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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en"><italic>Leucaena leucocephala</italic> and adjacent native limestone forest habitats contrast in soil properties on Tinian Island</title><author><name sortKey="Marler, Thomas E" sort="Marler, Thomas E" uniqKey="Marler T" first="Thomas E." last="Marler">Thomas E. Marler</name><affiliation><nlm:aff id="af0001"></nlm:aff></affiliation></author><author><name sortKey="Dongol, Nirmala" sort="Dongol, Nirmala" uniqKey="Dongol N" first="Nirmala" last="Dongol">Nirmala Dongol</name><affiliation><nlm:aff id="af0001"></nlm:aff></affiliation></author><author><name sortKey="Cruz, Gil N" sort="Cruz, Gil N" uniqKey="Cruz G" first="Gil N." last="Cruz">Gil N. Cruz</name><affiliation><nlm:aff id="af0001"></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">27829978</idno><idno type="pmc">5100652</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5100652</idno><idno type="RBID">PMC:5100652</idno><idno type="doi">10.1080/19420889.2016.1212792</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">000033</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000033</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main"><italic>Leucaena leucocephala</italic> and adjacent native limestone forest habitats contrast in soil properties on Tinian Island</title><author><name sortKey="Marler, Thomas E" sort="Marler, Thomas E" uniqKey="Marler T" first="Thomas E." last="Marler">Thomas E. Marler</name><affiliation><nlm:aff id="af0001"></nlm:aff></affiliation></author><author><name sortKey="Dongol, Nirmala" sort="Dongol, Nirmala" uniqKey="Dongol N" first="Nirmala" last="Dongol">Nirmala Dongol</name><affiliation><nlm:aff id="af0001"></nlm:aff></affiliation></author><author><name sortKey="Cruz, Gil N" sort="Cruz, Gil N" uniqKey="Cruz G" first="Gil N." last="Cruz">Gil N. Cruz</name><affiliation><nlm:aff id="af0001"></nlm:aff></affiliation></author></analytic><series><title level="j">Communicative & Integrative Biology</title><idno type="eISSN">1942-0889</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><title>ABSTRACT</title><p>An ex situ germplasm collection of the endangered <italic>Cycas micronesica</italic> was established in a transition zone between biodiverse native forest and mature stands of the invasive species <italic>Leucaena leucocephala</italic>. Soil chemical properties were determined for the 2 tree cover types to inform management decisions. Total carbon, total nitrogen, calcium, and net ammonification were greater in native forest cover than in <italic>L. leucocephala</italic> patches. Net nitrification and net mineralization were greater under <italic>L. leucocephala</italic> cover. Trace metals also differed between the 2 forest cover types, with chromium, cobalt, and nickel accumulating to greater concentration under <italic>L. leucocephala</italic> cover and zinc accumulating to greater concentration under native forest cover. The results indicated that <italic>L. leucocephala</italic> cover generated substantial changes in soil chemical properties when compared with native forest tree cover, illuminating one means by which understory vegetation may be affected by changes in invasive tree cover.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Donnegan, Ja" uniqKey="Donnegan J">JA Donnegan</name></author><author><name sortKey="Butler, Sl" uniqKey="Butler S">SL Butler</name></author><author><name sortKey="Grabowiecki, W" uniqKey="Grabowiecki W">W Grabowiecki</name></author><author><name sortKey="Hiserote, Ba" uniqKey="Hiserote B">BA Hiserote</name></author><author><name sortKey="Limtiaco, D" uniqKey="Limtiaco D">D Limtiaco</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Marler, T" uniqKey="Marler T">T Marler</name></author><author><name sortKey="Haynes, J" uniqKey="Haynes J">J Haynes</name></author><author><name 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Commun Integr Biol</journal-id><journal-id journal-id-type="iso-abbrev">Commun Integr Biol</journal-id><journal-id journal-id-type="publisher-id">KCIB</journal-id><journal-id journal-id-type="publisher-id">kcib20</journal-id><journal-title-group><journal-title>Communicative & Integrative Biology</journal-title></journal-title-group><issn pub-type="epub">1942-0889</issn><publisher><publisher-name>Taylor & Francis</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">27829978</article-id><article-id pub-id-type="pmc">5100652</article-id><article-id pub-id-type="publisher-id">1212792</article-id><article-id pub-id-type="doi">10.1080/19420889.2016.1212792</article-id><article-categories><subj-group subj-group-type="heading"><subject>Short Communication</subject></subj-group></article-categories><title-group><article-title><italic>Leucaena leucocephala</italic> and adjacent native limestone forest habitats contrast in soil properties on Tinian Island</article-title><alt-title alt-title-type="running-authors">T. E. MARLER ET AL.