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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2018.00294</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Perspective</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Cuban Connection: Regional Role for Florida&#x00027;s Manatees</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Alvarez-Alem&#x000E1;n</surname> <given-names>Anmari</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/588941/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Austin</surname> <given-names>James D.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/588638/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Jacoby</surname> <given-names>Charles A.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/172246/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Frazer</surname> <given-names>Thomas K.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x0002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/166195/overview"/>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>School of Natural Resources and Environment, Institute of Food and Agricultural Sciences, University of Florida</institution>, <addr-line>Gainesville, FL</addr-line>, <country>United States</country></aff>
<aff id="aff2"><sup>2</sup><institution>Centro de Investigaciones Marinas, Universidad de la Habana</institution>, <addr-line>La Habana</addr-line>, <country>Cuba</country></aff>
<aff id="aff3"><sup>3</sup><institution>Department of Wildlife Ecology and Conservation, Institute of Food and Agricultural Sciences, University of Florida</institution>, <addr-line>Gainesville, FL</addr-line>, <country>United States</country></aff>
<aff id="aff4"><sup>4</sup><institution>Soil and Water Sciences Department, Institute of Food and Agricultural Sciences, University of Florida</institution>, <addr-line>Gainesville, FL</addr-line>, <country>United States</country></aff>
<aff id="aff5"><sup>5</sup><institution>Fisheries and Aquatic Sciences Program, School of Forest Resources and Conservation, Institute of Food and Agricultural Sciences, University of Florida</institution>, <addr-line>Gainesville, FL</addr-line>, <country>United States</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: E. Christien Michael Parsons, George Mason University, United States</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Laura J. May-Collado, University of Vermont, United States; Lorelei Diane Crerar, George Mason University, United States</p></fn>
<corresp id="c001">&#x0002A;Correspondence: Thomas K. Frazer <email>frazer&#x00040;ufl.edu</email></corresp>
<fn fn-type="other" id="fn001"><p>This article was submitted to Marine Conservation and Sustainability, a section of the journal Frontiers in Marine Science</p></fn></author-notes>
<pub-date pub-type="epub">
<day>21</day>
<month>08</month>
<year>2018</year>
</pub-date>
<pub-date pub-type="collection">
<year>2018</year>
</pub-date>
<volume>5</volume>
<elocation-id>294</elocation-id>
<history>
<date date-type="received">
<day>08</day>
<month>05</month>
<year>2018</year>
</date>
<date date-type="accepted">
<day>31</day>
<month>07</month>
<year>2018</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x000A9; 2018 Alvarez-Alem&#x000E1;n, Austin, Jacoby and Frazer.</copyright-statement>
<copyright-year>2018</copyright-year>
<copyright-holder>Alvarez-Alem&#x000E1;n, Austin, Jacoby and Frazer</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/"><p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p></license>
</permissions>
<abstract>
<p>The status of the West Indian manatee [<italic>Trichechus manatus</italic> (Linnaeus, 1758)] remains enigmatic. The International Union for Conservation of Nature classifies the species as &#x0201C;Vulnerable&#x0201D; and the two subspecies [<italic>T. m. manatus</italic> (Linnaeus, 1758) and <italic>T. m. latirostris</italic> (Harlan, 1824)] as &#x0201C;Endangered.&#x0201D; The U.S. Fish and Wildlife Service now classifies West Indian manatees as &#x0201C;Threatened&#x0201D; primarily due to increased numbers of <italic>T. m. latirostris</italic>. Fully resolving the status of the species will require a better understanding of its population biology throughout the broader Caribbean region, especially the form and strength of genetic and demographic connections. Genetic studies indicate limited interbreeding among recent generations of groups of manatees separated by unsuitable coastal habitat or large expanses of open water, such as the Florida Straits. However, documentation of two, independent immigrations from Florida to Cuba within one generation raises the possibility of important demographic connections, especially if the number of manatees in Cuba is small. In addition, these events may foreshadow a change in the role that Florida&#x00027;s manatees play in effective conservation of <italic>T. manatus</italic> throughout its Caribbean range, with the potential for further connections as numbers of manatees in Florida increase and the availability of suitable habitat and food decreases.</p>
</abstract>
<kwd-group>
