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<front>
<journal-meta>
<journal-id journal-id-type="publisher">glbot</journal-id>
<journal-title-group>
<journal-title>The Great Lakes Botanist</journal-title>
</journal-title-group>
<issn pub-type="epub"></issn>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">4114</article-id>
<article-id pub-id-type="manuscript">glbot4114.pdf</article-id>
<article-id pub-id-type="doi">10.3998/glbot.4114</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Feature Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Two Southern Plant Species, <italic>Nutallanthus Canadensis</italic> (L.) D.A. Sutton and <italic>Opuntia Cespitosa</italic> Raf., Discovered as Disjunct in the Huron Mountains, Marquette County, Michigan</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Rutherford</surname>
<given-names>Ryne</given-names>
</name>
<email>biophilianature@gmail.com</email>
<aff id="aff1">Michigan Technological University. U.J. Noblet Forestry Building 1400 Townsend Drive Houghton, MI 49931-1295. Biophilia, LLC. 32104 W State Highway M-64, Ontonagon, MI 49953.</aff>
</contrib>
</contrib-group>
<pub-date>
<day>27</day>
<month>06</month>
<year>2023</year>
</pub-date>
<volume>61</volume>
<permissions>
<license><license-p>CC BY-NC-ND 4.0</license-p></license>
</permissions>
<counts>
<fig-count count="2"/>
</counts>
</article-meta>
</front>
<body>
<p>The Great Lakes region harbors disjunct species from various floristic affinities including western cordilleran, Gulf Coast and Atlantic coastal plain, and the Central North American prairie. The northern Great Lakes region is well known for harboring populations of western cordilleran disjuncts (<xref ref-type="bibr" rid="ref5">Marquis and Voss 1981</xref>; <xref ref-type="bibr" rid="ref1">Drummond et al. 2022</xref>), and several hypotheses have been proposed as to how they were able to migrate to the region. Eastward migration during a cooler and moister postglacial period appears most likely. Coastal plain disjuncts may have arrived from the Atlantic and Gulf coasts by making short and moderate distance migrations along post-glacial wetlands (<xref ref-type="bibr" rid="ref7">Reznicek 1994</xref>). Prairie species expanded eastwards from central North America during a post-glacial warm and dry period and became stranded when conditions changed, resulting in disjunct populations (<xref ref-type="bibr" rid="ref4">Manogaran 1983</xref>; <xref ref-type="bibr" rid="ref8">Reznicek and Maycock 1983</xref>). Southern, or &#x201C;Carolinian,&#x201D; plants are less documented as disjuncts in the northern Great Lakes region. While well represented in the southern Great Lakes region, most regional species with southern affinities have a roughly continuous distribution to the south which is likely due to the continuity of suitable habitat. However, the complex terrain in the Huron Mountains in the western Upper Peninsula of Michigan may allow for enough climatic variation at a fine scale to support populations of species outside their normal ranges.</p>
<p>This article reports the first collection of <italic>Nutallanthus canadensis</italic> (L.) D.A.Sutton (Plantaginaceae) and <italic>Opuntia cespitosa</italic> Raf. (Cactaceae) from the Lake Superior drainage basin, and both appear to be the northernmost occur-rences of these species in eastern North America.</p>
<p>Dry, open, sandy or rocky ground, jack pine plains and the beds of dried lakes have been noted as habitats for <italic>Nutallanthus canadensis</italic> in Michigan. Sandy fields, open oak forests, stabilized open dunes and disturbed ground along roadsides have been listed as habitats for <italic>Opuntia cespitosa</italic> in Michigan (<xref ref-type="bibr" rid="ref6">MICHIGAN FLORA ONLINE 2011</xref>). Both species have broad distributions further south in eastern North America.</p>
<p>A population of <italic>Nutallanthus canadensis</italic> was discovered at the summit of Trout Mountain in the northern Huron Mountains on June 18, 2022 (<xref ref-type="fig" rid="f001">Figure 1</xref>). Fifty plants, nearly all of them in flower, were observed in a granite bedrock glade in openings and under a filtered canopy of <italic>Quercus rubra</italic>. The site was south-facing and on top of a fairly steep ledge. Other plants noted include <italic>Arctostaphylos uva-ursi</italic> (L.) Spreng., <italic>Melampyrum lineare</italic> Desr., <italic>Acer rubrum</italic> L., <italic>Woodsia ilvensis</italic> (L.) R. Br., <italic>Carex pensylvanica</italic> Lam., and <italic>Vaccinium myrtilloides</italic> Michx. A moss, <italic>Hedwigia ciliata</italic> (Hedw.) P. Beauv., was also present. An additional ten plants were seen at Mount Ives, 2.8 km to the east on June 19, 2022, and seven were observed at the summit of Breakfast Roll, 5.1 km to the east of the original site on June 20, 2022. Both the Mount Ives and the Breakfast Roll populations were in open locations in shallow soil with similar flora and site characteristics (<xref ref-type="table" rid="tab1">Table 1</xref>).</p>
