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Mercuria carrillorum Miller, García-Guerrero & Ramos sp. nov. urn:lsid:zoobank.org:act: E9ED5EC6-E8DC-42ED-8EDC-2D2322296196 Figs 20–22; Supp. file 2: Tables S14–S18 Diagnosis Shell ovate-conic; periostracum dark brown to blackish; protoconch microsculpture granulated; aperture obliquely ovate; central radular tooth formula (3)4-C-4(3)/1-1; female genitalia with bursa copulatrix elongate, ca 3 times as long as wide; seminal receptacle elongate; distal end of the penis ca 1.5 times as long as penial appendix, darkly pigmented; penial appendix ovate, pigmented to the distal part, medially positioned on the inner edge of the penis; nervous system pigmented, elongate (mean RPG ratio = 0.63); cerebral ganglia approximately equal in size. Etymology Named after the family of Miguel Carrillo Pacheco who instilled his scientific curiosity and passion for malacology and in recognition for his help with the fieldwork conducted across Spain, France and Portugal. Type material Holotype SPAIN • sex unknown (dry preserved); Cádiz, Jerez de la Frontera, Canuto de la Gallina, Montes de Propios; MNCN 15.05/200175H. Paratypes SPAIN • 33 specs (preserved in ethanol 80%, 10 dissected and 4 processed for DNA sequencing); same collection data as for holotype; MNCN 15.05/94748 • 16 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/94750 • 17 specs (preserved in ethanol 80%); same collection data as for holotype; MZB 2021-2796 • 2 specs (dry preserved); same collection data as for holotype; NHMW 113527 • 2 specs (dry preserved); same collection data as for holotype; RMNH. MOL.507924 • 1 spec. (dry preserved); same collection data as for holotype; JPM-586 • 2 specs (dry preserved); same collection data as for holotype; MCP • 2 specs (dry preserved); same collection data as for holotype; MCP. Additional material examined SPAIN – Cádiz • 30 specs; Jerez de la Frontera, Montes de Propios, stream in Canuto de las Palas; MNCN 15.05/94751 • 3 specs; same collection data as for preceding; MNCN 15.05/94749. – Málaga • 3 specs; Manantial de los Doce PIlares spring; MNCN-ADN 110673 to 110675. Additional locality information provided in Supp. file 1: Table S1. Type locality Stream in Canuto de la Gallina, Montes de Propios, Jerez de la Frontera, Cádiz, Spain. 36.587509° N, 5.574243° W; 379.72 m a.s.l. Description MEASUREMENTS. Holotype: SL = 3.04 mm, SW = 2.26 mm, SL/SW = 1.34, AH = 1.43 mm, SL-LBW = 0.73 mm, WBW = 1.78 mm, AL = 1.44 mm, AW = 1.07 mm, WPW = 1.03 mm, WAW = 0.52 mm. SHELL. Ovate-conic, whorls 4–5, height 2.4–3.8 mm, width 1.8–2.7 mm (Fig. 20A–F; Supp. file 2: Table S14); periostracum dark brown to blackish; protoconch of 1.5 whorls, ca 350 µm wide, nucleus ca 150 µm wide (Fig. 21A–B); protoconch microsculpture granulated (Fig. 21C). Teleoconch whorls slightly convex, separated by a deep suture; body whorl large and convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip slightly thicker than outer lip; aperture margin straight; outer lip slightly sinuated; umbilicus narrow, not covered by the inner lip (Fig. 20A, C–F). OPERCULUM. As for the genus, orangish to brown, about 2 whorls; muscle attachment oval, located near the nucleus (Fig. 20G–H). RADULA. Length intermediate, ca 600 µm long (20% of total shell length), containing about 70 rows of teeth. Central tooth formula (3)4-C-4(3)/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 21D–E). Lateral tooth formula (4)3-C-3(4), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 12–16 cusps; outer marginal teeth with 25–27 cusps (Fig. 21F). Radular data were collected from the following specimens: MNCN 15.05/94748 and MNCN 15.05/94750 from Canuto de la Gallina, Cádiz and MNCN 15.05/94749 and MNCN 15.05/94751 from Canuto de las Palas, Cádiz. PIGMENTATION