<mods:titleid="3BD9C50BDF3FF2EE28019FBC00F728C6">Systematic revision of the genus Peronia Fleming, 1822 (Gastropoda, Euthyneura, Pulmonata, Onchidiidae)</mods:title>
<mods:affiliationid="B1B27DEA48ADEB3358F9B1652B89C64B">Department of Biology, Pennsylvania State University, University Park, PA 16802, USA</mods:affiliation>
<mods:affiliationid="237B3633D7FADDCDFB74EE06E3944609">Department of Biology, Pennsylvania State University, University Park, PA 16802, USA</mods:affiliation>
<mods:affiliationid="CD8DEC8D8025B22B1AD2BB6F263338F3">Centre of Excellence in Marine Biology, University of Karachi, Karachi 75270, Pakistan</mods:affiliation>
<mods:affiliationid="4B3DB33406EFFE330C0C2C5CFC3360E8">College of Engineering, Information Technology and the Environment, Charles Darwin University, Ellengowan Dr, Casuarina, NT 0810, Australia</mods:affiliation>
<mods:affiliationid="EF67750EFBD0B55BFC019639697D2ACA">National Museum of the Philippines, Taft Ave, Ermita, Manila, 1000, Metro Manila, Philippines</mods:affiliation>
<mods:affiliationid="F3F80E8C254CF6D2F16F1BE90EC14D01">Department of Marine Science, Universitas Malikussaleh, Reuleut Main Campus, Kecamatan Muara Batu, North Aceh, Aceh, 24355, Indonesia</mods:affiliation>
<mods:namePartid="6AF0F23C95545FB97D1BBD28F6293ADE">Ngo, Xuan Qu ảng</mods:namePart>
<mods:affiliationid="9A3D8803786E9ED4D9CFF64C7636A429">Institute of Tropical Biology, Vietnam Academy of Science and Technology, 85 Tran Quoc Toan Street, District 3, Ho Chi Minh City, Vietnam & Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam</mods:affiliation>
<mods:affiliationid="B3FA9219BC88DB31C37ED2C3B4394EAE">Lee Kong Chian Natural History Museum, 2 Conservatory Dr, National University of Singapore, 117377, Singapore</mods:affiliation>
<mods:affiliationid="CB18763DC95BFA07E00DF0600B6BE8B0">Centre for Marine and Coastal Studies, Universiti Sains Malaysia, 11800, Minden Penang, Malaysia & Marine Science Laboratory, School of Biological Sciences, Universiti Sains Malaysia, 11800, Minden Penang, Malaysia</mods:affiliation>
<taxonomicNameid="CEDF37D2E9A3EA3693A5AF9409AE0ED2"ID-CoL="7SNKL"LSID="DA1276D8-4DD5-5C5A-86BB-6D024EAAC1F7"authority="Dayrat & Goulding"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi"status="sp. nov.">Peronia griffithsi Dayrat & Goulding</taxonomicName>
<figureCitationid="28F17C24477F9AE4FCD3A00FCF259ED7"captionStart="Figure 39"captionStartId="F41"captionText="Figure 39. Habitats, Peronia griffithsi A Indonesia, Kei Islands, rocks behind muddy Rhizophora mangrove (st 144) B Papua New Guinea, New Ireland, mixed hard platform and seagrass bed at outlet of rivulet (st KM 05)."figureDoi="10.3897/zookeys.972.52853.figure39"httpUri="https://binary.pensoft.net/fig/458339"pageId="0"pageNumber="1">Figs 39</figureCitation>
<figureCitationid="E3B94DBED6A3CD11D625FF0C1F08EED3"captionStart="Figure 40"captionStartId="F42"captionText="Figure 40. Live animals, Peronia griffithsi, Mauritius A dorsal view, 20 mm long [3153] (MNHN-IM- 2019 - 1608) B dorsal view, 17 mm long [3155] (MNHN-IM- 2019 - 1608) C dorsal view, 8 mm long [3606] (MNHN-IM- 2019 - 1608) D holotype, ventral view, 15 mm long [3157 H] (MNHN-IM- 2000 - 35265) E ventral view, 22 mm long [3154] (MNHN-IM- 2019 - 1608)."figureDoi="10.3897/zookeys.972.52853.figure40"httpUri="https://binary.pensoft.net/fig/458340"pageId="0"pageNumber="1">, 40</figureCitation>
<figureCitationid="C78B70D4FD22F4DB15F6F1FE5995E9F6"captionStart="Figure 41"captionStartId="F43"captionText="Figure 41. Live animals, Peronia griffithsi, Indonesia, Kei Islands A dorsal view, 18 mm long [2934] (UMIZ 00177) B dorsal view, 17 mm long [2936] (UMIZ 00176) C dorsal view, 25 mm long [3566] (UMIZ 00177) D ventral view, same as A."figureDoi="10.3897/zookeys.972.52853.figure41"httpUri="https://binary.pensoft.net/fig/458341"pageId="0"pageNumber="1">, 41</figureCitation>
