<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article
  PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "http://dtd.nlm.nih.gov/publishing/3.0/journalpublishing3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML"
	xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article"
	dtd-version="3.0" xml:lang="en">
	<front>
		<journal-meta>
			<journal-id journal-id-type="nlm-ta">PLoS ONE</journal-id>
			<journal-id journal-id-type="publisher-id">plos</journal-id>
			<journal-id journal-id-type="pmc">plosone</journal-id>
			<journal-title-group>
				<journal-title>PLoS ONE</journal-title>
			</journal-title-group>
			<issn pub-type="epub">1932-6203</issn>
			<publisher>
				<publisher-name>Public Library of Science</publisher-name>
				<publisher-loc>San Francisco, USA</publisher-loc>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="publisher-id">PONE-D-14-20725</article-id>
			<article-id pub-id-type="doi">10.1371/journal.pone.0115884</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>Research Article</subject>
				</subj-group>
				<subj-group subj-group-type="Discipline-v2">
					<subject>Biology and life sciences</subject>
					<subj-group>
						<subject>Evolutionary biology</subject>
						<subj-group>
							<subject>Organismal evolution</subject>
						</subj-group>
					</subj-group>
					<subj-group>
						<subject>Taxonomy</subject>
						<subj-group>
							<subject>Animal taxonomy</subject>
						</subj-group>
					</subj-group>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>A Novel Reproductive Mode in Frogs: A New Species of
					Fanged Frog with Internal Fertilization and Birth of Tadpoles</article-title>
				<alt-title alt-title-type="running-head">A New Tadpole-Bearing Frog</alt-title>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author" xlink:type="simple">
					<name name-style="western">
						<surname>Iskandar</surname>
						<given-names>Djoko T.</given-names>
					</name>
					<xref ref-type="aff" rid="aff1">
						<sup>1</sup>
					</xref>
				</contrib>
				<contrib contrib-type="author" xlink:type="simple">
					<name name-style="western">
						<surname>Evans</surname>
						<given-names>Ben J.</given-names>
					</name>
					<xref ref-type="aff" rid="aff2">
						<sup>2</sup>
					</xref>
				</contrib>
				<contrib contrib-type="author" xlink:type="simple">
					<name name-style="western">
						<surname>McGuire</surname>
						<given-names>Jimmy A.</given-names>
					</name>
					<xref ref-type="aff" rid="aff3">
						<sup>3</sup>
					</xref>
					<xref ref-type="corresp" rid="cor1">
						<sup>*</sup>
					</xref>
				</contrib>
			</contrib-group>
			<aff id="aff1">
				<label>1</label>
				<addr-line>School of Life Sciences and Technology, Institut
					Teknologi Bandung, Bandung, Indonesia</addr-line>
			</aff>
			<aff id="aff2">
				<label>2</label>
				<addr-line>Center for Environmental Genomics, Department of Biology,
					McMaster University, Hamilton, Ontario, Canada</addr-line>
			</aff>
			<aff id="aff3">
				<label>3</label>
				<addr-line>Museum of Vertebrate Zoology and Department of
					Integrative Biology, University of California, Berkeley,
					California, United States of America</addr-line>
			</aff>
			<contrib-group>
				<contrib contrib-type="editor" xlink:type="simple">
					<name name-style="western">
						<surname>Stöck</surname>
						<given-names>Matthias</given-names>
					</name>
					<role>Editor</role>
					<xref ref-type="aff" rid="edit1" />
				</contrib>
			</contrib-group>
			<aff id="edit1">
				<addr-line>Leibniz-Institute of Freshwater Ecology and Inland
					Fisheries, Germany</addr-line>
			</aff>
			<author-notes>
				<corresp id="cor1">
					* E-mail:
					<email xlink:type="simple">mcguirej@berkeley.edu</email>
				</corresp>
				<fn fn-type="conflict">
					<p>The authors have declared that no competing interests exist.</p>
				</fn>
				<fn fn-type="con">
					<p>Conceived and designed the experiments: DTI BJE JAM. Performed
						the experiments: DTI BJE JAM. Analyzed the data: DTI BJE JAM.
						Contributed reagents/materials/analysis tools: DTI BJE JAM. Wrote
						the paper: DTI BJE JAM.</p>
				</fn>
			</author-notes>
			<pub-date pub-type="collection">
				<year>2014</year>
			</pub-date>
			<pub-date pub-type="epub">
				<day>31</day>
				<month>12</month>
				<year>2014</year>
			</pub-date>
			<volume>9</volume>
			<issue>12</issue>
			<elocation-id>e115884</elocation-id>
			<history>
				<date date-type="received">
					<day>17</day>
					<month>5</month>
					<year>2014</year>
				</date>
				<date date-type="accepted">
					<day>23</day>
					<month>11</month>
					<year>2014</year>
				</date>
			</history>
			<permissions>
				<copyright-year>2014</copyright-year>
				<copyright-holder>Iskandar et al</copyright-holder>
				<license xlink:type="simple">
					<license-p>
						This is an open-access article distributed under the terms of the
						<ext-link ext-link-type="uri"
							xlink:href="http://creativecommons.org/licenses/by/4.0/"
							xlink:type="simple">Creative Commons Attribution License</ext-link>
						, which permits unrestricted use, distribution, and reproduction
						in any medium, provided the original author and source are
						credited.
					</license-p>
				</license>
			</permissions>
			<abstract>
				<p>
					We describe a new species of fanged frog (
					<italic>Limnonectes larvaepartus</italic>
					) that is unique among anurans in having both internal
					fertilization and birth of tadpoles. The new species is endemic to
					Sulawesi Island, Indonesia. This is the fourth valid species of
					<italic>Limnonectes</italic>
					described from Sulawesi despite that the radiation includes at
					least 15 species and possibly many more. Fewer than a dozen of the
					6455 species of frogs in the world are known to have internal
					fertilization, and of these, all but the new species either deposit
					fertilized eggs or give birth to froglets.
				</p>
			</abstract>
			<funding-group>
				<funding-statement>This research was funded by the United States
					National Science Foundation Division of Environmental Biology (DEB
					0328700, DEB 0640967) to JAM, the Natural Science and Engineering
					Research Council of Canada (RGPIN/283102-2012) to BJE, and the
					Indonesian Directorate General of Higher Education, Ministry of
					Education and Culture (Basic Science Grant No.
					60/P4M/DPPM/L-3311/MS/1993) to DTI. The funders had no role in
					study design, data collection and analysis, decision to publish, or
					preparation of the manuscript.</funding-statement>
			</funding-group>
			<counts>
				<page-count count="14" />
			</counts>
			<custom-meta-group>
				<custom-meta id="data-availability" xlink:type="simple">
					<meta-name>Data Availability</meta-name>
					<meta-value>The authors confirm that all data underlying the
						findings are fully available without restriction. All relevant
						data are within the paper and its Supporting Information files.</meta-value>
				</custom-meta>
			</custom-meta-group>
		</article-meta>
	</front>
	<body>
		<sec id="s1">
			<title>Introduction</title>
			<p>
				The Indonesian island of Sulawesi in central Indonesia is home to a
				largely unstudied radiation of fanged frogs in the genus
				<italic>Limnonectes</italic>
				Fitzinger, 1843 (Anura: Dicroglossidae Anderson, 1871). To date,
				only three species are recognized from Sulawesi, but at least 15
				species occur on the island
				<xref ref-type="bibr" rid="pone.0115884-Emerson1">[1]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Evans1">[2]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Setiadi1">[3]</xref>
				. It was recently proposed that Sulawesi fanged frogs represent an
				adaptive radiation based on patterns of morphological
				diversification among co-distributed species, and correspondence of
				morphological features (body size, webbing) with habitat utilization
				<xref ref-type="bibr" rid="pone.0115884-Setiadi1">[3]</xref>
				. Adult body size ranges from ∼2 g to more than 900 g in members of
				the Sulawesi assemblage, with large-bodied species found in close
				association with fast moving rivers, and small-bodied species
				occupying more terrestrial niches in forest leaf-litter and on the
				banks of slow moving streams. Iskandar and Tjan
				<xref ref-type="bibr" rid="pone.0115884-Iskandar1">[4]</xref>
				first documented that there are multiple undescribed species of
				<italic>Limnonectes</italic>
				on Sulawesi, and indicated that one of the constituent species
				utilizes an unusual ovoviviparous reproductive strategy. Here we
				describe that species, and provide additional details on its
				reproductive mode.
