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Emmericia Brusina, 1870 is the only genus of the family Emmericiidae Brusina, 1870, and it inhabits springs and rivers from north-east Italy to southern Croatia, Bosnia and Herzegovina, and Montenegro. Their variable shells have resulted in taxonomic chaos with from one to sixty seven species recognised in Emmericia. Recently four species have been distinguished: E. patula (Brumati, 1838), E. expansilabris Bourguignat, 1881, E. narentana Bourguignat, 1881, and E. ventricosa Brusina, 1870. In the present study, Emmericia from 13 localities were collected. Principal component analysis on seven biometric characters measured on shells from three localities inhabited by two nominal species demonstrated the distinctiveness of these three studied populations. In contrast, the differences between the two populations representing E. patula were not smaller than between either of them and E. expansilabris. Similar remarks apply to the shells of the other two species. The sequences of the cytochrome oxidase subunit I did not confirm that E. expansilabris is a distinct species, so it is regarded as a synonym of E. patula, but did confirm the distinctness of E. ventricosa, which inhabits the localities higher above sea level. No sequences of E. narentana were obtained.

Litthabitella levkadensis Jaszczyńska sp. nov. (Figure 3 (c)) urn:lsid:zoobank.org:act: 714ABC27-992E-454F-B8A6-1006A5BD7D2A Diagnosis Shell small, ovate conical, smaller, more slender and more thin-walled than L. chilodia. H3 sequences: KY215964–KY215967; 18S sequences: KY215952. Etymology The specific name levkadensis reflects the geographic localisation of the type locality at the island Levkada (Levkas in dimōtiki Greek). Type material Holotype. GREECE: Ethanol-fixed specimen (Figure 3 (c)); Levkas (Levkada) Island, Sivros, small spring Piges Kerasias; 38.6708°N, 20.6503°E, our locality 10 (Figure 2 (e,f)), on the gravel and fallen leaves; September 2003; Magdalena Szarowska, Andrzej Falniowski; MNHW-1441. Paratypes. No preserved paratypes, all specimens were destroyed for DNA extraction. Description Shell (Figure 3 (c)) ovate conical up to 2.38 mm tall, growing regularly. Holotype measurements: see Table 4. Teleoconch whorls slightly convex, evenly rounded. Aperture narrow, ovate, angled adapically, peristome continuous, in its upper part fitting tight to body whorl. Parietal lip complete, adnate, umbilicus nearly completely covered by lip. Outer lip fluted, orthocline. Shell white with very slightly marked growth lines, thin-walled and translucent. Measurements of the shell of holotype and sequenced specimens: see Table 4. Morphology and anatomy of soft parts. Unknown. Remarks Litthabitella levkadensis may represent the same species as the unnamed Lithabiella in paratypes of Schütt (1980). The p-distances for the histone H3 sequences (Table 3) between L. levkadensis and each of the other three species of Litthabitella are more than twice those between these three species. Distribution and habitat Known from the type locality only: the small spring Piges Kerasias, Sivros, Levkada Island (Figure 2 (e,f))., Published as part of Falniowski, Andrzej, Jaszczyńska, Aleksandra, Osikowski, Artur & Hofman, Sebastian, 2023, Litthabitellidae: a new family of the Truncatelloidea (Mollusca: Caenogastropoda), pp. 299-329 in Journal of Natural History 57 (5 - 8) on pages 311-314, DOI: 10.1080/00222933.2023.2168573, http://zenodo.org/record/7778206, {"references":["Schutt H. 1980. Zur Kenntnis griechischer Hydrobiiden. Archiv fur Molluskenkunde. 110: 115 - 149."]}

