identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
447D878FFFDEFFB1FEE9982AFEE0FA57.text	447D878FFFDEFFB1FEE9982AFEE0FA57.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gastropteron Kosse 1813	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> GASTROPTERON KOSSE, 1813</p>
            <p>Type species</p>
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	https://treatment.plazi.org/id/447D878FFFDEFFB1FEE9982AFEE0FA57	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFDEFFB1FF7A98AAFD27F9D6.text	447D878FFFDEFFB1FF7A98AAFD27F9D6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gastropteron meckeli Blainville 1825	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Gastropteron meckeli Blainville, 1825 , by monotypy. </p>
            <p> GASTROPTERON MINUTUM ONG &amp; GOSLINER SP.</p>
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	https://treatment.plazi.org/id/447D878FFFDEFFB1FF7A98AAFD27F9D6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFD3FFBCFCEA9CE4FBF7FDAC.text	447D878FFFD3FFBCFCEA9CE4FBF7FDAC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gastropteron multo Ong & Hallas & Gosliner 2017	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> GASTROPTERON MULTO ONG &amp; GOSLINER SP. NOV.</p>
            <p>(FIGS 1C, D, 4B, 5, 6)</p>
            <p>LSID urn:lsid:zoobank.org:act: 025F1253-A0D7-4D2A- B332-CE30DF011436</p>
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	https://treatment.plazi.org/id/447D878FFFD3FFBCFCEA9CE4FBF7FDAC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFD3FFBCFCF09F22FBABFBB2.text	447D878FFFD3FFBCFCF09F22FBABFBB2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gastropteron undefined-3	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Gastropteron sp. 3 Gosliner et al., 2015: 55, lower right fig. </p>
            <p>Type material</p>
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                  Holotype, dissected, NMP 041181, originally CASIZ 199130,  
                <a title="Search Plazi for locations around (long 120.89564/lat 13.68688)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.89564&amp;materialsCitation.latitude=13.68688">Mainit Bubbles</a>
                 (13.68688°N 120.89564°E),  
                <a title="Search Plazi for locations around (long 120.89564/lat 13.68688)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.89564&amp;materialsCitation.latitude=13.68688">Mabini</a>
                 , Batangas, Luzon, Philippines, 5 m depth, 3 May 2014, Alexis Principe  .   Paratype, CASIZ 186048, one specimen,  
                <a title="Search Plazi for locations around (long 120.92615/lat 13.76001)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.92615&amp;materialsCitation.latitude=13.76001">Anilao Pier</a>
                 (13.76001°N 120.92615°E), Mabini, Batangas, Luzon, Philippines, 5 m depth, 30 April 2011  , T. Gosliner. 
            </p>
            <p>Type locality</p>
            <p>
                  Mainit Bubbles (13.68688°N 120.89564°E),  
                <a title="Search Plazi for locations around (long 120.89564/lat 13.68688)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.89564&amp;materialsCitation.latitude=13.68688">Mabini</a>
                 , Batangas, Luzon, Philippines  . 