			</p>
			<p>
				Anurans exhibit tremendous diversity in mode of reproduction. Not
				only have there been many independent origins of direct development,
				the condition in which a free-living tadpole stage is bypassed and
				froglets emerge from the egg capsules
				<xref ref-type="bibr" rid="pone.0115884-Wake1">[5]</xref>
				, but a bewildering array of mechanisms has evolved wherein parents
				care for their developing offspring. Examples of parental care
				include guarding of terrestrial eggs, transport of tadpoles on the
				back following hatching, carrying developing eggs in pouches or
				depressions on the back and flanks, males carrying developing
				tadpoles in the vocal sac, and even the now-extinct species of
				gastric brooding frogs in which the female carried her developing
				tadpoles in her stomach (see
				<xref ref-type="bibr" rid="pone.0115884-Haddad1">[6]</xref>
				for a summary). Despite this extreme reproductive diversity,
				internal fertilization has evolved only a few times among the 6455
				known species of anurans
				<xref ref-type="bibr" rid="pone.0115884-AmphibiaWeb1">[7]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-McDiarmid1">[8]</xref>
				. Only the African bufonid genera
				<italic>Nectophrynoides</italic>
				and
				<italic>Nimbaphrynoides</italic>
				and the now extinct Puerto Rican species,
				<italic>Eleutherodactylus jasperi</italic>
				, are documented to combine internal fertilization with egg
				retention and subsequent birth of froglets
				<xref ref-type="bibr" rid="pone.0115884-Wells1">[9]</xref>
				. To this group, we can now add a species of Sulawesi fanged frog in
				the genus
				<italic>Limnonectes</italic>
				with internal fertilization and live birth of tadpoles, a
				reproductive mode that is unique among anurans.
			</p>
		</sec>
		<sec id="s2" sec-type="materials|methods">
			<title>Materials and Methods</title>
			<p>
				We analyzed more than 100 specimens representing the new species
				from 23 localities on the island of Sulawesi. This study was
				conducted under an approved Institutional Animal Care and Use
				Committee protocol issued to JAM by the University of California at
				Berkeley (Protocol #R279). Fieldwork conducted in Indonesia was
				undertaken under research permits issued by LIPI and RISTEK, with
				specimen exportation authorized under permits or loans issued by
				LIPI, the Museum Zoologicum Bogoriense, and Pusat Studi Biologi.
				Specimens were captured by hand in the field, sacrificed via
				emersion in an aqueous solution of MS-222 (tricaine methane
				sulfonate) buffered to neutral pH, and prepared as formalin-fixed
				specimens deposited primarily in the Museum Zoologicum Bogoriense
				(the national museum of Indonesia) or Museum of Vertebrate Zoology
				at UC Berkeley. As part of a comprehensive study of Sulawesi
				<italic>Limnonectes</italic>
				diversification, we obtained morphometric data for the full Sulawesi
				assemblage. Sex was determined by inspection of gonads. The
				following measurements were made to the nearest 0.1 mm using digital
				calipers: snout-vent length (SVL) – from cloaca to tip of snout;
				head width (HW) – widest distance between posterior end of lower
				jaw; head length (HL) – from posterior end of lower jaw to tip of
				snout; femur length (FE) – from cloaca to distal end of femur; tibia
				length (TI) – from knee to distal end of tibia; foot length (FL) –
				from distal end of tibia-fibula to tip of 4th toe; inner metatarsal
				tubercle (IM); humerus length (UA) – from proximal end of upper arm
				to elbow; lower arm length (LA) – from elbow joint to base of middle
				palmar tubercle; hand length (HA) – from lower border of middle
				tubercle to distal tip of third finger; snout length (SL) – from
				bony border of eye socket to tip of snout; snout width at eye level
				(SWE); snout width at nostril level (SWN); eye diameter (EY) – outer
				diameter, measured from bones bordering eye; interorbital distance
				(IO) – distance between orbits ( = width of frontoparietal bones);
				eye-tympanum distance (ET); eye-narial distance (EN) – eye socket
				border to posterior border of nostril; Internarial distance (IN) –
				shortest distance between nostrils; nostril to tip of snout (NT);
				tympanum diameter (TY); odontoid process length (OP) –length of
				fang-like odontoid process of lower jaw; tibia diameter measured at
				its widest point (TD). Digital webbing formulae from
				<xref ref-type="bibr" rid="pone.0115884-Guayasamin1">[10]</xref>
				.
			</p>
			<sec id="s2a">
				<title>Nomenclatural acts</title>
				<p>
					The electronic edition of this article conforms to the requirements
					of the amended International Code of Zoological Nomenclature, and
					hence the new names contained herein are available under that Code
					from the electronic edition of this article. This published work
					and the nomenclatural acts it contains have been registered in
					ZooBank, the online registration system for the ICZN. The ZooBank
					LSID (Life Science Identifier) can be resolved and the associated
					information viewed through any standard web browser by appending
					the LSID to the prefix “
					<ext-link ext-link-type="uri" xlink:href="http://zoobank.org/"
						xlink:type="simple">http://zoobank.org/</ext-link>
					”. The LSID for this publication is: urn:lsid:zoobank.org:pub:
					A758E03A–C054–4193–9768–D0C4AD588AEA. The electronic edition of
					this work was published in a journal with an ISSN, and has been
					archived and is available from the following digital repositories:
					PubMed Central, LOCKSS.
				</p>
			</sec>
		</sec>
		<sec id="s3">
			<title>Results</title>
			<sec id="s3a">
				<title>Taxonomy</title>
				<p>
					<italic>Limnonectes </italic>
					<bold>
						<italic>larvaepartus</italic>
					</bold>
					new species (
					<xref ref-type="fig" rid="pone-0115884-g001">Figs. 1</xref>
					,
					<xref ref-type="fig" rid="pone-0115884-g002">2</xref>
					)
				</p>
				<fig id="pone-0115884-g001" position="float">
					<object-id pub-id-type="doi">10.1371/journal.pone.0115884.g001</object-id>
					<label>Figure 1</label>
					<caption>
						<title>
							Images of the holotype of
							<italic>Limnonectes larvaepartus</italic>
							(MZB.Amph.23755) in (
							<italic>a</italic>
							) lateral, (
							<italic>b</italic>
							) dorsal, and (
							<italic>c</italic>
							) ventral view.
						</title>
						<p>
							Ventral views of the right foot (
							<italic>d</italic>
							) and right hand (
							<italic>e</italic>
							) are also presented.
						</p>
					</caption>
					<graphic mimetype="image"
						xlink:href="info:doi/10.1371/journal.pone.0115884.g001" position="float"
						xlink:type="simple" />
				</fig>
				<fig id="pone-0115884-g002" position="float">
					<object-id pub-id-type="doi">10.1371/journal.pone.0115884.g002</object-id>
					<label>Figure 2</label>
					<caption>
						<title>
							Images of
							<italic>Limnonectes larvaepartus</italic>
							.
						</title>
						<p>
							(
							<italic>a</italic>
							) MVZ 268323 (
							<bold>male, left</bold>
							) and MVZ 268307 (
							<bold>female, right</bold>
							) collected from Desa Uaemate along the Tasio-Tibo Road,
							Kabupatan Mamuju, Provinsi Sulawesi Barat, Sulawesi Island
							(02.61287S, 119.14238 E, 89 m elev.); (
							<italic>b</italic>
							)
							<italic>Limnonectes larvaepartus</italic>
							female (MVZ 268426) with tadpoles removed from the oviduct. Note
							the large yolk reserves available to the tadpoles; (
							<italic>c</italic>
							) An
							<italic>in situ</italic>
							adult male
							<italic>L. larvaepartus</italic>
							(JAM 14234) observed calling while perched on the edge of a small
							pool 2 m away from a 2 m wide stream; several
							<italic>L. larvaepartus</italic>
							tadpoles were present in the pool including the two visible
							within the yellow circle; (
							<italic>d</italic>
							) dorsal and ventral views of ∼stage 25
							<italic>L. larvaepartus</italic>
							tadpoles (JAM 14271) released by a pregnant female (JAM 14237) at
							the moment of capture.