Family LITTHABITELLIDAE Falniowski and Hofman, fam. nov. urn:lsid:zoobank.org:act: A8E1F6A7-EEAE-4B8D-A82D-19A7144283C6 Type genus Litthabitella Boeters, 1970 Diagnosis Minute truncatelloid snails with the following synapomorphic character states: osphradium broadly ovate; pallial oviduct gland complex short; gonoporus deep inside the mantle cavity; the ventral channel not on the ventral side of the capsule gland, but somewhat lateral; thickened oviduct circular; the distal termination of the penis wide and blunt, with sharp and short filament harbouring the vas deferens situated laterally, and with two flat, cockscomb-like, four-folded lobes. Description Shell (Figures 3 and 4) ovate conical, convex in outline, thick-walled; the border between the proto- and teleoconch is easily discernible, the initial part of the protoconch narrow, on the protoconch surface only delicate irregularities (Figure 5 (a–c)), aperture pyriform, teleoconch smooth with delicate growth lines. Operculum (Figure 6) thin and horny, colourless and translucent, elongate-ellipsoidal, spiral, paucispiral, with submarginal nucleus. Head and tentacles white, with no pigment, no hyperciliation (long and dense cilia at one or two cephalic tentacles, characteristic of e.g. some Hydrobiidae); mantle partly black pigmented. Osphradium (Figure 7 (a)) broadly ovate; ctenidium well developed. In the radula (Figures 5 (d–i) and 8(b)) the central tooth with deep sinus at the proximal part of the cutting plate and one pair of large and prominent basal cusps. The median cusp narrow and slender, more than twice as long as the adjacent cusps, similar in shape. On each side of the median cusp are four to five cusps; the cutting edge is straight, with no cusps. In the lateral tooth, the biggest cusp is triangular, twice as long and broad as the adjacent ones. The formula of the tooth is (5)4–1–5; on the inner marginal tooth 26–28 long and slender cusps. The caecum is absent (Figure 8 (c)). The pallial oviduct gland complex (Figure 7 (a)) is relatively short, lying far posteriorly of the mantle margin, thus the gonopore is at the terminal part of the capsule gland and situated deep inside the mantle cavity, far behind the anus. The ventral channel is not on the ventral side of the capsule gland, but lies somewhat laterally (Figure 7 (b,d)). There is a bursa copulatrix and two receptacula typical of the Truncatelloidea: a distal one (rs 1 after Radoman 1983) situated near the junction of the oviduct and the duct of the bursa, and a proximal one (rs 2 after Radoman 1983) situated at the proximal end of thickened oviduct; in Litthabitella the proximal receptaculum is vestigial. The thickened oviduct is circular, forming neither a spiral nor a horseshoe-like loop (Figure 7 (c)). The prostate is small, nearly vestigial, observable as slight outgrowth of the vas deferens. The penis is big and massive (Figures 9 and 10), filled with glandular tissue (Figure 10), with a distal termination that is wide and blunt, with a sharp and short filament harbouring the vas deferens, which is situated laterally, and two flat, cockscomb-like, four-folded lobes. Molecular phylogeny We obtained 26 new sequences of COI (457 bp, GenBank accession numbers ON661272–ON661297), 16 of H3 (310 bp, GenBank accession numbers ON661565– ON661580) and 24 of 18S (401 bp, GenBank accession numbers ON667998– ON668023). The tests by Xia et al. (2003) revealed no saturation. Results from the substitution saturation analysis showed an Index of substitution saturation (ISS) (0.71 for COI; 0.48 for H3) significantly smaller than the critical ISS value (ISSC: 0.97 for COI; 0.59 for H3), indicating that all sequences are useful in phylogenetic reconstruction. In all analyses, the topologies of the resulting phylograms were identical in the ML and BI phylogram analyses; thus, we present the phylogram computed with RAxML. The COI phylogram (Figure 11) clearly shows all the populations of Litthabitella as a distinct, highly supported group (bootstrap 99%). As usual with COI, deeper nodes are not well supported; thus, the relationships among the Litthabitella remain unresolved. In the 18S phylogram of Szarowska (2006), Emmericia was a sister clade to all the Hydrobiidae (including Hydrobiinae and Sadlerianinae