<fig id="f001" position="anchor">
<label>Figure 1.</label>
<caption>
<title><italic>Nutallanthus canadensis</italic> in flower under <italic>Quercus rubra</italic> on a granite bedrock glade at the summit of Trout Mountain on June 18, 2022. Photo by Ryne D. Rutherford.</title>
</caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="glbot-4114-f001.jpg"/>
</fig>
<table-wrap position="float" id="tab1">
<label>Table 1.</label>
<caption>
<p>Habitat characteristics for the three populations of <italic>Nutallanthus canadensis</italic> in the northern Huron Mountains. The values given are mean values recorded in 12 one-meter square plots along 100-meter transects recorded in September 2022.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th valign="bottom" align="left">Site</th>
<th valign="bottom" align="center">Elevation (m)</th>
<th valign="bottom" align="center">Canopy Cover (%)</th>
<th valign="bottom" align="center">Vascular Plant Coverage (%)</th>
<th valign="bottom" align="center">Exposed Bedrock (%)</th>
<th valign="bottom" align="center">Distance to Lake Superior (km)</th>
<th valign="bottom" align="center">Slope Azimuth</th>
<th valign="bottom" align="center">Aspect</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Trout Mountain</td>
<td valign="top" align="center">391</td>
<td valign="top" align="center">25</td>
<td valign="top" align="center">25</td>
<td valign="top" align="center">38</td>
<td valign="top" align="center">3.2</td>
<td valign="top" align="center">18</td>
<td valign="top" align="center">170</td>
</tr>
<tr>
<td valign="top" align="left">Mount Ives</td>
<td valign="top" align="center">331</td>
<td valign="top" align="center">24</td>
<td valign="top" align="center">10</td>
<td valign="top" align="center">68</td>
<td valign="top" align="center">3.5</td>
<td valign="top" align="center">15</td>
<td valign="top" align="center">200</td>
</tr>
<tr>
<td valign="top" align="left">Breakfast Roll</td>
<td valign="top" align="center">354</td>
<td valign="top" align="center">9</td>
<td valign="top" align="center">31</td>
<td valign="top" align="center">47</td>
<td valign="top" align="center">3.9</td>
<td valign="top" align="center">5</td>
<td valign="top" align="center">180</td>
</tr>
</tbody>
</table>
</table-wrap>
<p><italic>Opuntia cespitosa</italic> was found on a small (10&#x00B4;14 m) south-facing glade on the north side of the Little Garlic River in the central Huron Mountains at 418 m elevation (<xref ref-type="fig" rid="f002">Figure 2</xref>). Plants were found in small pockets of soil and on bare meta-morphic rock of mostly mafic composition on August 1, 2022, on an overcast raining morning with an air temperature of 16&#x00B0;C. A total of six plants were found, one patch occupied about four square meters, and another occupied two square meters. Two fresh blooms and fifteen spent flowers were observed along with a single bud. No developing fruits were found, and no pollinators were present. The glade, which contained a few small openings, occurred along a ridge that trended east-west and was mostly dominated by <italic>Pinus strobus</italic> L. The plants occurred under a filtered canopy, about 70% open. Other plants included <italic>Vaccinium angustifolium</italic> Aiton, <italic>Diervilla lonicera</italic> Mill., <italic>Dicanthelium xanthophyllum, Capnoides sempervirens</italic> (L.) Borkh., <italic>Hieracium piloselloides</italic> Vill., <italic>Quercus rubra</italic> L., <italic>Juniperus communis</italic> L., <italic>Danthonia spicata</italic> (L.) Roem. &#x0026; Schult., and <italic>Amelanchier</italic> sp. The open patches of rock were lichen-dominated, including the following species: <italic>Stereocaulon saxatile</italic> H. Magn., <italic>Cladonia mitis</italic> Sandst., <italic>Cladonia rangiferina</italic> (L.) F. H. Wigg., <italic>Cladonia turgida</italic> Ehrh. Ex Hoffm., <italic>Cladonia gracilis</italic> (L.) Willd., <italic>Lepraria neglecta</italic> (Nyl.) Erichsen, <italic>Diploschistes scruposus</italic> (Schreber) Norman, <italic>Umbilicaria deusta</italic> (L.) Baumg., <italic>Acarospora fuscata</italic> B. de Lesd. and <italic>Porpidia</italic> sp. One moss species, <italic>Polytrichum juniperum</italic> Hedw., was noted.</p>
