciences, 73, 2086&amp;ndash;2090, 1976. </mixed-citation>
</ref>
<ref id="ref63">
<label>63</label><mixed-citation publication-type="other" xlink:type="simple"> Levis, S. and Foley, J. A.: CO&lt;sub&gt;2&lt;/sub&gt;, climate, and vegetation feedbacks at the Last Glacial Maximum, J. Geophys. Res., 104, 31 191&amp;ndash;31 198, 1999. </mixed-citation>
</ref>
<ref id="ref64">
<label>64</label><mixed-citation publication-type="other" xlink:type="simple"> Maher Jr., L.J., Miller, N. G., Baker, R. G., Curry, B. B., and Mickelson, D. M.: Paleobiology of the sand beneath the Valders diamicton at Valders, Wisconsin, Quaternary Res., 49, 208&amp;ndash;221, 1998. </mixed-citation>
</ref>
<ref id="ref65">
<label>65</label><mixed-citation publication-type="other" xlink:type="simple"> Maxwell, J. A. and Davis, M. B.: Pollen evidence of Pleistocene and Holocene vegetation on the Allegheny Plateau, Maryland, Quaternary Res., 2, 506&amp;ndash;530, 1972. </mixed-citation>
</ref>
<ref id="ref66">
<label>66</label><mixed-citation publication-type="other" xlink:type="simple"> McLachlan, J. S. and Clark, J. S.: Reconstructing historical ranges with fossil data at continental scales, Forest Ecol. Manage., 197, 139&amp;ndash;147, 2004. </mixed-citation>
</ref>
<ref id="ref67">
<label>67</label><mixed-citation publication-type="other" xlink:type="simple"> McLachlan, J. S., Clark, J. S., and Manos, P. S.: Molecular indicators of tree migration capacity under rapid climate change, Ecology, 86, 2088&amp;ndash;2098, 2005. </mixed-citation>
</ref>
<ref id="ref68">
<label>68</label><mixed-citation publication-type="other" xlink:type="simple"> Mickelson, D. M., Clayton, L., Fullerton, D. S., and Borns Jr., H. W.: The late Wisconsin glacial record of the Laurentide ice sheet in the United States, 3-37 in: Late-Quaternary environments of the United States, edited by: Wright Jr., H. E., Vol. 1, The late Pleistocene, S.C. Porter (Ed.), University of Minnesota Press, Minneapolis, 1983. </mixed-citation>
</ref>
<ref id="ref69">
<label>69</label><mixed-citation publication-type="other" xlink:type="simple"> Nakagawa, T., Tarasov, P. E., Nishida, K., Gotanda, K., and Yasuda, Y.: Quantitative pollen-based climate reconstruction in central Japan: application to surface and Late Quaternary spectra, Quaternary Sci. Rev., 21, 2099&amp;ndash;2113, 2002. </mixed-citation>
</ref>
<ref id="ref70">
<label>70</label><mixed-citation publication-type="other" xlink:type="simple"> Overpeck, J. T., Webb, R. S., and Webb III, T.: Mapping eastern North American vegetation change of the past 18 ka: No-analogs and the future, Geology, 20, 1071&amp;ndash;1074, 1992. </mixed-citation>
</ref>
<ref id="ref71">
<label>71</label><mixed-citation publication-type="other" xlink:type="simple"> Parker, K. C., Hamrick, J. L., Parkers, A. J., and Stacy, E. A.: Allozyme diversity in \textitPinus virginiana (Pinaceae): Intraspecific and interspecific comparisons, Am. J. Botany, 84, 1372&amp;ndash;1382, 1997. </mixed-citation>
</ref>
<ref id="ref72">
<label>72</label><mixed-citation publication-type="other" xlink:type="simple"> Péwé, T. L.: The periglacial environment in North America during Wisconsin time, 157&amp;ndash;189 in: Late-Quaternary environments of the United States, edited by: Wright Jr., H. E., Vol. 1, The late Pleistocene, S.C. Porter (Ed.), University of Minnesota Press, Minneapolis, 1983. </mixed-citation>
</ref>
<ref id="ref73">
<label>73</label><mixed-citation publication-type="other" xlink:type="simple"> Peyron, O., Guiot, J., Cheddadi, R., Tarasov, P., Reille, M., de Beaulieu, Bottema, S., and Andrieu, V.: Climatic reconstruction in Europe for 18 000 yr B.P. from pollen data, Quaternary Res., 49, 183&amp;ndash;196, 1998. </mixed-citation>
