Five times over: 42 new Angustopila species highlight Southeast Asia's rich biodiversity (Gastropoda, Stylommatophora, Hypselostomatidae) Author Pall-Gergely, Barna https://orcid.org/0000-0002-6167-7221 Centre for Agricultural Research, Plant Protection Institute, Eoetvoes Lorand Research Network, Herman Otto ut 15, H- 1022 Budapest, Hungary pallgergely2@gmail.com Author Hunyadi, Andras Adria setany 10 G 2 / 5., H- 1148 Budapest, Hungary Author Vermeulen, Jaap J. JK Art and Science, Lauwerbes 8, 2318 AT Leiden, Netherlands Author Grego, Jozef Horna Micina 219, SK- 97401 Banska Bystrica, Slovakia Author Sutcharit, Chirasak Animal Systematic Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand Author Reischuetz, Alexander Puechhaimgasse 52, A- 3580 Horn, Austria Author Dumrongrojwattana, Pongrat Department of Biology, Faculty of Science, Burapha University, 169 Longhardbangsaen Road, Muang District, Chonburi, 20131, Thailand Author Botta-Dukat, Zoltan Centre for Ecological Research, Institute of Ecology and Botany, Alkotmany 2 - 4, H- 2600, Vacratot, Hungary Author Oerstan, Aydin 12501 Milestone Manor Lane, Germantown, Maryland, 20876, USA Author Fekete, Judit University of Pannonia, Centre of Natural Science, Research Group of Limnology, Egyetem u. 10, H- 8200 Veszprem, Hungary & Centre for Ecological Research, Institute of Aquatic Ecology, Department of Tisza Research, 18 / c Bem square, H- 4026 Debrecen, Hungary Author Jochum, Adrienne https://orcid.org/0000-0002-6624-6412 Naturhistorisches Museum der Burgergemeinde Bern, CH- 3005 Bern, Switzerland & Institute of Ecology and Evolution, University of Bern, CH- 3012 Bern, Switzerland & Senckenberg Forschungsinstitut und Naturmuseum, 60325 Frankfurt am Main, Germany text ZooKeys 2023 2023-02-13 1147 1 177 http://dx.doi.org/10.3897/zookeys.1147.93824 journal article http://dx.doi.org/10.3897/zookeys.1147.93824 1313-2970-1147-1 9BB9881B0076473D8E53155D37CA1F50 FF2B6B317B505F9EA0E1000BDCD16CE7 Angustopila fratermajor Pall-Gergely & Vermeulen sp. nov. Fig. 43 Type material. Holotype : Vietnam • 1 empty shell (H: 0.84 mm, D: 0.8 mm); Haiphong Province, Cat Ba Island, large, somewhat disturbed, active cave with speleothems; 20°50.06'N , 106°55.91'E ; 0-50 m a.s.l.; 7 Jun. 2017; J.J. Vermeulen & K. Anker leg.; HNHM 105286 (original inventory number: JJV 16605). Paratypes : Vietnam • 68 shells; same data as for holotype; JJV 17658 (ex JJV 17605) • 10 shells; Quang Ninh Province, Halong-Campha area, 4.5 km SW Quang Hanh, limestone hill with regrowth; 20°58.98'N , 107°11.83'E ; 29 Sep. 1998; J.J. Vermeulen & A.J. Whitten leg.; JJV 6267 • 37 shells (one of them is figured: Fig. 99E ); Haiphong Province, Cat Ba Island, Cave Qua Vang, inside cave, large, ecologically intact active cave with speleothems; 20°48.64'N , 107°04.64'E ; 60 m a.s.l.; 6 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 16600 • 8 shells; Haiphong Province, Cat Ba Island, Cave Qua Vang, around cave entrance, rocky limestone slope with low, somewhat mature forest; 20°48.64'N , 107°04.64'E ; 100 m a.s.l.; 6 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 18866 (ex JJV 16604) • 41 shells; Haiphong Province, Cat Ba Island, Cave Hoa Cuong, inside cave, polluted cave disturbed by tourism, with concrete paths; 20°50.41'N , 106°59.15'E ; 50 m a.s.l.; 5 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 16593 • 3 shells (one of them is corroded); Haiphong Province, Cat Ba Island, Cave Xa Bac, inside cave; 20°50.07'N , 106°58.61'E ; 9 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 16606 • 13 shells; Quang Ninh Province, Halong Bay area, unnamed island 1.8 km west of the southernmost point of Cong Tai Island, Steep limestone slope bordering beach, dense vegetation; 20°52.43'N , 107°18.32'E ; 3 Oct. 1998; J.J. Vermeulen & A.J. Whitten leg.; JJV 6265 • 1 shell; Haiphong Province, Cat Ba Island, limestone cliff at SE end of Viet Hai polje, base of cliff, surrounded by degraded woodland; 20°47.50'N , 107°02.90'E ; 30 m a.s.l.; 10 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 16608 • 1 shell; Haiphong Province, Cat Ba Island, Cave Trung Trang, around cave entrance, steep limestone cliffs with vegetated ledges; 20°47.30'N , 106°59.84'E ; 50 m a.s.l.; 6 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 18867 (ex JJV 16617) • 2 shells; Haiphong Province, Cat Ba Island, Cave Minh Chou, inside cave, cave with freshwater stream ending in the sea, much disturbed by water extraction and concrete paths; 20°45.21'N , 107°00.75'E ; 50 m a.s.l.; 5 Jun. 2017; J.J. Vermeulen & K. Anker leg.; JJV 16594 • 533 shells; Quang Ninh Province, Halong Bay, Cap La Cave, deposit of soil fallen in through roof in pristine cave, vegetation outside cave tall and woody; 20°51.79'N , 107°13.54'E ; 7 Mar. 2018; J.J. Vermeulen & K. Anker leg.; JJV 17634 • 10 shells, same data as for