<fig id="f002" position="anchor">
<label>Figure 2.</label>
<caption>
<title><italic>Opuntia cespitosa</italic> in a small glade on the north side of the Little Garlic River on August 1, 2022. Photo by Ryne D. Rutherford.</title>
</caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="glbot-4114-f002.jpg"/>
</fig>
<p>The nearest record of <italic>Nutallanthus canadensis</italic> is 120 km to the south along the Menominee River on the Michigan/Wisconsin border and the closest continuous populations are 300 km SW in central Wisconsin along the Wisconsin River and 300 km SE in the northwestern Lower Peninsula of Michigan. The nearest apparently natural populations of <italic>Opuntia cespitosa</italic> are in the west-central Lower Peninsula of Michigan (370 km SSE) and from south-central Wisconsin (400 km SSW). (<xref ref-type="bibr" rid="ref2">GBIF 2023</xref>; <xref ref-type="bibr" rid="ref6">MICHIGAN FLORA ONLINE 2011</xref>). This is the first time <italic>Opuntia cespitosa</italic> has been collected in the Upper Peninsula of Michigan and the first reported collection from a rock outcrop in Michigan.</p>
<p>While the origin of these southern disjuncts cannot be known with certainty, natural colonization during the post-glacial hypsithermal warm period ca. 5,000 years BP is the most likely origin. The warm, dry conditions during the hypsithermal are known to have allowed for the northward and eastward colonization of many species and suitable local conditions allowed persistence of populations as the climate cooled (<xref ref-type="bibr" rid="ref8">Reznicek and Maycock 1983</xref>; <xref ref-type="bibr" rid="ref9">Strong and Hills 2003</xref>; <xref ref-type="bibr" rid="ref3">Hamilton and Eckert 2007</xref>). The local conditions on granite bedrock glades in the Huron Mountains are warmer than the surroundings, which may allow the persistence of species that are otherwise poorly suited to the regional climate in the Lake Superior region (<xref ref-type="table" rid="tab2">Table 2</xref>).</p>
<table-wrap position="float" id="tab2">
<label>Table 2.</label>
<caption>
<p>Ground surface temperature metrics for granite bedrock glades (n=2) and the surrounding forest (within 50 meters of the glade edge) (n=4) from May 7 to September 14, 2021, recorded with HOBO&#x2122; data loggers. All the loggers were placed directly on the ground.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th valign="bottom" align="left">Temperature Metrics</th>
<th valign="bottom" align="center">Glade</th>
<th valign="bottom" align="center">Matrix</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Mean Max &#x00B0;C</td>
<td valign="top" align="center">52</td>
<td valign="top" align="center">32</td>
</tr>
<tr>
<td valign="top" align="left">Mean Min &#x00B0;C</td>
<td valign="top" align="center">0</td>
<td valign="top" align="center">1</td>
</tr>
<tr>
<td valign="top" align="left">Mean &#x00B0;C</td>
<td valign="top" align="center">23.4</td>
<td valign="top" align="center">17.8</td>
</tr>
<tr>
<td valign="top" align="left">Hours above 30 &#x00B0;C</td>
<td valign="top" align="center">750</td>
<td valign="top" align="center">7.8</td>
</tr>
</tbody>
</table>
</table-wrap>
</body>
<back>
<ref-list>
<title>Specimen Citations</title>
<ref id="ref_sec0003_1"><mixed-citation publication-type="other"><person-group person-group-type="author"><collab><italic>Opuntia cespitosa</italic>: MICHIGAN</collab></person-group>. <article-title>Marquette County. 46.64256 -87.64070</article-title>. <month>August</month> <day>1</day>, <year>2022</year>. <source><italic>Ryne D. Rutherford 200</italic> (MICH), <italic>201</italic> (FLAS)</source>.</mixed-citation></ref>
<ref id="ref_sec0003_2"><mixed-citation publication-type="other"><person-group person-group-type="author"><collab><italic>Nutallanthus canadensis</italic>: MICHIGAN</collab></person-group>. <article-title>Marquette County. 46.86539 -87.88969</article-title>. <month>August</month> <day>1</day>, <year>2022</year>. <source><italic>Ryne D. Rutherford 202</italic> (MICH)</source>.</mixed-citation></ref>
</ref-list>
<ack>
<title>Acknowledgments</title>
<p>Many thanks to Kerry Woods, and the Huron Mountain Wildlife Foundation, my advisor David Flaspohler, and Michigan Technological University for supporting this research. Thanks to Susan Fawcett for excellent work at specimen preparation. Many thanks to Michael Huft for his timely and thoughtful edits of this manuscript.</p>
</ack>
<ref-list>
<title>Literature Cited</title>