sensu Szarowska 2006). The time of divergence of the Litthabitellidae was estimated as 8.69 ± 1.87 Mya in the COI phylogram. Our 18S phylogram (Figure 12) shows a similar picture: a well-supported clade (bootstrap 83%) includes all the Hydrobiidae and Litthabitella, with the latter as the sister clade of all the Hydrobiidae. The set of available sequences of histone H3 was very restricted, but the relationships (Figure 13) are similar to those in the 18S phylogram; the well-supported clade (bootstrap 97%) includes Litthabitella as a sister clade of all the Hydrobiidae, but also of Heleobia (Cochliopidae). The phylogram inferred for the three concatenated loci (Figure 14) also shows the Litthabitellidae as a sister taxon to all the Hydrobiidae, but the clade joining the Litthabitellidae and Hydrobiidae is not well supported (only 62%); the clade Litthabitellidae is well supported (bootstrap 100%)., Published as part of Falniowski, Andrzej, Jaszczyńska, Aleksandra, Osikowski, Artur & Hofman, Sebastian, 2023, Litthabitellidae: a new family of the Truncatelloidea (Mollusca: Caenogastropoda), pp. 299-329 in Journal of Natural History 57 (5 - 8) on pages 305-307, DOI: 10.1080/00222933.2023.2168573, http://zenodo.org/record/7778206, {"references":["Boeters HD. 1970. Die Gattung Microna Clessin, 1890 (Prosobranchia, Hydrobiidae). Archiv fur Molluskenkunde. 100: 113 - 145.","Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Serbian Academy of Sciences and Arts, Monograph 547, Department of Sciences. 57: 1 - 256.","Xia X, Xie Z, Salemi M, Chen L, Wang Y. 2003. An index of substitution saturation and its application. Mol Phylogenet Evol. 26: 1 - 7. doi: 10.1016 / S 1055 - 7903 (02).","Szarowska M. 2006. Molecular phylogeny, systematics and morphological character evolution in the Balkan Rissooidea (Caenogastropoda). Folia Malacol. 14 (3): 99 - 168. doi: 10.12657 / folmal. 014.014."]}

Litthabitella cetinensis Osikowski et Jaszczyńska sp. nov. (Figures 3 (b), 4 (t–v), 5 (c,f,i), 9(f)) urn:lsid:zoobank.org:act: 10089647-36C6-4773-8E7F-AA0D5BB02833 Diagnosis Shell small, ovate conical, distinguished from L. chilodia and L. aquaadcrucem by smaller and less thick-walled shell and penis with less broad tip and longer and broader filament. COI sequences: ON661294–ON661297; H3 sequences: KY215954–KY215958; 18S sequences: ON668020–ON668023. COI MDCs: 6, A; 21, G; 39, A; 51, C; 84, G; 90, C; 111, A; 132, G; 135, A; 144, T; 153, C; 171, G; 174, C; 183, G; 219, G; 276, C; 309, C; 369, G; 399, T; 402, C; 436, C; 438, A. Etymology The specific name cetinensis reflects the geographic localisation of the type locality within the valley of Cetina River. Type material Holotype. CROATIA: ethanol-fixed specimen (Figure 3 (b)); Kostanje, a medium-sized spring at the right bank of the Cetina River; 43.4411°N, 16.8228°E, 133 m asl; our locality 1 (Figure 2 (a)); June 2011; leg. Magdalena Szarowska, Andrzej Falniowski; MNHW-1440. Paratypes. CROATIA • 10 specs (ethanol-fixed); same collection data as for holotype, DMJU 2212. Description Shell (Figure 3 (b)) ovate conical up to 2.02 mm tall, growing regularly. Holotype measurements: see Table 4. Teleoconch whorls slightly convex, evenly rounded. Aperture narrow, ovate, weakly angled adapically, peristome continuous, in its upper part fitting tight to body whorl. Parietal lip complete, adnate, umbilicus nearly completely covered by lip. Outer lip fluted, orthocline. Shell white and glossy with very slightly marked growth lines, moderately thick-walled and translucent, periostracum white. Initial part of the protoconch narrow, on the protoconch surface only delicate irregularities (Figure 5 (c)). Measurements of holotype and sequenced and illustrated shells: see Table 4. Shell variability slight (Figure 4 (t–v)). Morphology and anatomy of soft parts. The head with eyes and no pigment; mantle pigmented less intensively than in the other Litthabitella. Ctenidium with 14 