AND ANATOMY. Animal brownish to pale brown (Fig. 22E); head and tentacles brown, eye lobes unpigmented; snout and neck weakly pigmented; snout longer than wide, approximately parallel-sided, with medium distal lobation. Pallial tentacle present. Ctenidium occupying almost the total length of the pallial cavity; 22–26 gill filaments, broad, triangular, and fused at the base by an epithelium (Fig. 22G). Osphradium elongate, more than 3 times as long as wide, positioned opposite to the middle of the ctenidium. Stomach almost as long as wide with two chambers about equal in size (Supp. file 2: Table S16); style sac longer than wide, with the unpigmented intestine surrounding its distal part before continuing on as a straight rectum (Fig. 22H). MALE GENITALIA. Prostate gland small, bean-shaped, about 2 times as long as wide, connected by the posterior vas deferens to a convoluted seminal vesicle and the testis (Fig. 22F). The distal end of the penis ca 1.5 times as long as penial appendix (Supp. file 2: Table S18), darkly pigmented, gradually tapering, attached to the neck behind the right eye; penial appendix ovate, pigmented on the distal part, medially positioned on the inner edge of the penis (Fig. 22C–E). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide; albumen gland longer than capsule gland (Fig. 22A); bursa copulatrix elongate, ca 3 times as long as wide (Supp. file 2: Table S17); bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle elongate, with a short duct, positioned on the distal part of the renal oviduct at the junction with the bursal duct (Fig. 22B). NERVOUS SYSTEM. Pigmented, elongate (mean RPG ratio = 0.63; Supp. file 2: Table S15); cerebral ganglia approximately equal in size; pleuro-supraoesophageal connective ca 9 times as long as pleuro-suboesophageal one (Fig. 22I). Ecology and distribution Restricted, as the species has been found at only three localities, all in Cádiz and Málaga, Spain. Very small specimens (probably juveniles) of the species were collected for the first time at Manantial de los Doce Pilares Spring, Málaga in 2015. However, we were unable to find any specimens at this locality during a subsequent sampling trip in 2017. Contrary to other species of Mercuria, M. carrillorum sp. nov. lives in streams with very low conductivities, ranging from 80–179 µS/cm. Co-occurring species, though at low abundances, are Ancylus fluviatilis O.F. Müller, 1774 and Galba truncatula (O.F. Müller, 1774), except the stream in Canuto de la Gallina in which only M. carrillorum is found (Félix Ríos Jiménez pers. com.). Remarks Mercuria carrillorum sp. nov. differs from the phylogenetically closely related species M. egarensis sp. nov. by having a more ovate than a high-spired shell, a smaller and more pigmented penial appendix, an elongated bursa copulatrix (from pyriform to elongate in M. egarensis), a smaller radular ribbon, a larger number of cusps on the central radular tooth and the presence of a pallial tentacle. Mercuria carrillorum sp. nov. resembles M. felixi sp. nov. and M. balearica in terms of shell shape (Fig. 3) but differs from them in other features. The new species has an ovate, pigmented penial appendix that is shorter than the distal end of the penis, whereas in M. felixi, the distal end of the penis and appendix are about equal in length. In M. balearica, the penial appendix is longer than the distal end of the penis, which is triangular and extremely pigmented. These species also differ in the lengths of the bursa copulatrix (longer in Mercuria carrillorum than in M. balearica) and the pleuro-supraoesophageal connective (longer in M. felixi). The major difference in the radula is the number of cusps on the outer marginal teeth: M. carrillorum has considerably more cusps (25–27) than M. balearica