<figureCitationid="1310C6D469DB113D93D7E873780CD42B"captionStart="Figure 42"captionStartId="F44"captionText="Figure 42. Digestive system, dorsal view, Peronia griffithsi, with intestinal loops of type V A holotype, Mauritius [3157 H] (MNHN-IM- 2000 - 35265) B Mauritius [3153] (MNHN-IM- 2019 - 1608) C Indonesia, Kei Islands [2936] (UMIZ 00176). Scale bars: 2 mm (A-C). Abbreviations: ddg dorsal digestive gland, i intestine, pdg posterior digestive gland, st stomach."figureDoi="10.3897/zookeys.972.52853.figure42"httpUri="https://binary.pensoft.net/fig/458342"pageId="0"pageNumber="1">, 42</figureCitation>
<figureCitationid="7519A7B66C6790AF83EFE1F11F0CEC79"captionStart="Figure 43"captionStartId="F45"captionText="Figure 43. Radula, Peronia griffithsi, Indonesia, Kei Islands, [3566] (UMIZ 00177) A rachidian and innermost lateral teeth B rachidian and lateral teeth C lateral teeth D outermost lateral teeth. Scale bars: 20 μm (A), 30 μm (B), 40 μm (C), 10 μm (D)."figureDoi="10.3897/zookeys.972.52853.figure43"httpUri="https://binary.pensoft.net/fig/458343"pageId="0"pageNumber="1">, 43</figureCitation>
<figureCitationid="98DA2522EB45B4CFF428CE9DFB98180B"captionStart="Figure 44"captionStartId="F46"captionText="Figure 44. Radula, Peronia griffithsi, Mauritius A holotype [3157 H] (MNHN-IM- 2000 - 35265) B-D [3153] (MNHN-IM- 2019 - 1608) A right half rows of teeth B rachidian and lateral teeth C lateral teeth D outermost lateral teeth. Scale bars: 100 μm (A), 10 μm (B, D), 60 μm (D)."figureDoi="10.3897/zookeys.972.52853.figure44"httpUri="https://binary.pensoft.net/fig/458344"pageId="0"pageNumber="1">, 44</figureCitation>
<figureCitationid="F157637E3F68FB0BFF2BFAEFA4319064"captionStart="Figure 45"captionStartId="F47"captionText="Figure 45. Posterior, hermaphroditic (female) reproductive system, Peronia griffithsi A holotype, Mauritius [3157 H] (MNHN-IM- 2000 - 35265) B Indonesia, Kei Islands [2936] (UMIZ 00176). Scale bars: 1 mm (A), 2 mm (B). Abbreviations: dd deferent duct, fgm female gland mass, hg hermaphroditic gland, ov oviduct, rs receptaculum seminis, sp spermatheca."figureDoi="10.3897/zookeys.972.52853.figure45"httpUri="https://binary.pensoft.net/fig/458345"pageId="0"pageNumber="1">, 45</figureCitation>
<figureCitationid="E1C40077BDAE1C71AD081DAF7DC3DF87"captionStart="Figure 46"captionStartId="F48"captionText="Figure 46. Anterior, male, copulatory apparatus, Peronia griffithsi A holotype, Mauritius [3157 H] (MNHN-IM- 2000 - 35265) B Indonesia, Kei Islands [2936] (UMIZ 00176). Scale bars: 1 mm (A, B). Abbreviations: ag accessory penial gland, dd deferent duct, ms muscular sac, ps penial sheath, rm retractor muscle, v vestibule."figureDoi="10.3897/zookeys.972.52853.figure46"httpUri="https://binary.pensoft.net/fig/458346"pageId="0"pageNumber="1">, 46</figureCitation>
<figureCitationid="A372F94241000EAEAB8C7201C3AF0F0F"captionStart="Figure 48"captionStartId="F50"captionText="Figure 48. Accessory penial gland spine, Peronia griffithsi A holotype, Mauritius [3157 H] (MNHN-IM- 2000 - 35265) B Mauritius [3153] (MNHN-IM- 2019 - 1608) C Indonesia, Kei [2934] (UMIZ 00177) D same as A; E Mauritius [3156] (MNHN-IM- 2019 - 1608) F same as C. Scale bars: 100 μm (A-C), 5 μm (D-F)."figureDoi="10.3897/zookeys.972.52853.figure48"httpUri="https://binary.pensoft.net/fig/458348"pageId="0"pageNumber="1">, 48</figureCitation>
<paragraphid="406D1BDE6267571D083AB39ABF0110AF"pageId="0"pageNumber="1">Additional material examined.</paragraph>
<paragraphid="C7FEC7062F2D8000B97E9F36B7ED2499"pageId="0"pageNumber="1">Mauritius • 7 specimens 8/5 mm [3606], 20/15 mm [3153], 22/17 mm [3154], 17/15 mm [3155], 20/14 mm [3156], 7/4 mm [3607], and 6/4 mm [3608]; same collection data as for the holotype; MNHN-IM-2019-1608.</paragraph>
