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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.2024.1401258</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Brief Research Report</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Observational data on the reproductive condition of female Oceania fantail rays, <italic>Taeniura lessoni</italic>, from Drawaqa Island, Fiji</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Glaus</surname>
<given-names>Kerstin</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2685268"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Vierus</surname>
<given-names>Tom</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2773616"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/validation/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Macfarlane</surname>
<given-names>Robert</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
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</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Institute of Marine Resources, School of Agriculture, Geography, Environment, Ocean and Natural Sciences (SAGEONS), The University of the South Pacific</institution>, <addr-line>Suva</addr-line>, <country>Fiji</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Independent Researcher</institution>, <addr-line>Suva</addr-line>, <country>Fiji</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Barefoot Manta Island Resort</institution>, <addr-line>Drawaqa Island</addr-line>, <country>Fiji</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Drawaqa Marine Conservation Trust</institution>, <addr-line>Drawaqa Island</addr-line>, <country>Fiji</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Brendan Shea, Beneath the Waves, Inc., United States</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Alfonsina E. Romo-Curiel, The University of Texas at Austin, United States</p>
<p>Daniel Fahy, Nova Southeastern University, United States</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Kerstin Glaus, <email xlink:href="mailto:kerstin.glaus@usp.ac.fj">kerstin.glaus@usp.ac.fj</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>05</day>
<month>07</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1401258</elocation-id>
<history>
<date date-type="received">
<day>15</day>
<month>03</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>14</day>
<month>06</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Glaus, Vierus and Macfarlane</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Glaus, Vierus and Macfarlane</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>Data on the reproductive biology of elasmobranchs are essential for understanding their life history. Published studies on batoid ray reproductive biology are comparatively scarce, leading to limited understanding and data gaps. The Oceania fantail ray, <italic>Taeniura lessoni</italic>, is a good example. This Data Deficient nearshore stingray is restricted to Melanesia, with lacking biological and ecological data, including reproduction. To expand upon the limited life-history data for this species, this short paper provides observational data on the reproductive condition in female <italic>T. lessoni</italic>, at Drawaqa Island, Fiji. Field work involved direct observations and ocean temperature measurements. Over 40 days spanning three months, 105 surveys were conducted across five sites, resulting in 71 sightings of the species. Based on spot patterns and body markings, four female individuals were identified. Between January and March 2024, these females exhibited convex dorsa indicating advanced gestation, transitioning to concave dorsa suggesting parturition. The presence of neonates from early March onwards coincided with the estimated parturition period inferred from the rays' condition. Furthermore, a female previously pregnant was photographed with a dermal abrasion around her pectoral fin, possibly indicating pre-copulatory biting, suggesting a continuous reproductive cycle. The average monthly water temperature at the surveyed sites remained relatively stable throughout the study. Collectively, our findings suggest that Drawaqa Island provides suitable habitat niches for reproductive activities in female <italic>T. lessoni</italic>. Repeated and long-term data is certainly needed to confirm either a continuous reproductive cycle or seasonal peaks. While preliminary, our observational data represents the first documentation on female reproductive condition in a stingray in Fiji.</p>
