identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03D887F6FFE9FFFFFE209B37FD5FB753.text	03D887F6FFE9FFFFFE209B37FD5FB753.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onchidella J. E. Gray 1850	<div><p>Onchidella J.E. Gray</p> <p>Onchidella J.E. Gray in M.E. Gray, 1850.</p> <p>Arctonchis Dall, 1910 (a section name).</p> <p>Occidentella Hoffmann, 1929.</p> <p>Peroniella Starobogatov, 1976.</p> <p>For more information, see Dayrat 2009.</p> <p>Type species</p> <p>Onchidella nigricans (Quoy and Gaimard, 1832), by subsequent designation (Fischer and Crosse 1878). Dall (1910) thought that Hermannsen (1852) had subsequently designated the type species. However, Hermannsen only listed O. nigricans as one of the species belonging to the genus, whereas Fischer and Crosse unambiguously cited O. nigricans as the type species.</p> <p>Remarks</p> <p>The generic name Oncidiella Fischer and Crosse, 1878 is an unjustified emendation of Onchidella.</p> <p>Diagnosis</p> <p>Dorsal notum without branchial gills. Dorsal papillae with no dorsal eyes. Two oral lobes not fused medially. Hyponotal line present. Male genital opening on right lateral side of right oral lobe. Intestine usually of type IV. Reno-pulmonary complex bilaterally symmetrical. Marginal glands in body wall. Visceral cavity not divided longitudinally by a septum. Accessory gland in the penial complex absent. Rectal gland absent.</p> <p>Remarks</p> <p>Given that the onchidiid supra-specific phylogenetic relationships are still unknown (Dayrat 2009; see Discussion), it is difficult to delineate and define Onchidella so that it could correspond to a clade (although the identification of Onchidella has remained much less problematic than for other onchidiid taxa, especially Onchidium). In particular, it is difficult to evaluate whether Hoffmannola is clustered within Onchidella, or if both taxa are separate. At this stage, it also is unclear whether the characters that have been traditionally accepted for Onchidella (e.g. Labbé 1934; Britton 1984) and are provided in the above diagnosis are synapomorphies, autapomorphies, or symplesiomorphies. However, the characters provided in the diagnosis above are shared by Onchidella nigricans (Quoy and Gaimard, 1832), the type species of the genus, as well as all valid species described below, which one can thus reasonably continue to classify in Onchidella, at least for the time being.</p> </div>	https://treatment.plazi.org/id/03D887F6FFE9FFFFFE209B37FD5FB753	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFE8FFE2FE249CF7FED6B556.text	03D887F6FFE8FFE2FE249CF7FED6B556.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onchidella binneyi Stearns 1894	<div><p>Onchidella binneyi Stearns, 1894</p> <p>(Figures 2–10)</p> <p>Onchidella binneyi Stearns 1894: 342–343, plate L, figs 1–2. — Watson 1925: 301. — Marcus and Marcus 1967: 227–230, fig. 83.</p> <p>Onchidella carpenteri. – Stearns 1879: 399–401, pl. VII, figs 7–8. — Semper 1885: 281–282, plate XXI, figs 14, 25–26. — Binney 1890: 224, pl. VI, figs D–E, [not carpenteri Binney, 1861].</p> <p>Onchidium carpenteri. – Semper 1885: 281–282, plate XXI, figs 14, 25–26, [not carpenteri Binney, 1861].</p> <p>Onchidella hildae Hoffmann 1928: 35–38, 69, 94, pl. 2, figs 1–2 [as Oncidiella]. — Labbé 1934: 240. — Marcus and Marcus 1967: 230–231, figs 84–86. — Marcus and Marcus 1970: 214. syn. nov.</p> <p>Type material Onchidella binneyi Stearns, 1894</p> <p>Two syntypes (NMNH 58824): two specimens 17/12 and 14/ 9 mm preserved, leg. Fisher, [no collecting date]. Type locality: not indicated on the current label, but indicated in the original description as “ San Francisquita Bay, Los Animas Bay, and Angeles Bay in the Gulf of California ”. Type material condition: Stearns mentions “several specimens” but only two are currently held at the NMNH. Both syntypes are still entire but they were probably dried for a while because the mantle is very hard and dark brown. Both syntypes are very poorly preserved, at least externally, and were not dissected for the present study.</p> <p>Type material (Onchidella hildae Hoffmann, 1928)</p> <p>The type material includes a total of 41 syntypes. Twenty-nine syntypes (SMNH type-957): 29 specimens 12/12 to 6/ 6 mm preserved, leg. Eugenie Expedition station 656-9, 1851–1853. Eleven syntypes (SMNH type-7521): 11 specimens 15/11 to 9/ 9 mm preserved, leg. Eugenie Expedition station 766(a), 1851–1853. One syntype (SMNH type-7522): 1 specimen 7/ 4 mm preserved, leg. Eugenie Expedition station, 20 March 1852. Type locality: Insel Puna, Ecuador (SMNH 957) and Panama [Pacific Ocean] (SMNH 7521 and 7522). Type material condition: specimens are well-preserved (SMNH 957, 7521 and 7522). Four specimens were dissected prior to the present study, likely by Hoffmann: two from Ecuador (SMNH 957), and two from Panama (SMNH 7521 and 7522). Six syntypes were dissected here, four from Ecuador and two from Panama: 12/12 (#1), 11/9 (#3), 10/12 (#4) and 6/6 (#2) from Ecuador (SMNH 957); 14/11 (#1) and 12/11(#2) from Panama (SMNH 7521).</p> <p>Remarks on the original description (Onchidella binneyi)</p> <p>Stearns’s original description contains observations on the external morphology as well as the natural history. However, the internal anatomy is not described.</p> <p>Remarks on the original description (Onchidella hildae)</p> <p>Hoffmann’s original description of O. hildae is quite detailed, illustrated with several figures. Hoffmann (1928) even included a schematic diagram of the posterior reproductive organs which is quite accurate (e.g. he did not miss the distal, vaginal, accessory gland, or oviduktdrüse). Although Hoffmann had access to 40 syntypes from two localities, he only dissected four of them (two from Ecuador and two from Panama), which did not allow him to address individual variation of some key features, such as the shape of the penial caecum, which Hoffmann called the blindsack and which he thought was the main difference between O. binneyi and O. hildae.</p> <p>Distribution</p> <p>From the Gulf of California to Ecuador: Gulf of California (type locality of O. binneyi; Stearns 1879, 1894; Semper 1885; Binney 1890; Marcus and Marcus 1967, 1970; present study), Nicaragua (present study), Costa Rica (present study), Panama (type locality of O. hildae; present study), Colombia (present study), Ecuador (type locality of O. hildae; present study).</p> <p>Habitat</p> <p>Intertidal, on rocks.</p> <p>Additional material examined and dissected</p> <p>A total of 319 non-type specimens were examined, 68 of which were dissected to study their internal anatomy. Those specimens were collected from 33 localities from 2.3 ◦ S on the Pacific coast of Ecuador to the northern end of the Gulf of California. ECUADOR, South Pacific Ocean, Punta Carnero, 4 March 1967, seven specimens 8/7 to 2/ 1 mm preserved [leg. D. Kirtley], identified as Onchidium sp. [unknown identifier], (NMNH 706696) [two specimens dissected: 8/7 (#1) and 2.5/2.5 (#2)]; COLOMBIA, [Pacific Ocean], El Morro, Tumaco, 29 August 1948, 25 specimens 16/20 to 9/ 11 mm preserved [leg. E.M. Poulsen], identified as Onchidiidae [unknown identifier], (ZMUC) [five specimens dissected: 16/20 (#1), 16/20 (#2), 13/15 (#3), 10/12 (#5) and 9/11 (#4)]; PANAMA, North Pacific coast, Gulf of Panama, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.01967&amp;materialsCitation.latitude=8.572667" title="Search Plazi for locations around (long -79.01967/lat 8.572667)">Perlas Archipelago</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.01967&amp;materialsCitation.latitude=8.572667" title="Search Plazi for locations around (long -79.01967/lat 8.572667)">Pajaro Island</a>, NE cove, 8 ◦ 34.36’ N, 79 ◦ 01.18’ W, 1 May 1971, one specimen 7/ 7 mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 733425) [not dissected]; North Pacific coast, Gulf of Panama, Panama, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.02067&amp;materialsCitation.latitude=8.589334" title="Search Plazi for locations around (long -79.02067/lat 8.589334)">Perlas Archipelago</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.02067&amp;materialsCitation.latitude=8.589334" title="Search Plazi for locations around (long -79.02067/lat 8.589334)">Chapera Island</a>, small cove N of E point, 8 ◦ 35.36’ N, 79 ◦ 01.24’ W, 1 May 1971, five specimens 7/7 to 1.5/1.5 mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 733414) [two specimens dissected: 7/7 (#1) and 3/3 (#2)]; North Pacific coast, Gulf of Panama, Panama, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.53433&amp;materialsCitation.latitude=8.918167" title="Search Plazi for locations around (long -79.53433/lat 8.918167)">Canal Zone</a>, Panama <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.53433&amp;materialsCitation.latitude=8.918167" title="Search Plazi for locations around (long -79.53433/lat 8.918167)">Bay</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.53433&amp;materialsCitation.latitude=8.918167" title="Search Plazi for locations around (long -79.53433/lat 8.918167)">Culebra Island</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.53433&amp;materialsCitation.latitude=8.918167" title="Search Plazi for locations around (long -79.53433/lat 8.918167)">Fort Amador</a>, Causeway, E side of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.53433&amp;materialsCitation.latitude=8.918167" title="Search Plazi for locations around (long -79.53433/lat 8.918167)">Naos Island</a>, 8 ◦ 55.09’ N, 79 ◦ 32.06’ W, 28 April 1971, one specimen 10/ 10 mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 733262) [not dissected]; North Pacific coast, Gulf of Panama, Panama, Canal Zone, Fort Amador, NW end of Culebra Island, 28 April 1971, nine specimens 13/16 to 3/ 4 mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 733240) [two specimens dissected: 13/16 (#1) and 5/6 (#2)]; North Pacific coast, Gulf of Panama, Panama, Panama Bay, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.51783&amp;materialsCitation.latitude=8.808666" title="Search Plazi for locations around (long -79.51783/lat 8.808666)">Taboguilla Island</a>, N shore, 8 ◦ 48.52’ N, 79 ◦ 31.07’ W, 7 November 1971, one specimen 8/ 9 mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 733282) [not dissected]; North Pacific coast, Gulf of Panama, Panama, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.01967&amp;materialsCitation.latitude=8.572667" title="Search Plazi for locations around (long -79.01967/lat 8.572667)">Perlas Archipelago</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.01967&amp;materialsCitation.latitude=8.572667" title="Search Plazi for locations around (long -79.01967/lat 8.572667)">Pajaro Island</a>, NE cove, 8 ◦ 34.36’ N, 79 ◦ 01.18’ W, 7 November 1971, four specimens 8/10 (#1) to 6/7 (#2) mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 733459); North Pacific coast, Gulf of Panama, Panama, Canal Zone, Perico Island, SE side of island, 12 April 1972, 11 specimens 12/10 (#1) to 5/5 (#2) mm preserved [leg. STRI Survey], identified as Onchidium sp. [unknown identifier], (NMNH 734256); Panama, Pacific coast, Bahia de Panama, Taboga Island, 2 km SE of “Hotel Taboga”, 20 February 1975, three specimens 13/11 (#1) to 6/6 (#2) mm preserved [leg. A.J. Ferreira], identified as Onchidella binneyi by A.J. Ferreira, (CASIZ 001430); Panama, Pacific coast, Gulf of Panama, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.03333&amp;materialsCitation.latitude=8.633333" title="Search Plazi for locations around (long -79.03333/lat 8.633333)">Archipelago</a> de las <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.03333&amp;materialsCitation.latitude=8.633333" title="Search Plazi for locations around (long -79.03333/lat 8.633333)">Perlas</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-79.03333&amp;materialsCitation.latitude=8.633333" title="Search Plazi for locations around (long -79.03333/lat 8.633333)">Contadora Island</a>, N side, 08 ◦ 38’ N, 79 ◦ 02’ W, 17–19 February 1975, one specimen 7/ 8 mm preserved [leg. A.J. Ferreira], identified as Onchidiidae by T. M. Gosliner, (CASIZ 071739) [not dissected]; COSTA RICA, SW corner of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-85.71917&amp;materialsCitation.latitude=10.551666" title="Search Plazi for locations around (long -85.71917/lat 10.551666)">Bahia del Coco</a>, 10 ◦ 33.10’ N, 85 ◦ 43.15’ W, 23 April 1982, nine specimens 11/12 (#1) to 7/7 (#2) mm preserved [leg. V. Maes and C. Skoglund], identified as Onchidella sp. [unknown identifier], (ANSP A9852); NICARAGUA, Pacific coast, Dept of Leon, immediately south of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-86.75&amp;materialsCitation.latitude=12.166667" title="Search Plazi for locations around (long -86.75/lat 12.166667)">Puerto Somozo</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-86.75&amp;materialsCitation.latitude=12.166667" title="Search Plazi for locations around (long -86.75/lat 12.166667)">El Velero</a>, 12 ◦ 10’ N, 86 ◦ 45’ W, 22 January 1974, one specimen 10/ 7 mm preserved [leg. A.J. Ferreira], identified as Onchidella hildae by A.J. Ferreira, (CASIZ 001445) [dissected]; MEXICO, Baja California Norte [north], Gulf of California, Bahia de Los Angeles, 7 June 1974, four specimens 20/17 (#1) to 15/ 13 mm preserved, leg. A.J. Ferreira, identified as Onchidella binneyi by A.J. Ferreira, (CASIZ 078717); North Pacific Ocean, Gulf of California, Mexico, Sonora, San Augustin, 12 November 1966, five specimens 9/10 (#1) to 8/ 7 mm preserved, leg. P. Pickens, identified as Onchidella hildae, [identifier unknown], (NMNH 753652) [those specimens are likely vouchers of Marcus and Marcus (1970)]; Mexico, Gulf of California, Sonora, Bahia San Carlos, 30–31 March 1940, 43 specimens 8/7 (#1) to 5/4 (#2) mm preserved [leg. unknown], identified as Onchidella binneyi by A.G. Smith, (CASIZ 