						</p>
					</caption>
					<graphic mimetype="image"
						xlink:href="info:doi/10.1371/journal.pone.0115884.g002" position="float"
						xlink:type="simple" />
				</fig>
				<p>Urn:lsid:zoobank.org:act: 60AA7136-89A0-4DBB-9FBC-BD0FAF8A214C</p>
				<p>
					This species has been referred to in the literature under the names
					<italic>Limnonectes larviparus</italic>
					<xref ref-type="bibr" rid="pone.0115884-Inger1">[11]</xref>
					and
					<italic>Limnonectes</italic>
					“ovovivipar”
					<xref ref-type="bibr" rid="pone.0115884-Iskandar1">[4]</xref>
					, both of which created nomina nuda. This species also corresponds
					to
					<italic>Limnonectes</italic>
					sp. V in
					<xref ref-type="bibr" rid="pone.0115884-Evans1">[2]</xref>
					,
					<xref ref-type="bibr" rid="pone.0115884-Setiadi1">[3]</xref>
					.
				</p>
			</sec>
			<sec id="s3b">
				<title>Etymology</title>
				<p>
					The species name
					<italic>larvaepartus</italic>
					(from ‘larvae’, plural of larva, the early form of an animal, and
					‘partus’, to give birth to) reflects the unique reproductive mode
					of this tadpole-bearing species.
				</p>
			</sec>
			<sec id="s3c">
				<title>Holotype</title>
				<p>
					MZB.Amph.23755 (Field Number: BSI 0605, see
					<xref ref-type="fig" rid="pone-0115884-g001">Fig. 1</xref>
					), an adult male, collected from Dunu Village, (0.92353
					<sup>o</sup>
					N; 122.64386
					<sup>o</sup>E)
					at 189 m elevation, Kecamatan Anggrek, Kabupaten Gorontalo,
					Provinsi Gorontalo, Sulawesi, Indonesia by J.A. McGuire &amp; team,
					18 October 2004.
				</p>
			</sec>
			<sec id="s3d">
				<title>Paratypes</title>
				<p>
					Paratypes (n = 30) are from Sulawesi Utara, Gorontalo, Sulawesi
					Tengah, and Sulawesi Barat Provinces: MZB 2834, a gravid female
					with 33 translucent tadpoles (Gosner stage 23) from the left
					oviduct and 2 from outside the body, from Toraut (00′33.72
					<sup>o</sup>
					N, 123′54.23
					<sup>o</sup>E),
					Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow,
					Sulawesi Utara at 370 m elevation, by D. T. Iskandar, 15 August
					1991; FMNH 252453, a female, from Toraut, Bogani Nani Wartabone
					National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara by D. T.
					Iskandar, August 1991. MVZ 255545, 256009-11, 256013 from Desa
					Lombongo (−1.43346, 120.30800), Kecamatan Suwawa, Kabupaten Bone
					Bolango, Bogani Nani Warta Bone National Park, Provinsi Gorontalo
					at 75 m elevation by J. A McGuire and team, 20 October 2004. ZRC
					1.3258 (left femur removed) from Toraut, Bogani Nani Wartabone
					National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara at 370 m
					elevation, by D. T. Iskandar, 15 August 1991. MZB 2835–2841 from
					Toraut, Bogani Nani Wartabone National Park, Kabupaten Bolaang
					Mongondow, Sulawesi Utara at 370 m elevation, by D. T. Iskandar, 15
					August 1991 &amp; 12 July 1992. MVZ 255548–49 from Desa Pontak
					(−2.62910, 118.99300), Kecamatan Motoling, Kabupaten Minahasa
					Selatan, Provinsi Sulawesi Utara at 285 m elevation by J. A.
					McGuire and team, 13 October 2004. MZB 3117, near Potolok river,
					Lolak, Bogani Nani Wartabone National Park at 350 m elevation,
					Kabupaten Bolaang Mongondow, Sulawesi Utara; MZB 3118, male from
					seashore forest near Bungbungan River, Lolak, Bogani Nani Wartabone
					National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara. MZB
					3120 from Tangkorak River, Desa Pindol, Kecamatan Lolak, Bolaang
					Mongondow, Sulawesi Utara, by Mumpuni, 26 June 1995; MZB 3121 from
					Tangaga Forest, Dudepo, Bolaang Mongondow, Sulawesi Utara, by I.
					Maryanto, 20 October 1995; MZB 3122 from Potolok River, Bogani Nani
					National Park, Lolak, Sulawesi Utara by Mumpuni, 19 June 1995; MZB
					3124 from Bungbungan River, Bogani Nani National Park, Lolak,
					Sulawesi Utara by Mumpuni, 19 June 1995; MZB Amph.8108 from Toraut,
					near Bogani Nani Wartabone National Park, Sulawesi Utara. LSUMZ
					84209, 84214, 84221, 84224 from Desa Torout, Bogani Nani Wartabone
					National Park, Kabupaten Bolaang Mongondow, Provinsi Sulawesi Utara
					at 267 m elevation by J. A. McGuire on 6 and 11 September 2001.
				</p>
			</sec>
			<sec id="s3e">
				<title>Other referred specimens</title>
				<p>
					LSUMZ 84218, 84219 from Desa Torout, Bogani Nani Wartabone National
					Park, Kabupaten Bolaang Mongondow, Provinsi Sulawesi Utara by J.A.
					McGuire. FMNH 130991, 106930 from Buang-buang Island, Sulawesi
					Utara. MZB 3119, juvenile from Seashore forest near Bungbungan
					River, Lolak, Bogani Nani Wartabone National Park, Kabupaten
					Bolaang Mongondow, Provinsi Sulawesi Utara. AMNH 167199 from
					Tangkoko National Park (1.570083
					<sup>o</sup>
					N, 125.156933
					<sup>o</sup>
					E), Kabupaten Minahasa, Provinsi Sulawesi Utara. MVZ 255546 from
					Desa Salumpaku (−1.60757, 119.29900), Kecamatan Banawa, Kabupaten
					Donggala, Provinsi Sulawesi Tengah. MVZ 255547 from Desa Kelapa Dua
					(−1.60757, 119.29900), Kecamatan Anreapi, Kabupaten Polewali
					Mandar, Provinsi Sulawesi Barat. MVZ 268426, 268428–30, 268432 from
					Polewali-Masawa Road (River 1) (−2.65490, 118.93300), Kecamatan
					Polewali, Kabupaten Polewali Mandar, Provinsi Sulawesi Barat. MVZ
					268309–10, 268313, 268317–19, 268322, 268325, MZB.Amph. 20675,
					20677–80 from Desa Uaemate (Tasiu-Tibo Road; S02.61550,
					E119.14417), Kecamatan Kaluku, Kabupaten Mamuju, Provinsi Sulawesi
					Barat. MZB.Amph.20663 from Desa Kabiraan (−2.62460, 119.14700),
					Kecamatan Ulumunda, Kabupaten Majene, Provinsi Sulawesi Barat.
				</p>
			</sec>
			<sec id="s3f">
				<title>Distribution</title>
				<p>
					<italic>Limnonectes larvaepartus</italic>
					occurs across the Northern Peninsula, as well as on the western
					margin of Sulawesi's Central Core (
					<xref ref-type="fig" rid="pone-0115884-g003">Fig. 3</xref>
					). We do not know the full extent of the species' range in the
					Central Core because the central highlands of Sulawesi remain
					poorly explored herpetologically. Several genera have species range
					boundaries in this same general region (e.g. the flying lizards
					<italic>Draco spilonotus</italic>
					and
					<italic>D. walker</italic>
					<xref ref-type="bibr" rid="pone.0115884-McGuire1">[12]</xref>
					, the fanged frogs
					<italic>Limnonectes</italic>
					sp.
					<italic>I</italic>
					and
					<italic>L.</italic>
					sp.
					<italic>D</italic>
					<xref ref-type="bibr" rid="pone.0115884-Setiadi1">[3]</xref>
					, and the tarsiers
					<italic>Tarsius lariang</italic>
					and
					<italic>T. dentatus</italic>
					<xref ref-type="bibr" rid="pone.0115884-Merker1">[13]</xref>
					.
				</p>
				<fig id="pone-0115884-g003" position="float">
					<object-id pub-id-type="doi">10.1371/journal.pone.0115884.g003</object-id>
					<label>Figure 3</label>
					<caption>
						<title>
							Distribution map (left panel) depicting the range of
							<italic>Limnonectes larvaepartus</italic>
							.