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	https://treatment.plazi.org/id/447D878FFFD3FFBCFCF09F22FBABFBB2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFD0FFBFFF609C40FD34FDDC.text	447D878FFFD0FFBFFF609C40FD34FDDC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Sagaminopteron Tokioka & Baba 1964	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> GENUS  SAGAMINOPTERON TOKIOKA &amp; BABA, 1964</p>
            <p>Type species</p>
            <p> Sagaminopteron ornatum Tokioka &amp; Baba, 1964 , by original designation. </p>
            <p> SAGAMINOPTERON MULTIMACULATUM ONG &amp; </p>
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	https://treatment.plazi.org/id/447D878FFFD0FFBFFF609C40FD34FDDC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFD6FFB9FF229870FD31FA46.text	447D878FFFD6FFB9FF229870FD31FA46.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Siphopteron Gosliner 1989	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> GENUS  SIPHOPTERON GOSLINER, 1989</p>
            <p>Type species</p>
            <p> Siphopteron tigrinum Gosliner, 1989 , by original designation </p>
            <p> SIPHOPTERON VERMICULUM ONG &amp; GOSLINER</p>
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	https://treatment.plazi.org/id/447D878FFFD6FFB9FF229870FD31FA46	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFD6FFB9FF7A9BD4FB59FDB4.text	447D878FFFD6FFB9FF7A9BD4FB59FDB4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Siphopteron undefined-4	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Siphopteron sp. 4 , Gosliner et al., 2015: 59, middle left fig. </p>
            <p>Type material</p>
            <p>
                  Holotype, dissected, NMP 041183, originally CASIZ 199134,  
                <a title="Search Plazi for locations around (long 120.83024/lat 13.68428)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.83024&amp;materialsCitation.latitude=13.68428">Cemetery Beach</a>
                 (13.68428°N 120.83024°E), Tingloy, Batangas, Luzon, Philippines, 7 m depth, 2 May 2014, T. Gosliner  .   Paratypes: CASIZ 199126, one specimen, Cemetery Beach (13.68428°N 120.83024°E),  
                <a title="Search Plazi for locations around (long 120.83024/lat 13.68428)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.83024&amp;materialsCitation.latitude=13.68428">Tingloy</a>
                 , Batangas, Luzon, Philippines, 7 m depth, 2 May 2014  ,   T.  Gosliner .   CASIZ 199129, three specimens, Cemetery Beach (13.68428°N 120.83024°E),  
                <a title="Search Plazi for locations around (long 120.83024/lat 13.68428)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.83024&amp;materialsCitation.latitude=13.68428">Tingloy</a>
                 , Batangas, Luzon, Philippines, 7 m depth, 6 May 2014  ,   T.  Gosliner . 
            </p>
            <p>Type locality</p>
            <p>
                  Cemetery Beach (13.68428°N 120.83024°E),  
                <a title="Search Plazi for locations around (long 120.83024/lat 13.68428)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.83024&amp;materialsCitation.latitude=13.68428">Tingloy</a>
                 , Batangas, Luzon, Philippines  . 
            </p>
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	https://treatment.plazi.org/id/447D878FFFD6FFB9FF7A9BD4FB59FDB4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFCDFFA3FCEC98F9FD7CFEBE.text	447D878FFFCDFFA3FCEC98F9FD7CFEBE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Siphopteron makisig Ong & Hallas & Gosliner 2017	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> SIPHOPTERON MAKISIG ONG &amp; GOSLINER SP. NOV.</p>
            <p>(FIGS 14C, 17C, D, 18, 19)</p>
            <p>LSID urn:lsid:zoobank.org:act: 9E247ACD-9C5B-49A4-BE53-B8A62C7C6961</p>
            <p> Siphopteron sp. 1 Lange et al., 2013: fig. 1b.  Siphopteron sp. 5 Gosliner et al., 2015: 59, middle right fig. </p>
            <p>Type material</p>
            <p>
                  Holotype, dissected, NMP 041186, ex CASIZ 199113, one specimen,  
                <a title="Search Plazi for locations around (long 120.6123/lat 13.929165)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.6123&amp;materialsCitation.latitude=13.929165">Bilbago Reef South</a>
                 , Calatagan (13.929165°N 120.612299°E), Batangas, Luzon, Philippines, 7 m depth, 17 May 2014, T. Gosliner. 
            </p>
            <p>Type locality</p>
            <p>
                  B i l b a g o R e e f S o u t h, C a l a t a g a n (1 3.9 2 9 1 6 5°N 120.612299°E), Batangas,  
                <a title="Search Plazi for locations around (long 120.6123/lat 5.0)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.6123&amp;materialsCitation.latitude=5.0">Luzon</a>
                 , Philippines  . 