078725) [very poorly preserved internally]; Mexico, Gulf of California, Sonora, Puerto Penasco, in front of desalting plant, 5 August 1964, 11 specimens 8/5 (#1) to 5/4 (#2) mm preserved [leg. R. Ohmart and R. Crossin], identified as Onchidella hildae by A.G. Smith, (CASIZ 078739) [very poorly preserved internally]; Mexico, Baja California Norte [north], Gulf of California, Isla San Lorenzo del Norte Isla Partida de Lorenzo, 29 April 1966, six specimens 13/13 (#1) to 6/6 (#2) mm preserved [leg. D. Chivers], identified as Onchidella binneyi by A.G. Smith, (CASIZ 078724); Mexico, Baja California, [no collecting date], 23 specimens 16/12 (#1), 10/7 (#4), 7/6 (#3), to 5/4 (#4) mm preserved [leg. H. Hemphill], identified as Onchidella carpenteri by A.G. Smith, (CASIZ 021518); Mexico, Gulf of California, Sonora, Puerto Penasco, Norse Beach, 6–9 June 1963, 16 specimens 16/13 to 6/ 6 mm preserved [leg. E. Coan], identified as Onchidella binneyi, [identifier unknown], (CASIZ 078721) [two specimens dissected: 9/6 (#1) and 11/7 (#2) mm]; Mexico, Baja California Norte [north], Gulf of California, Sonora, Puerto Refugio, NW end of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-113.416664&amp;materialsCitation.latitude=29.333334" title="Search Plazi for locations around (long -113.416664/lat 29.333334)">Isla Angel de la Guarda</a>, 29 ◦ 20’ N, 113 ◦ 25’ W, 23 April 1953, 22 specimens 24/17 to 16/ 14 mm preserved [leg. J.W. Sefton Jr], identified as Onchidiidae by T. M. Gosliner, (CASIZ 082076) [10 specimens dissected: 24/17 (#4), 23/19 (#10), 22/17 (#3), 21/20 (#6), 21/15 (#1), 21/17 (#7), 21/19 (#8), 19/15 (#5), 16/14 (#2), and 16/14 (#9) mm]; North Pacific Ocean, Gulf of California, Mexico, Sonora, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-113.583336&amp;materialsCitation.latitude=31.333334" title="Search Plazi for locations around (long -113.583336/lat 31.333334)">Puerto Penasco</a>, 31 ◦ 20’ N, 113 ◦ 35’ W, 7 May 1966, 12 specimens 23/15 to 15/ 10 mm preserved [leg. P. Pickens], identified as Onchidella binneyi [unknown identifier], (NMNH 753656) [two specimens dissected: 23/15 (#1) and 18/15 (#2)]; North Pacific Ocean, Gulf of California, Mexico, Sonora, San Francisquito Bay, 9 April 1911, 23 specimens 16/12 (#1) to 7/5 (#2) mm preserved [leg. unknown], identified as Onchidiidae [unknown identifier], (NMNH 805160); North Pacific Ocean, Gulf of California, Mexico, Sonora, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-112.36667&amp;materialsCitation.latitude=28.75" title="Search Plazi for locations around (long -112.36667/lat 28.75)">Tiburon Island</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-112.36667&amp;materialsCitation.latitude=28.75" title="Search Plazi for locations around (long -112.36667/lat 28.75)">Red Bluff Bay</a>, 28 ◦ 45’ N, 112 ◦ 22’ W, 6 April 1934, four specimens 14/8 (#1) to 5/5 (#2) mm preserved [leg. F. Lewis], identified as Onchidium carpenteri [unknown identifier], (NMNH 805170); North Pacific Ocean, Gulf of California, Mexico, Sonora, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-113.583336&amp;materialsCitation.latitude=31.333334" title="Search Plazi for locations around (long -113.583336/lat 31.333334)">Puerto Penasco</a>, 31 ◦ 20’ N, 113 ◦ 35’ W, 2 April 1966, one specimen ∼15/ 10 mm preserved [leg. P. Pickens], identified as Onchidella hildae [unknown identifier], (NMNH 753654) [specimen in pieces, dissected prior to the present study, most internal organs missing; specimen is very likely a voucher of Marcus and Marcus (1967)]; Mexico, Baja California Norte [north], Gulf of California, Bahia San Francisquito, in inner bay, about halfway down its eastern side, 8 June 1973, 22 specimens 17/15 to 8/ 7 mm preserved [leg. A.J. Ferreira], identified as Onchidella hildae by A.J. Ferreira, (CASIZ 010107) [four specimens dissected: 17/15 (#1), 12/11 (#4), 11/10 (#2), and 8/7 (#3) mm]; Mexico, Baja California Norte [north], Gulf of California, Puertecitos, 1957, eight specimens 15/10 to 8/ 5 mm preserved [leg. E.P. Chase], identified as Onchidella binneyi by A.G. Smith, (CASIZ 078719) [two specimens dissected: 11/9 (#2) and 10/8 (#1) mm]; Mexico, Baja California Norte [north], Bahia de Las Animas, 23 December 1977, 32 specimens 11/9 to 9/ 7 mm preserved [leg. Baumback], identified as Onchidella by E. Naranjo-Garcia, (CASIZ 080045) [three specimens dissected: 11/9 (#1), 11/9 (#3) and 10/8 (#2) mm]; Mexico, Baja California Norte [north], Gulf of California, Bahia de Los Angeles, 1 April 1940, four specimens 16/14 (#1) to 15/10 (#2) mm preserved [leg. E.F. Ricketts], identified as Hoffmannola lesliei by A.G. Smith, (CASIZ 021492); North Pacific Ocean, Gulf of California, Mexico, Baja California, Cape Pulmo, 26 December 1966, one specimen ∼8/ 6 mm preserved [leg. P. Vreeland], identified as Onchidella hildae [unknown identifier], (NMNH 753655) [specimen in pieces, dissected prior to the present study, most internal organs missing; this specimen is likely a voucher of Marcus and Marcus (1970)]; Mexico, Sinaloa [Pacific Coast], Mazatlan, 14 October 1975, four specimens 9/9 (#1) to 8/ 8 mm preserved [leg. D. Summers], identified as Onchidium sp. [unknown identifier], (FMNH 195976); Mexico, Nayarit [Pacific Coast], Manzanilla, 10 miles north of Puerto Vallarta, March 1971, four specimens 11/4 (#1), 8/7 (#2), 8/8 (#3) and 6/ 6 mm preserved, leg. A.J. Ferreira, identified as Onchidella by A.J. Ferreira, (CASIZ 006137); Mexico, Baja California Sur [south], Gulf of California, about four miles NE of La Paz, January 1959, 22 specimens 20/16 to 8/ 9 mm preserved [leg. A.G. Smith], identified as Onchidella binneyi [unknown identifier], (CASIZ 078722) [four specimens dissected: 20/16 (#1), 12/11 (#2), 10/8 (#3), and 8/9 (#4) mm]; Mexico, Baja California Sur [south], Gulf of California, Isla Coronados, 5 November 1973, six specimens 7/5 (#1) to 4/4 (#2) mm preserved, leg. A.J. Ferreira, identified as Onchidella by A.G. Smith, (CASIZ 078710).</p> <p>Description of new specimens</p> <p>The description below is based on the non-type specimens examined and dissected as well as the type material, especially the six syntypes of O. hildae dissected for the present study.</p> <p>External Morphology (Figures 2 and 3). Background colour of dorsal notum of live animals brown to black-brown; papillae covering notum slightly lighter. In preserved animals, dorsal colour from homogeneously white to dark brownish grey. Hyponotum and pedal sole whitish tan both in live and preserved animals. Dorsal colour of preserved animals from tan to reddish brown or dark brownish grey. In most individuals, centre of notum lighter than its side.</p> <p>Maximum size observed in preserved specimens 24 mm in length by 17 mm in width (CASIZ 082076). All specimens outside Gulf of California less than 13 mm long, with the exception of a few specimens 16 mm long (Colombia, ZMUC, no catalogue number). Smallest specimens 2 mm long. Body high, not flattened. Hyponotum horizontal. Dorsal notum oval, i.e. longer than wide. In most individuals, notum granular, covered by papillae of various sizes (usually all &lt;1 mm high), more abundant on the side of the notum than in its centre (although papillae evenly distributed in some individuals). Generally, larger papillae surrounded by smaller ones. Size, number and distribution of the papillae on the notum likely affected by preservation. Margin of the notum not smooth, but characterized by warts of various sizes (&lt;1.5 mm long). In some individuals, larger warts (locations with repugnatory glands) regularly separated by shorter ones (from three to nine). Distinguishing marginal warts based on their size not possible because most warts look similar in many individuals. Dorsal papillae with dorsal eyes absent. Dorsal gills absent.</p> <p>In most individuals, left side of hyponotum, right side of hyponotum and pedal sole approximately same width: total width of the hyponotum (left and right sides) greater than pedal sole. Occasionally, total width of hyponotum same as that of pedal sole (e.g. 3/6/ 3 mm, NMNH 805160; 2/4/ 2 mm, CASIZ 021492; 2/4/ 2 mm, SMNH-type-957; 3/5/ 2 mm, NMNH 734256). On hyponotum, hyponotal line surrounds pedal sole and separates hyponotum into an inner area (close to the foot) and an outer area. Distance between hyponotal line and pedal sole from one fifth to one third of width of hyponotum (hyponotal line closer to pedal sole than to hyponotum margin) in specimens from Gulf of California. Outside Gulf of California, hyponotal line even closer to pedal sole (distance between hyponotal line and pedal sole from one sixth to one quarter of width of hyponotum). All openings within smooth area delimited by hyponotal line (pneumostome, male opening, eye tentacles).</p> <p>Pedal sole surrounded by two grooves on left and right sides. However, left groove shallow. Peripodial groove present on right side, from posterior genital openings (female opening and anus) to opening of pedal gland in buccal area. Anus posterior, median, very close to pedal sole. Occasionally, however, slightly on right of median line (e.g. SMNH 7522, CASIZ 021492, CASIZ 021518). Posterior female genital opening very close to the anus. Position of pneumostome on hyponotum relative to notum margin and pedal sole varies among and within lots. In most individuals from Gulf of California, pneumostome at equal distance from pedal sole and notum margin; occasionally, pneumostome closer to pedal sole (e.g. CASIZ 75362, CASIZ 021492), although pneumostome sometimes closer to notum margin (e.g. CASIZ 0805170, CASIZ 021519). In specimens outside Gulf of California, position of pneumostome varies from close to pedal sole to about halfway between pedal sole and notum margin. Generally, pneumostome located slightly on right of median line, but, occasionally, median (e.g. NMNH 753656, CASIZ 021518, SMNH 957). In anterior region, head covered dorsally by notum. Head bears pair of retractile, ocular tentacles, with eyes at tip. In all preserved specimens, however, tentacles deeply retracted. Left and right oral lobes distinct, i.e. not fused medially, superior to mouth, but inferior to ocular tentacles. Opening of pedal gland median, inferior to mouth. Male genital opening on right lateral side of right oral lobe.</p> <p>Marginal glands (Figure 4). Marginal glands in body wall, around visceral cavity, and open in margin of notum. In most individuals, between seven and ten glands on each side. Glands about 1 mm in diameter. Adjacent glands separated by small gap (from 0.5 to 1.25 mm). Glands all on same frontal plane and single longitudinal row, although posterior glands slightly more ventral.</p> <p>Digestive system (Figures 4–6). Jaw-like structure present on the surface of oral tube. Thin, inconspicuous (between 300 and 500 µm long and less than 50 µm wide), whitish, and easily missed. Left and right salivary glands, heavily branched, join buccal mass dorsally, on either side of oesophagus. Radula in between two large posterolateral muscular masses. Radular maximum size 8/ 3 mm, when flattened. Each radular row with a rachidian tooth and two half rows of lateral teeth of similar size and shape (exception of first few innermost lateral teeth, smaller). Radular formulae vary a great deal among individuals (Table 1). In specimens from Gulf of California, from 63 to 108 rows, and from 38 to 200 teeth per half row. In specimens from outside Gulf of California, from 42 to 80 rows, and from 29 to 112 teeth per half row. Larger specimens generally with radulae with more rows and more teeth per half row, but no strict correlation between size of specimens and radular formula. Rachidian tooth tricuspid: median cusp always present; two lateral cusps on two lateral branches of base of rachidian tooth, may be absent (especially in small rachidian teeth). Rachidian teeth tend to be smaller than lateral teeth: total length of base of rachidian tooth usually &lt;50 µm, and median cusp usually &lt;20 µm in length. Posterior-lateral aspect of base of rachidian teeth concave. Half rows of lateral teeth at an angle of approximately 45 ◦ to rachidian axis. Lateral teeth characterized by base, attached to radular membrane, with a dorsal, strong, flattened hook forming a 45 ◦ angle with radular membrane; triangular structure in between base and hook supports hook but does not reach its tip. In dorsal view, unless teeth misplaced, one can only see dorsal hooks: length of hook &lt;80 µm but on average about 50 µm. Smaller (&lt;20 µm) pointed cusp on outer, lateral expansion of base. In most cases, lateral cusp not observed on SEM picture because hidden below hook of adjacent, outer tooth; however, lateral cusps conspicuous when teeth are not too close (such as in innermost and outermost regions). With exception of, sometimes, first two to five innermost lateral teeth, size of lateral teeth even across half row, and among half rows. Shape of tip of hook from clearly truncate</p> <p>of California; (C) buccal mass, nervous system, and male copulatory organ, ZMUC, no catalogue number #1, Colombia; (D) visceral cavity, dorsalview, syntype SMNH 957 #1, Ecuador; (E) stomach, dorsal view, CASIZ 021518 #4, Gulf of California. Scale bar: A, 250 µm; B, 3 mm; C, 1.3 mm; D, 2 mm; E, 1.3 mm. Abbreviations: au, auricle; bm, buccal mass; cr, crop; dd, deferent duct; dg, digestive gland; e, oesophagus; i, intestine; mgg, marginal gland; ps, penial sheath; rm, retractor muscle; rps, reproductive system; st, stomach; st1, stomach chamber #1; st2, stomach chamber #2; st3, stomach chamber #3; st4, stomach chamber #4; v, ventricle.</p> <p>to clearly rounded; shape of tip of hook varies among individuals and between rows of same radula (e.g. hooks in oldest rows eroded, which give teeth an artificially different shape). Hooks not straight (slightly curved inward). Finally, although half rows at 45 ◦ angle with rachidian axis, hooks almost parallel to the rachidian axis.