						</title>
						<p>
							The right panel shows the phylogenetic position of
							<italic>L. larvaepartus</italic>
							, with different symbol shapes denoting regional genetic
							structure in the species.
						</p>
					</caption>
					<graphic mimetype="image"
						xlink:href="info:doi/10.1371/journal.pone.0115884.g003" position="float"
						xlink:type="simple" />
				</fig>
			</sec>
			<sec id="s3g">
				<title>Diagnosis</title>
				<p>
					Prior workers have recognized substantial species diversity in the
					genus
					<italic>Limnonectes</italic>
					on Sulawesi. However, diagnosing many of these lineages on the
					basis of morphology is challenging, and several authors have
					instead opted to apply names to Sulawesi specimens representing
					species from outside Sulawesi. Consequently, the following names
					have all been incorrectly applied to Sulawesi populations: the
					Lesser Sundas species (type locality: Flores Island)
					<italic>L. dammermani</italic>
					(Mertens, 1927), the Bornean species
					<italic>L. finchii</italic>
					(Inger, 1966), the Mollucan species (type locality: Ambon)
					<italic>L. grunniens</italic>
					(Daudin, 1801), and the Philippine taxa
					<italic>L. leytensis</italic>
					(Boettger, 1893),
					<italic>L. magnus</italic>
					(Stejneger, 1909), and
					<italic>L. palavanensis</italic>
					(Boulenger, 1894). These names should not be applied to Sulawesi
					populations, as was verified phylogenetically for several of these
					taxa
					<xref ref-type="bibr" rid="pone.0115884-Setiadi1">[3]</xref>
					. Only four Sulawesi species have been described:
					<italic>L. arathooni</italic>
					(Smith, 1927),
					<italic>L. heinrichi</italic>
					(Ahl, 1933),
					<italic>L. modestus</italic>
					(Boulenger, 1882), and
					<italic>L. microtympanum</italic>
					(van Kampen, 1909). However, we will show elsewhere that
					<italic>L. heinrichi</italic>
					is a junior synonym of
					<italic>L. modestus</italic>
					and the species complex that we have referred to previously
					<xref ref-type="bibr" rid="pone.0115884-Evans1">[2]</xref>
					,
					<xref ref-type="bibr" rid="pone.0115884-Setiadi1">[3]</xref>
					as
					<italic>L. modestus</italic>
					remains undescribed – thus, at present there are but three valid
					described species of Sulawesi
					<italic>Limnonectes</italic>
					.
				</p>
				<p>
					<italic>Limnonectes larvaepartus</italic>
					can be distinguished from all other described species of
					<italic>Limnonectes</italic>
					by its reproductive mode (
					<xref ref-type="fig" rid="pone-0115884-g002">Fig. 2</xref>
					). It can be further differentiated from all described Sulawesi
					species by its combination of body size (mean male SVL = 37.4;
					female SVL = 40.2 mm), coloration, tympanum size, and degree of
					hind foot webbing, as well as on the basis of phylogenetic
					placement (
					<xref ref-type="fig" rid="pone-0115884-g003">Fig. 3</xref>
					).
					<italic>Limnonectes arathooni</italic>
					is endemic to Sulawesi's Southwestern (SW) Peninsula south of the
					Tempe Depression, and thus does not occur within the range of
					<italic>L. larvaepartus</italic>
					. It is similar in size to the new species (male SVL = 36.6 mm,
					females = 39.6 mm), but differs in having substantially reduced
					webbing (extending to penultimate phalange of fourth toe vs. to toe
					disc), in lacking fine granular dorsal tubercles, and in having
					melanic spots above the forelimb insertion, a fine ridge extending
					posteriorly behind each eye, and an alternative derived
					reproductive mode in which males guard clutches of terrestrial eggs
					that hatch into tadpoles that then make their own way to an
					adjacent stream by sliding down steep stream-side embankments
					<xref ref-type="bibr" rid="pone.0115884-Brown1">[14]</xref>
					.
					<italic>Limnonectes microtympanum</italic>
					, like
					<italic>L. arathooni</italic>
					, is restricted to the SW Peninsula south of the Tempe Depression,
					and thus does not overlap in geographical range with
					<italic>L. larvaepartus</italic>
					.
					<italic>Limnonectes microtympanum</italic>
					is moderately large (male SVL = 78.4; female SVL = 72.4 mm), and
					thus much larger than the new species.
					<italic>Limnonectes microtympanum</italic>
					also differs from the new species in having proportionally smaller
					tympana (TY/SVL = 0.05
					<underline>+</underline>
					0.01 in males, 0.06
					<underline>+</underline>
					0.01 in females versus 0.08
					<underline>+</underline>
					0.01 in both sexes in
					<italic>L. larvaepartus</italic>
					). The new species occurs in broad sympatry with
					<italic>L. modestus</italic>
					, which is a moderate sized (male SVL = 70.2 mm; female SVL = 64.0
					mm) inhabitant of fast-moving streams and substantially larger than
					<italic>L. larvaepartus</italic>
					. Like
					<italic>L. larvaepartus</italic>
					,
					<italic>L. modestus</italic>
					has nearly complete hindfoot webbing (slightly more extensive in
					<italic>L. modestus</italic>
					than in
					<italic>L. larvaepartus</italic>
					but reaching the toe disc in both species), a dusky throat with
					melanic pigments extending onto the pectoral region (in a clear
					wedge shape in
					<italic>L. modestus</italic>
					, more randomly distributed in
					<italic>L. larvaepartus</italic>
					), and skin with extensive fine granular tubercles.
					<italic>Limnonectes modestus</italic>
					also exhibits a derived reproductive mode involving production of a
					relatively small number of large (10 mm diameter) eggs that are
					deposited along the edge of fast moving streams.
				</p>
			</sec>
			<sec id="s3h">
				<title>Description of the holotype</title>
				<p>
					An adult male (
					<xref ref-type="fig" rid="pone-0115884-g001">Fig. 1</xref>
					) 48 mm SVL, body moderately robust, head not broader than body,
					head about 65% longer than wide, length 45% of snout-vent length,
					snout 17% of snout-vent length, moderately pointed, projecting
					above lower jaw, nostril lateral, closer to tip of snout than to
					eye, lore essentially straight, canthus rostralis distinct, eye
					about equal to snout length, pupil diamond-shaped, upper eyelid
					with tubercles; interorbital region smooth, width 69% of
					internarial distance, tympanum moderate, slightly wider than
					interorbital distance, supratympanic fold distinct, extending from
					posterior corner of eye to supra-axillary region, in contact with
					tympanic annulus, temporal muscle slightly enlarged; odontoid
					process 2.1 mm. Dentigerous process of vomer distinct, angled
					anterolaterally, approximately at 45
					<sup>o</sup>
					angle, posterior ends separated by distance approximately equal to
					one-third diameter of choanae. Limbs relatively slender, tibia
					width at thickest part 7.5 mm; femur length 52% of snout-vent
					length, heels moderately overlapping when placed perpendicular to
					body axis; tibia length 64.9% of foot length, 48.9% of snout-vent
					length; foot length 71% of snout-vent length; tarsal fold
					indistinct, only evident as a ridge; toe discs moderately expanded,
					circum-marginal groove horseshoe-shaped, pointed anteriorly.
					Plantar surface of foot smooth, subarticular tubercle rounded,
					relative length of toes 4&gt;3&gt;5&gt;2&gt;1. Inner metatarsal
					tubercle prominent, elongate, ovoid with a sharp spade like ventral
					edge; outer metatarsal tubercle absent; hind foot webbing full,
					extending to toe discs, slightly emarginated. Manus length 51.3% of
					foot length, fingers slender, terminal discs slightly expanded,
					length formula 3&gt;1≥4≥2, with slight differences in length,
					subarticular tubercle rounded, convex; supernumerary tubercles
					absent; inner and outer metacarpal tubercles enlarged, nuptial pads
					and webbing absent, forearm muscle not enlarged. See
					<xref ref-type="table" rid="pone-0115884-t001">Table 1</xref>
					for measurements and variation.