</abstract>
<kwd-group>
<kwd>batoids</kwd>
<kwd>reproductive biology</kwd>
<kwd>Melanesia</kwd>
<kwd>data deficient species</kwd>
<kwd>range restricted</kwd>
<kwd>
<italic>Dasyatidae</italic>
</kwd>
</kwd-group>
<counts>
<fig-count count="3"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="55"/>
<page-count count="7"/>
<word-count count="2622"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Discoveries</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Elasmobranchs (sharks, skates, and rays) are an evolutionarily conserved, diverse, and threatened vertebrate group (<xref ref-type="bibr" rid="B9">Dulvy et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B46">Sherman et&#xa0;al., 2023</xref>). Threats encompass direct and indirect fisheries activities (<xref ref-type="bibr" rid="B55">Worm et&#xa0;al., 2024</xref>), ocean warming (<xref ref-type="bibr" rid="B41">Rosa et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B36">Osgood et&#xa0;al., 2021</xref>), and habitat degradation, including disturbances to coastal breeding and pupping grounds (<xref ref-type="bibr" rid="B45">Sherman et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B47">Simpfendorfer et&#xa0;al., 2023</xref>).</p>
<p>Elasmobranchs have developed nine distinct reproductive strategies (<xref ref-type="bibr" rid="B3">Awruch, 2015</xref>) over their 400-million-year evolutionary history (<xref ref-type="bibr" rid="B26">Kriwet et&#xa0;al., 2008</xref>). Batoids, the most diverse group of cartilaginous fishes (<xref ref-type="bibr" rid="B2">Aschliman et&#xa0;al., 2012</xref>), comprise four orders, 23 families and 663 species, with many more yet to be described (<xref ref-type="bibr" rid="B28">Last et&#xa0;al., 2016a</xref>). While the largest order, Rajiformes (skates), is strictly oviparous, the remaining orders (<italic>Myliobatiformes, Rhinopristiformes</italic>, and <italic>Torpediniformes</italic>) are viviparous, with lecithotrophic and matrotrophic modes of reproduction (<xref ref-type="bibr" rid="B6">Conrath and Musick, 2012</xref>; <xref ref-type="bibr" rid="B3">Awruch, 2015</xref>). Myliobatiformes, including the family <italic>Dasyatidae</italic> (stingrays), rely on lipid histotrophy via trophonemata. Stingrays contain 19 genera, 86 extant species (<xref ref-type="bibr" rid="B28">Last et&#xa0;al., 2016a</xref>), and represent the most abundant group of rays occurring in tropical and subtropical coastal waters (<xref ref-type="bibr" rid="B52">White and Dharmadi, 2007</xref>). Stingrays produce broods of one to 10 with gestation periods up to 11 months, while smaller tropical species have shorter gestation periods of three to six months (<xref ref-type="bibr" rid="B11">Fahy et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B38">Pierce et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B32">Mull et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B13">Furumitsu et&#xa0;al., 2019</xref>). The duration of reproductive cycles varies among stingrays (<xref ref-type="bibr" rid="B51">Walker, 2020</xref>), and annual and biannual cycles have been confirmed in wild populations (<xref ref-type="bibr" rid="B39">Ram&#xed;rez-Mosqueda et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B42">Schieber et&#xa0;al., 2023</xref>). However, batoid ecology and life history are comparably less understood than in sharks (<xref ref-type="bibr" rid="B5">Br&#xe4;utigam et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B30">Martins et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B23">Jorgensen et&#xa0;al., 2022</xref>), with more than 250 Data Deficient species in the International Union for Conservation of Nature (IUCN) Red List (<xref ref-type="bibr" rid="B20">IUCN, 2023</xref>).</p>