</p> <p>Oesophagus narrow and straight at its proximal end but enlarged into wide crop before stomach. Oesophagus enters stomach anteriorly, close to connection of stomach with dorsal and lateral lobes of digestive gland. Only small portion of posterior aspect of stomach seen in dorsal view, because largely covered by three lobes of digestive gland: dorsal lobe mainly located on right aspect of visceral mass; left, lateral lobe mainly ventral; posterior lobe covers posterior aspect of visceral mass. Stomach U-shaped sac divided into three chambers: first chamber (receives oesophagus) delimited by thin layer of tissue, and receives ducts of dorsal and left lateral lobes of digestive gland; second chamber, posterior, delimited by thick muscular tissue and receives duct of posterior lobe of digestive gland; third chamber delimited by thin layer of tissue; in most individuals, third chamber funnel-shaped. Short pouch in proximal region of intestine, just distal to third chamber, regarded by some authors as fourth stomach chamber. No strong ridges found on internal surface of stomach. Intestine long, narrow, of type IV (Labbé, 1934). No rectal gland.</p> <p>Nervous system (Figure 7). Central nervous system densely compact, often embedded and protected within layer of connective tissue. Circum-oesophageal nerve ring post-pharyngeal. Two cerebral ganglia not fused, separated by short commissure. Pleural and pedal ganglia distinct. Visceral ganglion not median, located slightly on left side of visceral loop. Cerebro-pleural and pleuro-pedal connectives very short: pleural, cerebral, and pedal ganglia touch each other. Nerves from cerebral ganglia innervate buccal area and ocular tentacles, and male anterior genital organs (penial complex) on right side. Nerves from pedal ganglia innervate foot. Nerves from pleural ganglia innervate lateral and dorsal regions of mantle. Nerves from visceral ganglia innervate visceral organs.</p> <p>Pallial complex (Figure 4). Heart enclosed in pericardium, in posterior half of right side of visceral cavity. Large, anterior ventricle becomes large aorta branching into smaller vessels delivering blood to visceral organs. Posterior auricle significantly smaller than the ventricle. Pericardium communicates through small hole with right portion of renal-pulmonary complex. Kidney intricately attached to pulmonary cavity and forms two symmetrical, left and right parts. Pulmonary cavity characterized by complex folds of internal lining (likely to increase tissue surface participating in gas exchange).</p> <p>Reproductive system: posterior parts (Figure 8). Hermaphroditic gland forms single mass subdivided into acini. Hermaphroditic duct, highly coiled, conveys gametes (eggs and autosperm) up to spermoviduct. Small branch of hermaphroditic duct goes directly to female gland mass. Another branch of hermaphroditic duct leads to small, coiled, finger-shaped pouch, the receptaculum seminis (= vesicula seminalis). From receptaculum seminis, duct goes back toward female gland mass. Latter includes mucous and albumen glands with ducts that open near where hermaphroditic duct becomes spermoviduct. Proximally, spermoviduct embedded within the female gland mass and not divided externally. Prostate not distinct externally, may be located within walls of the spermoviduct or vas deferens. Distally, male and female ducts separate: vas deferens conveys autosperm up to cephalic region; free oviduct conveys eggs up to female opening and exosperm from female opening up to fertilization chamber, near proximal end of spermoviduct. Distally, oviduct becomes vagina. Spherical spermatheca (which stores exosperm) connects to distal region of oviduct through short duct. Accessory vaginal gland (long and highly coiled to short and straight) opens into distal portion of the oviduct.</p> <p>Reproductive system: male, anterior parts (Figures 9 and 10). Male anterior organs only include penial complex (penial sheath, deferent duct, retractor muscle) because accessory penial gland absent. No distinct penis inside penial sheath. No conspicuous, solid papilla at distal end of deferens duct. Penial sheath not covered internally by folds. In most individuals, diameter of penial sheath about 500 µm. Exceptionally, in large specimens (&gt; 20 mm long) from Gulf of California, diameter of penial sheath up to 2 mm. Length of penial sheath depends on sexual maturity and size of individual. In most specimens, penial sheath less than 2 mm long. In immature specimens or even specimens not fully mature, penial sheath usually not longer than buccal mass, although, in some cases, almost as long as visceral cavity: penial sheath 1.5 mm in length in 2.5 mm-long specimen from Ecuador (NMNH 706696, #2). However, in largest specimens from Gulf of California (e.g. NMNH 753656 #1) or Colombia (e.g. ZMUC, #1) penial sheath up to twice as long as buccal mass and up to 8 mm long.</p> <p>(Continued)</p> <p>Caecum, digitiform pouch, joins penial sheath laterally. Caecum straight or curved (varies within lots). Filled by fewer than 10 small concretions that could be seen due to transparency of caecum. Function of concretions unknown. Caecum usually present and occasionally absent (but often absent in specimens from Panama). Presence /absence of caecum varied within and among lots, and not strictly correlated with sexual maturity: for instance, caecum found in male organs of two immature (no posterior female organs) specimens from Gulf of California (CASIZ 021518 #2 and #3), but caecum absent from fully mature specimen (#4) from same lot. In Panama, most specimens dissected lacked caecum, although it was occasionally present (NMNH 733882). In specimens from Panama lacking caecum, concretions could be seen within lumen of penial sheath, by transparency.</p> <p>Deferent duct straight to loosely convoluted (up to five loops) from where it enters visceral cavity to where it joins penial sheath. Number of loops varied among individuals, even within same lot. Deferent duct with fewer loops in small but mature specimens (with all female organs developed). Immature specimens (with no or poorly-developed female organs) have short, straight deferent duct. Exceptionally, deferent duct found extremely convoluted, but only in two specimens from two different lots while regular deferent duct found in all other specimens from same lots (NMNH 734256 #2, Panama; NMNH 706696 #1, Ecuador). Variation regarding deferent duct observed consistently within each geographical area represented in samples. However, deferent duct, straight, nearly straight, or with just one or two loose loops in specimens from Panama. Diameter of deferent duct more or less constant along its whole length, about 200 µm maximum.</p> <p>Retractor muscle anchors on penial sheath near where deferent duct enters penial sheath and inserts at end of second posterior third of floor of visceral cavity. In small, immature specimens with small penial sheath, retractor muscle can be quite weak, even occasionally absent (e.g. NMNH 733425, Panama).</p> <p>Discussion</p> <p>A synonymy between O. binneyi (type locality: Gulf of Mexico) and O. hildae (type localities: Ecuador and Panama) has never been proposed, mainly because few specimens from few localities have been studied and compared so far. All descriptions of O. binneyi (as O. carpenteri or O. binneyi) by Stearns (1879, 1894), Semper (1885) and Binney (1890) were based on the material collected by W.J. Fisher in the Gulf of California: Stearns (1894) mentioned he examined “several examples” (only two syntypes are now held at the NMNH) from three localities from the Gulf of California; Semper (1885) mentioned 17 individuals from one locality (Las Animas Bay); and Binney (1890) did not give any specific information on the material he studied, but he did receive it from W.J. Fisher (see Stearns 1894). As for O. hildae, Hoffmann had access to the 40 type specimens, but he only dissected four of them: two from Ecuador and two from Panama. Watson (1925) and Labbé (1934) did not study any new material. Finally, Eveline and Ernst Marcus redescribed O. hildae and O. binneyi based on 15 specimens from three localities: they identified seven specimens as O. binneyi and one specimen as O. hildae from Puerto Penasco, Gulf of California (Marcus and Marcus 1967); they also identified seven other specimens as O. hildae from two other localities from the Gulf of California (Marcus and Marcus 1970). Vouchers are available for some of the material studied by Marcus and Marcus, but those were entirely dissected and are now largely destroyed (NMNH 753652, 753654, and 753655). It is important to emphasize that Marcus and Marcus only examined material from the Gulf of California, i.e. they did not examine material of O. hildae from Panama or Ecuador. Before Marcus and Marcus (1967, 1970), it was thought that O. hildae was restricted to Panama and Ecuador, and O. binneyi was restricted to the Gulf of California. Marcus and Marcus (1967, 1970) thought that both species were found in the Gulf of California.</p> <p>The abundant material studied here offers for the first time the opportunity to address individual variation of important taxonomic characters, including in relation to geography. Indeed, a total of 319 non-type specimens were examined (68 of which were dissected), collected from 33 localities from six different countries (Ecuador, Colombia, Panama, Costa Rica, Nicaragua, Mexico), from 2.3 ◦ S’ on the Pacific coast of Ecuador to the northern end of the Gulf of California. All type material available was also examined: two syntypes of O. binneyi from the Gulf of California (not dissected here); 41 syntypes of O. hildae from Ecuador and Panama (four specimens from Ecuador and two from Panama dissected here).</p> <p>According to Hoffmann (1928), the main difference between O. binneyi and O. hildae was the shape of the penial caecum: curved and straight, respectively. After Hoffmann, the differences between O. binneyi and O. hildae were only discussed by Marcus and Marcus (1967, 1970) who added the following differences: course of the deferent duct in the body cavity (runs straight from the body wall to the penial sheath in O. binneyi; forms a free spiral in O. hildae); size of the vaginal gland (very long and coiled in O. binneyi; short and blunt in O. hildae); width of the pedal sole relative to the hyponotum (narrow in O. hildae); maximum size of the smallest papillae (60 µm in O. binneyi; 200 µm in O. hildae); position of the hyponotal line (very close to the foot in O. hildae).</p> <p>However, observations made by Marcus and Marcus were also based on few specimens dissected from few localities. Our present data show that the individual variation is much higher than expected and that it is not possible to find discrete differences between two groups of populations or individuals. The penial caecum is not either curved or straight, but its shape (which is likely affected by preservation) varies from curved to straight. The number of loops, the arrangement and the length of the deferent duct vary from straight and short with no loop, to highly convoluted and long. The size of the vaginal gland varies from short and slightly curved to long and highly convoluted. The width of the pedal sole may be greater than, equal to, or less than the width of the hyponotum. The size of the dorsal papillae varies a great deal (from ∼ 40 µm up to 1 mm) within the same individual. Finally, the position of the hyponotal line varies from one sixth to one third closer to the foot than the notal margin. More importantly, beyond their variation, those features are not combined in two distinct sets: a specimen can have a short and blunt vaginal gland (supposedly diagnostic of O. hildae) and a curved penial caecum (supposedly diagnostic of O. binneyi). Finally, all the specimens of O. binneyi and O. hildae seem to share a diagnostic feature that is not found in other species from the north-eastern Pacific: the presence of a penial caecum hosting some concretions; in most specimens in which the caecum is lacking, the concretions are found in the penial sheath itself.</p> <p>As a result, O. binneyi and O. hildae are regarded as two synonyms. Although only one species of Onchidella is thought to inhabit the eastern Pacific coasts from Ecuador to the Gulf of California (excluding the Galapagos), there are some variation patterns which suggest that there might be more than one species. For instance, large specimens (&gt; 20 mm) are common in the Gulf of California, whereas the vast majority of the specimens outside the Gulf of California measure less than 13 mm long. Also, a penial caecum is only exceptionally found in specimens from Panama, whereas it is exceptionally missing in specimens from the Gulf of California. Thus, it is possible that cryptic species might be found in the future. At this stage however, given the data available, all the specimens studied here are regarded as part of one variable species.</p> <p>The fact that binneyi belongs to Onchidella has never been challenged. Semper classified binneyi in Onchidium, but he did not consider the genus Onchidella to be a valid genus and classified all Onchidella in Onchidium.</p> <p>Pepe and Pepe (1985) showed that the activities of O. binneyi are synchronous with the tides. Also, O. binneyi has been studied for its pharmacology (Abramson et al. 1989).