				</p>
				<table-wrap id="pone-0115884-t001" position="float">
					<object-id pub-id-type="doi">10.1371/journal.pone.0115884.t001</object-id>
					<label>Table 1</label>
					<caption>
						<title>
							Summary of univariate morphological variation among mensural
							characters in
							<italic>Limnonectes larvaepartus</italic>
							.
						</title>
					</caption>
					<alternatives>
						<graphic id="pone-0115884-t001-1" position="float"
							mimetype="image" xlink:href="info:doi/10.1371/journal.pone.0115884.t001"
							xlink:type="simple" />
						<table>
							<colgroup span="1">
								<col align="left" span="1" />
								<col align="center" span="1" />
								<col align="center" span="1" />
								<col align="center" span="1" />
							</colgroup>
							<thead>
								<tr>
									<td align="left" rowspan="1" colspan="1" />
									<td colspan="3" align="left" rowspan="1">
										<italic>Limnonectes larvaepartus</italic>
									</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1" />
									<td align="left" rowspan="1" colspan="1">Males (31)</td>
									<td align="left" rowspan="1" colspan="1">Females (30)</td>
									<td align="left" rowspan="1" colspan="1">Holotype</td>
								</tr>
							</thead>
							<tbody>
								<tr>
									<td align="left" rowspan="1" colspan="1">Snout-Vent Length (SVL)</td>
									<td align="left" rowspan="1" colspan="1">39.8+4.7</td>
									<td align="left" rowspan="1" colspan="1">41.2+3.1</td>
									<td align="left" rowspan="1" colspan="1">48.0</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Range</td>
									<td align="left" rowspan="1" colspan="1">31.3−48.3</td>
									<td align="left" rowspan="1" colspan="1">35.7−47.5</td>
									<td align="left" rowspan="1" colspan="1">-</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Head Length (HL)</td>
									<td align="left" rowspan="1" colspan="1">17.6+2.0</td>
									<td align="left" rowspan="1" colspan="1">17.9+1.3</td>
									<td align="left" rowspan="1" colspan="1">21.5</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Head Width (HW)</td>
									<td align="left" rowspan="1" colspan="1">15.4+2.1</td>
									<td align="left" rowspan="1" colspan="1">15.1+1.5</td>
									<td align="left" rowspan="1" colspan="1">18.9</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Snout Length (SL)</td>
									<td align="left" rowspan="1" colspan="1">7.0+0.9</td>
									<td align="left" rowspan="1" colspan="1">7.2+0.5</td>
									<td align="left" rowspan="1" colspan="1">8.2</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Eye-Narial Distance
										(EN)</td>
									<td align="left" rowspan="1" colspan="1">3.6+0.5</td>
									<td align="left" rowspan="1" colspan="1">3.9+0.4</td>
									<td align="left" rowspan="1" colspan="1">4.0</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Nostril-Tip of Snout
										(NT)</td>
									<td align="left" rowspan="1" colspan="1">3.6+0.4</td>
									<td align="left" rowspan="1" colspan="1">3.8+0.4</td>
									<td align="left" rowspan="1" colspan="1">4.3</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Internarial Distance
										(IN)</td>
									<td align="left" rowspan="1" colspan="1">4.5+0.6</td>
									<td align="left" rowspan="1" colspan="1">4.4+0.4</td>
									<td align="left" rowspan="1" colspan="1">4.9</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Snout Width at Eye
										(SWE)</td>
									<td align="left" rowspan="1" colspan="1">10.5+1.2</td>
									<td align="left" rowspan="1" colspan="1">10.6+1.0</td>
									<td align="left" rowspan="1" colspan="1">12.2</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Snout Width at Nostril
										(SWN)</td>
									<td align="left" rowspan="1" colspan="1">6.0+0.8</td>
									<td align="left" rowspan="1" colspan="1">5.8+0.7</td>
									<td align="left" rowspan="1" colspan="1">7.5</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Eye-Tympanum Distance
										(ET)</td>
									<td align="left" rowspan="1" colspan="1">1.8+0.5</td>
									<td align="left" rowspan="1" colspan="1">1.6+0.4</td>
									<td align="left" rowspan="1" colspan="1">1.7</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Odontoid Process Length
										(OP)</td>
									<td align="left" rowspan="1" colspan="1">2.0+0.3</td>
									<td align="left" rowspan="1" colspan="1">1.6+0.2</td>
									<td align="left" rowspan="1" colspan="1">2.1</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Tympanum Diameter (TY)</td>
									<td align="left" rowspan="1" colspan="1">3.1+0.5</td>
									<td align="left" rowspan="1" colspan="1">3.3+0.4</td>
									<td align="left" rowspan="1" colspan="1">3.6</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Interorbital Distance
										(IO)</td>
									<td align="left" rowspan="1" colspan="1">3.0+0.5</td>
									<td align="left" rowspan="1" colspan="1">3.0+0.4</td>
									<td align="left" rowspan="1" colspan="1">3.9</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Eye Diameter (EY)</td>
									<td align="left" rowspan="1" colspan="1">7.2+0.8</td>
									<td align="left" rowspan="1" colspan="1">7.4+0.5</td>
									<td align="left" rowspan="1" colspan="1">8.7</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Femoral Length (FE)</td>
									<td align="left" rowspan="1" colspan="1">21.1+2.6</td>
									<td align="left" rowspan="1" colspan="1">22.0+2.4</td>
									<td align="left" rowspan="1" colspan="1">23.7</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Tibial Length (TI)</td>
									<td align="left" rowspan="1" colspan="1">22.7+2.8</td>
									<td align="left" rowspan="1" colspan="1">24.1+2.3</td>
									<td align="left" rowspan="1" colspan="1">23.5</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Foot Length (FL)</td>
									<td align="left" rowspan="1" colspan="1">32.1+3.6</td>
									<td align="left" rowspan="1" colspan="1">33.5+3.0</td>
									<td align="left" rowspan="1" colspan="1">36.2</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Humeral Length (UA)</td>
									<td align="left" rowspan="1" colspan="1">11.5+1.0</td>
									<td align="left" rowspan="1" colspan="1">11.4+1.1</td>
									<td align="left" rowspan="1" colspan="1">13.5</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Lower Arm Length (LA)</td>
									<td align="left" rowspan="1" colspan="1">8.4+1.0</td>
									<td align="left" rowspan="1" colspan="1">9.1+1.0</td>
									<td align="left" rowspan="1" colspan="1">9.5</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">Hand Length (HA)</td>
									<td align="left" rowspan="1" colspan="1">9.7+1.2</td>
									<td align="left" rowspan="1" colspan="1">9.9+1.0</td>
									<td align="left" rowspan="1" colspan="1">11.8</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">HL/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.44+0.02</td>
									<td align="left" rowspan="1" colspan="1">0.44+0.03</td>
									<td align="left" rowspan="1" colspan="1">0.45</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">HW/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.39+0.02</td>
									<td align="left" rowspan="1" colspan="1">0.37+0.02</td>
									<td align="left" rowspan="1" colspan="1">0.39</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">SL/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.18+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.18+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.17</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">EN/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.09+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.09+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.08</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">IN/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.11+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.11+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.10</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">SWE/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.26+0.02</td>
									<td align="left" rowspan="1" colspan="1">0.26+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.25</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">SWN/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.15+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.14+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.16</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">ET/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.04+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.04+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.04</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">OP/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.05+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.04+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.04</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">TY/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.08+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.08+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.08</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">IO/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.08+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.07+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.08</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">EY/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.18+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.18+0.01</td>
									<td align="left" rowspan="1" colspan="1">0.18</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">FE/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.53+0.03</td>
									<td align="left" rowspan="1" colspan="1">0.53+0.03</td>
									<td align="left" rowspan="1" colspan="1">0.49</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">TI/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.57+0.04</td>
									<td align="left" rowspan="1" colspan="1">0.59+0.04</td>
									<td align="left" rowspan="1" colspan="1">0.49</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">PL/SVL</td>
									<td align="left" rowspan="1" colspan="1">0.81+0.03</td>
									<td align="left" rowspan="1" colspan="1">0.81+0.04</td>
									<td align="left" rowspan="1" colspan="1">0.75</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">SL/SWE</td>
									<td align="left" rowspan="1" colspan="1">0.67+0.07</td>
									<td align="left" rowspan="1" colspan="1">0.69+0.07</td>
									<td align="left" rowspan="1" colspan="1">0.67</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">IN/SWN</td>
									<td align="left" rowspan="1" colspan="1">0.74+0.08</td>
									<td align="left" rowspan="1" colspan="1">0.76+0.08</td>
									<td align="left" rowspan="1" colspan="1">0.65</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">HW/HL</td>
									<td align="left" rowspan="1" colspan="1">0.87+0.07</td>
									<td align="left" rowspan="1" colspan="1">0.84+0.08</td>
									<td align="left" rowspan="1" colspan="1">0.88</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">SL/HW</td>
									<td align="left" rowspan="1" colspan="1">0.46+0.05</td>
									<td align="left" rowspan="1" colspan="1">0.48+0.03</td>
									<td align="left" rowspan="1" colspan="1">0.43</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">IO/IN</td>
									<td align="left" rowspan="1" colspan="1">0.68+0.07</td>
									<td align="left" rowspan="1" colspan="1">0.67+0.08</td>
									<td align="left" rowspan="1" colspan="1">0.80</td>
								</tr>
								<tr>
									<td align="left" rowspan="1" colspan="1">TI/TD</td>
									<td align="left" rowspan="1" colspan="1">3.6+0.4</td>
									<td align="left" rowspan="1" colspan="1">3.8+0.2</td>
									<td align="left" rowspan="1" colspan="1">3.6</td>
								</tr>
							</tbody>
						</table>
					</alternatives>
				</table-wrap>
			</sec>
			<sec id="s3i">
				<title>Coloration</title>
				<p>
					The dorsal coloration is highly variable, typically brownish-grey,
					but can be darker brown on the dorsolateral region, and some
					individuals are reddish-brown or golden-tan (see
					<xref ref-type="fig" rid="pone-0115884-g002">Fig. 2</xref>
					). ∼23% of specimens have a bold mid-dorsal stripe. The venter is
					either yellowish or cream colored, with the upper end of the tibia
					usually bearing a prominent dark spot. A light bar is often present
					in the interorbital region, and the coloration of the snout to
					interorbital region may be lighter than the remainder of the
					dorsum. The tympanum is often masked in black leaving only the
					lower rim sharing the predominant body coloration. The gular region
					is usually darker in males and may have a finely mottled
					wedge-shaped melanic patch. The dorsal half of the iris is
					golden-orange in coloration in at least some individuals (we do not
					know of exceptions, but have not documented iris coloration for
					most specimens).