<p>The Oceania fantail ray, <italic>Taeniura lessoni</italic>, (Last, White &amp; Naylor, 2016) was described in 2016 (<xref ref-type="bibr" rid="B29">Last et&#xa0;al., 2016b</xref>), marking the second species within the genus alongside the widely-distributed bluespotted lagoon ray, <italic>T. lymma</italic>, (Forssk&#xe5;l, 1775). The disc width (DW), defined as the maximum distance between the wingtips (<xref ref-type="bibr" rid="B44">Serra-Pereira et&#xa0;al., 2010</xref>), ranges between 18 cm to 22 cm in female <italic>T. lessoni</italic> paratypes (<xref ref-type="bibr" rid="B29">Last et&#xa0;al., 2016b</xref>). The mature male holotype has a 20.9 cm DW, with large claspers measuring 21.2% of this width, while the immature paratype male measured 18.5 cm (<xref ref-type="bibr" rid="B29">Last et&#xa0;al., 2016b</xref>). <italic>T. lessoni</italic> is smaller than <italic>T. lymma</italic> and lacks the pair of vivid blue longitudinal stripes found along the tail. Also, the species appears to be restricted to Melanesia, including Papua New Guinea, Solomon Islands, Vanuatu, and Fiji (<xref ref-type="bibr" rid="B28">Last et&#xa0;al., 2016a</xref>; <xref ref-type="bibr" rid="B19">Hylton et&#xa0;al., 2017</xref>). <italic>T. lessoni</italic> inhabits shallow-water coral reefs, usually at depths of 20 m and less, typically shelters in caves during the day, and forages at night (<xref ref-type="bibr" rid="B28">Last et&#xa0;al., 2016a</xref>). In <italic>T. lymma</italic>, the embryos are initially nourished by the yolk sac and subsequently feed on uterine secretions (<xref ref-type="bibr" rid="B17">Hamlett et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B35">Musick et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B1">Abel and Grubbs, 2020</xref>). Fecundity has been reported with brood sizes of one to seven in wild individuals (<xref ref-type="bibr" rid="B12">Ferreira, 2013</xref>; <xref ref-type="bibr" rid="B37">Pereira et&#xa0;al., 2017</xref>). However, reproductive biology data for <italic>T. lessoni</italic> is lacking, and this species is classified as Data Deficient by the IUCN (<xref ref-type="bibr" rid="B27">Kyne and Finucci, 2018</xref>). In Fiji, <italic>T. lessoni</italic> is regularly observed during snorkeling trips and dives. Sightings are reported from across the archipelago (<xref ref-type="bibr" rid="B14">Glaus et&#xa0;al., 2024a</xref>), and the species is caught in small-scale fishing activities (<xref ref-type="bibr" rid="B15">Glaus et&#xa0;al., 2024b</xref>). To expand upon the limited life-history data for this range-restricted species, this study represents preliminary observational data focusing on the distribution and reproductive condition of female <italic>T. lessoni</italic>.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Study area</title>
<p>The study area was located approximately between 17&#xb0; South latitude and 177&#xb0; East longitude (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>) and focused on the coastal waters off Drawaqa Island within the Yasawa Island Group in western Fiji. Besides a single tourism operator (Barefoot Manta Island Resort), Drawaqa Island is uninhabited and belongs to the traditional landowners of Mua-ira on the nearby island of Naviti (<xref ref-type="bibr" rid="B34">Murphy et&#xa0;al., 2018</xref>). The northern point of Drawaqa Island includes five beaches, which were surveyed: Goat, Lagoon, Manta, Sunrise, and Sunset Beach (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Location of the five beaches surveyed in Drawaqa Island, in Western Fiji.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1401258-g001.tif"/>
</fig>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Field work</title>
<p>Since January 2024, research on Drawaqa Island has been exploring environmental factors influencing spatiotemporal variation in ray species abundance and distribution, with the data presented here being part of a larger dataset. Fieldwork involved direct observations and ocean temperature measurements using three HOBO water temperature Pro V2 (U22) data loggers deployed on-site. Over 40 days across January to March, 105 surveys were conducted to observe rays. Each survey lasted 45 min for standardization, and were conducted as roving explorations, including 53 snorkel surveys, 32 beach walks, and 20 SCUBA dives: Sunrise Beach (68 surveys: 39 snorkels, 20 dives, 9 walks), Lagoon Beach (11 surveys: 10 walks, 1 snorkel), Goat Beach (10 snorkels), Manta Beach (9 surveys: 2 snorkels, 7 walks), and Sunset Beach (7 surveys: 6 walks, 1 snorkel). Snorkel surveys were conducted at all sites along 100 m line transects parallel to the shoreline. Maximum depth was 12 m, with visibility extending to the seafloor. SCUBA dives, performed only at Sunrise Beach due to sufficient depth, followed predetermined 100 m line transects perpendicular to the shore, from the shallows to a maximum depth of 21 m, with visibility ranging from 8 to 20m. For each line transect, the surrounding visibility was scanned for the presence of <italic>T. lessoni</italic> within the maximum visible range on either side of the line. The line transects followed the reef structure along the same route for each snorkel and dive survey. Beach walks were carried out at four of the five sites.</p>
