taxonID	type	description	language	source
03F3175FEF6E5E46FC2F3B56FDAFF7EB.taxon	diagnosis	Diagnosis. Shell small to medium-sized (D = 9 – 25 mm), subglobose with moderately to strongly elevated, exceptionally also low spire; umbilicus frequently concealed by greatly expanded callus, occasionally forming a chink or narrowly open; protoconch smooth to weakly sculptured by radial ridgelets or ribs; teleoconch with faint to well-developed sculpture of axial ridgelets or ribs, mostly on upper sector of whorls; whorls separated by shallow suture; periphery wellrounded to weakly angulated. Shell colour generally variable, background whitish or brownish horn, with well-developed, red subsutural, peripheral and frequently additional, thinner spiral bands. Eversible head wart between ommatophores. Penis without penial sheath, with well-developed, long and coiled epiphallus with flagellum of variable length; penial retractor muscle attached at epiphallus; epiphallus opens to lumen of penial chamber through large, cone-shaped verge with characteristic sperm groove. Bursa copulatrix short to long, simple to kinked (based on Köhler, 2011).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E46FC7A3C94FC3AF9F2.taxon	materials_examined	Material examined. AM C. 471154 (Indonesia, Solor Island, 3 wet, 2 dry).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E46FC7A3C94FC3AF9F2.taxon	discussion	Taxonomic history. Representing the first named non-Australian species, R. solorensis has continuously been maintained as a distinct species in previous taxonomic treatments. Vermeulen & Whitten (1998) and Maassen (2009) reported R. solorensis from Bali and Nusa Penida, apparently unaware of the description of R. marghitae from these islands. In fact, Maassen’s (2009) report of R. solorensis is entirely based on material of R. marghitae. Rensch (1931) and Maassen (2009) assigned R. floresiana as a second subspecies for its similar shell. However, Rensch (1931) indiscriminately lumped together most Lesser Sunda species while Maassen (2009) confused R. solorensis and R. marghitae. The proposals of both authors are therefore not well-substantiated. In light of generally widespread narrowrange endemism in Rhagada (Köhler, 2011; Johnson et al., 2012), I prefer maintaining the original treatment of Martens (1863, 1891) and consider R. solorensis as endemic to Solor Island. Populations from other islands should not be lumped with R. solorensis in absence of evidence from comparative studies. Consequently, R. floresiana is also maintained as a distinct species endemic to Flores.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E46FC7A3C94FC3AF9F2.taxon	diagnosis	Diagnosis. Shell comparatively large (D = 16 – 19 mm; Table 1), smooth or with fine axial ribs only, umbilicus forming a chink or narrowly open, peripheral band particularly welldeveloped (Fig. 1 A) (Martens, 1891). Penis with large, smooth penial verge with rounded tip, epiphallus with short flagellum (Fig. 2 A).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E46FC7A3C94FC3AF9F2.taxon	discussion	Comparative remarks. Shells figured by Maassen (2009) from Nusa Penida are misidentified specimens of R. marghitae. Most typical characteristic is smooth penial verge with rounded tip.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E41FC683A1FFE75FD29.taxon	discussion	Taxonomic history. While Rensch (1931) treated R. colona as subspecies of the Australian species Plectorhagada plectilis based on a similar shell, Maassen (2009) maintained it as a distinct species. Museum material from Savu Island has also been assigned to this species under an unpublished variety name (ZMB Moll. 52292).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E41FC683A1FFE75FD29.taxon	diagnosis	Diagnosis. Shell comparatively small, smooth, umbilicus forming a chink (Martens, 1877; Maassen, 2009). Genitalia unknown.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6C5E41FC683A1FFE75FD29.taxon	discussion	Remarks. Poorly characterised by reference to shell features; comparative study of anatomical and molecular characters required to confirm status.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E41FF213ED4FEFAF9B3.taxon	diagnosis	Diagnosis. Shell comparatively large (D = 16 – 19 mm), smooth or with fine axial ribs only, umbilicus forming a chink or narrowly open, peripheral band particularly welldeveloped (Martens, 1891). Comparative remarks. Shell similar to R. solorensis; species poorly defined by shell characters only; comparative study of anatomical and molecular characters required to confirm status.