</p> </div>	https://treatment.plazi.org/id/03D887F6FFE8FFE2FE249CF7FED6B556	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFF5FFE7FE1C9EF8FC73B7F0.text	03D887F6FFF5FFE7FE1C9EF8FC73B7F0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onchidella borealis Dall 1872	<div><p>Onchidella borealis Dall, 1872</p> <p>(Figures 11 and 12)</p> <p>Onchidella borealis Dall 1872: 135. — Binney 1876: 184–185, pl. VI, figs BB, E, EE. — Binney 1885: 162–163, figs 148–149. – von Plate 1893: 206–207 [as Oncidiella]. — Dall 1910: 112–115. — Watson 1925: 301. — Hoffmann 1928: 35. — Labbé 1934: 241. — Abbott 1954: 275. — Marcus 1961: 3–4, pl. I, figs 1–2. — Morris et al. 1980: 342. — Weiss and Wägele 1998: 77–80, figs 22–33, 47, 50–51, 61–63.</p> <p>Onchidium boreale. – Semper 1885: 282–283, pl. XXI, fig. 13.</p> <p>Onchidium borealis. – Dall 1910: 112–115.</p> <p>Type material</p> <p>Dall did not specify the number of specimens he used for the original description. Also, the type material could not be located and is very likely lost. Type locality: Sitka, Alaska Territory. Type material condition unknown.</p> <p>Remarks on the original description</p> <p>The brief original description is reproduced here in its entirety:</p> <p>Animal small, black with dots and streaks of yellowish white, foot lightcoloured, also muzzle and tentacles. Back regularly rounded, but a little pointed in the middle; smooth or very finely granulose, tough and coriaceous. Eyes globular, blue, on very short constricted tentacles. Muzzle short, rounded-transverse. Head not produced beyond the anterior edge of the mantle. Sexual appendages on the right side, near the head. Foot ovate, narrow, rather roundly pointed behind. Lon.. 3 in. Habitat, Sitka, Alaska Territory, on the rocks near tide marks, especially on the small islets in the Bay. Dall, August, 1866.</p> <p>Distribution</p> <p>Most authors have studied only some Alaskan material (type locality; Binney 1876; Semper 1885; von Plate 1893; Weiss and Wägele 1998). However, a wide distribution has been recognized for a long time: e.g. from Prince William Sound to Vancouver Island (Binney 1885); from Californa to the Bering Sea (Dall 1910); Alaska and west coast of Canada and the United States (Watson 1925); from Alaska to Oregon (Abbott 1954); from Alaska to California (Marcus 1961). The southernmost record is San Luis Obispo, California (Morris et al. 1980).</p> <p>Habitat</p> <p>Intertidal. On rocks, although they can also be found on algae (Morris et al. 1980).</p> <p>Additional material examined and dissected</p> <p>A total of 213 specimens were examined, 19 of which were dissected to study their internal anatomy. Specimens examined were collected from 11 localities from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Alaska</a> to the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">San Francisco Bay</a>, covering the entire range of the species. ALASKA, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Kodiak Island</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Three Saints Bay</a>, 57 ◦ 07.8’ N, 154 ◦ 28.7’ W, 29 May 1975, nine specimens 5/3 to 2/ 1 mm preserved [leg. Sears], identified as Onchidella borealis by <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Mueller</a>, (CASIZ 013314) [two specimens dissected: 5/3 (#1) and 3/2 (#2)]; CANADA, British Columbia, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Pacific</a> coast of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Vancouver Island</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Clayoquot Sound</a>, February 1965, four specimens 6/5 to 5/ 5 mm preserved [leg. A. G. Smith], identified as Onchidella by A.G. Smith, (CASIZ 021510) [two specimens dissected: 6/5 (#1) and 5/5 (#2)]; OREGON, Lincoln County, Road’s End, zone 2, 16 June 1962, five specimens 9/5 (#1) to 4/ 3 mm preserved [leg. Giles], identified as Onchidella carpenteri by M.G. Harasewych, (NMNH 1102498); Oregon, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Coos County</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Coos Bay</a>, February 1965, 26 specimens 7/4 to 3/ 2 mm preserved [leg. H. Hemphill], identified as Onchidella borealis by A.G. Smith, (CASIZ 021557) [two specimens dissected: 7/4 (#1), 5/3 (#2), and 6/4 (#3)]; Oregon, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">South Bay</a>, Cape Arago, 24 June 1951, one specimen 8/ 5 mm preserved [leg. J. McCauley], identified as Onchidella borealis [unknown identifier], (NMNH 574669) [not dissected]; Oregon, Cape Arago, [collecting date unknown], nine specimens from 9/5 (#1) to 5/4 (#2) mm preserved [leg. unknown], identified as Onchidium sp. [unknown identifier], (NMNH 836671); CALIFORNIA, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Del Norte County</a>, Crescent City, Battery Point Lighthouse, S of lighthouse, N of breakwater, 41.75 ◦ N, 124.20 ◦ W, 27 July 2007, two specimens from 6/3 to 5/3 (#1) mm preserved [leg. T. M. Gosliner, R. Johnson and V. Smith], identified as Onchidella borealis by T. M. Gosliner, (CASIZ 174474); California, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Mendocino County</a>, mouth of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Buckhorn Creek</a>, 1 mile S of Little River, 4 May 1946, 23 specimens from 9/5 to 4/ 3 mm preserved [leg. A.G. Smith], identified as Onchidella [unknown identifier], (CASIZ 081437) [two specimens dissected: 8/5 (#1) and 5/3 (#2)]; California, [Mendocino County], Noyo, [collecting date unknown], two specimens 4/3 and 3/ 2 mm preserved [leg. Harford], identified as Onchidella borealis [unknown identifier], (NMNH 047483) [not dissected]; California, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Humboldt County</a>, two miles S of Cape Mendocino, February 1965, 92 specimens 12/7 to 3/ 2 mm preserved [leg. G.D. Hanna and A.G. Smith], identified as Onchidella by A.G. Smith, (CASIZ 021519) [three specimens dissected: 7/5 (#1), 7/5 (#2), and 6/4 (#3)]; California, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">San Francisco Bay</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Marin County</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-124.2&amp;materialsCitation.latitude=41.75" title="Search Plazi for locations around (long -124.2/lat 41.75)">Angel Island</a>, wind swept point, 1 November 1975, 40 specimens 12/7 to 7/ 5 mm preserved [leg. S. Gray], identified as Onchidella borealis by T. M. Gosliner, (CASIZ 057732) [three specimens dissected: 9/6 (#1), 9/6 (#4), 9/5 (#2), and 6/4 (#3)].</p> <p>Description of new specimens</p> <p>Several anatomical systems of O. borealis are very similar to O. binneyi (nervous system, pallial organs, digestive system except for the radular formulae, and the posterior genital organs). Their description is not provided here again. Given that the type material of O. borealis could not be located, the description below is exclusively based on the non-type specimens examined and dissected here.</p> <p>External Morphology (Figure 11). Background colour of dorsal notum of live animals quite variable: reddish brown, brown, dark grey, or black, mottled with white and yellowish dots and streaks. In preserved animals, dorsal colour from homogeneously white to dark brownish grey. Hyponotum and pedal sole whitish tan both in live and preserved animals.</p> <p>Size from 12/ 7 mm (CASIZ 021519, California) to 2/ 1 mm (CASIZ 013314, Alaska). Body high, not flattened. Hyponotum horizontal. Dorsal notum oval, longer than wide. Notum finely granular with small papillae of various sizes (usually &lt;0.7 mm high), covering mainly sides of notum; centre covered by smaller papillae, occasionally smooth. Margin of notum not smooth: characterized by warts of various sizes (&lt;1 mm long). In some individuals, larger warts (locations with repugnatory glands) regularly separated by shorter ones (from two to seven). However, distinguishing marginal warts based on their size not possible in many individuals, because most warts similar. Marginal warts hosting marginal glands more prominent in posterior region of body. Papillae with dorsal eyes absent. Dorsal gills absent.</p> <p>In most individuals, pedal sole wider than the left and right hyponotum added (H &lt;S). Exceptionally, total width of hyponotum (left and right side added) equal to the width of pedal sole (H = S). Exceptionally, left and right hyponotum added together wider than pedal sole (H&gt; S) (e.g. CASIZ 057732). On hyponotum, hyponotal line around pedal sole which separates hyponotum into inner area (close to the foot) and outer area. Distance between hyponotal line and pedal sole from one fifth to one third of width of hyponotum (hyponotal line closer to pedal sole than to hyponotum margin). Openings (pneumostome, male opening, eye tentacles) within smooth area delimited by hyponotal line.</p> <p>Pedal sole surrounded by two grooves on left and right sides. However, the left groove is shallow. Peripodial groove present on right side, from buccal area (pedal gland opening) to posterior openings (anus and female opening). Anus posterior, slightly on right of median line, very close to pedal sole. Anus occasionally median (e.g. CASIZ 081437, CASIZ 057732). Posterior female genital opening very close to anus. Position of pneumostome on hyponotum relative to notum margin and pedal sole varies among and within lots. However, pneumostome close to anus in most individuals. Generally pneumostome slightly on right side but occasionally median (e.g. NMNH 836651, CASIZ 081437). In anterior region, head covered dorsally by the notum. Head bears pair of retractile, ocular tentacles, with eyes at tip. In most specimens, tentacles deeply retracted. Left and right oral lobes distinct, i.e. not fused medially, superior to mouth and inferior to ocular tentacles. Opening of pedal gland is median, inferior to mouth. Male genital opening on right of lateral side of (or behind) right oral lobe.</p> <p>Marginal glands. Marginal glands similar to those found in O. binneyi, except between eight and twelve glands on each side.</p> <p>Digestive system (Figures 11 and 12). Radular maximum size 5/ 2 mm, when flattened. Radular formulae vary among individuals (Table 1). From 44 to 78 rows, and from 32 to 58 teeth per half row.</p> <p>Reproductive system (Figure 11). No accessory penial gland. No distinct penis inside penial sheath. No conspicuous papilla at distal end of deferens duct. Penial sheath not covered internally by thick folds. Length of penial sheath rarely&gt; 2 mm. No accessory caecum filled with white concretions opening into penial sheath. Deferent duct convoluted and coiled but loops always easy to count (fewer than 10). The diameter of deferent duct, more or less constant, is ∼ 250 µm maximum. Retractor muscle anchors on penial sheath near where deferent duct joins penial sheath and inserts on floor of visceral cavity between one third and one half of its length, just posterior to nervous system. Length of retractor muscle rarely exceeds 3 mm.</p> <p>Discussion</p> <p>Of all the species described here, O. borealis is relatively well known. Dall (1910) described in detail its habitat and colour variation. Binney (1876) illustrated accurately the radular teeth. Its anatomy was described by Marcus (1961) based on a few specimens from Oregon and California, and by Weiss and Wägele (1998) based on a few specimens from Alaska. Semper’s (1885) and von Plate’s (1893) anatomical notes are less detailed. Its dorsal colour variation and its habitat have been mentioned in field guides (e.g. Morris et al. 1980).</p> <p>The present description of O. borealis is in agreement with past descriptions. However, because it is based on a total of 213 specimens examined and 19 specimens dissected, covering the entire range of the species (Alaska to California), it provides new insight on individual variation, such as, for instance, the anterior male organs. The maximum size found here (12 mm) is the maximum size found in the literature. The range of radular formulae found here of 44/78 × (32/58–1–32/58) is compatible with the literature, although Binney (1876) found a few more teeth per half row (61– 1–61) without specifying the number of rows; Weiss and Wägele (1998) found 47/49 × (40–1–40); Marcus (1961) found 50/52 × (45/53–1–45/53). Von Plate (1893) found a number of teeth per half row that is beyond all other observations: 88-1-88, in a specimen 11 mm long (which is not longer than other specimens dissected).</p> <p>The taxonomy and nomenclature of O. borealis would be quite simple without O. carpenteri. The latter, originally described from the Strait of Juan de Fuca, i.e. within the range of O. borealis, has remained very confused in the literature because many authors have erroneously claimed that O. carpenteri had been described from the Gulf of California. Ultimately, it led authors to believe that there were two distinct, sympatric species of Onchidella from California to Alaska: for instance, Abbott (1954) mentioned O. carpenteri from Puget Sound to Lower California, and O. borealis from Alaska to Oregon. Onchidella carpenteri is regarded here as a nomen dubium although it was very likely based on young specimens of O. borealis (for more information, see the detailed discussion on O. carpenteri below).</p> <p>Binney (1876) described the radular teeth of Onchidella borealis from Alaska, but he did not discuss the possible synonymy between his O. carpenteri, which he had described in 1861, and Dall’s O. borealis, described in 1872. Later, Binney (1885) thought there were two species in the Pacific North-west, Onchidella borealis, which he mentioned from Prince William Sound (Alaska) to Vancouver Island (i.e. Juan de Fuca), and Onchidella carpenteri, from the Strait of Juan de Fuca to the Gulf of California. According to Binney, a jaw was present in O. borealis and absent in O. carpenteri. However, this difference might have simply been due to the fact that jaws are quite inconspicuous. In any case, Onchidella borealis remains the valid name because Onchidella carpenteri is a nomen dubium (see below).