<p>Whenever possible, sighted <italic>T. lessoni</italic> were photographed using either a GoPro12 or Olympus TG7. Images of <italic>T. lessoni</italic> were recorded to identify distinct spotting patterns and body markings (<xref ref-type="bibr" rid="B31">McIvor et&#xa0;al., 2023</xref>). Photographs were also inspected for signs of advanced gestation (i.e., convex dorsum), recent parturition (i.e., concave dorsum) (<xref ref-type="bibr" rid="B18">Henningsen, 2000</xref>; <xref ref-type="bibr" rid="B50">Spieler et&#xa0;al., 2013</xref>), and evidence of mating behaviour (i.e., fresh bite wounds) (<xref ref-type="bibr" rid="B24">Kajiura et&#xa0;al., 2000</xref>). The sex of rays was determined by the presence or absence of claspers (<xref ref-type="bibr" rid="B4">Awruch et&#xa0;al., 2008</xref>), and the DW was estimated <italic>in situ</italic> and from images (<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure&#xa0;1</bold>
</xref>). <italic>T. lessoni</italic> with a DW greater than 20 cm were classified as mature, and rays less than 20 cm DW were considered immature (<xref ref-type="bibr" rid="B29">Last et&#xa0;al., 2016b</xref>). Size at birth for <italic>T. lymma</italic> is 13 cm to 14 cm DW (<xref ref-type="bibr" rid="B29">Last et&#xa0;al., 2016b</xref>). As <italic>T. lessoni</italic> is smaller and due to the uncertainty associated with <italic>in situ</italic> estimates, only the smallest individuals, with an estimated DW of 10 cm (roughly palm-sized), were considered as neonates.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Data handling</title>
<p>The data were recorded, with individuals whose sex could not be determined denoted as &#x201c;nA&#x201d;. Plots were generated using the ggplot2 package (<xref ref-type="bibr" rid="B53">Wickham, 2011</xref>; <xref ref-type="bibr" rid="B54">Wickham and Bryan, 2023</xref>) in R (<xref ref-type="bibr" rid="B40">R Development Core Team, 2005</xref>). Catch per unit effort (CPUE) was calculated by dividing total <italic>T. lessoni</italic> sightings at a site by the number of surveys conducted there.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<p>
<italic>T. lessoni</italic> were observed in 71 surveys (67.6% of the total number surveys). Number of sightings by location varied: Sunrise Beach (<italic>n</italic>=47), Goat Beach (<italic>n</italic>=9), Lagoon Beach (<italic>n</italic>=6), Manta Beach (<italic>n</italic>=6), and Sunset Beach (<italic>n</italic>=3). At Sunrise Beach, mature females were sighted 38 times, six times at Goat Beach, and once each at Manta Beach and Sunset Beach. Mature males were observed three times each at Sunrise Beach and Goat Beach, and once at Sunset Beach. Neonates were sighted from early March onwards: six times at Lagoon Beach, two times at Manta Beach, and one time each at Sunrise and Sunset Beach (<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure&#xa0;2</bold>
</xref>). CPUE was highest at Goat Beach (0.9 <italic>T. lessoni</italic> observed per survey), followed by Sunrise Beach (0.7 <italic>T. lessoni</italic> observed per survey), Manta Beach (0.7 <italic>T. lessoni</italic> observed per survey), Lagoon Beach (0.6 <italic>T. lessoni</italic> observed per survey), and Sunset Beach (0.4 <italic>T. lessoni</italic> observed per survey).</p>
<p>Snorkelling recorded the highest number of sightings (<italic>n</italic>=48), followed by beach walks (<italic>n</italic>=15), and dives (<italic>n</italic>=8) (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). Mature females and males were mostly observed during snorkelling surveys, while immature specimens were predominantly sighted during beach walks (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Sightings of <italic>T. lessoni</italic> across different survey methods, including the number of sightings per sex, and likely maturity level based on <italic>in situ</italic> DW estimates.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Survey type</th>
<th valign="top" align="center">#Sightings</th>
<th valign="top" align="center">#Mature F &gt; 20 cm</th>
<th valign="top" align="right">#Immature F<break/>15-20 cm</th>