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E41FC663ED5FBD0F9D3.taxon	materials_examined	Material examined. AM C. 327851 (Indonesia, Nusa Penida, Goa Giri Putri, 8 ° 45 ' N, 115 ° 30 ' E, 5 wet).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E41FC663ED5FBD0F9D3.taxon	discussion	Taxonomic history. Apparently not aware of Falconieri’s description, this species was subsumed under R. solorensis by Vermeulen & Whitten (1998) and Maassen (2009). The original description focuses on characters exhibited by several species but provides little information permitting the recognition of this species.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E41FC663ED5FBD0F9D3.taxon	diagnosis	Diagnosis. Shell moderately large (D = 14 – 15.5 mm), with fine axial growth lines, umbilicus forming a chink or narrowly open. Penial verge smooth with pointed tip and well-developed sperm groove, inner penial wall smooth, epiphallus with moderately long flagellum (Fig. 2 B).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E41FC663ED5FBD0F9D3.taxon	discussion	Comparative remarks. Confused with R. solorensis by Vermeulen & Whitters (1998) and Maassen (2009), who depicted shells of this species as R. solorensis. Most typical characteristics are smooth, pointed penial verge and smooth inner penial wall.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E40FC733A7EFDF3FE8B.taxon	materials_examined	Material examined. AM C. 471156 (Indonesia, Alor Island, 3 dry).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E40FC733A7EFDF3FE8B.taxon	diagnosis	Diagnosis. Shell comparatively large (D = 16 – 19 mm), with usually well-developed axial ribs, umbilicus closed. Penis with long epiphallus, short flagellum, inner penial anatomy unknown (based on Maassen, 2009). Comparative remarks. Comparatively large shell with well-developed axial sculpture and closed umbilicus is most characteristic. Well-differentiated from R. solorensis and R. abbasi n. sp. in a molecular phylogeny based on mitochondrial COI and 16 S sequences (Fig. 3). Specimens from Atauro Island differ by consistently smaller shell, weaker sculpture, narrowly open umbilicus but are differentiated by comparatively low genetic distances (Table 2); therefore being recognised as distinct subspecies.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6B5E41FF073A5EFB05FD28.taxon	diagnosis	Diagnosis. Shell comparatively small, umbilicus forming a chink, most specimens with distinct basal ridge. Genitalia unknown (Maassen, 2009). Comparative remarks. Poorly defined by shell characters only; comparative study of anatomical and molecular characters required to confirm status.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6A5E40FF013DB6FC13FCEA.taxon	materials_examined	Type locality. Timor-Leste, Atauro Island, 2.6 km NW of Beloi, 8 ° 12 ' 08.3 " S, 125 ° 35 ' 31.9 " E. Holotype. AM C. 478282 (dissected specimen, wet, shell destroyed). Paratypes. AM C. 468712 (type locality, 23 wet), AM C. 468711 (near Makadade, 8 ° 12 ' 42.3 " S, 125 ° 34 ' 54.9 " E, 20 wet, 5 dry), AM C. 468709 (northern tip of Atauro, 8 ° 07 ' 56.3 " S, 125 ° 37 ' 59.4 " E, 7 wet, 8 dry), AM C. 470222 (2.6 km NNW of Beloi, 8 ° 12 ' 30.3 " S, 125 ° 36 ' 19.9 " E, 4 dry).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6A5E40FF013DB6FC13FCEA.taxon	etymology	Etymology. Named for Atauro Island.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF6A5E40FF013DB6FC13FCEA.taxon	description	Description. Shell moderate in size (D = 13 – 19 mm; Table 1), with faint to well-developed axial ribs, umbilicus forming a chink or narrowly open. Penial verge with circular folds, penial wall with weakly developed longitudinal pilasters, epiphallus with short flagellum (Fig. 2 C). Comparative remarks. Differs from the nominate form by smaller shell, weaker ribs, more vivid colouration, not completely closed umbilicus. Both island forms are separated by comparatively low genetic distances of about 4 % in COI and 2 % in 16 S (Table 2). Because low genetic differentiation might indicate incomplete reproductive isolation, the Atauro Island populations are described as a subspecies of R. setzeri rather than being recognised as a full species. Rhagada setzeri is well-differentiated from R. solorensis and R. abbasi n. sp. by means of genetic distances of 12 – 16 % in COI and 8 – 9 % in 16 S.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF695E42FE923F2DFD36FECA.taxon	description	(Figs 1 E – H, 2 D)	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF695E42FE923F2DFD36FECA.taxon	materials_examined	Type locality. Indonesia, Nusa Tenggara Timur Province, Lembata Island (= Lomblem), – 8 ° 25 ' S, 123 ° 28 ' E. Holotype. AM C. 478280 (dissected specimen, wet). Paratypes. AM C. 471153 (4 wet, 7 dry), AM C. 468979 (2 wet, 5 dry), AM C. 478281 (50 dry), ZMB 117967 (50 dry).