<figureCitationid="EF9FE0EBA2EE5AE283FF95BD088E9835"captionStart="Figure 6"captionStartId="F8"captionText="Figure 6. Geographical distribution of the Peronia species A distribution of all Peronia species except for P. peronii B distribution of P. peronii. The colors are the same as those used in Figs 2 - 5. Colored areas correspond to hypothetical geographical ranges based on confirmed records only. Distinct colors are used for each unit of P. verruculata. The distribution of P. verruculata in the Indo-West Pacific is actually continuous. However, because it is unclear which units are present in regions from where we have no fresh material of P. verruculata (red areas), no unit of P. verruculata is shown there. For P. peronii, black dots correspond to material identified based on anatomical characters and blue dots correspond to material with DNA sequences. Details on species distribution can be found in each species description."figureDoi="10.3897/zookeys.972.52853.figure6"httpUri="https://binary.pensoft.net/fig/458306"pageId="0"pageNumber="1">6</figureCitation>
<taxonomicNameid="CE050D9FB33141CD32ABAC499A7AB7E9"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<figureCitationid="7BDE1810B5DEE395C52BDB7B0D42882A"captionStart="Figure 39"captionStartId="F41"captionText="Figure 39. Habitats, Peronia griffithsi A Indonesia, Kei Islands, rocks behind muddy Rhizophora mangrove (st 144) B Papua New Guinea, New Ireland, mixed hard platform and seagrass bed at outlet of rivulet (st KM 05)."figureDoi="10.3897/zookeys.972.52853.figure39"httpUri="https://binary.pensoft.net/fig/458339"pageId="0"pageNumber="1">39</figureCitation>
<taxonomicNameid="DAACC9EC4CBB4717A508854EE01A4018"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<taxonomicNameid="7DCB45E15E4D476BC3789FC480448D7F"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<figureCitationid="6205737C7D90C14208F1D64FC1634D83"captionStart="Figure 40"captionStartId="F42"captionText="Figure 40. Live animals, Peronia griffithsi, Mauritius A dorsal view, 20 mm long [3153] (MNHN-IM- 2019 - 1608) B dorsal view, 17 mm long [3155] (MNHN-IM- 2019 - 1608) C dorsal view, 8 mm long [3606] (MNHN-IM- 2019 - 1608) D holotype, ventral view, 15 mm long [3157 H] (MNHN-IM- 2000 - 35265) E ventral view, 22 mm long [3154] (MNHN-IM- 2019 - 1608)."figureDoi="10.3897/zookeys.972.52853.figure40"httpUri="https://binary.pensoft.net/fig/458340"pageId="0"pageNumber="1">40</figureCitation>
<figureCitationid="C68A787779B3E33E9125E312FF65BEF2"captionStart="Figure 41"captionStartId="F43"captionText="Figure 41. Live animals, Peronia griffithsi, Indonesia, Kei Islands A dorsal view, 18 mm long [2934] (UMIZ 00177) B dorsal view, 17 mm long [2936] (UMIZ 00176) C dorsal view, 25 mm long [3566] (UMIZ 00177) D ventral view, same as A."figureDoi="10.3897/zookeys.972.52853.figure41"httpUri="https://binary.pensoft.net/fig/458341"pageId="0"pageNumber="1">41</figureCitation>
). No picture of live animals was available for specimens from Kavieng. The description of the color of live animals is based on Mauritius and Kei individuals. The dorsal notum is greenish brown, light to dark, mottled with darker and lighter areas. The color of the dorsal papillae varies as that of the background itself, but dorsal papillae can also be yellowish-greenish. The ventral surface (foot and hyponotum) varies from whitish to yellowish and can change rapidly in any given individual. The ocular tentacles are brown-grey, like the head. The dorsal notum of live animals is covered by dozens of papillae of various sizes. Some papillae bear black dorsal eyes at their tip. The number of papillae with dorsal eyes is variable (from 6 to 10). The largest specimens are 25 mm long.