</p> <p>All authors have agreed that borealis belongs to Onchidella, with the exception of Semper (1885) who classified borealis in Onchidium. However, Semper (1885) only accepted two genera in Onchidiidae: Onchidina (then a monotypic genus) and Onchidium (for all other onchidiid species). In total, Semper (1885) redescribed five species that are now classified in Onchidella, and he considered them all to be in the same “group” within Onchidium. So, although he did not regard Onchidella as a valid name, he did acknowledge the close relationship of all Onchidella species.</p> <p>Onchidella borealis was designated by Dall (1910) the type species for Arctonchis, a section of a subgenus (Onchidella), of the genus Onchidium. The name Arctonchis has never, to our knowledge, been raised to the generic rank. According to Dall, this section included both Onchidella borealis and O. celtica, both characterized by a thin jaw in the oral tube. Dall’s internal classification of Onchidella has not been adopted, especially after Watson (1925) demonstrated that the main feature used by Dall (the thin jaw) to separate O. borealis and O. celtica from the rest of the Onchidella species, might actually be shared by all Onchidella species (see below, Discussion). However, the supra-specific name Arctonchis will remain available if a name is needed for a clade including O. borealis, when the phylogeny of Onchidella is known.</p> </div>	https://treatment.plazi.org/id/03D887F6FFF5FFE7FE1C9EF8FC73B7F0	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFF0FFDAFE769B5AFE35B770.text	03D887F6FFF0FFDAFE769B5AFE35B770.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onchidella carpenteri (Binney 1861)	<div><p>Onchidella carpenteri (Binney, 1861) nom. dub.</p> <p>Onchidium carpenteri Binney 1861: 154. — Tryon 1868: 317, pl. 18, fig. 39. — Binney and Bland 1869: 307–308, figs 544–545.</p> <p>Onchidella carpenteri. – Fischer and Crosse 1878: 697–698 [as Oncidiella (?) carpenteri]. — Binney 1885: 163, fig. 150.</p> <p>Type material</p> <p>The original description mentioned five syntypes: maximum size (preserved) of 5/ 3 mm, leg. and collecting date unknown. The syntypes were supposedly from the collections of the National Museum of Natural History, Washington, DC, but they could not be found there. They are likely lost. Type locality: Strait of Juan de Fuca (see Discussion). Type material condition unknown, although Binney and Bland (1869) mentioned that the syntypes were dry and destroyed.</p> <p>Remarks on the original description</p> <p>The brief original description is reproduced here in its entirety: ONCHIDIUM CARPENTERI. Among the mollusca from the Straits of De Fuca, Mr Carpenter has detected five specimens of a shelless mollusk, which evidently belong to the genus Onchidium. Being preserved in alcohol it is difficult to obtain any more satisfactory specific characters than the following: The body is oblong, with its extremities circularly rounded; the upper surface is regularly arched; below, quite near the edge, the border of the mantle is readily distinguished, most of the under surface is occupied by the broad, distinct, locomotive disk; the body is uniformly smoke-coloured; in size the individuals vary considerably, the length of the largest being 5, the extreme breadth 3 millimetres.</p> <p>Discussion</p> <p>To say the least, the case of O. carpenteri is complicated. Here it is regarded as a nomen dubium. In order to justify this nomenclatural decision, two key issues need to be addressed: (1) the number of specimens identified by Binney as O. carpenteri; (2) the type locality of O. carpenteri.</p> <p>In the brief original publication, the type locality is unambiguously cited as the Strait of Juan de Fuca, which is at the north-western border between Canada and the United States, more specifically between Vancouver Island and Washington State. Based on the original description, it might seem that P.P. Carpenter collected the specimens. However, P.P. Carpenter was actually working at the Smithsonian, and it is more likely that he studied some material available in the collections. In fact, Binney and Bland (1869) indicated that it was unknown who had sent the specimens to the Smithsonian.</p> <p>The problems start with Tryon (1868), who mentioned O. carpenteri from “Lower California ”, i.e. Baja California, without any explanation. Tryon seems to be the first author who made a mistake about the known distribution of O. carpenteri (the Strait of Juan de Fuca is not in Baja California). Maybe the mistake came from the fact that, in the same one-page article where he described carpenteri, Binney (1861) also described a new species of Pedipes from “St Lucas Peninsulae Californiae” which is, indeed, in Baja California. Another explanation is that Tryon actually had access to Binney and Bland’s (1869) manuscript because Tryon’s figure of O. carpenteri (Tryon 1868) is the same as Binney and Bland’s (1869) figure 544. Tryon could have been influenced by Binney and Bland’s (1869) mention of Cape San Lucas as a locality for O. carpenteri.</p> <p>Indeed, Binney and Bland (1869) reproduced the original description in its entirety. They did not mention any new material, but added two black-and-white drawings of the original specimens (their figures 544–545). As material available in the collections of the Smithsonian, they mentioned four specimens from “De Fuca”. However, they also indicated that their figure 545 was drawn from “one of specimens collected at Cape San Lucas”, which is in Baja California. Binney (1883) later addressed the confusion generated by the mention of Cape San Lucas for a species then supposedly only known from the frontier between Canada and the US, by admitting that “The locality, Cape San Lucas, is doubtful. It is so referred, probably by mistake...”</p> <p>Two years later, however, Binney (1885) wrote that Onchidella carpenteri was distributed from “Strait of Fuca to Gulf of California ”. It could mean one of two things. First, that Binney had changed his mind and thought that Cape San Lucas was indeed one of the type localities or even the type locality. Second, that he had received new material from the Gulf of California which he had identified as O. carpenteri. The correct answer very likely is the second one. Indeed, to our knowledge, the only specimens that had become available in the last years of the 1870s from the western coast of North America were those collected by W.J. Fisher from the Gulf of California. Stearns (1879) originally described those specimens as O. carpenteri, which explains why, in 1885, Binney could assume that his species O. carpenteri was distributed from the Strait of Juan de Fuca all the way down to the Gulf of California.</p> <p>A few years later, Binney (1890) published two new figures of an alcohol-preserved specimen of Onchidella carpenteri with no locality data. Those two drawings look completely different from the drawings published earlier by Binney (1885) for O. carpenteri. According to Stearns (1894), Binney made those two new drawings based on a specimen collected by W.J. Fisher (not P. Fischer) from the Gulf of California (and which Binney had received from Dall, who had also received some specimens from W.J. Fisher).</p> <p>Cooper (1892) mentioned some specimens of Onchidella carpenteri collected by [John] Xantus from Cape San Lucas. However, his identification was based on the assumption that O. carpenteri was “doubtfully reported from lat. 48 ◦ north ”. However, according to Binney (1861, 1883), O. carpenteri was originally described from there.</p> <p>Unfortunately, Stearns (1894) changed his mind and decided that the material collected by W.J. Fisher from the Gulf of California was a different species, mainly because O. carpenteri, for which only the five original specimens had ever been recorded, was a very tiny species (5 mm maximum in length) compared to the specimens collected from the Gulf of California (17.2/12.2 mm on average). Stearns decided to refer to those specimens as Onchidella binneyi, to honour Binney.</p> <p>No other mention of O. carpenteri found in the literature was based on new material. According to P. Fischer [not W.J. Fisher] and Crosse (Fischer and Crosse 1878, translated here): “Only one species is cited from the Pacific coast of Mexico: Oncidiella Carpenteri, W. G. Binney. ” Also, in their translation of Binney’s description of O. carpenteri, Fischer and Crosse (1878) claimed that its habitat was “Cape San Lucas, at the extremity of Lower California (P. Carpenter)”. However, Carpenter was most likely not the collector (see above) and the locality mentioned in the original description was Juan de Fuca, which is not in Lower California. Finally, the long anatomical description of Onchidella by Fischer and Crosse is based on specimens of Onchidella celtica, not O. carpenteri (for which Fischer and Crosse did not have any additional material).</p> <p>The erroneous idea that O. carpenteri was distributed in the Gulf of California was perpetuated in all subsequent literature, and, again, without new material (Watson, 1925; Hoffmann, 1928; Labbé, 1934). Hoffmann (1928) even wrote that the type locality was “Straits of De Fuca [Golf von Californien]”. Watson (1925) made the same mistake in his map of all known Onchidella species, and placed O. carpenteri off southern California (and not in the Strait of Juan de Fuca). Labbé (1934) erroneously cited O. binneyi Stearns, 1894 as a synonym of O. carpenteri Binney, 1894 (with a wrong date), but he did not examine any specimens and only mentioned it from the Gulf of California.</p> <p>Later, Abbot (1954) still regarded O. carpenteri as a valid species distributed from Puget Sound to Lower California. Marcus (1961) indicated that O. carpenteri differed from O. borealis, but could not be distinguished from O. binneyi. However, this statement was due to the fact that Marcus (1961) regarded O. carpenteri mainly as a “southern species”, and he was still referring to specimens of O. binneyi as O. carpenteri. Finally, Marcus and Marcus (1967) suggested that, perhaps, O. carpenteri (type locality Strait of Juan de Fuca) could be a synonym of O. borealis (type locality Alaska), but still thought that O. carpenteri was present in the Gulf of California.</p> <p>In summary, Binney’s type locality of O. carpenteri was the Strait of Juan de Fuca, at the border between Canada and the United States. None of the records of O. carpenteri from the Gulf of California were based on new material, but simply on an erroneous placement of the Strait of Juan de Fuca on a map. The fact that Binney and Bland (1869) listed, by mistake, Cape San Lucas (which is in the Gulf of California) as a locality for O. carpenteri did not help, although Binney (1883) clearly corrected that mistake. Finally, the identification of the material collected by W.J. Fisher from the Gulf of California as O. carpenteri was a misidentification, and all those specimens were later referred to as O. binneyi (see that species).</p> <p>For several reasons, O. carpenteri is a nomen dubium. First, the type material is lost. Second, the original description is extremely brief, largely incomplete, and no additional material was described. Third, given their size (maximum 5 mm long), those specimens were very young and likely immature. Fourth, the type locality, although clearly in the Strait of Juan de Fuca, has remained extremely confused in the literature (because authors erroneously thought that Juan de Fuca was in Baja California). In any case, there is only one species of Onchidella from California to Alaska (Onchidella borealis), and it is very likely that the description of O. carpenteri was simply based on young specimens of O. borealis.</p> </div>	https://treatment.plazi.org/id/03D887F6FFF0FFDAFE769B5AFE35B770	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFCDFFDCFE4B9CDAFD8BB21B.text	03D887F6FFCDFFDCFE4B9CDAFD8BB21B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onchidella steindachneri (Semper 1885)	<div><p>Onchidella steindachneri (Semper, 1885)</p> <p>(Figures 1, 13–18)</p> <p>Onchidium steindachneri Semper 1885: 280, plate XIX, figs 7–8, plate XXI, fig. 15, plate XXIII, fig. 14.</p> <p>Onchidella steindachneri. – Stearns 1894: 384, plate LI, figs 4–5. — Watson 1925: 304. — Hoffmann 1928: 38–44, 70, 94–95, plate 2, figs 3, 5–6, plate 3, figs 4, 7. — Labbé 1934: 241.</p> <p>Type material</p> <p>One syntype (ZMB 39.038): one specimen ∼ 25/ 10 mm preserved [leg. and collecting date unknown]. Type locality: Galapagos Inseln. Type material condition: the original description was based on two syntypes, but there seems to be only one of them held at the ZMB, which was dissected and cut in pieces prior to the present study (so the size indicated above is tentative).