<kwd>West Indian manatee</kwd>
<kwd><italic>Trichechus manatus</italic></kwd>
<kwd>genetic connectivity</kwd>
<kwd>demographic connectivity</kwd>
<kwd>conservation</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="16"/>
<page-count count="4"/>
<word-count count="2249"/>
</counts>
</article-meta>
</front>
<body>
<p>The status of the West Indian manatee [<italic>Trichechus manatus</italic> (Linnaeus, 1758)] is a concern. The species is classified as &#x0201C;Vulnerable&#x0201D; by the International Union for Conservation of Nature, and two subspecies [<italic>T. m. manatus</italic> (Linnaeus, 1758) and <italic>T. m. latirostris</italic> (Harlan, 1824)] are classified as &#x0201C;Endangered&#x0201D; (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.2305/IUCN.UK.2008.RLTS.T22103A9356917.en">https://doi.org/10.2305/IUCN.UK.2008.RLTS.T22103A9356917.en</ext-link>). Recently, the U.S. Fish and Wildlife Service reclassified the West Indian manatee as &#x0201C;Threatened&#x0201D; rather than &#x0201C;Endangered&#x0201D; primarily due to a marked increase in numbers of <italic>T. m. latirostris</italic>, the Florida subspecies, over the last 25 years (Federal Register, <xref ref-type="bibr" rid="B3">2017</xref>). Overall, the status of <italic>T. manatus</italic> throughout its broader Caribbean range remains uncertain because estimates of numbers and demographic trends for many populations are based on anecdotal information or gray literature (Self-Sullivan and Mignucci-Giannoni, <xref ref-type="bibr" rid="B13">2012</xref>; Alvarez-Alem&#x000E1;n et al., <xref ref-type="bibr" rid="B1">2018</xref>). One key uncertainty is whether manatees in different Caribbean regions should be managed as distinct populations because unsuitable coastal habitat or large expanses of marine water may preclude movements that establish genetic and demographic connectivity (Garcia-Rodriguez et al., <xref ref-type="bibr" rid="B4">1998</xref>; Lowe and Allendorf, <xref ref-type="bibr" rid="B9">2010</xref>).</p>
<p>Studies of genetic markers document several phylogeographic patterns for <italic>T. manatus</italic> among and within regions of the Caribbean. Sequences in mitochondrial deoxyribonucleic acids (mtDNA) identified three distinct lineages occupying (i) Florida and the West Indies, (ii) the Gulf of Mexico to the northern Caribbean coast of South America, and (iii) the northeast Atlantic coast of South America (Garcia-Rodriguez et al., <xref ref-type="bibr" rid="B4">1998</xref>). Examination of mtDNA from additional manatees identifies a similar set of lineages and suggests historical barriers to dispersal that separate the Dominican Republic and Puerto Rico from other parts of the Greater Antilles (Vianna et al., <xref ref-type="bibr" rid="B16">2006</xref>). Variation in microsatellites also differentiates manatees in Mexico from those in Florida, with minimal evidence of interbreeding, and the results further identify two relatively distinct groups within Mexico, a mixing zone with alleles from both of those groups, and the potential influence of migrants from Belize (Nourisson et al., <xref ref-type="bibr" rid="B10">2011</xref>). In Belize, mtDNA and microsatellite markers from groups of manatees separated by &#x0003C;100 km indicate genetic differentiation characteristic of isolated populations (Hunter et al., <xref ref-type="bibr" rid="B6">2010</xref>). Furthermore, manatees in Belize appear unrelated to manatees in Florida (Hunter et al., <xref ref-type="bibr" rid="B6">2010</xref>). Within the eastern Caribbean, a study that combined mtDNA and nuclear microsatellites provides support for the separation of <italic>T. m. latirostris</italic> in Florida from <italic>T. m. manatus</italic> in Puerto Rico (Hunter et al., <xref ref-type="bibr" rid="B7">2012</xref>). In contrast, a study of mtDNA in manatees from Cuba suggests historical links to manatees in Florida and the Greater Antilles and the possibility of a link to Belize (Hern&#x000E1;ndez Mart&#x000ED;nez et al., <xref ref-type="bibr" rid="B5">2013</xref>). Within Florida, microsatellite markers yield very weak evidence of genetic differentiation among the four recognized management units (Tucker et al., <xref ref-type="bibr" rid="B15">2012</xref>). Overall, the results of existing genetic analyses indicate reduced dispersal among many parts of the Caribbean and levels of dispersal within specific localities that appear to exhibit a positive relationship with increased abundance of manatees and availability of suitable corridors for dispersal. However, these studies of genetics do not assess demographic connectivity as defined by the effect of immigrants on vital rates, which is an important element in effective conservation of <italic>T. manatus</italic> (Lowe and Allendorf, <xref ref-type="bibr" rid="B9">2010</xref>).</p>