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	https://treatment.plazi.org/id/447D878FFFCDFFA3FCEC98F9FD7CFEBE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFC2FFADFF6C983DFD64F8CE.text	447D878FFFC2FFADFF6C983DFD64F8CE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Siphopteron dumbo Ong & Hallas & Gosliner 2017	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> SIPHOPTERON DUMBO ONG &amp; GOSLINER SP. NOV.</p>
            <p>(FIGS 14D, 20, 21, 22)</p>
            <p>LSID urn:lsid:zoobank.org:act: C4E5B8E4-5C13- 4D08-9E7C-CCC19FCDC1F0</p>
            <p>Type material</p>
            <p>
                  Holotype, dissected, NMP 041187, ex CASIZ 204849, one specimen, La Laguna,  
                <a title="Search Plazi for locations around (long 120.97016/lat 13.525953)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.97016&amp;materialsCitation.latitude=13.525953">Puerto Galera</a>
                 (13.525953°N 120.970160°E), Mindoro Oriental, Philippines, 20 m depth, 26 April 2015, T. Gosliner. 
            </p>
            <p>Type locality</p>
            <p> L a L a g u n a, P u e r t o G a l e r a (1 3. 5 2 5 9 5 3 ° N 120.970160°E), Mindoro Oriental, Philippines . </p>
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	https://treatment.plazi.org/id/447D878FFFC2FFADFF6C983DFD64F8CE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
447D878FFFC5FFABFF4898DFFB24F9C1.text	447D878FFFC5FFABFF4898DFFB24F9C1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Sagaminopteron nigropunctatum	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> sympatric  Sagaminopteron nigropunctatum and  S.</p>
            <p> ornatum are 11.0% divergent. Geographically isolated sister species such as  S. leah and  S. quadrispinosum are 9.3–9.9% divergent, whereas  S. michaeli and  S.</p>
            <p> tigrinum are 10.2% divergent. In contrast, allopatric conspecific Hawai’ian and Philippine specimens of  Gastropteron minutum are 3.2% divergent. Widely separated specimens of some apparent conspecifics exhibit some intriguing patterns of divergence. A Red Sea specimen of  Sagaminopteron nigropunctatum is 8.8% different to a specimen of what has been considered the same species from the Philippines. Three western Pacific specimens of  S. psychedelicum from Australia, Papua New Guinea and the Philippines are 1.1–1.4% divergent to each other, whereas the specimen from Madagascar is 11.1–12.3% divergent to the western Pacific specimens. The data for both  S. nigropunctatum and  S. psychedelicum strongly suggest that the Red Sea and western Indian Ocean specimens of these species represent distinct cryptic species and are in need of more detailed study. The western Pacific and western Indian Ocean specimens of  S. tigrinum are strongly divergent (12.3%). In fact,  S. michaeli is more similar to the western Indian Ocean specimen of  S. tigrinum (8.1%) than the two  S. tigrinum specimens are related to each other. It appears that the species listed as  S. brunneomarginatum by Anthes et al. (2008) represents a distinct taxon from the specimen identified as  S. brunneomarginatum and sequenced in this study as these two taxa are 14.8% divergent in their COI sequences and 10.5% divergent from its closest relative,  S. tigrinum . The identity of this taxon remains questionable and no voucher specimens exist. </p>
            <p>RELATIONSHIPS BETWEEN MORPHOLOGY AND MOLECULAR PHYLOGENY</p>
            <p> Much of the comparative anatomy and phylogenetic relationships based on morphological characters were presented by Gosliner (1989). In that study, four genera were distinguished:  Gastropteron ,  Enotepteron ,  Sagaminopteron and  Siphopteron with  Gastropteron and  Enotepteron as sister taxa and with  Sagaminopteron and  Siphopteron as sister taxa. Anthes et al. (2008) found a similar arrangement of taxa but did not include any species of  Enotepteron in their molecular analysis. In their study,  S. pohnpei nested with species of  Sagaminopteron rather than with species of  Siphopteron . In this study, we find a similar result with both  S. pohnpei and  S. multimaculatum nesting within  Sagaminopteron and have placed these taxa within this genus. No suitably preserved specimens of  Enotepteron were available to include in the molecular analysis and the specimen initially identified as  Enotepteron sp. is nested within  Siphopteron , as clade 2, and was likely misidentified. This species is the sister to the remainder of  Siphopteron , but was not observed alive. </p>