</alt-title><alt-title alt-title-type="running-title">COMMUNICATIVE & INTEGRATIVE BIOLOGY</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Marler</surname><given-names>Thomas E.</given-names></name><xref ref-type="aff" rid="af0001"></xref><xref ref-type="corresp" rid="an0001"></xref></contrib><contrib contrib-type="author"><name><surname>Dongol</surname><given-names>Nirmala</given-names></name><xref ref-type="aff" rid="af0001"></xref></contrib><contrib contrib-type="author"><name><surname>Cruz</surname><given-names>Gil N.</given-names></name><xref ref-type="aff" rid="af0001"></xref></contrib><aff id="af0001"><institution>Western Pacific Tropical Research Center, University of Guam, UOG Station</institution>, Mangilao, Guam,<country>USA</country></aff></contrib-group><author-notes><corresp id="an0001"><bold>CONTACT</bold> Thomas E. Marler <email xlink:href="thomas.marler@gmail.com">thomas.marler@gmail.com</email><institution>Western Pacific Tropical Research Center, 303 University Drive, University of Guam</institution>, Mangilao, Guam 96923, <country>USA</country></corresp></author-notes><pub-date pub-type="collection"><year>2016</year></pub-date><pub-date pub-type="epub"><day>18</day><month>8</month><year>2016</year></pub-date><pub-date pub-type="pmc-release"><day>18</day><month>8</month><year>2016</year></pub-date><pmc-comment> PMC Release delay is 0 months and 0 days and was based on the
<volume>9</volume><issue>5</issue><elocation-id seq="1">e1212792</elocation-id><history><date date-type="received"><day>16</day><month>5</month><year>2016</year></date><date date-type="rev-recd"><day>6</day><month>7</month><year>2016</year></date><date date-type="accepted"><day>8</day><month>7</month><year>2016</year></date></history><permissions><pmc-comment> © Thomas E. Marler, Nirmala Dongol, and Gil N. Cruz </pmc-comment>
<copyright-statement>© 2016 The Author(s). Published with license by Taylor & Francis</copyright-statement><copyright-year>2016</copyright-year><copyright-holder>The Author(s)</copyright-holder><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc/3.0/"><license-p>This is an Open Access article distributed under the terms of the Creative Commons Attribution-Non-Commercial License <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by-nc/3.0/">http://creativecommons.org/licenses/by-nc/3.0/</ext-link>, which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. The moral rights of the named author(s) have been asserted.</license-p></license></permissions><self-uri content-type="pdf" xlink:href="kcib-09-05-1212792.pdf"></self-uri><abstract><title>ABSTRACT</title><p>An ex situ germplasm collection of the endangered <italic>Cycas micronesica</italic> was established in a transition zone between biodiverse native forest and mature stands of the invasive species <italic>Leucaena leucocephala</italic>. Soil chemical properties were determined for the 2 tree cover types to inform management decisions. Total carbon, total nitrogen, calcium, and net ammonification were greater in native forest cover than in <italic>L. leucocephala</italic> patches. Net nitrification and net mineralization were greater under <italic>L. leucocephala</italic> cover. Trace metals also differed between the 2 forest cover types, with chromium, cobalt, and nickel accumulating to greater concentration under <italic>L. leucocephala</italic> cover and zinc accumulating to greater concentration under native forest cover. The results indicated that <italic>L. leucocephala</italic> cover generated substantial changes in soil chemical properties when compared with native forest tree cover, illuminating one means by which understory vegetation may be affected by changes in invasive tree cover.