</p> <p>Additional material examined and dissected</p> <p>A total of 95 non-type specimens were examined, 29 of which were dissected to study their internal anatomy. Examined specimens were collected from 12 localities on six of the Galapagos Islands, four islands being new records (see Distribution). Ecuador, Galapagos Islands, Isla Cristóbal [Chatham Island], 11 May 1852, 21 specimens 28/20 to 20/ 12 mm preserved [leg. Eugenie Expedition, station 838], identified as Onchidella steindachneri [unknown identifier], (SMNH 103166) [one specimen dissected here: 25/10 (#1); the jar contains pieces of possibly four additional specimens dissected prior to the present study; those specimens are the vouchers of Hoffmann’s (1928) description of steindachneri]; Ecuador, Galapagos Islands, Santa Cruz, Academy Bay, 1934, three specimens 23/18 to 20/16 (#1) mm preserved [leg. Blomberg], identified as Onchidella steindachneri [unknown identifier], (SMNH 103167); Ecuador, Galapagos Islands, Santa Cruz, Academy Bay, 1934, two specimens 24/17 and 16/15 (#1) mm preserved [leg. Blomberg], identified as Onchidella steindachneri [unknown identifier], (SMNH 103168); south Pacific coast, Ecuador, Galapagos Islands, Sulivan Bay, James Island, 24 July 1938, one specimen 17/ 13 mm preserved [leg. unknown], identified as Onchidiidae [unknown identifier], (NMNH 574530) [dissected]; south Pacific coast, Ecuador, Galapagos Islands, Elizabeth Bay, Albemarle Island, 26 July 1938, two specimens 25/10 (#1) and 25/ 20 mm preserved [leg. unknown], identified as Onchidiidae [unknown identifier], (NMNH 574379); Galapagos Islands, W coast of Isla Santa Cruz, opposite Guy Fawkes Island, “Venice” cove, 19 February 1964, three specimens 13/7 to 10/9 (#1) mm preserved [leg. D.P. Abbott and J.W. Durham], identified as Onchidella steindachneri by B. Roth (CASIZ 017922); Galapagos Islands, Isla Santa Cruz, Academy Bay, February 1964, nine specimens 45/25 to 15/ 11 mm preserved [leg. A.G. Smith], identified as Onchidella steindachneri by A.G. Smith (CASIZ 078745) [four specimens dissected: 45/25 (#2), 30/24 (#5), 20/17 (#4), 18/16 (#3) and 15/11 (#1)]; Galapagos Islands, Isla Pinta [Abingdon Island], SW coast, 25 May 1964, three specimens 16/14 (#1) to 13/11 (#2) mm preserved [leg. D.Q. Cavagnaro], identified as Onchidella steindachneri by A.G. Smith (CASIZ 078743); Galapagos Islands, Isla Pinzon [Duncan Island], 7 June 1932, two specimens 12/11 (#1) and 13/ 10 mm preserved [leg. T. Crocker and F. Taiga], identified as Onchidiidae by L. Borock (CASIZ 077057); Galapagos Islands, Isla San Salvadore [James Island], N end James Bay, 2 February 1967, seven specimens 18/15 to 11/ 10 mm preserved [leg. I. Wiggins], identified as Onchidiidae [unknown identifier], (CASIZ 079303) [four specimens dissected: 18/15 (#1), 16/12 (#4), 12/10 (#2) and 11/10 (#3)]; Galapagos Islands, Isla Santa Cruz, Academy Bay, 18 July 1906, four specimens 18/23 (#1) to 18/15 (#2) mm preserved [leg. W.H. Oschner], identified as Onchidella steindachneri by A.G. Smith, (CASIZ 078749); Galapagos Islands, Isla Santa Cruz [Indefatigable Island], Dove Roads, 1905–1906, 37 specimens 26/22 to 12/ 14 mm preserved [leg. W.H. Oschner], identified as Onchidella [unknown identifier], (CASIZ 078354) [10 specimens dissected: 26/22 (#1), 20/23 (#7), 20/21 (#8), 20/18 (#10), 19/24 (#4), 18/15 (#2), 16/21 (#5), 13/15 (#6), 13/10 (#3), and 12/14 (#9)].</p> <p>Distribution</p> <p>Endemic to the Galapagos Islands: San Cristobal (Hoffmann 1928; present study); Santa Maria (= Charles Island; Stearns 1894); Isabella (= Albemarle; Stearns 1894; present study); San Salvadore (present study); Pinzon (present study); Santa Cruz (present study); Pinta (present study). Semper (1885) recorded O. steindachneri from the Galapagos Islands, without citing any specific island. Watson (1925) and Labbé (1934) only mentioned O. steindachneri from the Galapagos, without new material.</p> <p>Habitat</p> <p>Intertidal. On rocks.</p> <p>Remarks on the original description</p> <p>The dorsal colour of the two syntypes was described as blackish-greenish, and the ventral surface as yellowish. Semper also described some of the internal anatomy, especially the reproductive system: the insertion of the retractor muscle is at the very end of the visceral cavity, near the anus; also, the vas deferens was described as highly convoluted; finally, there is no penial accessory gland. The drawings of the rachidian and lateral teeth are poorly informative.</p> <p>Description of new specimens</p> <p>Several anatomical systems of O. steindachneri are similar to O. binneyi (nervous system, pallial organs, digestive system except for the radular formulae, and the posterior genital organs). Their written description is not provided here again, although some organs, which have not – or only poorly – been illustrated, are illustrated here. Given that the only syntype available has been completely destroyed, the description below is exclusively based on non-type specimens.</p> <p>External Morphology (Figures 1 and 13). Dorsal notum of live animals blackish. In preserved animals, dorsal colour from dark grey or bluish to dark brownish; centre of notum light tan in most individuals; hyponotum and pedal sole whitish.</p> <p>Size from 45/25 (CASIZ 078745) to 10/ 9 mm (CASIZ 017922). Body high, not flattened. Hyponotum horizontal. Dorsal notum oval, longer than wide. In most individuals, entire surface of dorsal notum granular with papillae of various sizes (usually &lt;2 mm high). However, notum of some individuals with smooth centre (e.g. CASIZ 078745, CASIZ 078749, SMNH 103166). Margin of notum is not smooth: characterized by digitiform warts of various sizes up to 2 mm long. In large individuals, many marginal warts much larger (up to 4 mm long) and subdivided (one main, central wart with several, smaller digitiform warts). Papillae with dorsal eyes absent. Dorsal gills absent.</p> <p>In most individuals, total width of hyponotum (left and right sides added) superior to width of pedal sole (H&gt; S): left side of hyponotum, pedal sole, and right side of the hyponotum of equal width. However, exceptionally, pedal sole wider than left and right hyponotum added (e.g., left hyponotum/pedal sole/right hyponotum as 5/10/4 and 4/10/ 3 mm, CASIZ 078745). On hyponotum, hyponotal line around pedal sole separating hyponotum into inner area (close to foot) and outer area. Distance between hyponotal line and pedal sole varies from one fifth to one third of width of</p> <p>ventral view, SMNH 103167 #1; (E) posterior (female) genital organs, SMNH 103167 #1; (F) posterior (female) genital organs, SMNH 103167 #1. Scale bar: A, 5 mm; B, 4 mm; C, 3.3 mm; D, 2 mm; E, 2 mm; F, 1.7 mm. Abbreviations: bm, buccal mass; cr, crop; dd, deferent duct; dg, digestive gland; e, oesophagus; fgm, female gland mass; hd, hermaphroditic duct; hg, hermaphroditic gland; hl, hyponotal line; i, intestine; lrpc, left reno-pulmonary complex; mgg, marginal gland; mo, male opening; ol, oral lobe; ov, oviduct; pgo, pedal gland opening; ppg, peripodal groove; rs, receptaculum seminis; sp, spermatheca; st, stomach; st2, stomach chamber #2; vg, vaginal gland.</p> <p>hyponotum (hyponotal line closer to pedal sole than to hyponotum margin). Openings (pneumostome, male opening and eye tentacles) are within smooth area delimited by hyponotal line.</p> <p>Pedal sole surrounded by two grooves on left and right sides. Left groove shallow. Peripodial groove present on right side, from buccal area (pedal gland opening) to posterior openings (anus and female opening). Anus posterior, median, close to pedal sole. Exceptionally, anus slightly on right side (CASIZ 078749 #2 and #9). Posterior female genital opening is very close to anus. Position of pneumostome on the hyponotum relative to notum margin and lateral side of the pedal sole varies among and within lots. Pneumostome can be: at an equal distance between pedal sole and hyponotal margin (e.g. CASIZ 078745 #2 and #3); one third closer to sole than hyponotal margin (e.g., CASIZ 078749 #2 and #9) or very close to anus, almost touching it (e.g. CASIZ 078354, two spms). Pneumostome generally close to median line, but slightly to right, occasionally median (e.g. SMNH 103166, three spms; NMNH 574379, two spms). Head bears pair of retractile, ocular tentacles, with eyes at tip. In all preserved specimens, ocular tentacles deeply retracted inside body. Left and right oral lobes distinct, not fused medially, superior to mouth but inferior to ocular tentacles. Opening of pedal gland median, inferior to mouth. Male genital opening on the right lateral side of right oral lobe.</p> <p>Marginal glands (Figures 13 and 14). Marginal glands are similar to those found in O. binneyi. However, between eight and fourteen glands on each side. Glands about 1 mm in diameter, although some smaller glands occasionally found in between larger glands.</p> <p>Digestive system (Figures 13, 15 and 16). Radular maximum size 10/ 4 mm, when flattened. Radular formulae vary among individuals (Table 1). From 66 to 121 rows, and from 63 to 114 teeth per half row.</p> <p>Reproductive system (Figs 13, 17 and 18). No accessory penial gland. No distinct penis inside penial sheath. No distinct, conspicuous papilla at distal end of deferens duct. Penial sheath covered internally by thick folds. Length of penial sheath up to 15 mm in large specimens. No accessory caecum filled with white concretions opening into penial sheath. Deferent duct highly convoluted and coiled. In most specimens, many loops that can hardly be counted. Number of loops varies among individuals: young, small specimens generally with deferent duct with fewer loops; deferent duct never straight and short. Diameter of deferent duct more or less constant (∼ 300 µm maximum). Retractor muscle anchors where deferent duct enters penial sheath and runs to the end of visceral cavity to insert near posterior openings (female opening and anus). Length of retractor muscle depends on size of penial sheath (long if short penial sheath and short if long penial sheath). However, retractor muscle inserts at posterior end of visceral cavity in all specimens dissected.</p> <p>Discussion</p> <p>Prior to the present study, only six specimens of O. steindachneri had been dissected: two by Semper (1885) and four by Hoffmann (1928); Stearns (1894) only described the external morphology. The present study, based on 29 specimens dissected (for 95 specimens examined in total) from 12 localities from six of the Galapagos Islands, provides us with the first opportunity to address the intra-specific variation of some of the characters. In particular, the variation of the male copulatory organs, which clearly distinguish O. steindachneri from other species in the north-eastern Pacific, is described here for the first time: the deferent duct is highly coiled, and, although the number of loops may vary among individuals, the deferent duct is never straight. Also, the variation of the radular formula is now better known: 66/121 × 63/114-1-63/114 (neither Semper nor Hoffmann gave specific radular formulae).</p> <p>Overall, the taxonomy of O. steindachneri is not problematic. It is a species that clearly differs from other Onchidella from the north-eastern Pacific and for which only one name has ever been proposed in the literature. As for its supra-specific relationships, it clearly does not belong to Onchidium, as originally proposed by Semper, who actually only used one generic name for all onchidiids, with the exception of one species which he classified in Onchidina.</p> </div>	https://treatment.plazi.org/id/03D887F6FFCDFFDCFE4B9CDAFD8BB21B	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFCBFFD3FE01994FFDF3B19F.text	03D887F6FFCBFFD3FE01994FFDF3B19F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hoffmannola Strand 1932	<div><p>Hoffmannola Strand, 1932</p> <p>Hoffmannola Strand, 1932: 194.</p> <p>Type species</p> <p>Hoffmannola lesliei (Stearns, 1892) by monotypy.</p> <p>Remarks</p> <p>Hoffmannola was created by Strand (1932: 194) as a new replacement name for Watsoniella Hoffmann, 1928, a permanently invalid name because it is preoccupied by Watsoniella Berg, 1898 (Lepidoptera).</p> <p>Diagnosis</p> <p>Dorsal notum without branchial gills. Dorsal papillae with no dorsal eyes. Two oral lobes fused medially. Hyponotal line present. Male genital opening median, superior to fused oral lobes. Intestine of type IV. Reno-pulmonary complex asymmetrical (left part significantly longer than right part). Marginal glands in body wall. Visceral cavity</p> <p>divided longitudinally by a septum. Accessory gland in the penial complex absent. Rectal gland absent.</p> <p>Remarks</p> <p>Given that the onchidiid supra-specific phylogenetic relationships are still unknown (Dayrat 2009; see Discussion), it is unclear whether the characters that have been traditionally accepted for Hoffmannola (e.g. Labbé 1934; Britton 1984) and are provided in the above diagnosis are synapomorphies, autapomorphies, or symplesiomorphies. Also, it is difficult to evaluate whether Hoffmannola is clustered within Onchidella, or whether both taxa are separate.</p> </div>	https://treatment.plazi.org/id/03D887F6FFCBFFD3FE01994FFDF3B19F	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFC4FFC8FE2B9ABBFDC1B536.text	03D887F6FFC4FFC8FE2B9ABBFDC1B536.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hoffmannola lesliei (Stearns 1892)	<div><p>Hoffmannola lesliei (Stearns, 1892)</p> <p>(Figures 19–21)</p> <p>Onchidium lesliei Stearns 1892: 87 [as Orchidium, a spelling mistake of Onchidium].</p> <p>Watsoniella lesliei. – Hoffmann 1928: 56–68, figs 11–15, plate 4, figs 14–18. — Labbé 1934: 245.</p> <p>Type material</p> <p>The three syntypes could not be located. Type locality: Galapagos Islands, Charles Island (one specimen, collected 8 April 1888, US Steamer Albatross) and Albermarle Island (two specimens, collected 10 April 1888, US Steamer Albatross).