            <p> Three of the four genera of  Gastropteridae recognized by Gosliner (1989) were studied here. Of those, only  Sagaminopteron is strongly supported as monophyletic, while  Gastropteron and  Siphopteron are not. Nevertheless, they have morphological synapomorphies that support their continued use until more robust phylogenies are developed. Gosliner (1989) noted that species of  Gastropteron had a synapomorphy of possessing a triangular thickening on the inner lateral tooth that is also present in  G. multo and  G. minutum , described here (Figs 3C, 6C, indicated by arrow). The same is true for  Siphopteron . While we transferred  Sagaminopteron pohnpei together with its sister taxon,  S. multimaculatum to  Sagaminopteron , the remaining species of  Siphopteron all have a complex penis and narrow, triangular inner lateral teeth with a reduced number of denticles, attributes that are considered to be synapomorphies in the morphological phylogeny of Gosliner (1989). </p>
            <p> While the monophyly of  Gastropteron and  Siphopteron was not supported, the various well-supported clades recovered in our molecular phylogeny have strong morphological correlates. In  Gastropteron , the strongly supported subclade that includes  G. multo and  G. bicornutum , both sister taxa have a pair of posterior flagella, whereas in the strongly supported subclade of  G. minutum and  G. rubrum have only a single flagellum. Unfortunately, molecular data are not available for other species of  Gastropteron to ascertain, for example, whether the Indo-Pacific  G. minutum is indeed most closely related to the Atlantic  G. rubrum or whether other taxa from different geographical regions are more closely related to these two taxa. Inclusion of these taxa in the future is necessary for further testing the monophyly of  Gastropteron , which was not supported in this study. </p>
            <p> In  Sagaminopteron , the clade that includes  S. ornatum ,  S. nigropunctatum and  S. psychedelicum all have a simple penis and inner lateral teeth with two large cusps (Tokioka &amp; Baba, 1964; Gosliner, 1989), while  S. pohnpei and  S. multimaculatum have a more complex penis with a secondary duct and have an inner lateral tooth with many elongate denticles (Gosliner, 1989; this study). Only when  S. pohnpei and  S. multimaculatum are added to  Sagaminopteron , is the monophyly of this genus strongly supported. </p>
            <p> The three clades of Siphoptero n recovered here also have some unifying morphological features. In the first well-supported clade of  Siphopteron (Fig. 23, clade 1) that includes,  S. tigrinum ,  S. michaeli and  S. brunneomarginatum , the inner radular teeth have a weakly developed masticatory margin with a series of small denticles. The penial papilla of  S. tigrinum and  S. michaeli have cuticular spines, whereas the penis of their sister species,  S. brunneomarginatum , lacks spines. The strongly supported large clade 3 that includes  S. nakakatuwa ,  S. flavolineatum ,  S. vermiculum , S. sp. 1, S. sp. 2,  S. nigromarginatum ,  S. dumbo ,  S. quadrispinosum ,  S. leah and  S. makisig , the radula has an inner tooth with two large, triangular denticles. These are the only species of  Siphopteron with this form of dentition. In the well-supported subclade of clade 3 that includes  S. quadrispinosum ,  S. leah and  S. makisig , the penis is complex with a few large basal cuticular spines on the penial papilla that has spirally arranged spines or tubercles and a cuticular stylet present on a secondary papilla. In the remaining species in the second subclade of clade 3, there are multiple rows of cuticular spines associated with the penis and the penial papilla is not elongate with a series of spirally arranged spines. The anatomy of S. sp. CASIZ 181575 (clade 2) that is the sister taxon to the remain- der of  Siphopteron remains unstudied. </p>
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	https://treatment.plazi.org/id/447D878FFFC5FFABFF4898DFFB24F9C1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ong, Elise;Hallas, Joshua M.;Gosliner, Terrence M.	Ong, Elise, Hallas, Joshua M., Gosliner, Terrence M. (2017): Like a bat out of heaven: the phylogeny and diversity of the bat-winged slugs (Heterobranchia: Gastropteridae). Zoological Journal of the Linnean Society 180: 755-789