<taxonomicNameid="DC3966070651A35B3A19EE62E5CA519A"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<taxonomicNameid="7DFE2866F644E15C1BFEB960A9812334"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<tableCitationid="93FBD4AA9E552E7F0F3DA5303A547916"captionStart="Table 5"captionStartId="T5"captionText="Table 5. Radular formulae in Peronia species. Each formula follows the same format: number of rows x number of lateral teeth per left half row - 1 (rachidian tooth) - number of lateral teeth per right half row. Each DNA extraction number corresponds to one individual. The letter H next to an extraction number indicates a holotype."httpUri="http://table.plazi.org/id/F6A603768F442AB5CA201CDCEC46CA6A"pageId="0"pageNumber="1"tableUuid="F6A603768F442AB5CA201CDCEC46CA6A">5</tableCitation>
<taxonomicNameid="BF2E37C149C0AF113CFD18F47B501D50"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<taxonomicNameid="9DBFB441A2BE5875F481A90AF5ED5271"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<taxonomicNameid="B76F3FEEB116F03B2B803BBE4F6111EB"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<figureCitationid="EEE7B7D05981341282FF7249E4F851DE"captionStart="Figure 48"captionStartId="F50"captionText="Figure 48. Accessory penial gland spine, Peronia griffithsi A holotype, Mauritius [3157 H] (MNHN-IM- 2000 - 35265) B Mauritius [3153] (MNHN-IM- 2019 - 1608) C Indonesia, Kei [2934] (UMIZ 00177) D same as A; E Mauritius [3156] (MNHN-IM- 2019 - 1608) F same as C. Scale bars: 100 μm (A-C), 5 μm (D-F)."figureDoi="10.3897/zookeys.972.52853.figure48"httpUri="https://binary.pensoft.net/fig/458348"pageId="0"pageNumber="1">48</figureCitation>
). In the anterior (male) parts, the muscular sac of the accessory penial gland is less than 5 mm long. The hollow spine of the accessory penial gland is narrow, elongated, and straight or slightly curved, and its shape (including at its tip) varies between individuals. Its length is 0.62 mm ([2934] UMIZ 00177) in unit Kei and ranges from 0.5 mm ([3157 H] MNHN-IM-2000-35265) to 0.61 mm ([3153] MNHN-IM-2019-1608) in Mauritius. Its diameter at the conical base ranges from 60 to 65
. The retractor muscle is shorter or longer than the penial sheath and inserts near the heart. Inside the penial sheath, the penis is a narrow, elongated, soft, hollow tube. Its distal end bears conical hooks which are less than 25
<taxonomicNameid="3161FA0120D841D1F271C6E80AF5E5AD"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<taxonomicNameid="4ED3FEDF6043B0731638358B0568AD2E"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<tableCitationid="3A1FD9F77C6F5844078D96178A31EF1D"captionStart="Table 4"captionStartId="T4"captionText="Table 4. Summary of traits that can help identify Peronia species, based exclusively on specimens examined for the present revision. Species are arranged in a phylogenetic order (Figs 2 - 4). Traits are described in detail in species descriptions."httpUri="http://table.plazi.org/id/8BDBED7FCCE9A8E1C67A716B088DDD7D"pageId="0"pageNumber="1"tableUuid="8BDBED7FCCE9A8E1C67A716B088DDD7D">4</tableCitation>
<taxonomicNameid="DF5EF70A43C85401C6C2C8295FDC0BC1"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
is characterized by a unique combination of anatomical traits: intestinal loops of type V, muscular sac of the accessory penial gland less than 5 mm long, spine of the accessory penial gland less than 0.62 mm long. In
A new species name is needed because no existing name applies to the species described here. A large population (161 specimens) from Kei Islands and identified by Hoffmann as
because their intestinal loops are of type V (only a few individuals were dissected). Also, the retractor muscle of the few individuals dissected inserts near the end of the visceral cavity, as in specimens from Mauritius, suggesting that an insertion near the heart is not as common. Interestingly,
<taxonomicNameid="4FD544C2A4CE8478E46C527CD4803A78"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">
<taxonomicNameid="613ED695750C77023627C4B38C38FDDF"authorityName="Dayrat & Goulding"authorityYear="2020"class="Gastropoda"family="Onchidiidae"genus="Peronia"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Peronia griffithsi"order="Systellommatophora"pageId="0"pageNumber="1"phylum="Mollusca"rank="species"species="griffithsi">