<p>Initial evidence of a potentially important connection comes from photographs of a female and calf in Florida in 2006 and a subsequent sighting of these manatees in Cuban waters near Havana in 2007 (Alvarez-Alem&#x000E1;n et al., <xref ref-type="bibr" rid="B2">2010</xref>). In November 2017, another adult and a calf were photographed along Cuba&#x00027;s northwestern coast near Havana (Figure <xref ref-type="fig" rid="F1">1A</xref>), and the U.S. Geological Survey&#x00027;s Manatee Individual Photo-identification System positively identified the adult&#x00027;s two sets of scars as belonging to FM582 (Figure <xref ref-type="fig" rid="F1">1B</xref>). Records of FM582 in southwest Florida span 11 years, and it was sighted near Fort Myers, Florida in January 2017. Calves were near FM582 in multiple years, which suggests she is a female. Both she and the calf likely traveled from Florida to Cuba because calves depend on their mothers for 1&#x02013;2 winters (O&#x00027;Shea et al., <xref ref-type="bibr" rid="B11">1995</xref>). These sightings represent direct evidence of connectivity between two geographically separated populations, and two, independent immigrations within a decade (i.e., less than a generation) suggest that such movements may have demographic import. In conjunction with shared mtDNA haplotypes (Hern&#x000E1;ndez Mart&#x000ED;nez et al., <xref ref-type="bibr" rid="B5">2013</xref>), such sightings suggest that populations of manatees in Florida, Cuba, and other Caribbean regions may not remain evolutionarily or demographically distinct if manatees successfully cross expanses of open water, such as the Florida Straits. Increased connectivity in the Caribbean region engenders important ramifications for conservation, especially selecting the optimal number of management units and incorporating source-sink dynamics into decisions about levels of protection (Pulliam, <xref ref-type="bibr" rid="B12">1988</xref>; Lowe and Allendorf, <xref ref-type="bibr" rid="B9">2010</xref>).</p>
<fig id="F1" position="float">
<label>Figure 1</label>
<caption><p>Photographs of a manatee taken in Cuba <bold>(A)</bold> and in southwest Florida <bold>(B)</bold> showing the scars that identified it as FM582. Permissions shown on photographs.</p></caption>
<graphic xlink:href="fmars-05-00294-g0001.tif"/>
</fig>
<p>Several challenges prevent a full interpretation of the importance of these sightings. Demographic connectivity requires the migrants to exert an effect on vital rates in the population they enter (Lowe and Allendorf, <xref ref-type="bibr" rid="B9">2010</xref>). Unfortunately, there is no information on the fate of the manatees that reached Cuba. In addition, the size of the Cuban population of <italic>T. manatus</italic> is uncertain (Alvarez-Alem&#x000E1;n et al., <xref ref-type="bibr" rid="B1">2018</xref>), so an estimate of the relative contribution of these immigrants cannot be derived. Programs to address these issues would advance conservation of the Antillean manatee. For example, rigorous censuses of manatees in Cuba would provide an estimate of the size of the population, and a program that involves professional and citizen scientists in the development, maintenance, and application of a photographic identification system would help identify additional immigrants and clarify their fates. Regardless of the fate of the recent immigrants, the two sightings suggest that the regional significance of Florida&#x00027;s population of manatees may be increasing as numbers around the state rise and the availability of suitable habitat and food decreases (Littles, <xref ref-type="bibr" rid="B8">2016</xref>). Overall, effective conservation of <italic>T. manatus</italic> will be difficult without close cooperation among researchers in all relevant countries and a supportive political environment (Stone, <xref ref-type="bibr" rid="B14">2017</xref>).</p>
<sec id="s1">
<title>Ethics statement</title>
<p>This article was based on photo-documented observations only. There was no direct handling or manipulation of animals. As such, the work is exempt from IACUC oversight.</p>
</sec>
<sec id="s2">
<title>Author contributions</title>
<p>AA-A, JA, CJ, and TF contributed equally to the preparation of this manuscript.</p>
<sec>
<title>Conflict of interest statement</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The reviewer LC and handling editor declared their shared affiliation at the time of the review.</p>
</sec>
</sec>
</body>
<back>
<ack><p>Many thanks to the people from the Almendares River community who informed us about the manatee sighting promptly. We are grateful to A Ruiz Abierno who obtained and provided the video taken in Cuba. Thanks to C Beck who verified the match to FM582 and to K Rood and S Barton who scoured the Manatee Individual Photo-Identification system, provided the image from southwest Florida, and supplied the history of sightings in Florida. We also thank Sea to Shore Alliance for supporting manatee conservation in Cuba.</p>
</ack>
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<fn-group>
<fn fn-type="financial-disclosure"><p><bold>Funding.</bold> Funding for this work was provided by the University of Florida&#x00027;s Institute of Food and Agricultural Sciences.</p>
</fn>
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