and M. felixi. The sequence divergence of COI for Mercuria carrillorum sp. nov. and the other Iberian congeners ranged between 1.3% (with M. egarensis sp. nov.) and 8% (with M. felixi sp. nov.). With a divergence of 8.9%, the most distantly related species was the Tunisian M. saharica Letourneux & Bourguignat, 1887 (Miller et al. 2022), Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 33-38, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Letourneux A. & Bourguignat J. R. 1887. Prodrome de la Malacologie terrestre et fluviatile de la Tunisie. Imprimerie nationale, Paris. https: // doi. org / 10.5962 / bhl. title. 132280","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

Mercuria similis (Draparnaud, 1805) Figs 4–7; Supp. file 2: Tables S2–S5 Cyclostoma simile Draparnaud, 1805: 34, pl. 1 fig 15. Bythinia meridionalis Risso, 1826: 100. Amnicola confusa Frauenfeld, 1863: 1029. Amnicola emiliana Paladilhe, 1869: 229, pl. 19 figs 22–23; 106, pl. 5. Amnicola compacta Paladilhe, 1869: 110, pl. 19 figs 14–15. Paludina cerulea Massot, 1872: 128. Amnicola roigiana Salvañá, 1887: 141. Amnicola monjoi Chia, 1887: 14. Amnicola vallensana Almera & Bofill, 1898: 83; pl. III fig. 23. Paludina similis – Turton & Gray 1840. Hydrobia similis – Dupuy 1851: 553. Pseudamnicola confusa – Adam 1940: 1–7, figs 1–2. Mercuria confusa – Boeters 1971: 178–179, fig. 10. Pseudamnicola similis – Gasull 1971: 45; pl. II fig. 8. Mercuria emiliana – Boeters 1988: 208, figs 92–93; 210, figs 118, 125; 211, pl. 3 fig. 34. non Mercuria meridionalis (Risso, 1826) – Girardi 2003: 83. non Pseudamnicola emilianus (Paladilhe, 1869) – Boeters & Falkner 2017: 251, fig 11i–j. Revised diagnosis Shell ovate-conic; aperture obliquely broad ovate; periostracum whitish to grey; protoconch microsculpture granulated; central radular tooth formula (2)3-C-3(2)/1-1; female genitalia with bursa copulatrix pyriform to elongate, ca 3 times as long as wide; seminal receptacle elongate with a short duct; penis darkly pigmented, gradually tapering; penial appendix triangular, variable in length (equal to or shorter/longer than the distal end of the penis), strongly pigmented at the junction with the penis; distal end of the penis triangular; nervous system pigmented, elongate (mean RPG ratio = 0.63); cerebral ganglia approximately equal in size. Type material (not examined) Neotype FRANCE • sex unknown; “Gallia meridionalis” [South France]; NHMW-MO 92596; designated by Boeters & Falkner (2000). Type locality According to original description: “Gallia meridionalis”. Material examined FRANCE – Languedoc-Roussillon • 30 specs; Salses-le-Château, Font Dame Spring; MNCN 15.05/94776 • 30 specs; Salses-le-Château, Font d‘Estramar Spring; MNCN 15.05/94777 • 30 specs; pond in La Palme; MNCN 15.05/94778. – Provence-Alpes-Côte d’Azur • 30 specs; Étang de Berre, stream near La Suriane; MNCN 15.05/94789 • 30 specs; Étang de Berre, Arc River near Les Cabanes; MNCN 15.05/94790 • 30 specs; Var, La Foux-de-Draguignan Spring; MNCN 15.05/94791. SPAIN – Granada Prov. • 30 specs; La Malahá Spring; MNCN 15.05/94758 • La Pucha Spring; MNCN 15.05/94759. – Cádiz • 30 specs; Prado del Rey, Pilar de los Playeros Spring; MNCN 15.05/94760 • 30 specs; Villamartín, La Zarza Spring; MNCN 15.05/94761 • 30 specs; Medina-Sidonia, El Berrueco Spring; MNCN 15.05/94803 • 30 specs; Arcos de la Frontera, Platero Spring; MNCN 15.05/94795 • 30 specs; Puerto Serrano, Pozo Amargo Spring; MNCN 15.05/94796 • 30 specs; spring in Alcalá de los Gazules; MNCN 15.05/94797 • 30 specs; Algodonales, El Algarrobo Spring; MNCN 15.05/94799 • 30 specs; Medina-Sidonia, La Salá Spring; MNCN 15.05/94800-2546; 30 specs; Alcalá de los Gazules, Las Presillas Spring; MNCN 15.05/94802. – Guadalajara • 30 specs; saltings of Riba de Santiuste; MNCN 