</p></abstract><kwd-group kwd-group-type="author"><title>KEYWORDS</title><kwd><italic>Cycas micronesica</italic> conservation</kwd><kwd>invasive species</kwd><kwd>mineralization</kwd><kwd>nitrification</kwd><kwd>nitrogen</kwd></kwd-group><counts><fig-count count="1"></fig-count><table-count count="2"></table-count><ref-count count="41"></ref-count><page-count count="6"></page-count></counts></article-meta></front><body><sec sec-type="intro" id="s0001"><title>Introduction</title><p><italic>Cycas micronesica</italic> transitioned from the most abundant tree on the island of Guam in 2002<xref rid="cit0001" ref-type="bibr"><sup>1</sup></xref> to Endangered status under IUCN<xref rid="cit0002" ref-type="bibr"><sup>2</sup></xref> and Threatened status under the United States Endangered Species Act<xref rid="cit0003" ref-type="bibr"><sup>3</sup></xref> mainly as a result of exotic phytophagous insects that invaded the island.<xref rid="cit0004" ref-type="bibr"><sup>4,5</sup></xref> Plant mortality following the invasions was epidemic, and population size and structure dramatically changed.<xref rid="cit0006" ref-type="bibr"><sup>6</sup></xref> Ex situ and in situ conservation programs were subsequently initiated, including a living collection of Guam germplasm established on the island of Tinian and funded by the United States Department of the Navy.<xref rid="cit0007" ref-type="bibr"><sup>7</sup></xref> The germplasm collection was situated in a transition zone with a portion of the plants established beneath a diverse native forest canopy, and a portion established beneath adjacent mature stands of non-native <italic>Leucaena leucocephala</italic>. Growth and health of the <italic>C. micronesica</italic> plants have been highly variable (see Fig. 2 in ref. <xref rid="cit0008" ref-type="bibr">8</xref>), with larger plants under the <italic>L. leucocephala</italic> cover than under native tree cover.</p><p>Conservation and restoration projects are often initiated in the absence of relevant publications to inform decisions. An adaptive management approach calls for persistent pursuit of new evidence-based outcomes. Therefore, the observations in Tinian reveal a need to identify which factors may be mediating the disparity in <italic>C. micronesica</italic> plant performance. Invasive plant species often exert profound changes in the habitats they invade.<xref rid="cit0009" ref-type="bibr"><sup>9,10</sup></xref> The literature on this subject reveals a bias toward several well-known invasive plant species, which limits a comprehensive evaluation of how invasive plants fit into global change issues.<xref rid="cit0011" ref-type="bibr"><sup>11</sup></xref><italic>Leucaena leucocephala</italic> has been extensively exploited in agroforestry settings as a source of nitrogen-rich green mulch to improve soil quality for cash crops.<xref rid="cit0012" ref-type="bibr"><sup>12</sup></xref> Nitrogen inputs, soil nitrification, and soil ammonification are among the components of the nitrogen cycle<xref rid="cit0013" ref-type="bibr"><sup>13</sup></xref> that may be influenced by <italic>L. leucocephala</italic>. Therefore, one potential factor that could influence understory <italic>C. micronesica</italic> plant growth is the difference in nutrient relations of the soils beneath the native forest cover versus the <italic>L. leucocephala</italic> patches.</p><p>Long-standing patches of vegetation contribute to system spatial heterogeneity through chronic influences on biogeochemical cycling and through interactions with microbial communities associated with the vegetation.<xref rid="cit0014" ref-type="bibr"><sup>14</sup></xref> These plant-soil feedbacks are orchestrated by complex integrated relationships among many biotic and abiotic factors.<xref rid="cit0015" ref-type="bibr"><sup>15</sup></xref> Influences of tree genotype on these processes include prolonged extraction and sequestration of soil elements in plant organs, extent of element resorption prior to leaf senescence then abscission, litter quality effects on organic matter lability, and local amplification of root-associated microorganisms and specialist saprophytic microorganisms. These phenomena are one means by which invasive plant species can affect native plants through changes in soil properties.