<th valign="top" align="right">#Neonate F<break/>10 cm</th>
<th valign="top" align="right">#Mature M &gt;20 cm</th>
<th valign="top" align="right">#Neonate M<break/>10 cm</th>
<th valign="top" align="right">#Mature nA<break/>&gt;20 cm</th>
<th valign="top" align="left">#Immature nA<break/>15-20 cm</th>
<th valign="top" align="left">#Neonate nA<break/>10 cm</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Snorkel</td>
<td valign="top" align="center">48</td>
<td valign="top" align="center">36</td>
<td valign="top" align="right">0</td>
<td valign="top" align="right">1</td>
<td valign="top" align="right">6</td>
<td valign="top" align="right">1</td>
<td valign="top" align="right">3</td>
<td valign="top" align="left">0</td>
<td valign="top" align="left">1</td>
</tr>
<tr>
<td valign="top" align="left">Beach walk</td>
<td valign="top" align="center">15</td>
<td valign="top" align="center">4</td>
<td valign="top" align="right">1</td>
<td valign="top" align="right">2</td>
<td valign="top" align="right">1</td>
<td valign="top" align="right">0</td>
<td valign="top" align="right">1</td>
<td valign="top" align="left">1</td>
<td valign="top" align="left">5</td>
</tr>
<tr>
<td valign="top" align="left">Dive</td>
<td valign="top" align="center">8</td>
<td valign="top" align="center">6</td>
<td valign="top" align="right">0</td>
<td valign="top" align="right">0</td>
<td valign="top" align="right">0</td>
<td valign="top" align="right">0</td>
<td valign="top" align="right">2</td>
<td valign="top" align="left">0</td>
<td valign="top" align="left">0</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>At Sunrise Beach, four females in advanced gestation with conspicuously convex dorsa (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>) were recorded, occurring in shallow waters down to eight m depth. These four females were collectively recorded 24 times at the same site from the beginning of January until the end of March 2024. Individual 1 was recorded 13 times, Individual 2 six times, Individual 3 four times, and Individual 4 once, accounting for 63.2% of all female <italic>T. lessoni</italic> sightings at Sunrise Beach (<italic>n</italic>=38, <xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure&#xa0;2</bold>
</xref>). The DW of Individual 1 was estimated to be 24 cm to 25 cm, DW of the remaining three individuals in advance gestation was estimated between 21 to 25 cm.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Photographs of individual <italic>T. lessoni</italic> in advanced gestation observed at Sunrise Beach, Drawaqa Island, with convex dorsa, indicative for advanced gestation. Encircled in red are selected differentiating features: Individual 1 displays single antorbital blue spots, and a row of three prominent vertical blue spots followed by a row of horizontal blue spots posterolateral; Individual 2 has two antorbital blue spots; Individual 3 features a plain antorbital area, including a tail partly cut (shown in <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3F</bold>
</xref>), and Individual 4 has scapular and posterolateral rows of three blue spots.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1401258-g002.tif"/>
</fig>
<p>The first indication of parturition was noted on February 29, 2024, when Individual 1 was photographed with a concave dorsum (<xref ref-type="fig" rid="f3">
<bold>Figures&#xa0;3A, B</bold>
</xref>). Over the following two weeks, Individual 3 (March 4) and Individual 2 (March 12) followed suit (<xref ref-type="fig" rid="f3">
<bold>Figures&#xa0;3C-F</bold>
</xref>). Additionally, Individual 1 exhibited a visible dermal abrasion on the left pectoral fin (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>; <xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure&#xa0;3</bold>
</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Visual comparison of Individuals 1, 2, and 3 during advanced gestation <bold>(A, C, E)</bold> and post-partum <bold>(B, D, F)</bold> stages. Image <bold>(A)</bold> of individual 1 was taken on February 3, 2024, during advanced gestation, while <bold>(B)</bold> was captured on February 29, 2024, post-partum. Image <bold>(C)</bold> shows the individual 2 on January 13, 2024, during advanced gestation, and <bold>(D)</bold> on March 12, 2024, post-partum. Image <bold>(E)</bold> shows the individual 3 on February 11, 2024, during pregnancy, and <bold>(F)</bold> on March 4, 2024, post-partum.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1401258-g003.tif"/>