15.05/94762 • 30 specs; Alcolea Stream, tributary of Salado River, road from Imón to Santamera; MNCN 15.05/94763 • 30 specs; Salado River at Santamera; MNCN 15.05/94764, 15.05/94765 • 30 specs; La Vega Stream at Saelices de la Sal; MNCN 15.05/94766. – Almería • 30 specs; Las Negras ravine; MNCN 15.05/94767. – Albacete • 30 specs; Cordovilla saltings; MNCN 15.05/94768 • 30 specs; Casas de Ves, La Salaboreja Spring; MNCN 15.05/94769 • 30 specs; Casas de Ves, La Cañada Stream near La Salaboreja Spring; MNCN 15.05/94770 • 30 specs; Fuentealbilla, stream feeding Galayo’s Pond; MNCN 15.05/94771. – Zaragoza • 30 specs; Gelsa, stream in Barranco del Agua Salada; MNCN 15.05/94772. – Huesca • 30 specs; saltwater stream near Aguinaliu; MNCN 15.05/94773 • 30 specs; stream crossing the town of Peralta de la Sal; MNCN 15.05/94774 • 30 specs; Torres del Obispo Spring; MNCN 15.05/94794. – Tarragona • 30 specs; Amposta, Ullals de Baltasar; MNCN 15.05/94780. – Murcia • 30 specs; Chícamo Stream near La Umbría; MNCN 15.05/94781 • 30 specs; La Mula River; MNCN 15.05/94782 • 30 specs; river in Sierra de la Muela; MNCN 15.05/94742 • 30 specs; Puerto de la Cadena; MNCN 15.05/94793. – Castellón • 30 specs; Fuente Amarga Spring; MNCN 15.05/94783 • 30 specs; marsh at Peñíscola; MNCN 15.05/94786 • 30 specs; irrigation ditch in Moli de la Font; MNCN 15.05/94787 • 30 specs; irrigation ditch in Cirat; MNCN 15.05/94788. – Valencia • 30 specs; Sax, Vinalopó River; MNCN 15.05/94784. – Alicante • 30 specs; Braña’s Ravine, near L’Olla Beach; MNCN 15.05/94785. – Islas Baleares • 30 specs; Majorca, Albufera of Majorca, Siquia de Son Senyor; MNCN 15.05/94805 • 30 specs; Majorca, Albufera of Majorca, Siquia d’en Moix; MNCN 15.05/94806 • 30 specs; Majorca, Muro, Font de Son Sant Joan; MNCN 15.05/94804. – Barcelona • 30 specs; El Prat de Llobregat, Estany de la Ricarda Pond; MNCN 15.05/94779. Additional locality information provided in Supp. file 1: Table S1. Description SHELL. Ovate-conic, whorls 4–5, height 3.5–5.7 mm, width 1.5–3.3 mm (Fig. 4 A–O; Supp. file 2: Table S2); periostracum whitish to grey; protoconch of 1.5 whorls, ca 400 µm wide, nucleus ca 200 µm wide (Fig. 5A–B); protoconch microsculpture granulated (Fig. 5C); teleoconch whorls very convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin straight, inner lip touching the shell wall; umbilicus narrow, not covered by the inner lip. OPERCULUM. As for the genus, orange to brown, sometimes yellowish, about two whorls; muscle attachment oval, located near the nucleus (Fig. 4P–Q). RADULA. Length intermediate, ca 800 µm long (35% of total shell length), containing about 65 rows of teeth. Central tooth formula (2)3-C-3(2)/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 5D–F). Lateral tooth formula (3)2-C-2(3), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 11–15 cusps (Fig. 5E); outer marginal teeth with 12–25 cusps (Fig. 5F). Radular data were collected from the following specimens: MNCN 15.05/94760 – Spring Pilar de los Playeros, Prado del Rey, Cádiz, Spain; MNCN 15.05/94767 – Las Negras ravine, Almería, Spain; MNCN 15.05/94768 – Cordovilla Saltings, Albacete, Spain; MNCN 15.05/94776 – Fonte Dame Spring, Salses-le-Château, Aude, France; MNCN 15.05/94777 – Estramar Spring, Salses-le-Château, Aude, France; MNCN 15.05/94778 – pond in La Palme, Aude, France; MNCN 15.05/94791 – La Foux-de-Draguignan Spring, France and 15.05/94804 – Font de Son Sant Joan, Muro, Majorca, Spain. PIGMENTATION AND ANATOMY. Animal darkly pigmented, although unpigmented specimens were also found (Fig. 4E); head and tentacles black, pigmentation lighter on eye lobes, snout and neck; snout about as long as wide, approximately parallel-sided, with medium distal lobation (Fig. 7F). Ctenidium occupying almost the total length of the pallial cavity; 22–27 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 6E). Pallial tentacle present. Osphradium elongate, more than 3 times as long as wide (Supp. file 2: Table S3), positioned opposite middle of ctenidium. Stomach almost as long as wide with two chambers almost equal in size; style sac longer than wide (Supp. file 2: Table S3), with the unpigmented intestine surrounding its distal end before continuing on as a straight rectum (Fig. 6F). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide; albumen gland longer than capsule gland (Fig. 6A–D; Supp. file 2: Table S4); bursa copulatrix pyriform to elongate, ca 3 times as long as wide; bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle elongate, with a short duct, positioned at the distal end of the renal oviduct just above the junction with the bursal duct (Fig. 6A–D). MALE GENITALIA. Penis darkly pigmented, gradually tapering, attached to the neck behind the right eye; penial appendix longer (Fig. 7A–C) or shorter (Fig. 7D–F) than the distal end of the penis, triangular, strongly pigmented at the junction with the penis, pigmentation gradually weakens from the junction to the middle of the penial appendix where it is very weak. Penial appendix base narrow, medially positioned on the inner edge of the penis. Prostate gland bean-shaped, about 2 times as long as wide, connected by the posterior vas deferens to a convoluted seminal vesicle and the testis (Fig. 7G–H; Supp. file 2: Table S5). NERVOUS SYSTEM. Pigmented, elongate (mean RPG ratio = 0.63; Supp. file 2: Table S15); cerebral ganglia approximately equal in size; pleuro-supraoesophageal connective ca 9 times as long as pleuro-suboesophageal one (Fig. 6G). Ecology and distribution Most of the examined specimens were found in waters with very high conductivities (592–28 900 µS/cm, average 6364 µS/cm). These localities are affected by the Keuper facies, Mesozoic evaporitic deposits that contain high levels of NaCl and CaSO 4, among others, that dissolve into the superficial waters inhabited by the species. Specimens were most often found in the mud among the lower parts of the shoreline vegetation. Co-occurring gastropod species are Melanopsis spp., Theodoxus spp., Belgrandia gibba (Draparnaud, 1805), Hydrobia acuta (Draparnaud, 1805), Pseudamnicola subproductus (Paladilhe, 1869), Diegus gasulli (Boeters, 1981) and Potamopyrgus antipodarum (Gray, 1843). The species is distributed in springs, streams, small rivers, coastal lakes (étangs) and ditches near the Mediterranean coast of southern France, the Iberian Peninsula and North Africa (Boulaassafer et al. 2018) and on the island of Majorca (Fig. 1). It has also been reported near the Atlantic coast of the Iberian Peninsula, northern France, Apennine peninsula, Great Britain and various Atlantic islands (Wollaston 1878; Kerney 1999; Bodon et al. 2005; Kadolsky 2011). We were unable to confirm previous records of the species for Great Britain (Baker et al. 1999; Abrehart & Forster 2012), the Azores islands (Wollaston 1878) and North Africa (Taybi et al. 2017). Some of these citations may have been incorrectly attributed to M. similis instead of M. tachoensis, which is found in, at least, Great Britain (see below). Remarks The material here referred to as M. similis from southern France (Fig. 4I–K, N–O) closely conforms to the neotype of this species in terms of shell shape, size and colouration (for comparison with the type material, see Boeters & Falkner (2017: fig. 9) or Eschner et al. (2020: fig. 1t)). Based on our morphological and previous phylogenetic results (Miller et al. 2022), we suggest that several populations previously assigned to other species of Mercuria should now be considered as members of M. similis (see discussion), which increases the known geographic