</p> <p>Additional material examined and dissected</p> <p>A total of 57 specimens was examined, 15 of which were dissected to study their internal anatomy. Examined specimens were collected from nine localities on five of the Galapagos Islands, four islands being new records (see Distribution). Galapagos Islands, Isla San Salvadore [James Island], N end of James Bay, 2 February 1967, three specimens 20/20 (#1) to 14/ 12 mm preserved [leg. I. Wiggins], identified as Onchidiidae [unknown identifier], (CASIZ 009777); Galapagos Islands, Isla Santa Cruz, Academy Bay, January 1974, one specimen 22/ 19 mm preserved [leg. unknown], identified as Hoffmannola lesliei [unknown identifier], (CASIZ 078708) [dissected]; Galapagos Islands, Isla Santa Cruz, Academy Bay, 25 January 1964, three specimens 24/18 (#2) to 18/12 (#1) mm preserved [leg. D.P. Abbott], identified as Hoffmannola lesliei by D.P. Abbott, (CASIZ 078709); Galapagos Islands, Isla Santa Cruz, Academy Bay, 25 January 1964, eight specimens 20/14 to 15/ 12 mm preserved [leg. D.P. Abbott], identified as Onchidiidae by T. M. Gosliner, (CASIZ 072952) [three specimens dissected: 20/14 (#1), 20/17 (#3) and 15/12 (#2)]; Galapagos Islands, Isla Santa Cruz, Academy Bay, 12 February 1964, 18 specimens 28/23 (#1) to 20/20 (#2) mm preserved [leg. A.G. Smith], identified as Hoffmannola lesliei by A.G. Smith, (CASIZ 078368); Galapagos Islands, Garden [possibly for Gardner] Island, [collecting date unknown], 13 specimens 20/18 (#1) to 10/8 (#2) mm preserved [leg. unknown], identified as Onchidium [unknown identifier], (ANSP A7486 d) [specimens poorly preserved]; Galapagos Islands, Albemarle Island, Iguana cove, [collecting date unknown], three specimens 25/25 (#1) to 11/ 16 mm preserved [leg. unknown], identified as Onchidium [unknown identifier], (ANSP 81945) [specimens poorly preserved]; Galapagos Islands, Marborough Island [Isabela], Iguana cove, 26 January 1899, four specimens 25/20 (#1) to 15/12 (#2) mm preserved [leg. unknown], identified as Onchidium lesliei [unknown identifier], (ANSP 81946) [specimens poorly preserved]; Galapagos Islands, Albemarle Island, three specimens 36/25 (#1) to 15/ 10 mm preserved [leg. USFC Albatross], identified as Onchidium binneyi ? [unknown identifier], (ANSP 111156) [specimens poorly preserved].</p> <p>Distribution.</p> <p>Endemic to the Galapagos Islands: Santa Maria (= Charles Island, type locality; Stearns 1892); Isabella (= Albemarle, type locality; Stearns 1892; Hoffmann, 1928; present study); San Salvadore (present study); Santa Cruz (present study); Garden [possibly for Gardner] Island (present study); Isabela Island (present study). Labbé (1934) only mentioned H. lesliei from the Galapagos, without new material.</p> <p>Habitat</p> <p>Intertidal. On rocks.</p> <p>Remarks on the original description</p> <p>The original description, quite short, is reproduced here in its entirety:</p> <p>Form rounded ovate, nearly as broad as long. Dorsum coriaceum, nearly black, shiny, closely irregularly reticulated, with finely incised lineation, and otherwise characterized by somewhat distant tiatly rounded papillae. Underside, dingy yellowish white; margin of mantle wide, nearly smooth; edge of same simple. Anal opening posterior, near edge of mantle and somewhat produced. Respiratory orifice smaller, in median line with and infront of anus. Sexual orifice anterior on the right side, under the edge of the large oval hood or collar. Labial palpi thin, largely expanded.</p> <p>Stearns only described the external morphology, which, at the time, was enough to distinguish H. lesliei from all other known onchidiid species, because of the fused oral lobes, which Stearns referred to as the “collar”. However, his description of the position of the male anterior opening is erroneous. As pointed out by Hoffmann (1928), the male opening is median, superior to the oral “collar”.</p> <p>Description of new specimens</p> <p>Because the type material could not be located, the description below is based exclusively on non-type specimens.</p> <p>External Morphology (Figure 19). Background colour of dorsal notum of live animals brown to blackish, slightly mottled with lighter areas around small papillae. In preserved animals, dorsal colour from homogeneously whitish tan to dark brown. Hyponotum and pedal sole whitish tan or, exceptionally, light brown.</p> <p>Maximum size of preserved specimens from 36/ 25 mm (ANSP 81946 #1) to 10/ 8 mm (ANSP A7486d #2). Body high, not flattened. Hyponotum horizontal. Dorsal notum oval (slightly longer than wide) or round. In most individuals, notum almost smooth to finely granular covered by distant, low, rounded papillae (&lt;1 mm high); papillae more abundant on side of notum than in centre. Margin of notum smooth. Dorsal papillae with dorsal eyes absent. Dorsal gills absent.</p> <p>In most individuals, total width of hyponotum (left and right sides added) equal to width of pedal sole (H = S). Exceptionally, pedal sole wider than total width of hyponotum (H &lt;S). On hyponotum, hyponotal line around pedal sole separating hyponotum into inner area (close to foot) and outer area. Distance between hyponotal line and pedal sole from one sixth to one third of width of hyponotum (hyponotal line closer to the pedal sole than to hyponotum margin). Openings (pneumostome, male opening and eye tentacles) within area delimited by hyponotal line.</p> <p>Peripodial groove present on right side, from posterior genital openings (female opening and anus) to opening of pedal gland in buccal area. Anus posterior, close to pedal sole, slightly on right of median line. Anus occasionally median. Posterior female genital opening close to anus. Pneumostome very close to anus, or one third closer to pedal sole than hyponotal margin. Pneumostome elongated and slightly on the right of median line (although exceptionally median). In anterior region, head covered dorsally by notum. Head bears pair of retractile, ocular tentacles, with eyes at tip. In most specimens, tentacles deeply retracted. Left and right oral lobes not distinct (fused medially), superior to mouth, inferior to ocular tentacles. Opening of pedal gland median, inferior to mouth. Male genital opening median, superior to fused oral lobes.</p> <p>Marginal glands (Figure 19). From 36 to 44 glands on each side. Glands very close to each other, in approximately three longitudinal rows, and on different frontal planes. Glands from 0.5 to 1 mm in diameter.</p> <p>Visceral cavity (Figure 19). In its two posterior thirds, visceral cavity divided by strong, longitudinal septum. Septum attached ventrally to floor of visceral cavity and dorsally to dorsal notum. Heart and part of the digestive system (intestine loop and digestive gland) on right side of septum. Septum divided ventrally into two walls separated by tiny space (retractor muscle inserts between these two walls of septum).</p> <p>Digestive system (Figures 19 and 20). No jaw-like structure present on surface of oral tube. Left and right salivary glands heavily branched, join buccal mass dorsally, on either side of oesophagus. Radula in between two large postero-lateral muscular masses. Radular maximum size 10/ 6 mm, when flattened. Each radular row with one rachidian tooth and two half rows of lateral teeth of similar size and shape (except first and second inner lateral teeth). Radular formulae vary among individuals (Table 2). From 85 to 160 rows, and from 150 to 515 teeth per half row. No strict correlation between size of specimens and radular formulae. Rachidian tooth tricuspid: median cusp always present; two lateral cusps on two lateral branches of base of rachidian tooth may be absent (especially in small rachidian teeth). Rachidian teeth slightly smaller than lateral teeth: total length of base of rachidian tooth &lt;40 µm, median cusp ∼ 10 µm in length. Posterior-lateral aspect of base of rachidian teeth only slightly concave. Half rows of lateral teeth at an angle of aproximiately 45 ◦ to rachidian axis. Lateral teeth characterized by base attached to radular membrane, with dorsal, strong, flattened hook at 45 ◦ angle to radular membrane; triangular structure between base and hook supports hook but does not reach tip. Length of dorsal hook usually is ∼ 20 µm and &lt;40 µm. Also, smaller (&lt;10 µm) pointed cusp on outer, lateral expansion of base. In most cases, lateral cusp cannot be observed on SEM picture because hidden below hook of adjacent, outer tooth; however, lateral cusps conspicuous when teeth not too close (such as in the innermost and outermost regions). With exception of first two to five innermost lateral teeth, size of lateral teeth does not vary across half row, nor does it among half rows. Shape of tip of hook from truncate to rounded, varies among individuals and between rows of same radula. Hooks not straight, slightly curved inward. Finally, half rows at a 45 ◦ angle to rachidian axis, but hooks almost parallel to rachidian axis.</p> <p>Oesophagus narrow and straight at proximal end but enlarges into wide crop covering almost entire posterior end of buccal mass. Oesophagus enters stomach anteriorly, close to connection of stomach with dorsal and lateral lobes of digestive gland. Only small portion of posterior aspect of stomach can be seen in dorsal view, because largely covered by three lobes of digestive gland: dorsal lobe mainly located on right aspect of visceral mass; left, lateral lobe mainly ventral; posterior lobe covers posterior aspect of visceral mass. U-shaped sac stomach divided into three chambers: first chamber, delimited by a thin layer of tissue, receives oesophagus and ducts of dorsal and left lateral lobes of digestive gland; second chamber, posterior, delimited by thick muscular tissue and receives duct of posterior lobe of digestive gland; third chamber delimited by thin layer of tissue; in some individuals, third chamber funnel-shaped. No strong ridges on internal surface of stomach. Intestine long, narrow, and of type IV (Labbé, 1934). No rectal gland.</p> <p>Nervous system (Figure 19). Central nervous system densely compact, often embedded and protected within layer of connective tissue. Circum-oesophageal nerve ring post-pharyngeal. Two cerebral ganglia not fused, separated by short commissure. Pleural and pedal ganglia distinct. Cerebro-pleural and pleuro-pedal connectives very short: pleural, cerebral, and pedal ganglia touch each other. Visceral loop very short, and visceral ganglion touches both left and right pleural ganglia. Nerves from cerebral Note: H = hyponotum total width (left and right); S = pedal sole width; radular formula format as range of rows × (range of number of left lateral teeth per half row – one rachidian tooth – range of number of left lateral teeth per half row); patterns of attachment of retractor muscle follows Plate’s (1893) classification.</p> <p>ganglia innervate buccal area and ocular tentacles, and male anterior genital organs (penial complex) on right side. Nerves from pedal ganglia innervate foot. Nerves from pleural ganglia innervate lateral and dorsal regions of mantle. Nerves from visceral ganglia innervate visceral organs.</p> <p>Pallial complex (Figure 19). Heart enclosed in pericardium, located on right portion of visceral cavity, i.e. on right side of septum. Large anterior ventricle becomes large aorta that branches into smaller vessels delivering blood to visceral organs. Posterior auricle significantly smaller than ventricle. Pericardium communicates through small hole with right portion of renal-pulmonary complex. Kidney intricately attached to pulmonary cavity and forms two, left and right, parts, left part significantly longer than right part (length of left pulmonary cavity about same as that of septum). Pulmonary cavity characterized by complex folds of internal lining, likely to increase tissue surface participating in gas exchange.</p> <p>Reproductive system: posterior parts (Figure 21). Hermaphroditic gland forms single mass sub-divided into acini. Hermaphroditic duct highly-coiled, conveys gametes (eggs and autosperm) up to spermoviduct. Small branch of hermaphroditic duct goes directly to female gland mass. Another branch of hermaphroditic duct leads to small, coiled, finger-shaped pouch (receptaculum seminis = vesicula seminalis). From receptaculum seminis, narrow duct goes back toward female gland mass (mucous and albumen glands with ducts opening near where hermaphroditic duct becomes spermoviduct). Proximal spermoviduct embedded within the female gland mass and not divided, at least externally. Prostate not distinct externally may be within walls of spermoviduct or vas deferens. Distally, male and female ducts separate: vas deferens conveys autosperm up to cephalic region; free oviduct conveys eggs up to female opening and exosperm from female opening up to fertilization chamber, near proximal end of spermoviduct. Distally, oviduct becomes vagina. Spherical spermatheca (which stores exosperm) connects to distal region of oviduct through short duct. Short, accessory, vaginal gland opens into distal portion of oviduct.</p> <p>Reproductive system: male, anterior parts (Figure 21). No accessory penial gland. No distinct penis inside penial sheath. No papilla at distal end of deferens duct. Penial sheath not covered internally by folds. Penial sheath &lt;500 µm in diameter and 6 mm in length. No caecum and no concretions in penial sheath. Deferent duct loosely convoluted (fewer than 10 loops) from entrance in visceral cavity up to where it joins penial apparatus. Number of loops varies among individuals. Diameter of deferent duct more or less constant along its whole length (∼ 200 µm maximum). After it joins penial sheath, deferent duct does not directly join male opening but forms several loops inside proximal portion of penial sheath, makes U-turn, and then goes back distally toward male opening. Retractor muscle anchors on penial sheath near where deferent duct makes a U-turn inside penial sheath, and inserts in between two walls of septum, close to floor of visceral cavity. Retractor muscle very thin, inconspicuous, even missing in many specimens.</p> <p>Discussion</p> <p>Prior to the present study, the internal anatomy was only described by Hoffmann (1928), based on material from only one locality. Hoffmann provided a detailed description of the septum dividing the visceral cavity, based on a series of drawings of transversal cuts of the body. Generally speaking, his description is quite detailed and long. However, most systems were not illustrated, which makes the description difficult to read: the drawings of the radular teeth, penial sheath and posterior genital organs are not satisfactory. Also, Hoffmann, who had access to material from only one station, did not address individual variation.</p> <p>Hoffmann’s description fits well within our description: for instance, the radular formula provided by Hoffmann (120 × 360-0-360) is within the range observed here (85/160 × 150/515-150/515). However, the present study, based on 15 specimens dissected (for 57 specimens examined in total), from nine localities and five of the Galapagos Islands, provides us with the first opportunity to address the character intra-specific variation, which is already given above and is not repeated here. Also, the anatomy of most systems is illustrated here for the first time. Only two nominal species exist in Hoffmannola: H. lesliei and H. hansi. Their differences are discussed under the next species (H. hansi).</p> </div>	https://treatment.plazi.org/id/03D887F6FFC4FFC8FE2B9ABBFDC1B536	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