</fig>
<p>During the study period, the water temperature ranged from 28.22&#xb0;C to 33.08&#xb0;C. The lowest temperature was recorded at Sunrise Beach in January with an average of 29.94&#xb0;C, while the highest temperature was measured at Sunset Beach with an average of 30.52&#xb0;C in March.</p>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>This study represents the first documentation of advanced gestation in a stingray species in Fiji. We recorded four pregnant <italic>T. lessoni</italic> (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), of which three gave birth between the end of February and mid-March, based upon photographic evidence (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). The presence of neonates from early March onwards coincided with the estimated parturition period inferred from the rays&#x2019; condition. We were unable to determine the periodicity of the reproductive cycle and the gestation period, but <italic>T. lessoni</italic> may reproduce asynchronously and aseasonally. In a captive stingray species (formerly known as <italic>Dasyatis kuhlii</italic> and <italic>Neotrygon kuhlii</italic>) mating was observed immediately after parturition, suggesting the absence of a specific breeding season (<xref ref-type="bibr" rid="B21">Janse and Schrama, 2010</xref>). Based on bycatch data from eastern Indonesia, neither distinct seasonal reproductive cycles nor synchronicity in three stingray species could be determined, indicating a continuous reproductive cycle (<xref ref-type="bibr" rid="B52">White and Dharmadi, 2007</xref>). The dermal wound around the pectoral fin in Individual 1 (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>; <xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure&#xa0;3</bold>
</xref>) suggests a bite or abrasion from a pectoral grip or pre-copulatory biting, indicating that copulation may occur soon or immediately after parturition, as observed in captive <italic>T. lymma</italic> (<xref ref-type="bibr" rid="B49">Smith et&#xa0;al., 2017</xref>). Therefore, a continuous cycle with the potential for multiple pregnancies annually seems likely. However, long-term data, ideally combined with ultrasound diagnostics (<xref ref-type="bibr" rid="B33">Murakumo et&#xa0;al., 2020</xref>), are required to document gestation and to determine whether <italic>T. lessoni</italic> follows a continuous reproductive cycle or exhibits seasonal peaks around Drawaqa Island. To ensure practicality and minimal invasiveness, we determined maturity levels based solely on <italic>in situ</italic> size estimates. Assaying sex steroid hormones instead, enables accurate determination of maturity levels and depiction of cumulative proportions across different developmental stages (<xref ref-type="bibr" rid="B32">Mull et&#xa0;al., 2010</xref>). The surveyed sites on Drawaqa Island exhibit similar average temperatures ranging from 29.94&#xb0;C to 30.52&#xb0;C. Neonates and presumably immature <italic>T. lessoni</italic> were mostly observed during beach walks (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>), and at Manta Beach and Lagoon Beach (<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure&#xa0;2</bold>
</xref>). Manta Beach and Lagoon Beach have a similar habitat composition with areas of sandy patches, rubbles, and seagrass assemblages, which could provide shelter for neonate <italic>T. lessoni</italic> (<xref ref-type="bibr" rid="B7">Dabruzzi et&#xa0;al., 2013</xref>). Sunrise Beach, predominantly frequented by mature females, has narrower expanses of sandy patches. Except for Individual 4, which was sighted only once, the other three females remained at Sunrise Beach, indicating repeated use of the area. Interestingly, mature males were only encountered seven times (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). One possibility to explain the skewed sex ratio towards females could be attributed to the increased energetic requirements of females during gestation (<xref ref-type="bibr" rid="B22">Jirik and Lowe, 2012</xref>), and thus the selection of areas that offer favourable conditions, including prey availability (<xref ref-type="bibr" rid="B8">Delpiani et&#xa0;al., 2013</xref>) and protection from predators (<xref ref-type="bibr" rid="B30">Martins et&#xa0;al., 2018</xref>).</p>