<xref rid="cit0016" ref-type="bibr"><sup>16,17</sup></xref></p><p>Our objective was to use paired sampling sites throughout the <italic>C. micronesica</italic> germplasm to determine how forest tree cover type influenced soil chemical traits and nitrogen mineralization dynamics. The results may improve management decisions in our ex situ conservation program. Moreover, the information will increase knowledge of broader topics of tropical invasive species management, rare plant conservation, and integrative biological influences on soil chemical properties.</p></sec><sec sec-type="results" id="s0002"><title>Results</title><p>Total carbon concentration (<xref ref-type="table" rid="t0001">Table 1</xref>) and total nitrogen concentration (<xref ref-type="fig" rid="f0001">Fig. 1</xref>) were less in soils from <italic>L. leucocephala</italic> sites than soils from native forest sites. The quotient carbon:nitrogen did not differ between the 2 tree cover types. The pH of soils beneath <italic>L. leucocephala</italic> tree cover was slightly less than that of soils from native tree cover (<xref ref-type="table" rid="t0001">Table 1</xref>). From highest to lowest, the concentrations of extractable macronutrients in the soils from both tree cover types were in the following order: calcium>magnesium>potassium>phosphorus (<xref ref-type="table" rid="t0001">Table 1</xref>). Calcium concentration was greater under diverse native tree cover than under <italic>L. leucocephala</italic> cover, but the other nutrients were not influenced by tree cover type (<xref ref-type="table" rid="t0001">Table 1</xref>).
<fig id="f0001" orientation="portrait" position="float"><label>Figure 1.</label><caption><p>Available nitrate, available ammonium, net nitrification rate, net ammonification rate, net mineralization rate, and total nitrogen of soils as influenced by <italic>Leucaena leucocephala</italic> tree cover versus biodiverse native tree cover. D = Kolmogorov-Smirnov statistic. Mean + SE, n = 10.</p></caption><graphic specific-use="web-only" xlink:href="kcib-09-05-1212792-g001"></graphic></fig><table-wrap id="t0001" orientation="portrait" position="float"><label>Table 1.</label><caption><p>Soil properties as influenced by <italic>Leucaena leucocephala</italic> tree cover vs. native forest cover on Tinian Island. Mean + SE, n = 10. D = Kolmogorov-Smirnov statistic.</p></caption><pmc-comment>OASIS TABLE HERE</pmc-comment><table frame="hsides" rules="groups"><colgroup><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col></colgroup><thead><tr><th align="left"> </th><th align="center">Native</th><th align="center"><italic>Leucaena</italic></th><th align="left"> </th><th align="left"> </th></tr><tr><th align="left">Trait</th><th align="center">tree cover</th><th align="center">tree cover</th><th align="center">D</th><th align="center">P</th></tr></thead><tbody><tr><td align="left">pH</td><td align="char" char="±">7.44 ± 0.06</td><td align="char" char="±">7.15 ± 0.07</td><td align="char" char=".">0.600</td><td align="char" char=".">0.031</td></tr><tr><td align="left">Total Carbon (mg·g<sup>−1</sup>)</td><td align="char" char="±">130.4 ± 3.5</td><td align="char" char="±">73.7 ± 7.4</td><td align="char" char=".">0.900</td><td align="char" char=".">0.001</td></tr><tr><td align="left">Carbon/Nitrogen</td><td align="char" char="±">11.1 ± 0.3</td><td align="char" char="±">10.5 ± 0.3</td><td align="char" char=".">0.300</td><td align="char" char=".">0.675</td></tr><tr><td align="left">Phosphorus (μg·g<sup>−1</sup>)</td><td align="char" char="±">37.7 ± 6.3</td><td align="char" char="±">49.7 ± 4.7</td><td align="char" char=".">0.400</td><td align="char" char=".">0.313</td></tr><tr><td align="left">Potassium (μg·g<sup>−1</sup>)</td><td align="char" char="±">130.1 ± 19.8</td><td align="char" char="±">100.0 ± 8.4</td><td align="char" char=".">0.400</td><td align="char" char=".">0.313</td></tr><tr><td align="left">Magnesium (μg·g<sup>−1</sup>)</td><td align="char" char="±">218.1 ± 25.2</td><td align="char" char="±">237.6 ± 15.2</td><td align="char" char=".">0.300</td><td align="char" char=".">0.675</td></tr><tr><td align="left">Calcium (μg·g<sup>−1</sup>)</td><td align="char" char="±">4957 ± 374</td><td align="char" char="±">3316 ± 281</td><td align="char" char=".">0.700</td><td