03D887F6FFDFFFCDFEB59E05FE5CB263.text	03D887F6FFDFFFCDFEB59E05FE5CB263.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hoffmannola hansi Marcus and Marcus 1967	<div><p>Hoffmannola hansi Marcus and Marcus, 1967</p> <p>(Figures 22 and 23)</p> <p>Hoffmannola hansi Marcus and Marcus 1967: 232–236, figs 87–95. — Marcus and Marcus 1970: 214, figs 92–93.</p> <p>Type material</p> <p>A total of 10 syntypes. One syntype (NMNH 678419): one specimen 25/ 25 mm preserved [leg. and collecting date unknown, but likely P. Pickens or M.A. Hill in 1966]; a label printed on 10 July 2007 indicates that this specimen is the “ holotype ”, however Marcus and Marcus did not designate a holotype in the original description, and the other, older label in the jar does not indicate that this specimen is the holotype, thus, this specimen is regarded here as one of the syntypes. Five syntypes (NMNH 753653): five specimens 28/23 to 20/ 17 mm preserved, leg. M.A. Hill, 22 June 1966; both the old and recent labels indicate that these five specimens are paratypes, however, the old label indicates that these specimens were determined by Hans Bertsch as the paratypes; because Marcus and Marcus did not distinguish a holotype and paratypes in the original description, these specimens are regarded here as syntypes. Four syntypes (NMNH 753651): four specimens 28/23 to 22/ 18 mm preserved, leg. P. Pickens, 12 November 1966; both the old and recent labels indicate that these four specimens are paratypes, however, the old label indicates that they were determined by Hans Bertsch as paratypes; because Marcus and Marcus did not distinguish a holotype and paratypes in the original description, these specimens are regarded here as syntypes. Type locality: El Sahuaral, Sonora, [Gulf of California], Mexico [all syntypes come from El Sahuaral, although the specimen catalogued as NMNH 678415 was collected more specifically from Kino Bay, in El Sahuaral]. Type material condition: all specimens are well preserved; only two specimens (NMNH 753651) were dissected prior to the present study, likely by Marcus and Marcus; two specimens were dissected for the present study [27/ 22 mm, labelled as syntype # 1 in the jar, NMNH 753653; 28/ 23 mm, labelled as syntype # 2 in the jar, NMNH 753651].</p> <p>Additional material examined and dissected</p> <p>Mexico, Sinaloa, Mazatlan, N of Gauiola Beach, 1 December 1953, three specimens 28/23 (#1) to 23/14 (#2) mm preserved [leg. L.O. Miles], identified as Hoffmannola hansi by T. M. Gosliner, (CASIZ 081809); Mexico, Sonora, just N of Ensenada Lalo (a bay just N of San Carlos Bay), 24 March 1966, one specimen 28/ 25 mm preserved [leg. E. Coan and R. Szal], identified as Hoffmannola hansi by T. M. Gosliner, (CASIZ 073902) [dissected].</p> <p>Distribution</p> <p>Endemic to the Gulf of California: Sonora (original description, Marcus and Marcus 1967; present study); Sinaloa (present study). Marcus and Marcus (1970) only mentioned some material collected by P. Pickens on 12 November 1966 from Mexico, without additional locality information; however, Marcus and Marcus (1970), who only briefly commented on H. hansi and did not provide any new anatomical information, likely only referred to the type material. Marcus and Marcus (1967, 1970) also mentioned San Augustin and Angel de la Guarda Island as two other possible localities, but without any material or vouchers. In any case, H. hansi is known to inhabit the Gulf of California from its northern end (Sonora) to it southern end (Sinaloa).</p> <p>Habitat</p> <p>Intertidal. On rocks.</p> <p>Remarks on the original description</p> <p>The original description is quite detailed and accurate. Several illustrations help clarify the description: radular teeth, a dorsal view of the visceral cavity, and portions of the male, anterior apparatus are drawn. However, the original description of H. hansi was based on the dissection of only two specimens from one locality (Sonora), and could hardly document individual variation of features that seem to distinguish H. hansi from H. lesliei.</p> <p>Description of new specimens</p> <p>Several anatomical systems of H. hansi are similar to H. lesliei (nervous system, pallial organs, septum in visceral cavity, digestive system except for the radular formulae, and the posterior genital organs). Their written description is not provided here again, although some organs, which previously have been illustrated only poorly or not at all, are illustrated here. The description below is based on the non-type specimens as well as the type material, especially the two syntypes dissected for the present study.</p> <p>External Morphology (Figure 22). Background colour of dorsal notum of live animals brownish to black, with lighter pigments around low papillae. In preserved animals, dorsal colour varies from whitish tan to brown and dark grey. Colour of hyponotum and pedal sole varies from whitish tan to light brown. Size of preserved specimens from 28/23 to 20/ 17 mm. In most individuals, total width of pedal sole greater than total width of hyponotum, left and right sides added (H &lt;S), e.g. 1/15/ 1 mm (NMNH 753653). On hyponotum, hyponotal line around pedal sole separating hyponotum into inner area (close to foot) and outer area. Distance between hyponotal line and pedal sole varies from one sixth to one quarter of width of hyponotum.</p> <p>Digestive system (Figures 22 and 23). Radular maximum size 10/ 6 mm, when flattened. Radular formulae vary among individuals (Table 1). From 100 to 195 rows, and from 430 to 735 teeth per half row.</p> <p>Marginal glands (Figure 22). From 8 to 14 glands on each side. Glands do not touch each other and form only one row on single frontal plane. Glands from 0.5 to 1 mm in diameter.</p> <p>Reproductive system: male, anterior parts (Figure 22). No accessory penial gland. No distinct penis inside penial sheath. No distinct, conspicuous papilla at distal end of deferens duct. Penial sheath not covered internally by thick folds. Penial sheath &lt;500 µm in diameter and 6 mm in length. No caecum and no concretions in penial sheath. Deferent duct heavily convoluted (actual loops cannot be counted) from entrance in visceral cavity up to penial apparatus. Number of loops varies among individuals, but always many. Diameter of deferent duct more or less constant along its whole length (∼ 200 µm maximum). Deferent duct forms several loops inside the proximal portion of the penial sheath, makes a U-turn, and then goes back distally toward male opening. Anchor of retractor muscle on the penial sheath near U-turn of deferent duct inside penial sheath. Insertion of retractor muscle in between two walls of septum, close to floor of visceral cavity. Retractor muscle thick and strong, present in all specimens.</p> <p>Discussion</p> <p>So far, H. hansi has only been known from the original description, and a few additional notes by Marcus and Marcus (1967, 1970), who only dissected two specimens from the same locality. Five additional specimens were dissected for the present study, which significantly improves our knowledge: the distribution range is extended to the entire Gulf of California, with specimens described here from Sinaloa (southern part of the Gulf); the intra-specific variation within H. hansi, as well as the differences between H. lesliei and H. hansi are better known, especially because it seems that Marcus and Marcus never dissected any specimens of H. lesliei (at least they never indicated that they had done), and their knowledge of H. lesliei was based only on Hoffmann’s redescription of that species.</p> <p>The original description by Marcus and Marcus and ours are largely in agreement. Marcus and Marcus gave one radular formula (170 × 600–0–600) that fits within the range observed here (100/195 × 430/735–1–430/735). The teeth are illustrated here for the first time with SEM. The nervous system, the posterior genital organs and the course of the deferent duct within the visceral cavity are also illustrated here for the first time. Only large specimens were available for the present study; in all of them, the deferent duct was found to be very highly convoluted. However, given what we know about the variation of this character in other species, it will likely be found that, at least in smaller, younger specimens, the deferent duct is less coiled.</p> <p>The present study, in which intra-specific variation within both H. hansi and H. lesliei is evaluated, also helps refine our understanding of the key differences between H. hansi and H. lesliei. In particular, a major difference not mentioned by</p> <p>penial sheath; (E) posterior (female) genital organs; (F) anterior male genital organs. Scale bars: A, 2.4 mm; B, 9 mm; C, 5 mm; D, 2.4 mm; E, 4 mm; F, 0.75 mm. Abbreviations: bm, buccal mass; dd, deferent duct; dg, digestive gland; fgm, female gland mass; hd, hermaphroditic duct; hg, hermaphroditic gland; i, intestine; lc, left cerebral ganglion; lpl, left pleural ganglion; lrpc, left reno-pulmonary complex; mgg, marginal gland; ot, ocular tentacle; ov, oviduct; pc, pericardium; pg, pedal gland; ps, penial sheath; rm, retractor muscle; rrpc, right, reno-pulmonary complex rs, receptaculum seminis; sp, spermatheca; st, stomach; vg, vaginal gland; vs, visceral ganglion.</p> <p>Marcus and Marcus is the coiling of the deferent duct. In H. lesliei, the deferent duct of large specimens (which are sexually fully mature) is only loosely convoluted with a few loops. In H. hansi, the deferent duct of similar specimens is very highly convoluted. Although the variation of this character is poorly known in H. hansi, this difference will likely remain a valid, important difference. The deferent ducts of H. hansi (Gulf of California) and H. lesliei (Galapagos) differ as much as the deferent ducts differ between Onchidella steindachneri (Galapagos) and O. binneyi (Gulf of California to Ecuador), which have a highly-convoluted and poorly-convoluted deferent duct, respectively. Marcus and Marcus did not see that major difference, likely because they did not have access to any material of H. lesliei, and it is always very difficult to “see” organs in someone else’s descriptions, regardless of whether or not illustrations are available.</p> <p>Marcus and Marcus (1967) mentioned five differences between H. lesliei and H. hansi. They are commented on here.</p> <p>(1) According to Marcus and Marcus, the notal glands of H. lesliei are much smaller, more numerous, and on different levels, whereas in H. hansi, the notal glands are larger, less numerous, and not on different levels. Although the size, number, and position of notal (= repugnatory, marginal) glands vary among individuals (see descriptions above), the observation by Marcus and Marcus remains correct.</p> <p>(2) Marcus and Marcus also claimed that the reno-pulmonary complex is smaller in H. lesliei than in H. hansi. However, that is actually not the case because the width of the reno-pulmonary complex does vary among specimens, and its length relative to the length of the visceral cavity is quite similar between and within each species.</p> <p>(3) According to Marcus and Marcus, the radulae also differ. However, Marcus and Marcus only knew one formula for H. lesliei (120 × 360–1–360) and one formula for H. hansi (170 × 600–1–600), which might have explained why they differed, but, generally speaking, Marcus and Marcus paid little attention to the individual variation of radular formulae and their impact on taxonomy (see Dayrat 2010a, 2010b). We know now that the range of variation of radular formulae of H. hansi (100/195 × 430/735–1–430/735) and H. lesliei (85/160 × 150/515–1–150/515) overlap partly. So the ranges of radular formulae differ significantly, but must be used with caution because they do overlap.</p> <p>(4) According to Marcus and Marcus, the penial retractor originates under the nerve-ring in H. lesliei and deep between the two muscles of the septum in H. hansi. However, now that more specimens have been dissected, it seems that the retractor muscle attaches in between the two walls of the septum in both species. However, the retractor muscle of H. lesliei is thin and weak (and it is occasionally difficult to establish its course) and even often absent, whereas it is very strong and always present in H. hansi.</p> <p>(5) According to Marcus and Marcus, glandular villosities are present in the penial atrium of H. lesliei, which are absent in H. hansi. Such glandular villosities were not observed here, which might be due to the fact that no histology was used.</p> </div>	https://treatment.plazi.org/id/03D887F6FFDFFFCDFEB59E05FE5CB263	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	Dayrat, Benoît;Zimmermann, Sara;Raposa, Melissa	Dayrat, Benoît, Zimmermann, Sara, Raposa, Melissa (2011): Taxonomic revision of the Onchidiidae (Mollusca: Gastropoda: Pulmonata) from the Tropical Eastern Pacific. Journal of Natural History 45 (15 - 16): 939-1003, DOI: 10.1080/00222933.2010.545486, URL: http://dx.doi.org/10.1080/00222933.2010.545486