</ref>
<ref id="ref74">
<label>74</label><mixed-citation publication-type="other" xlink:type="simple"> Pielou, E. C.: After the Ice Age, University of Chicago Press, Chicago, 1991. </mixed-citation>
</ref>
<ref id="ref75">
<label>75</label><mixed-citation publication-type="other" xlink:type="simple"> Pinot, S., Ramstein, G., Harrison, S. P., Prentice, I. C., Guiot, J., Stute, M., and Joussaume, S.: Tropical paleoclimates at the Last Glacial Maximum: Comparison of Paleoclimate Modeling Intercomparison Project (PMIP) simulations and paleodata, Clim. Dyn., 15, 857&amp;ndash;874, 1999. </mixed-citation>
</ref>
<ref id="ref76">
<label>76</label><mixed-citation publication-type="other" xlink:type="simple"> Polley, H. W., Johnson, H. B., Marino, B. D., and Mayeux, H. S.: Increase in C&lt;sub&gt;3&lt;/sub&gt; plant water-use efficiency and biomass over glacial to present CO&lt;sub&gt;2&lt;/sub&gt; concentrations, Nature, 361, 61&amp;ndash;64, 1993. </mixed-citation>
</ref>
<ref id="ref77">
<label>77</label><mixed-citation publication-type="other" xlink:type="simple"> Polley, H. W., Johnson, H. B., and Mayeux, H. S.: Nitrogen and water requirements of C&lt;sub&gt;3&lt;/sub&gt; plants grown at glacial to present carbon dioxide concentrations, Functional Ecol., 9, 86&amp;ndash;96, 1995. </mixed-citation>
</ref>
<ref id="ref78">
<label>78</label><mixed-citation publication-type="other" xlink:type="simple"> Potts, R. and Behrensmeyer, A. K.: Late Cenozoic terrestrial ecosystems, 419&amp;ndash;436, in: Terrestrial ecosystems through time, edited by: Behrensmeyer, A. K., Damuth, J. D., DiMichele, W. A., Potts, R., Sues, H.-D., and Wing, S. L., University of Chicago Press, Chicago, 1992. </mixed-citation>
</ref>
<ref id="ref79">
<label>79</label><mixed-citation publication-type="other" xlink:type="simple"> Prentice, I. C., Bartlein, P. J., and Webb III, T.: Vegetation and climate change in eastern North America since the last glacial maximum, Ecology, 72, 2038&amp;ndash;2056, 1991. </mixed-citation>
</ref>
<ref id="ref80">
<label>80</label><mixed-citation publication-type="other" xlink:type="simple"> Qian, H.: Spatial pattern of vascular plant diversity in North America north of Mexico and its floristic relationship with Eurasia, Ann. Botany, 83, 271&amp;ndash;283, 1999. </mixed-citation>
</ref>
<ref id="ref81">
<label>81</label><mixed-citation publication-type="other" xlink:type="simple"> Ritchie, J. C.: Postglacial vegetation of Canada, Cambridge University Press, Cambridge, 1987. </mixed-citation>
</ref>
<ref id="ref82">
<label>82</label><mixed-citation publication-type="other" xlink:type="simple"> Robinson, J. M.: Speculations on carbon dioxide starvation, late Tertiary evolution of stomatal regulation and floristic modernization, Plant, Cell Environment, 17, 345&amp;ndash;354, 1994. </mixed-citation>
</ref>
<ref id="ref83">
<label>83</label><mixed-citation publication-type="other" xlink:type="simple"> Roy, K., Valentine, J. W., Jablonski, D., and Kidwell, S. M.: Scales of climatic variability and time averaging in Pleistocene biotas: Implications for ecology and evolution, Trends in Ecology and Evolution, 11, 458&amp;ndash;463, 1996. </mixed-citation>
</ref>
<ref id="ref84">
<label>84</label><mixed-citation publication-type="other" xlink:type="simple"> Royall, P. D., Delcourt, P. A., and Delcourt, H. R.: Late Quaternary paleoecology and paleoenvironments of the central Mississippi alluvial valley, Geol. Soc. Am. Bull., 103, 157&amp;ndash;170, 1991. </mixed-citation>
</ref>
<ref id="ref85">
<label>85</label><mixed-citation publication-type="other" xlink:type="simple"> Saxe, H., Ellsworth, D. S., and Heath, J.: Tree and forest functioning in an enriched CO&lt;sub&gt;2&lt;/sub&gt; atmosphere, New Phytologist, 139, 395&amp;ndash;436, 1998. </mixed-citation>