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF695E42FE923F2DFD36FECA.taxon	etymology	Etymology. In honour of John Abbas, Jakarta, who kindly provided material from Lembata and Solor.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF695E42FE923F2DFD36FECA.taxon	description	Description. Shell moderate in size (D = 15 – 15 mm; Table 1), broadly subglobose, with variable reddish-brown banding on whitish to creamish background, with sculpture of faint to well-developed axial ribs, umbilicus forming a chink, lip thickened, slightly reflected (Fig. 1 E – H). Penial verge smooth with pointed tip and well-developed sperm groove, inner penial wall with well-developed longitudinal pilasters, epiphallus with longitudinal pilasters along inner wall, short flagellum (Fig. 2 D). Comparative remarks. Shell smaller than R. solorensis, R. floresiana, and R. setzeri setzeri, but similar to. R setzeri atauroensis. Most typical features are smooth penial verge with pointed tip and well-developed longitudinal pilasters of penial wall. Well-differentiated from R. solorensis and R. setzeri in a molecular phylogeny based on mitochondrial COI and 16 S sequences (Fig. 3).	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
03F3175FEF695E42FE923F2DFD36FECA.taxon	discussion	Discussion. Previous taxonomic treatments of Lesser Sunda species were predominantly based on shell characters, which were usually inferred from small series of specimens. The better known Australian Rhagada species were shown to have rather conserved shells with regard to general shape and size while exhibiting considerable variation in shell sculpture, umbilical morphology and colour (Köhler, 2011; Johnson et al., 2012). Extreme amounts of intraspecific variation in shells were observed in some island populations, but are governed by strong selection in an extreme environment (Stankowski, 2011, 2013). By contrast, the species from the Lesser Sunda Islands remain poorly documented in terms of available study material. However, where large numbers of specimens are available for study (i. e., R. setzeri atauroensis), the intraspecific variation of shells was found to correspond well with levels observed in Australian species. There is no reason to assume that other Lesser Sunda species exhibit much lower amounts of intraspecific variability. Consequently, traditionally emphasised shell characters, such as sculpture, umbilical morphology and banding pattern, may be of little taxonomic utility. Recent studies in Australian camaenids, including Rhagada, showed that comparative penial anatomy (Fig. 2) may provide valuable taxonomic information even in species that possess conserved shells (e. g., Köhler & Johnson, 2012; Criscione & Köhler, 2013 a, 2013 b). The penial anatomy of two already described species (R. solorensis, R. marghitae) and two new taxa (R. setzeri atauroensis n. sp., R. abbasi n. sp.) provided useful taxonomic information complementing the shell-based taxonomy. Molecular phylogenetic analyses based on mitochondrial sequences of the COI and 16 S genes showed that Rhagada species from the Lesser Sunda Islands form a clade, which represents the sister group of all known species from the Australian Pilbara region (Fig. 3; Köhler & Criscione, 2013). Amongst these species, R. setzeri atauroensis forms the sister group of R. setzeri setzeri from Alor. Both species are differentiated by mean p-distances of 4 % in COI and 2 % in 16 S (Table 2). Other species are differentiated by p-distances of 11 – 16 % in COI and 8 – 9 % in 16 S. These distances correspond well with amounts of interspecific variation found in Australian Rhagada species (about 4 %; Johnson et al., 2012) as well as other camaenid genera, such as Exiligada (about 3 %; Criscione et al., 2012) and Amplirhagada (about 6 %; Köhler & Johnson, 2012). However, extreme intraspecific distances of up to 10 % have occasionally also been observed in camaenid snails (Chiba, 1999; Criscione & Köhler, 2013 a). Progress has been made towards a more complete documentation of the diversity of Rhagada across the Lesser Sunda Islands. However, complete sampling on all Lesser Sunda Islands is required to address the remaining taxonomic issues and to improve our understanding of the evolution of this group throughout the entire archipelago.	en	Köhler, Frank (2014): On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands. Raffles Bulletin of Zoology 62: 115-123, DOI: 10.5281/zenodo.4502132