<ref id="ref1"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Drummond</surname>, <given-names>C. P.</given-names></string-name>, <string-name><surname>Cochrane</surname>, <given-names>T. S.</given-names></string-name> and <string-name><surname>Sytsma</surname>, <given-names>K. J.</given-names></string-name></person-group> (<year>2022</year>). <article-title>Western North American plants disjunct in the Great Lakes region: 40 years after Marquis and Voss</article-title>. <source>International Journal of Plant Sciences</source> <volume>183</volume>: <fpage>691</fpage>&#x2013;<lpage>705</lpage>.</mixed-citation></ref>
<ref id="ref2"><mixed-citation publication-type="web"><person-group person-group-type="author"><collab>GBIF</collab></person-group>. (<year>2023</year>). <source>Global Biodiversity Information Facility</source>. Available at <ext-link xmlns:xlink="http://www.w3.org/1999/xlink" ext-link-type="uri" xlink:href="http://www.gbif.org">http://www.gbif.org</ext-link>. (Accessed <date-in-citation content-type="access-date">February 15. 2023</date-in-citation>).</mixed-citation></ref>
<ref id="ref3"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Hamilton</surname>, <given-names>J. A.</given-names></string-name>, and <string-name><surname>Eckert</surname>, <given-names>C. G.</given-names></string-name></person-group> (<year>2007</year>). <article-title>Population genetic consequences of geographic disjunction: A prairie plant isolated on Great Lakes alvars</article-title>. <source>Molecular Ecology</source> <volume>16</volume>: <fpage>1649</fpage>&#x2013;<lpage>1660</lpage>.</mixed-citation></ref>
<ref id="ref4"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Manogaran</surname>, <given-names>C.</given-names></string-name></person-group> (<year>1983</year>). <article-title>The prairie peninsula: A climatic perspective</article-title>. <source>Physical Geography</source>. <volume>4</volume>: <fpage>153</fpage>&#x2013;<lpage>166</lpage>.</mixed-citation></ref>
<ref id="ref5"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Marquis</surname>, <given-names>R. J.</given-names></string-name>, and <string-name><surname>Voss</surname>, <given-names>E. G.</given-names></string-name></person-group> (<year>1981</year>). <article-title>Distributions of some western North American plants disjunct in the Great Lakes region</article-title>. <source>The Michigan Botanist</source> <volume>20</volume>: <fpage>53</fpage>&#x2013;<lpage>82</lpage>.</mixed-citation></ref>
<ref id="ref6"><mixed-citation publication-type="web"><person-group person-group-type="author"><collab>MICHIGAN FLORA ONLINE</collab></person-group>. <person-group person-group-type="author"><string-name><given-names>A. A.</given-names> <surname>Reznicek</surname></string-name>, <string-name><given-names>E. G.</given-names> <surname>Voss</surname></string-name>, and <string-name><given-names>B. S.</given-names> <surname>Walters</surname></string-name></person-group>. (<year>2011</year>). <source>University of Michigan</source>. Available at <ext-link xmlns:xlink="http://www.w3.org/1999/xlink" ext-link-type="uri" xlink:href="http://michiganflora.net">http://michiganflora.net</ext-link>. (Accessed <date-in-citation content-type="access-date">February 15, 2023</date-in-citation>).</mixed-citation></ref>
<ref id="ref7"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Reznicek</surname>, <given-names>A. A.</given-names></string-name></person-group> (<year>1994</year>). <article-title>The disjunct coastal plain flora in the Great Lakes region</article-title>. <source>Biological Conservation</source> <volume>68</volume>: <fpage>203</fpage>&#x2013;<lpage>215</lpage>.</mixed-citation></ref>
<ref id="ref8"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Reznicek</surname>, <given-names>A. A.</given-names></string-name>, and <string-name><surname>Maycock</surname>, <given-names>P. F.</given-names></string-name></person-group> (<year>1983</year>). <article-title>Composition of an isolated prairie in central Ontario</article-title>. <source>Canadian Journal of Botany</source>. <volume>61</volume>: <fpage>3107</fpage>&#x2013;<lpage>3116</lpage>.</mixed-citation></ref>
<ref id="ref9"><mixed-citation publication-type="journal"><person-group person-group-type="author"><string-name><surname>Strong</surname>, <given-names>W. L.</given-names></string-name>, and <string-name><surname>Hills</surname>, <given-names>L. V.</given-names></string-name></person-group> (<year>2003</year>). <article-title>Post hypsithermal plant disjunctions in western Alberta, Canada</article-title>. <source>Journal of Biogeography</source> <volume>30</volume>: <fpage>419</fpage>&#x2013;<lpage>430</lpage>.</mixed-citation></ref>
</ref-list>
</back>
</article>