align="char" char=".">0.007</td></tr><tr><td align="left">Available nitrogen (μg·g<sup>−1</sup>)</td><td align="char" char="±">102.2 ± 14.9</td><td align="char" char="±">107.6 ± 15.2</td><td align="char" char=".">0.300</td><td align="char" char=".">0.675</td></tr></tbody></table></table-wrap></p><p>The differences in various components of nitrogen cycling were highly contrasting between the 2 forest tree cover types. Net nitrification rate of soils beneath <italic>L. leucocephala</italic> cover was 4.25-fold greater than soils beneath native tree cover (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). Net ammonification of soils was positive beneath native tree cover, but negative beneath <italic>L. leucocephala</italic> tree cover (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). These results caused the soils from <italic>L. leucocephala</italic> microsites to exhibit net mineralization that was 156% greater than that from the diverse native forest microsites (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). Available nitrate, available ammonium, and total available nitrogen did not differ between the 2 forest tree cover types (<xref ref-type="table" rid="t0001">Table 1</xref>, <xref ref-type="fig" rid="f0001">Fig. 1</xref>).</p><p>From highest to lowest, the concentrations of measured metals in the soils were in the following order: zinc>copper>chromium>cobalt>nickel>cadmium>selenium>lead (<xref ref-type="table" rid="t0002">Table 2</xref>). Chromium, cobalt, and nickel were greater in the <italic>L. leucocephala</italic> soils than in the native tree soils. Selenium and zinc were greater in the native tree soils than in the <italic>L. leucocephala</italic> soils. Cadmium, copper, and lead were not influenced by tree cover type.
<table-wrap id="t0002" orientation="portrait" position="float"><label>Table 2.</label><caption><p>Metal content of soils in Tinian Island as influenced by <italic>Leucaena leucocephala</italic> tree cover versus native forest cover. Mean + SE, n = 10. D = Kolmogorov-Smirnov statistic.</p></caption><pmc-comment>OASIS TABLE HERE</pmc-comment><table frame="hsides" rules="groups"><colgroup><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col><col width="86.4pt" align="left"></col></colgroup><thead><tr><th align="left"> </th><th align="center">Native</th><th align="center"><italic>Leucaena</italic></th><th align="left"> </th><th align="left"> </th></tr><tr><th align="left">Property (μg·g<sup>−1</sup>)</th><th align="center">cover</th><th align="center">cover</th><th align="center">D</th><th align="center">P</th></tr></thead><tbody><tr><td align="left">Cadmium</td><td align="char" char="±">2.97 ± 0.21</td><td align="char" char="±">3.14 ± 0.22</td><td align="char" char=".">0.200</td><td align="char" char=".">0.975</td></tr><tr><td align="left">Chromium</td><td align="char" char="±">32.32 ± 1.83</td><td align="char" char="±">53.21 ± 1.21</td><td align="char" char=".">1.000</td><td align="char" char=".">0.001</td></tr><tr><td align="left">Cobalt</td><td align="char" char="±">9.39 ± 0.64</td><td align="char" char="±">21.61 ± 2.39</td><td align="char" char=".">1.000</td><td align="char" char=".">0.001</td></tr><tr><td align="left">Copper</td><td align="char" char="±">49.20 ± 7.08</td><td align="char" char="±">50.81 ± 11.68</td><td align="char" char=".">0.500</td><td align="char" char=".">0.111</td></tr><tr><td align="left">Lead</td><td align="char" char="±">0.012 ± 0.004</td><td align="char" char="±">0.015 ± 0.002</td><td align="char" char=".">0.300</td><td align="char" char=".">0.675</td></tr><tr><td align="left">Nickel</td><td align="char" char="±">3.52 ± 0.28</td><td align="char" char="±">12.91 ± 0.88</td><td align="char" char=".">1.000</td><td align="char" char=".">0.001</td></tr><tr><td align="left">Selenium</td><td align="char" char="±">1.43 ± 0.04</td><td align="char" char="±">1.27 ± 0.03</td><td align="char" char=".">0.600</td><td align="char" char=".">0.031</td></tr><tr><td align="left">Zinc</td><td align="char" char="±">95.73 ± 3.85</td><td align="char" char="±">77.33 ± 2.78</td><td align="char" char=".">1.000</td><td align="left">0.001</td></tr></tbody></table></table-wrap></p></sec><sec sec-type="discussion" id="s0003"><title>Discussion</title><p>Insular habitats including islands may be more susceptible to plant invasions than continental habitats.