</ref>
<ref id="ref86">
<label>86</label><mixed-citation publication-type="other" xlink:type="simple"> Schofield, W. B.: Endemic genera of bryophytes of North America (north of Mexico), Preslia Praha, 76, 255&amp;ndash;277, 2004. </mixed-citation>
</ref>
<ref id="ref87">
<label>87</label><mixed-citation publication-type="other" xlink:type="simple"> Schoonmaker, P. K. and Foster, D. R.: Some implications of paleoecology for contemporary ecology, Botanical Rev., 57, 204&amp;ndash;245, 1991. </mixed-citation>
</ref>
<ref id="ref88">
<label>88</label><mixed-citation publication-type="other" xlink:type="simple"> Soltis, D. E., Morris, A. B., McLachlan, J. S., Manos, P. S., and Soltis, P. S.: Comparative phylogeography of unglaciated eastern North America, Molecular Ecol., 15, 4261&amp;ndash;4293, 2006. </mixed-citation>
</ref>
<ref id="ref89">
<label>89</label><mixed-citation publication-type="other" xlink:type="simple"> Stewart, J. R. and Lister, A. M.: Cryptic northern refugia and the origins of the modern biota, Trends Ecol. Evolution, 16, 608&amp;ndash;613, 2001. </mixed-citation>
</ref>
<ref id="ref90">
<label>90</label><mixed-citation publication-type="other" xlink:type="simple"> Street-Perrott, F. A., Huang, Y., Perrott, R. A., Eglinton, G., Barker, P., Khelifa, L. B., Harkness, D. D., and Olago, D. O.: Impact of lower atmospheric carbon dioxide on tropical mountain ecosystems, Science, 278, 1422&amp;ndash;1426, 1997. </mixed-citation>
</ref>
<ref id="ref91">
<label>91</label><mixed-citation publication-type="other" xlink:type="simple"> Svenning, J.-C.: Deterministic Plio-Pleistocene extinctions in the European cool-temperate tree flora, Ecol. Lett., 6, 646&amp;ndash;653, 2003. </mixed-citation>
</ref>
<ref id="ref92">
<label>92</label><mixed-citation publication-type="other" xlink:type="simple"> Tallis, J. H.: Plant community history, Chapman and Hall, New York, 1991. </mixed-citation>
</ref>
<ref id="ref93">
<label>93</label><mixed-citation publication-type="other" xlink:type="simple"> Tarasov, P. E., Peyron, O., Guiot, J., Brewer, S., Volkova, V. S., Bezusko, L. G., Dorofeyuk, N. I., Kvavadze, E. V., Osipova, I. M., and Panova, N. K.: Last Glacial Maximum climate of the former Soviet Union and Mongolia reconstructed from pollen and plant macrofossil data, Clim. Dyn., 15, 227&amp;ndash;240, 1999. </mixed-citation>
</ref>
<ref id="ref94">
<label>94</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A.: The full-glacial vegetation of northwestern Georgia, Ecology, 51, 17&amp;ndash;33, 1970. </mixed-citation>
</ref>
<ref id="ref95">
<label>95</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A.: Postglacial and interglacial vegetation history of southern Georgia and central Florida, Ecology, 52, 676&amp;ndash;689, 1971. </mixed-citation>
</ref>
<ref id="ref96">
<label>96</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A.: The vegetation record of a mid-Wisconsin interstadial in northwest Georgia, Quaternary Res., 3, 257&amp;ndash;268, 1973. </mixed-citation>
</ref>
<ref id="ref97">
<label>97</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A.: Late Quaternary vegetation of central Appalachia and the New Jersey coastal plain, Ecol. Monogr., 49, 427&amp;ndash;469, 1979. </mixed-citation>
</ref>
<ref id="ref98">
<label>98</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A.: The late Quaternary vegetation history of the southeastern United States, Ann. Rev. Ecol. Syst., 11, 387&amp;ndash;409, 1980a. </mixed-citation>
</ref>
<ref id="ref99">
<label>99</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A.: Late Quaternary vegetation history at White Pond on the inner coastal plain of South Carolina, Quaternary Res., 13, 187&amp;ndash;199, 1980b. </mixed-citation>