range and intraspecific variation of this species. The species’ moderate shell variation can be observed across its geographic range (Figs 1, 4). However, the PCA displayed all populations identified as M. similis in proximity, suggesting a low level of intraspecific variation in shell shape. Most of the variation is attributed to differences in colour pattern and shell size. Although some individuals are orangish-brown [e.g., those from the La Cañada Stream (MNCN 15.05/94770), stream feeding Galayo’s Pond (MNCN 15.05/94771), the stream in Peralta de la Sal (MNCN 15.05/94774) and the Arc River near Les Cabanes (MNCN 15.05/94790), Fig. 4B–C, G, N] the majority of the specimens have a whitish-grey periostracum. Another character that affects the perception of shell colour is the colouration of the epithelium. Most specimens have a darkly pigmented epithelium, though a few have an unpigmented epithelium [e.g., individuals from Barranco del Agua Salada (MNCN 15.05/94772), Fig. 4E]. Unpigmented individuals were found cohabiting with pigmented ones in MNCN 15.05/94772. Additionally, the species presents moderate variability in shell size both within and among populations (Supp. file 2: Table S2), varying from 3.00– 4.84 mm, although larger specimens (5.4–5.7 mm) were found in the saltings in Riba de Santiuste, Guadalajara (MNCN 15.05/94762). This variability is likely influenced by environmental factors, such as flow rate and turbidity (Verhaegen et al. 2018), and by the relative age of the animal. Hydrobiids are generally univoltine (i.e., have one generation per year) and semelparous (i.e., breed only once in a lifetime). In the case of the population from Riba de Santiuste, we found animals from a previous generation living together with adults from the current one. The older specimens (i.e., those with a greater number of whorls) collected from this population were indeed the larger-sized adults. Major anatomical differences are observed in penis features, with three penis morphotypes found in the populations examined: 1-distal end of the penis longer than the penial appendix (Fig. 7A–B), 2-distal end of the penis and penial appendix about the same length (Fig. 7C), 3-distal end of the penis shorter than the penial appendix (Fig. 7E–F). This variation was observed within individual populations and thus does not correspond with any geographic distribution pattern. Holyoak et al. (2017) described the same phenomenon for M. tachoensis. These authors attributed the variation to allometric changes in the sexual maturity of the animals that occur during the reproductive season. Boulaassafer et al. (2018) found high variability of penis features in parasitized specimens of M. tensiftensis. Nevertheless, in our case, none of the specimens of M. similis dissected were parasitized. Mercuria similis can be distinguished from the phylogenetically closest species M. egarensis sp. nov. and M. carrillorum sp. nov. by its larger, more globose shell (Fig. 3). It can be distinguished from the geographically closest species M. balearica by its larger, more globose shell and also its slightly smaller, often pigmented, penial appendix and from M. tachoensis by its distant geographic distribution (M. tachoensis lives in Atlantic coastal zones, whereas M. similis inhabits Mediterranean areas), granulated protoconch microsculpture and radula, which present one cusp less on the central and lateral teeth., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 8-15, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Draparnaud J. P. R. 1805. Histoire naturelle des Mollusques terrestres et fluviatiles de la France. Chez Louis Colas, Paris. https: // doi. org / 10.5962 / bhl. title. 