				</p>
			</sec>
			<sec id="s3j">
				<title>Eggs and tadpoles</title>
				<p>
					Females produce ∼100 non-pigmented eggs (see
					<xref ref-type="bibr" rid="pone.0115884-Kusrini1">[15]</xref>
					), though the most we have observed is 55, which possibly
					represents the contents of just one oviduct. The eggs lack a
					jelly-coat and reach at least ∼3 mm in diameter. These eggs develop
					within the oviducts into pigmented tadpoles that reach at least
					Gosner stage 35 prior to parturition (see
					<xref ref-type="bibr" rid="pone.0115884-Gosner1">[16]</xref>
					for staging). The gut is initially provisioned with substantial
					yolk (
					<xref ref-type="fig" rid="pone-0115884-g002">
						Fig. 2
						<italic>b</italic>
					</xref>
					), and developing tadpoles prematurely removed from the oviducts at
					approximately stage 21 progressed to approximately stage 25 over
					the course of two weeks in a water bottle without supplemental
					food, suggesting plasticity in terms of the timing of parturition.
					A detailed description of the tadpole is provided in the
					accompanying paper by Kusrini et al.
					<xref ref-type="bibr" rid="pone.0115884-Kusrini1">[15]</xref>
					.
				</p>
			</sec>
			<sec id="s3k">
				<title>Natural history</title>
				<p>
					<italic>Limnonectes larvaepartus</italic>
					occurs in natural and disturbed forest habitats of Sulawesi,
					generally in sympatry with at least one, and sometimes as many as
					five other
					<italic>Limnonectes</italic>
					species. In the western Central Core of Sulawesi, we have always
					found
					<italic>L. larvaepartus</italic>
					living in sympatry with a much larger species that has been
					referred to in the literature as
					<italic>L.</italic>
					sp.
					<italic>D</italic>
					(2,3). Whereas
					<italic>L.</italic>
					sp.
					<italic>D</italic>
					is generally found on rocks in fast-moving streams or within a
					meter of water on the banks of fast-flowing streams,
					<italic>L. larvaepartus</italic>
					is usually found further from the stream (2–10 meters from water)
					on rocky substrates, in leaf-litter, or secluded in grassy
					vegetation. Because we have observed that
					<italic>L.</italic>
					sp.
					<italic>D</italic>
					predates other frogs, including other
					<italic>Limonectes</italic>
					, it is possible that
					<italic>L. larvaepartus</italic>
					avoids large streams in response to predation pressure from larger
					<italic>Limnonectes</italic>
					species. Male
					<italic>L. larvaepartus</italic>
					typically call from the margins of seeps, puddles, or small pools
					away from the main stream. Notably, we have found many males
					calling from small pools that were already inhabited by
					<italic>L. larvaepartus</italic>
					tadpoles (
					<xref ref-type="fig" rid="pone-0115884-g002">
						Fig. 2
						<italic>c</italic>
					</xref>
					), with as many as three size-classes of tadpoles represented. It
					is unclear whether some or all of the observed tadpoles were sired
					by the accompanying male. We have furthermore collected at least
					one pregnant female from a small stream-side puddle already
					inhabited by tadpoles of two size classes, again suggesting the
					possibility that individual pools may be visited repeatedly by the
					same adult males and females during the reproductive season.
				</p>
			</sec>
		</sec>
		<sec id="s4">
			<title>Discussion</title>
			<p>
				We first became aware of the unusual reproductive mode of
				<italic>Limnonectes larvaepartus</italic>
				while conducting fieldwork on Sulawesi. In several instances, we
				discovered tadpoles in the oviducts while preparing specimens. In
				each case, having sacrificed frogs for preparation, the abdominal
				wall was observed to quiver, and incision resulted in living
				tadpoles emerging from the opening (see
				<xref ref-type="supplementary-material" rid="pone.0115884.s001">S1 Movie</xref>
				). On one occasion, a gravid female gave birth to tadpoles in-hand
				at the moment of capture. On four other occasions, field-collected
				<italic>L. larvaepartus</italic>
				in our possession gave birth to tadpoles while being held
				individually in collecting bags. In total, we have either observed
				tadpoles in the oviducts or direct-birth of tadpoles on 19
				occasions.
			</p>
			<p>
				Because we have not witnessed natural birth of tadpoles in
				free-living frogs, two possible alternative reproductive modes are
				possible for this species.