<p>Moving forward, a small-scale passive acoustic telemetry study, combined with ongoing environmental data collection, could help better understand presence, activity patterns, sexual segregation (<xref ref-type="bibr" rid="B48">Simpson et&#xa0;al., 2021</xref>), and seasonal or long-term site fidelity (<xref ref-type="bibr" rid="B43">Schlaff et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B10">Elston et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B25">Kraft et&#xa0;al., 2023</xref>). The waters surrounding Drawaqa Island are home to at least nine batoid species: <italic>T. lessoni</italic> (Data Deficient), <italic>Mobula birostris</italic> (Endangered), <italic>Aetobatus ocellatus</italic>, <italic>M. alfredi</italic>, <italic>Pateobatis fai</italic>, <italic>Taeniurops meyeni</italic> and <italic>Urogymnus asperrimus</italic> (all Vulnerable), <italic>Rhynchobatus australiae</italic> (Critically Endangered), and <italic>Neotrygon</italic> sp (<xref ref-type="bibr" rid="B16">Gordon and Vierus, 2022</xref>; <xref ref-type="bibr" rid="B14">Glaus et&#xa0;al., 2024a</xref>). Our preliminary data indicate that Drawaqa&#x2019;s nearshore waters likely serve as pupping grounds for <italic>T. lessoni</italic>. Considering this, along with the diversity of species, the easy accessibility, and the continued monitoring setup, this location is ideal for further studies to deepen our understanding of ray ecology and biology.</p>
</sec>
<sec id="s5" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation upon reasonable request.</p>
</sec>
<sec id="s6" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>Ethical approval was not required for the study involving animals in accordance with the local legislation and institutional requirements because the study was non-invasive and did not include any handling of living animals. The findings solely derive from visual observations.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>KG: Conceptualization, Funding acquisition, Investigation, Methodology, Project administration, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. TV: Conceptualization, Data curation, Validation, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. RM: Data curation, Funding acquisition, Investigation, Methodology, Project administration, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
</body>
<back>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. Funding was provided by the DrawaqaMarine Conservation Trust and by the Deutsche Stiftung Meeresschutz. The Funders are Drawaqa Marine Conservation Trust and Deutsche Stiftung Meeresschutz.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>Special thanks go to Victor Bonito of Reef Explorers Fiji for sharing the water temperature data, to James Szymankiewicz for the picture used in <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>, and to Peter Last for his expertise on the photographs of the individual female rays. We acknowledge the Directors of Barefoot Manta Island Resort, and Mirko Rossi, Rafal Jaklik, and Martin Eberle for their continuous support during the surveys. We thank the anonymous reviewers for their constructive comments and valuable suggestions, which greatly improved the quality of this paper.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>RM is a board member of the Drawaqa Marine Conservation Trust.</p>
<p>The remaining 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.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s11" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fmars.2024.1401258/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2024.1401258/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Image_1.jpeg" id="SF1" mimetype="image/jpeg">
<label>Supplementary Figure&#xa0;1</label>
<caption>
<p>
<italic>In situ</italic> DW estimate of Individual1.</p>
</caption>
</supplementary-material>
<supplementary-material xlink:href="Image_2.tif" id="SF2" mimetype="image/tiff">
<label>Supplementary Figure&#xa0;2</label>
<caption>
<p>Number of T. lessoni sightings and estimated maturity levels across the surveyed sites.</p>
</caption>
</supplementary-material>
<supplementary-material xlink:href="Image_3.jpeg" id="SF3" mimetype="image/jpeg">
<label>Supplementary Figure&#xa0;3</label>
<caption>
<p>Individual1 feeding, showing visible dermal abrasion on the left side.</p>
</caption>
</supplementary-material>
</sec>
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