<xref rid="cit0018" ref-type="bibr"><sup>18</sup></xref> Yet the Pacific islands have been under-represented in publications on invasive plant species.<xref rid="cit0011" ref-type="bibr"><sup>11</sup></xref> We have addressed this bias with a look at a widespread invasive woody tree species in a small oceanic island habitat.</p><p>This diagnosis of the integrative changes in soil chemical properties caused by persistent <italic>L. leucocephala</italic> tree cover is the first empirical look at how this invasive tree affects ecosystems in the Mariana Islands. We have shown that a high proportion of soil nutrients and metals were substantially changed by the <italic>L. leucocephala</italic> tree cover. Various components of the nitrogen cycle were among the soil properties that were most affected by tree cover type. That total nitrogen was greater under native tree cover may seem counterintuitive, as <italic>L. leucocephala</italic> is a well-known legume, its <italic>Rhizobium</italic> endosymbionts are prevalent in the calcareous soils of the Mariana Islands,<xref rid="cit0019" ref-type="bibr"><sup>19</sup></xref> and none of the prevalent native tree species at our site associate with nitrogen-fixing symbionts. Indeed, habitats dominated by legume species tend to exhibit greater soil nitrogen.<xref rid="cit0020" ref-type="bibr"><sup>20</sup></xref> These interesting relationships were reconciled by the acute differences in nitrification and ammonification. Net mineralization rates in the <italic>L. leucocephala</italic> microsites were 256% of those of native forest microsites. Moreover, net nitrification of the <italic>L. leucocephala</italic> soils exceeded net mineralization, but net nitrification of the native forest soils was only 87% of net mineralization. Clearly, nitrogen-fixing bacteria genera <italic>Nitrosomonas</italic> and <italic>Nitrobacter</italic> were highly active under <italic>L. leucocephala</italic> cover and less active under native tree cover.</p><p>We propose that the prodigious nitrification in soils beneath <italic>L. leucocephala</italic> patches causes excessive nitrogen losses from the system through leaching since nitrate is highly mobile.<xref rid="cit0021" ref-type="bibr"><sup>21</sup></xref> In contrast, the limited nitrification of the native forest microsites protects those soils from similar losses. Annual precipitation in Tinian is 204 cm (<ext-link ext-link-type="uri" xlink:href="http://www.wunderground.com">www.wunderground.com</ext-link>), evincing considerable leaching potential. As a result, the soil nitrogen pool is labile in the <italic>L. leucocephala</italic> patches and recalcitrant in the native forest patches.</p><p>Biodiversity is critical for sustaining many components of ecosystem services and maintaining forest productivity.<xref rid="cit0022" ref-type="bibr"><sup>22</sup></xref> Indeed, species mixtures and high plant biodiversity may increase nitrogen retention, reduce nitrogen losses, and decrease the potential for groundwater contamination due to nitrogen leaching.<xref rid="cit0023" ref-type="bibr"><sup>23,24</sup></xref> Therefore, our native forest may have exhibited greater retention of nitrogen simply because it had greater biodiversity than the <italic>L. leucocephala</italic> microsites, which were monospecific.</p><p>Two issues from our study are relevant for climate change research. First, climate change is predicted to increase nitrate leaching as extreme events increase in frequency.<xref rid="cit0025" ref-type="bibr"><sup>25</sup></xref> Therefore, in a climate change scenario nitrate leaching in <italic>L. leucocephala</italic> sites could be aggravated. Second, soil organic matter is the largest terrestrial carbon pool,<xref rid="cit0026" ref-type="bibr"><sup>26</sup></xref> and soil carbon and nitrogen mineralization are closely coupled.<xref rid="cit0027" ref-type="bibr"><sup>27,28</sup></xref> Our results indicate that monospecific <italic>L. leucocephala</italic> sites act as less effective carbon sinks than native forest sites.</p><p>Ecosystem changes caused by invasive species are some of the complex consequences of anthropogenic changes to the global environment. A full understanding of how to manage invasive species cannot develop in the absence of an all-inclusive viewpoint founded in empirical studies.