				<italic>Limnonectes larvaepartus</italic>
				reproduction may simply reflect what we have observed in the hand –
				direct birth of tadpoles. Alternatively, this species may be capable
				of retaining developing young in the oviducts through metamorphosis
				with subsequent birth of froglets, as is the case for
				<italic>Eleutherodactylus jasperi</italic>
				and members of the African bufonid genera
				<italic>Nectophrynoides</italic>
				and
				<italic>Nimbaphrynoides</italic>
				<xref ref-type="bibr" rid="pone.0115884-Wells1">[9]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Wake2">[17]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Wake3">[18]</xref>
				. The latter mode seems unlikely for several reasons. First, we have
				collected 19 pregnant individuals carrying tadpoles in the oviducts
				but none carrying froglets. These 19 females were collected across
				different months, years, and localities. Second, we have observed
				and collected at least four clutches of free-living
				<italic>L. larvaepartus</italic>
				tadpoles in small pools of water on the margins of streams, with
				each of these samples exhibiting two or three size classes and thus
				possibly representing multiple clutches. Three of these clutches
				were accompanied by males, some of which were calling, and one was
				found in association with the gravid female that gave birth to 55
				tadpoles (some in the hand of JAM, others subsequently deposited in
				a collecting bag). Finally, although adaptive plasticity has been
				demonstrated for many frog species, whereby tadpoles or froglets are
				capable of hatching from their eggs prematurely when attacked by
				predators
				<xref ref-type="bibr" rid="pone.0115884-Warkentin1">[19]</xref>
				, this has only been documented for a single direct-developer,
				<italic>Eleutherodactylus coqui</italic>
				<xref ref-type="bibr" rid="pone.0115884-Buckley1">[20]</xref>
				. In the case of
				<italic>E. coqui</italic>
				, the capacity for early hatching commenced at stages 13 or 14,
				whereas normal hatching occurred at stage 15 (see
				<xref ref-type="bibr" rid="pone.0115884-Townsend1">[21]</xref>
				for staging). Thus,
				<italic>E. coqui</italic>
				were capable of premature hatching only as froglets, with toe pads
				and eyelids present but before the tail was completely resorbed, a
				stage much later than would be required by
				<italic>L. larvaepartus</italic>
				if it were simultaneously capable of hatching as either a tadpole or
				full-term froglet. Most direct-developers, as well as the few frog
				species that give birth to froglets, pass through the tadpole stage
				in a form poorly suited for free-living. In many such species, the
				tadpole has a broad, highly vascularized ‘respiratory tail’
				specialized for intra-egg or intra-oviductal gas exchange rather
				than for swimming, as is the case for
				<italic>Eleutherodactylus jasperi</italic>
				<xref ref-type="bibr" rid="pone.0115884-Wake2">[17]</xref>
				, and may lack mouthparts and functional gills
				<xref ref-type="bibr" rid="pone.0115884-Lutz1">[22]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Lynn1">[23]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Thibaudeau1">[24]</xref>
				. In others, the tail remains rudimentary and limb buds appear early
				in development such that a typical tadpole phenotype never occurs
				<xref ref-type="bibr" rid="pone.0115884-Lutz1">[22]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Lynn1">[23]</xref>
				. Given these observations, it appears unlikely that any one species
				would be characterized by the combination of (1) internal
				fertilization, (2) complete metamorphosis within the oviducts and
				live-birth in the form of froglets, as well as (3) an oviductal
				tadpole stage that is capable of premature birth and free-living.
				Although we think that
				<italic>L. larvaepartus</italic>
				is much more likely to give birth to tadpoles as its sole mode of
				reproduction as opposed to exhibiting adaptive plasticity allowing
				for birth of either tadpoles or froglets depending on the
				circumstances confronting the frog, we note that either condition
				would be unique among Anura. In either case,
				<italic>L. larvaepartus</italic>
				requires internal fertilization, which is, itself, extremely rare
				among anurans
				<xref ref-type="bibr" rid="pone.0115884-McDiarmid1">[8]</xref>
				,
				<xref ref-type="bibr" rid="pone.0115884-Duellman1">[25]</xref>
				. The mechanism by which internal fertilization takes place is
				unknown, and there is no obvious intromittent organ present to
				facilitate sperm transfer. If
				<italic>L. larvaepartus</italic>
				reproduction always involves birth of tadpoles, it is likely that
				they are ovoviparous given that the tadpoles are well provisioned
				with yolk and appear fully capable of developing without nutrient
				transfer from the mother. Prior to parturition, the tadpoles have
				well-developed tails and pigmentation, and well developed
				mouthparts. Given this morphology, it is unlikely that these
				tadpoles are endotrophic (never feed before metamorphosing into
				froglets). It is more likely that the tadpoles are born after
				exhausting their yolk supply, and are subsequently self-feeding
				prior to metamorphosis. Nevertheless, it is clear that much remains
				to be learned about this unusual frog, the discovery of which brings
				to light yet another axis of diversity characterizing the remarkable
				Sulawesi fanged frog adaptive radiation.
			</p>
		</sec>
		<sec id="s5">
			<title>Supporting Information</title>
			<supplementary-material id="pone.0115884.s001"
				mimetype="video/quicktime" xlink:href="info:doi/10.1371/journal.pone.0115884.s001"
				position="float" xlink:type="simple">
				<label>S1 Movie</label>
				<caption>
					<p>
						<bold>Video showing the characteristic quivering abdomen caused by
							movement of tadpoles within a pregnant female </bold>
						<bold>
							<italic>Limnonectes larvaepartus.</italic>
						</bold>
					</p>
					<p>(MOV)</p>
				</caption>
			</supplementary-material>
		</sec>
	</body>
	<back>
		<ack>
			<p>We thank K. N. Tjan, U. Arifin, K. Laras, Suwatio, A. Rachmansah,
				R. M. Brown, and C. J. Hayden for assistance in the field. Boeadi
				and Mumpuni for MZB specimens; R. F. Inger for discussion and access
				to comparative materials from FMNH, USNM and MCZ; D. N. Cannatella,
				M. H. Wake, and members of the McGuire Lab for helpful discussion.</p>
		</ack>
		<ref-list>
			<title>References</title>
			<ref id="pone.0115884-Emerson1">
				<label>1</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Emerson</surname>
						<given-names>SB</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Inger</surname>
						<given-names>RF</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Iskandar</surname>
						<given-names>DT</given-names>
					</name>
					(
					<year>2000</year>
					)
					<article-title>Molecular phylogenetics and Evolution of fanged
						Frogs</article-title>
					.
					<source>Mol Phylogenet Evol</source>
					<volume>16</volume>
					:
					<fpage>131</fpage>
					–
					<lpage>142</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Evans1">
				<label>2</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Evans</surname>
						<given-names>BJ</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Brown</surname>
						<given-names>RM</given-names>
					</name>
					,
					<name name-style="western">
						<surname>McGuire</surname>
						<given-names>JA</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Supriatna</surname>
						<given-names>J</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Andayani</surname>
						<given-names>N</given-names>
					</name>
					,
					<etal>et al</etal>
					. (
					<year>2003</year>
					)
					<article-title>Phylogenetics of fanged frogs; testing
						biogeographical hypotheses at the interface of the Asian and
						Australian faunal zones</article-title>
					.
					<source>Syst Biol</source>
					<volume>526</volume>
					:
					<fpage>794</fpage>
					–
					<lpage>819</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Setiadi1">
				<label>3</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Setiadi</surname>
						<given-names>MI</given-names>
					</name>
					,
					<name name-style="western">
						<surname>McGuire</surname>
						<given-names>JA</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Brown</surname>
						<given-names>RM</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Zubairi</surname>
						<given-names>M</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Iskandar</surname>
						<given-names>DT</given-names>
					</name>
					,
					<etal>et al</etal>
					. (
					<year>2011</year>
					)
					<article-title>
						Adaptive radiation and ecological opportunity in Sulawesi and
						Philippine fanged frogs (
						<italic>Limnonectes</italic>
						) Communities
					</article-title>
					.
					<source>Amer Nat</source>
					<volume>178</volume>
					:
					<fpage>221</fpage>
					–
					<lpage>240</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Iskandar1">
				<label>4</label>
				<mixed-citation publication-type="other" xlink:type="simple">
					Iskandar DT, Tjan KN (1996) The Amphibians and Reptiles of
					Sulawesi: with notes on the distribution and chromosomal number of
					frogs. Proc. 1
					<sup>st</sup>
					Intl. Conf. Eastern Indonesian-Australian Vertebrates. Manado pp.
					39–46.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Wake1">
				<label>5</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Wake</surname>
						<given-names>MH</given-names>
					</name>
					(
					<year>2003</year>
					)
					<article-title>Reproductive modes, ontogenies, and the evolution of
						body form</article-title>
					.
					<source>Anim Biol</source>
					<volume>53</volume>
					:
					<fpage>209</fpage>
					–
					<lpage>223</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Haddad1">
				<label>6</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Haddad</surname>
						<given-names>CFB</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Prado</surname>
						<given-names>CPA</given-names>
					</name>
					(
					<year>2005</year>
					)
					<article-title>Reproductive modes in frogs and their unexpected
						diversity in the Atlantic Forest of Brazil</article-title>
					.