short lamellae, the osphradium broad. Radula (Figure 5 (f,i)) typical of Litthabitella. Penis (Figure 9 (f)) with rather narrow tip and longer and broader filament than in the other Litthabitella species. Distribution and habitat Known from the type locality only: Kostanje, a medium-sized spring at the right bank of the Cetina River (Figure 2 (a))., Published as part of Falniowski, Andrzej, Jaszczyńska, Aleksandra, Osikowski, Artur & Hofman, Sebastian, 2023, Litthabitellidae: a new family of the Truncatelloidea (Mollusca: Caenogastropoda), pp. 299-329 in Journal of Natural History 57 (5 - 8) on pages 309-311, DOI: 10.1080/00222933.2023.2168573, http://zenodo.org/record/7778206, {"references":["Biltekin D. 2010. Vegetation and climate of north anatolian and north aegean region since 7 Ma according to pollen analysis. Earth Sciences. Lyon: Universite Claude Bernard."]}

Litthabitella aquaadcrucem Jaszczyńska et Osikowski sp. nov. (Figures 3 (a), 4 (m–s), 5 (b,e,h), 7, 8, 9 (b–e), 10) urn:lsid:zoobank.org:act: 2ABE7402-A94B-4289-BD15-E98F5C147B2E Diagnosis Shell small, ovate conical, distinguished from similarly high and broad L. chilodia by a proportionally lower aperture and thus higher spire, less weakly angled adapically, and less broad tip of the penis. COI sequences: ON661272–ON661278; H3 sequences: ON661565–ON661570; 18S sequences: ON667998–ON668004. COI MDCs: 12, C; 33, C; 66, A; 93, A; 115, T; 120, G; 159, G; 174, T; 177, T; 222, T; 225, G; 354, C; 363, C; 393, G; 399, G; 402, A; 408, C; 432, G; 453, G. Etymology The specific name aquaadcrucem is the Latin phrase ′ aqua ad crucem ̍, which means ′water at the cross̍, and is a translation of the Serbian Voda u Križa, which is the name of the spring. It should be considered a noun in apposition. Type material Holotype. MONTENEGRO: ethanol-fixed specimen (Figure 3 (a)); W of Sotonići, small spring Voda u Križa near the restaurant of the same name; 42.2367°N, 19.0439°E; 183 m a.s.l.; our locality 8 (Figure 2 (d)), on the gravel; July 2005; Magdalena Szarowska, Andrzej Falniowski; MNHW-1439. Paratypes. MONTENEGRO: 20 specimens (ethanol-fixed); same collection data as for holotype, DMJU 2211. Description Shell. (Figure 3 (a)) Ovate conical up to 2.51 mm high, whorls growing regularly. Holotype measurements: see Table 4. Teleoconch whorls slightly convex, evenly rounded. Aperture narrow, ovate, weakly angled adapically, peristome continuous, in its upper part fitting tight to body whorl. Parietal lip complete, adnate, umbilicus nearly completely covered by lip. Outer lip fluted, orthocline. Shell glossy with very slightly marked growth lines, thickwalled and translucent, periostracum whitish. Initial part of the protoconch narrow, on the protoconch surface only delicate irregularities (Figure 5 (b)). Measurements of holotype and sequenced and illustrated shells: see Table 4. Shell variability slightly marked (Figure 4 (m–s)). Morphology and anatomy of soft parts. The head with eyes and no pigment; mantle intensively black pigmented. Ctenidium with 19 short lamellae (in five examined specimens), the osphradium broad (Figure 7 (a)). Radula (Figure 5 (e,h)) typical of Litthabitella. Anus long forward from the female gonopore (Figure 7 (a)). Female reproductive organs (Figure 7 (a–d)) as described above for the family. Penis (Figure 9 (b–e)) as described above for the family, with less broad tip than in L. chilodia (Figure 9 (a)). Distribution and habitat Known from the type locality (8), the small spring Voda u Križa near the restaurant of the same name (Figure 2 (d)), and localities 6 and 9 (Figure 1 and Table 1)., Published as part of Falniowski, Andrzej, Jaszczyńska, Aleksandra, Osikowski, Artur & Hofman, Sebastian, 2023, Litthabitellidae: a new family of the Truncatelloidea (Mollusca: Caenogastropoda), pp. 299-329 in Journal of Natural History 57 (5 - 8) on pages 308-309, DOI: 10.1080/00222933.2023.2168573, http://zenodo.org/record/7778206