<xref rid="cit0029" ref-type="bibr"><sup>29</sup></xref> Meta-analyses have shown that net primary production, litter decomposition, and altered nitrogen cycle are some of the most common ways that successful invasive plants modify nutrient cycling.<xref rid="cit0030" ref-type="bibr"><sup>30</sup></xref> Bardon et al.<xref rid="cit0031" ref-type="bibr"><sup>31</sup></xref> recently reported that root exudates of a successful invasive plant reduced metabolic activity of 2 denitrifying bacteria species, adding a previously unknown means by which invasive plants may directly influence the nitrogen cycle.</p><p>The differences in soil chemical components we have reported from <italic>L. leucocephala</italic> vs. diverse native tree forest cover may influence understory vegetation growth and health. In our experimental site, the main understory species of interest was the Guam-sourced <italic>C. micronesica</italic> germplasm that we introduced and managed. <italic>Cycas micronesica</italic> is one of more than 350 species of extant cycads.<xref rid="cit0032" ref-type="bibr"><sup>32</sup></xref> This plant group is among the most threatened groups of plants worldwide.<xref rid="cit0033" ref-type="bibr"><sup>33,34</sup></xref> Developing successful management strategies for cycad conservation is an urgent agenda. An ongoing refinement of our <italic>C. micronesica</italic> conservation program based on national threatened and international endangered listings<xref rid="cit0002" ref-type="bibr"><sup>2,3</sup></xref> fits into that international agenda. Although we have shown the soils in <italic>L. leucocephala</italic> microsites exhibited substantial differences from soils in biodiverse native forest microsites, the magnitude and direction of differences in macronutrients (<xref ref-type="table" rid="t0001">Table 1</xref>) and metals (<xref ref-type="table" rid="t0002">Table 2</xref>) were not likely to explain why <italic>C. micronesica</italic> plants have grown better as understory plants in the <italic>L. leucocephala</italic> microsites. Furthermore, the increased net nitrification and net mineralization of soils in <italic>L. leucocephala</italic> microsites are not likely to substantially benefit the understory <italic>C. micronesica</italic> plants, as available nitrogen did not differ between the soils from the 2 forest cover types. Additionally, all cycad plants associate with nitrogen-fixing cyanobacteria endosymbionts, therefore cycads may not be affected by ecologically relevant differences in nitrogen among various soils.</p><p>Several critical research needs are illuminated by this study. A greater understanding of litterfall quantity, seasonality, and quality may reveal some of the factors that mediate the differences in soil traits between the native forest cover and <italic>L. leucocephala</italic> cover. Reciprocal litter incubation studies would tease apart the influences of litter quality and soil microbes that influence decomposition speed. Which soil microbes are influential players in the changes that <italic>L. leucocephala</italic> imposes on soils may be identified by DNA sequencing of bulk soils or rhizosphere, and would greatly improve mechanistic insight. <italic>Cycas micronesica</italic> plant growth has been greater under <italic>L. leucocephala</italic> cover than under native tree cover in our ex situ germplasm, yet soil nutrition does not appear to mediate this response. Site differences in incident light, relative humidity, and temperature may be more important factors than the soil properties for explaining the disparity in growth and health of the <italic>C. micronesica</italic> germplasm.</p><p>Many Guam and Tinian habitats that have experienced past disturbance are characterized by <italic>L. leucocephala</italic> cover. Once established, this species effectively monopolizes emergent canopy cover. Future large-scale restoration plans for Guam and Tinian include landscape-scale efforts to remove invasive, non-native plant species.<xref rid="cit0035" ref-type="bibr"><sup>35</sup></xref> Our results indicate that forest restoration goals may require many years to achieve following the invasive tree removal, considering the transformed soil properties that need to be restored.