					<source>BioScience</source>
					<volume>55</volume>
					:
					<fpage>207</fpage>
					–
					<lpage>217</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-AmphibiaWeb1">
				<label>7</label>
				<mixed-citation publication-type="other" xlink:type="simple">
					AmphibiaWeb: Information on amphibian biology and conservation
					<bold>.</bold>
					Available:
					<ext-link ext-link-type="uri" xlink:href="http://amphibiaweb.org/"
						xlink:type="simple">http://amphibiaweb.org/</ext-link>
					. Accessed: October 13, 2014.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-McDiarmid1">
				<label>8</label>
				<mixed-citation publication-type="book" xlink:type="simple">McDiarmid
					RW, Altig R (1999) Tadpoles: the biology of anuran larvae. Chicago:
					University of Chicago Press. 444 p.</mixed-citation>
			</ref>
			<ref id="pone.0115884-Wells1">
				<label>9</label>
				<mixed-citation publication-type="book" xlink:type="simple">Wells
					KD (2008) The Ecology and Behavior of Amphibians. Chicago: The
					University of Chicago Press. 1148 p.</mixed-citation>
			</ref>
			<ref id="pone.0115884-Guayasamin1">
				<label>10</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Guayasamin</surname>
						<given-names>JM</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Bustamante</surname>
						<given-names>MR</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Almeida-Reinoso</surname>
						<given-names>D</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Funk</surname>
						<given-names>WC</given-names>
					</name>
					(
					<year>2006</year>
					)
					<article-title>Glass frogs (Centrolenidae) of Yanayacu Biological
						Station, Ecuador, with the description of a new species and
						comments on centrolenid systematics</article-title>
					.
					<source>Zool J Linn Soc</source>
					<volume>147</volume>
					:
					<fpage>489</fpage>
					–
					<lpage>513</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Inger1">
				<label>11</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Inger</surname>
						<given-names>RF</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Voris</surname>
						<given-names>HK</given-names>
					</name>
					(
					<year>2001</year>
					)
					<article-title>The biogeographical relations of the frogs and
						snakes of Sundaland</article-title>
					.
					<source>J Biogeogr</source>
					<volume>28</volume>
					:
					<fpage>863</fpage>
					–
					<lpage>891</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-McGuire1">
				<label>12</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>McGuire</surname>
						<given-names>JA</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Brown</surname>
						<given-names>RM</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Mumpuni, Riyanto</surname>
						<given-names>A</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Andayani</surname>
						<given-names>N</given-names>
					</name>
					(
					<year>2007</year>
					)
					<article-title>
						The flying lizards of the
						<italic>Draco lineatus</italic>
						group (Squamata: Iguania: Agamidae): A taxonomic revision with
						descriptions of two new species
					</article-title>
					.
					<source>Herp Monog</source>
					<volume>21</volume>
					:
					<fpage>179</fpage>
					–
					<lpage>212</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Merker1">
				<label>13</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Merker</surname>
						<given-names>S</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Driller</surname>
						<given-names>C</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Dahruddin</surname>
						<given-names>H</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Wirdateti, Sinaga</surname>
						<given-names>W</given-names>
					</name>
					,
					<etal>et al</etal>
					. (
					<year>2010</year>
					)
					<article-title>
						<italic>Tarsius wallacei</italic>
						: A new tarsier species from Central Sulawesi occupies a
						discontinuous range
					</article-title>
					.
					<source>Int J Primatol</source>
					<volume>31</volume>
					:
					<fpage>1107</fpage>
					–
					<lpage>1122</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Brown1">
				<label>14</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Brown</surname>
						<given-names>RM</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Iskandar</surname>
						<given-names>DT</given-names>
					</name>
					(
					<year>2000</year>
					)
					<article-title>
						Nest site Selection, larval hatching and advertisement calls of
						<italic>Rana arathooni</italic>
						from southwestern Sulawesi (Celebes) Island, Indonesia
					</article-title>
					.
					<source>J Herp</source>
					<volume>34</volume>
					:
					<fpage>404</fpage>
					–
					<lpage>413</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Kusrini1">
				<label>15</label>
				<mixed-citation publication-type="other" xlink:type="simple">
					Kusrini MD, Rowley JJL, Khairunnisa LR, Shea GM, Altig R (2014) A
					new breeding mode for anurans: reproductive biology and larvae of
					the Indonesian frog
					<italic>Limnonectes larvaepartus</italic>
					. PLOS ONE. In press Vol. 10, Issue 1.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Gosner1">
				<label>16</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Gosner</surname>
						<given-names>KL</given-names>
					</name>
					(
					<year>1960</year>
					)
					<article-title>A simplified table for staging anuran embryos and
						larvae with notes on identification</article-title>
					.
					<source>Herpetologica</source>
					<volume>16</volume>
					:
					<fpage>183</fpage>
					–
					<lpage>190</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Wake2">
				<label>17</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Wake</surname>
						<given-names>MH</given-names>
					</name>
					(
					<year>1978</year>
					)
					<article-title>
						The reproductive biology of
						<italic>Eleutherodactylus jasperi</italic>
						(Amphibia, Anura, Leptodactylidae), with comments on the evolution
						of live-bearing systems
					</article-title>
					.
					<source>J Herpetol</source>
					<volume>12</volume>
					:
					<fpage>121</fpage>
					–
					<lpage>133</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Wake3">
				<label>18</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Wake</surname>
						<given-names>MH</given-names>
					</name>
					(
					<year>1980</year>
					)
					<article-title>
						The reproductive biology of
						<italic>Nectophrynoides malcolmi</italic>
						(Amphibia: Bufonidae) with comments on the evolution of
						reproductive modes in the genus
						<italic>Nectophrynoides</italic>
					</article-title>
					.
					<source>Copeia</source>
					<volume>1980</volume>
					:
					<fpage>193</fpage>
					–
					<lpage>209</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Warkentin1">
				<label>19</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Warkentin</surname>
						<given-names>KM</given-names>
					</name>
					(
					<year>2011</year>
					)
					<article-title>Plasticity of hatching in amphibians: Evolution,
						trade-offs, cues and mechanisms</article-title>
					.
					<source>Integr Comp Biol</source>
					<volume>51</volume>
					:
					<fpage>111</fpage>
					–
					<lpage>127</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Buckley1">
				<label>20</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Buckley</surname>
						<given-names>CR</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Michael</surname>
						<given-names>SF</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Irschick</surname>
						<given-names>DJ</given-names>
					</name>
					(
					<year>2005</year>
					)
					<article-title>
						Early hatching decreases jumping performance in a
						direct-developing frog,
						<italic>Eleutherodactylus coqui</italic>
					</article-title>
					.
					<source>Funct Ecol</source>
					<volume>19</volume>
					:
					<fpage>67</fpage>
					–
					<lpage>72</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Townsend1">
				<label>21</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Townsend</surname>
						<given-names>DS</given-names>
					</name>
					,
					<name name-style="western">
						<surname>Stewart</surname>
						<given-names>MM</given-names>
					</name>
					(
					<year>1985</year>
					)
					<article-title>
						Direct development in
						<italic>Eleutherodactylus coqui</italic>
						(Anura: Leptodactylidae): a staging table
					</article-title>
					.
					<source>Copeia</source>
					<volume>1985</volume>
					:
					<fpage>423</fpage>
					–
					<lpage>436</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Lutz1">
				<label>22</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Lutz</surname>
						<given-names>B</given-names>
					</name>
					(
					<year>1948</year>
					)
					<article-title>Ontogenetic evolution in frogs</article-title>
					.
					<source>Evolution</source>
					<volume>2</volume>
					:
					<fpage>29</fpage>
					–
					<lpage>39</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Lynn1">
				<label>23</label>
				<mixed-citation publication-type="journal"
					xlink:type="simple">
					<name name-style="western">
						<surname>Lynn</surname>
						<given-names>WG</given-names>
					</name>
					(
					<year>1942</year>
					)
					<article-title>
						The embryology of
						<italic>Eleutherodactylus nubicola</italic>
						, an anuran which has no tadpole stage
					</article-title>
					.
					<source>Carn Inst Wash Publ</source>
					<volume>541</volume>
					:
					<fpage>27</fpage>
					–
					<lpage>62</lpage>
					.
				</mixed-citation>
			</ref>
			<ref id="pone.0115884-Thibaudeau1">
				<label>24</label>
				<mixed-citation publication-type="book" xlink:type="simple">Thibaudeau
					G, Altig R (1999) Endotrophic anurans – development and evolution.
					In: McDiarmid RW, R Altigeditors.Tadpoles. The biology of anuran
					larvae, Chicago: The University of Chicago Press pp. 170–188.</mixed-citation>
			</ref>
			<ref id="pone.0115884-Duellman1">
				<label>25</label>
				<mixed-citation publication-type="book" xlink:type="simple">Duellman
					WE, Trueb L (1986) Biology of Amphibians. New York: McGraw-Hill
					Publishing Company. 670 p.</mixed-citation>
			</ref>
		</ref-list>
	</back>
</article>