</ref>
<ref id="ref100">
<label>100</label><mixed-citation publication-type="other" xlink:type="simple"> Watts, W. A. and Stuvier, M.: Late Wisconsin climate of northern Florida and the origin of species-rich deciduous forest, Science, 210, 325&amp;ndash;327, 1980. </mixed-citation>
</ref>
<ref id="ref101">
<label>101</label><mixed-citation publication-type="other" xlink:type="simple"> Webb III, T., Anderson, K. H., Bartlein, P. J., and Webb, R. S.: Late Quaternary climate change in eastern North America: A comparison of pollen-derived estimates with climate model results, Quat. Sci. Rev., 17, 587&amp;ndash;606, 1997. </mixed-citation>
</ref>
<ref id="ref102">
<label>102</label><mixed-citation publication-type="other" xlink:type="simple"> Webb III, T., Bartlein, P. J., Harrison, S. P., and Anderson, K. H.: Vegetation, lake levels, and climate in eastern North America for the past 18 000 years, 415&amp;ndash;467, in: Global climates Since the Last Glacial Maximum, edited by: Wright Jr., H. E., Kutzbach, J. E., Webb III, T., Ruddiman, W. F., Street-Perrott, F. A., and Bartlein, P. J., University of Minnesota Press, Minneapolis, 1993. </mixed-citation>
</ref>
<ref id="ref103">
<label>103</label><mixed-citation publication-type="other" xlink:type="simple"> Webb III, T., Bartlein, P. J., and Kutzbach, J. E.: Climatic change in eastern North America during the past 18 000 years: Comparisons of pollen data with model results, 447&amp;ndash;462, in: The geology of North America, edited by: Ruddiman, W. F. and Wright Jr., H. E., , Vol. K-3, North American and adjacent oceans during the last deglaciation, Geological Society of America, 1988. </mixed-citation>
</ref>
<ref id="ref104">
<label>104</label><mixed-citation publication-type="other" xlink:type="simple"> Whitehead, D. R.: Late Wisconsin vegetational changes in unglaciated eastern North America, Quaternary Res., 3, 621&amp;ndash;631, 1973. </mixed-citation>
</ref>
<ref id="ref105">
<label>105</label><mixed-citation publication-type="other" xlink:type="simple"> Wilkins, G. R., Delcourt, P. A., Delcourt, H. R., Harrison, F. W., and Turner, M. R.: Paleoecology of central Kentucky since the Last Glacial Maximum, Quaternary Res., 36, 224&amp;ndash;239, 1991. </mixed-citation>
</ref>
<ref id="ref106">
<label>106</label><mixed-citation publication-type="other" xlink:type="simple"> Williams, J. W., Shuman, B. N., Webb III, T., Bartlein, P. J., and Leduc, P. L.: Late-Quaternary vegetation dynamics in North America: Scaling from taxa to biomes, Ecol. Monogr., 74, 309&amp;ndash;334, 2004. </mixed-citation>
</ref>
<ref id="ref107">
<label>107</label><mixed-citation publication-type="other" xlink:type="simple"> Willis, K. J. and van Andel, T. H.: Trees or no trees? The environments of central and eastern Europe during the Last Glaciation, Quaternary Sci. Rev., 23, 2369&amp;ndash;2387, 2004. </mixed-citation>
</ref>
<ref id="ref108">
<label>108</label><mixed-citation publication-type="other" xlink:type="simple"> Willis, K. J. and Whittaker, R. J.: The refugial debate, Science, 287, 1406&amp;ndash;1407, 2000. </mixed-citation>
</ref>
<ref id="ref109">
<label>109</label><mixed-citation publication-type="other" xlink:type="simple"> Wiser, S. K.: Comparison of southern Appalachian high-elevation outcrop plant communities with their northern Appalachian counterparts, J. Biogeogr., 25, 501&amp;ndash;513, 1998. </mixed-citation>
</ref>
<ref id="ref110">
<label>110</label><mixed-citation publication-type="other" xlink:type="simple"> Wolfe, J. N.: The possible role of microclimate, Ohio J. Sci., 51, 134&amp;ndash;138, 1951. </mixed-citation>
</ref>
</ref-list>
</back>
</article>