preceding, coll. HA • 22 shells; Haiphong Province, Cat Ba Island, Cave Uy Ban, inside small, disturbed cave; 20°46.39'N , 107°00.88'E ; 40 m a.s.l.; 9 Jun. 2015; J.J. Vermeulen & K. Anker leg.; JJV 16613. Additional material. Vietnam • 1 figured shell (Suppl. material 3: fig. S30) + 5 shells; Thanh Hoa Province , Pu Puong National Park , limestone hill near small native village Am (locality code: WMVT.0344); 20°27.39'N , 105°13.65'E ; 21 Sep. 2003 ; W.J.M. Maassen leg.; RMNH.5006717 • 1 shell; same data as for preceding; NMBE 550646 . Diagnosis. A small to large Angustopila species with a conical-globular shell, a relatively wide umbilicus, and a weak, deeply set parietal tooth. Description. Shell small to large for the genus, mostly higher than wide, rarely slightly wider than high; off-white, transparent, conical or more often conical-globular, only slightly higher than wide, body whorl widest from standard apertural view; protoconch consists of 1.5 whorls, with very weak spiral striation preceding the first teleoconch whorl; teleoconch finely ornamented with irregular radial growth lines crossed by fine rows of equidistantly-spaced microscopic spiral threads (ca. 14-17 on body whorl from apertural view); on frontal and ventral surfaces of body whorl spiral and radial lines dominant; whorls 4-4.5, rounded, slightly shouldered and/or pushed from basolateral direction; aperture oblique to shell axis from lateral view; umbilicus relatively wide; aperture teardrop shaped to ovoid with straight or convex parietal part; peristome slightly expanded, not reflected; parietal callus separated from penultimate whorl; aperture with a weak (low), deeply-set parietal tooth; some shells are toothless (probably representing immature individuals). Measurements (in mm). H = 0.8-1.11, D = 0.75-0.85, H/D*100 = 97.5-117.5 ( n = 16, specimens from the Halong Bay), RUD = 25.6-29.6 ( n = 3); H = 0.8-0.93, D = 0.8-0.85, H/D*100 = 97.6-114.8 ( n = 6, WMVT.0344), RUD = 30.2 ( n = 1). Differential diagnosis. Some specimens of A. fratermajor sp. nov. are almost as small and have a lower spire than A. maasseni sp. nov. However, the two species can be distinguished based on aperture shape. Namely, that of A. fratermajor sp. nov. is comparatively larger, and more rounded (more impressed from parietal direction in A. maasseni sp. nov.), and the parietal tooth is situated deeper (reaching or nearly reaching the peristome in A. maasseni sp. nov.). Angustopila fratermajor sp. nov. was found together with A. apiostoma sp. nov. in multiple samples. The two species are similar in size and general shape and therefore, careful attention needs to be given to distinguish them. The differences are the following: Angustopila fratermajor sp. nov. has a lower conical shell with a wider umbilicus, the body whorl is "sharper" from umbilical view (i.e., there is a tendency towards a blunt periumbilical keel); there is a distinct parietal tooth (however, it is situated deep within the shell and difficult to recognise in some specimens), which is rare and if present, very low in A. apiostoma sp. nov.; the parietal callus is straight or convex (concave in A. apiostoma sp. nov.), and the aperture is less protruding anteriorly than in the other species. Angustopila fabella shells of the Halong Bay area are concave-conical in shape (conical-globular in A. fratermajor sp. nov.). Angustopila babel sp. nov. possesses larger shells and a comparatively larger aperture with a more anteriorly situated parietal tooth than A. fratermajor sp. nov. Angustopila tonkinospiroides sp. nov. is superficially similar in shell shape, but it is larger, and has no parietal tooth. See also under A. fraterminor sp. nov. Etymology. The specific epithet Angustopila fratermajor (Latin: older brother) refers to the presence of two similar (a smaller and a larger) species in the Halong Bay area. Distribution. This species is known from several sites of the Halong Bay area, and the sample from the Pu Puong National Park, Thanh Hoa Province is also assigned to this species (see Remarks) (Fig. 44 ). Figure 43. Angustopila fratermajor Pall-Gergely & Vermeulen, sp. nov., paratype ( A ) and holotype (HNHM 105286), JJV 17605 ( B-I ). Apertural ( A, B ), lateral ( C ), apical ( D ) and ventral ( E ) sides of the shell; aperture ( G ), microstructure of the protoconch ( F ), ventral ( H ) and frontal ( I ) surface of the body whorl. Figure 44. Distribution of Angustopila species. Note that the occurrence of Angustopila fratermajor Pall-Gergely & Vermeulen, sp. nov. in the Pu Puong National Park (Thanh Hoa Province) is not indicated. Remarks. The shells from the Pu Puong National Park have a weaker parietal tooth and show a more precisely shaped conical shell form than those deriving from the Halong Bay area. Therefore, we have chosen against selecting their serving as paratypes.