</p></sec><sec sec-type="materials|methods" id="s0004"><title>Materials and methods</title><p>The experimental site was located on the island of Tinian in karsty outcrop soils (Loamy-skeletal, carbonatic, isohyperthermic Lithic Haplustolls).<xref rid="cit0036" ref-type="bibr"><sup>36</sup></xref> A robust <italic>C. micronesica</italic> planting is being maintained along a north-south oriented ecotone between diverse native tree cover and an adjacent belt of <italic>L. leucocephala</italic> cover. The soil samples were obtained within this ex situ germplasm. The <italic>L. leucocephala</italic> sites were primarily mono-specific for the emergent canopy cover, but contained a diverse understory vegetation palette. The native forest sites were highly diverse, and dominant tree species were <italic>Pisonia grandis, Psychotria mariana, Aglaia mariannensis, Cynometra ramiflora</italic>, and <italic>Eugenia palumbis</italic>.</p><p>We collected paired soil samples from 10 locations along the ecotone on 10 Sept 2014. Each sample from the <italic>L. leucocephala</italic> cover was located 25-35 m east of its paired sample from the native forest cover. The soil cores were collected from the A horizon from 0-10 cm. Rainfall for the 2 weeks prior to the soil harvests averaged 9.54 mm·d<sup>-1</sup> (<ext-link ext-link-type="uri" xlink:href="http://www.wunderground.com">www.wunderground.com</ext-link>).</p><sec><title>Analyses</title><p>A portion of each sample was dried at 50°C then total carbon and nitrogen were determined by dry combustion (Nelson and Sommers);<xref rid="cit0037" ref-type="bibr"><sup>37</sup></xref> extractable P, K, Mg, and Ca were determined by Mehlich-3 digestion (Mehlich 1984);<xref rid="cit0038" ref-type="bibr"><sup>38</sup></xref> and total metals were determined by nitric acid digestion.<xref rid="cit0039" ref-type="bibr"><sup>39</sup></xref> Nitrate and ammonium were determined colorimetrically from fresh moist soil samples following 2<sc>M</sc> KCl extraction. Soil was incubated using the buried bag method<xref rid="cit0040" ref-type="bibr"><sup>40</sup></xref> in a homogeneous site at 28°C (range 25°C – 31°C) soil temperature for 32 d. Nitrate and ammonium were determined at the end of the incubation period. Net nitrification rate was calculated by subtracting initial from final nitrate concentration and dividing by the incubation period. Net ammonification rate was calculated by subtracting initial from final ammonium concentration and dividing by the incubation period. Net mineralization was calculated as the sum of nitrification and ammonification.</p><p>The data did not meet requirements for parametric analysis, primarily because of unequal variances. We used the non-parametric and distribution-free Kolmogorov-Smirnov 2-sample test<xref rid="cit0041" ref-type="bibr"><sup>41</sup></xref> to test the null hypothesis that the 2 groups of soil samples were not different in each of the chemical properties that were quantified. This test does not require any assumption about the distribution of data. Levels of significance of at least P < 0.05 were considered significant.</p></sec></sec></body><back><sec><title>Disclosure of potential conflicts of interest</title><p>No potential conflicts of interest were disclosed.</p></sec><sec><title>Funding</title><p>Support provided by the United States Department of the Navy through N40192-12-2-8000 and N40192-13-2-8003, administered by Naval Facilities Engineering Command Marianas, Guam.</p></sec><ref-list><title>References</title><ref id="cit0001"><label>[1]</label><mixed-citation publication-type="book"><person-group person-group-type="author"><name name-style="western"><surname>Donnegan</surname><given-names>JA</given-names></name>, <name name-style="western"><surname>Butler</surname><given-names>SL</given-names></name>, <name name-style="western"><surname>Grabowiecki</surname><given-names>W</given-names></name>, <name name-style="western"><surname>Hiserote</surname><given-names>BA</given-names></name>, <name name-style="western"><surname>Limtiaco</surname><given-names>D</given-names></name></person-group><chapter-title>Guam's forest resources, 2002</chapter-title><source>Resource Bulletin PNW-RB-243</source>. <publisher-loc>Portland</publisher-loc>: <publisher-name>US. 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