12856","Risso A., 1826. Histoire naturelle des Principales Productions de l'Europe meridionale: Principalement de celles des Environs de Nice et des Alpes-Maritimes. Chez F. G. Levrault, Paris. https: // doi. org / 10.5962 / bhl. title. 58984","Frauenfeld G. 1863. Vorlaufige Aufzahlung der Arten der Gattungen Hydrobia Htm. und Amnicola Gld. Hldm. in der kaiserlichen und in Cuming's Sammlung. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 13: 1017 - 1030.","Paladilhe A. 1869. Nouvelles miscellanees malacologiques. Chez Savy, libraire-editeur, Paris. https: // doi. org / 10.5962 / bhl. title. 12995","Massot P. 1872. Enumeration des Mollusques terrestres et fluviatiles du departement des Pyrenees-Orientales. Bulletin de la Societe agricole, scientifique et litteraire des Pyrenees-Orientales 19: 1 - 116.","Salvana J. M. 1887. Moluscos nuevos de Espana. Cronica Cientifica de Barcelona 10: 137 - 142.","Chia M. 1887. Nota de los Moluscos Terrestres y de Agua Dulce de los Alrededores de Barcelona. Impr. y Libr. de Paciano Torres.","Almera J. & Bofill A. 1898, Moluscos fosiles recogidos en los terrenos pliocenicos de Cataluna. Boletin de la Comision del Mapa Geologico de Espana 24: 1 - 223.","Turton W. & Gray J. E. 1840. A Manual of the Land and Fresh-water Shells of the British Islands. Longman, Orme, Brown, Green, and Longmans, London. https: // doi. org / 10.5962 / bhl. title. 14403","Dupuy D. 1851. Histoire naturelle des Mollusques terrestres et d'eau douce qui vivent en France fasc. 5. V. Masson, Paris. https: // doi. org / 10.5962 / bhl. title. 46522","Adam W. 1940. Notes sur les Gasteropodes, 8. Sur la presence de Pseudamnicola confusa (Frauenfeld, 1863) en Belgique. Bulletin du Musee royal d'histoire naturelle de Belgique 16: 1 - 7.","Boeters H. 1971. Pseudamnicola Paulucci, 1878 und Mercuria n. gen. (Prosobranchia, Hydrobiidae). Archiv fur Molluskenkunde 101: 175 - 181.","Gasull L. 1971. Fauna malacologica de las aguas continentales dulces y salobres del sudeste iberico. Boletin de la Sociedad de Historia Natural de Baleares 16: 23 - 93.","Boeters H. 1988. Moitessieriidae und Hydrobiidae in Spanien und Portugal (Gastropoda: Prosobranchia). Archiv fur Molluskenkunde 118: 181 - 261.","Girardi H. 2003. Anatomie et biometrie de Mercuria similis (Draparnaud, 1805) (Gastropoda: Hydrobiidae) du Languedoc-Roussillon-France. Documents Malacologiques 4: 83 - 86.","Boeters H. D. & Falkner G. 2017. The genus Mercuria Boeters, 1971 in France (Gastropoda: Caenogastropoda: Hydrobiidae). West-European Hydrobiidae, Part 13. Zoosystema 39: 227 - 261. https: // doi. org / 10.5252 / z 2017 n 2 a 4","Boeters H. D. & Falkner G. 2000. Beitrage zur Nomenklatur der europaischen Binnenmollusken, XII. Cyclostoma simile Draparnaud 1805 and Amnicola confusa Frauenfeld 1863. Heldia 3: 37 - 40.","Boulaassafer K., Ghamizi M. & Delicado D. 2018. The genus Mercuria Boeters, 1971 in Morocco: first molecular phylogeny of the genus and description of two new species (Caenogastropoda, Truncatelloidea, Hydrobiidae). ZooKeys 782: 95 - 128. https: // doi. org / 10.3897 / zookeys. 779.26797","Wollaston T. V. 1878. Testacea Atlantica: or the Land and Freshwater Shells of the Azores, Madeiras, Salvages, Canaries, Cape Verdes, and Saint Helena. L. Reeve & Company, London. https: // doi. org / 10.5962 / bhl. title. 10705","Kerney M. P. 1999. Atlas of Land and Freshwater Molluscs of Britain and Ireland. Harley Books, Brill.","Bodon M., Cianfanelli S., Manganelli C., Pezzoli E. & Giusti F. 2005. Checklist e distribuzione della fauna italiana. 10.000 specie terrestri e delle acque interne. In: Ruffo S. S. (ed.) Memorie del Museo Civico di Storia Naturale di Verona, 2 ° Serie, Sezione Scienze della Vita: 79 - 81. Ministerio dell´Ambiente, Verona.","Kadolsky D. 2011. Nomenclatural comments on non-marine molluscs occurring in the British Isles. Journal of Conchology 41: 65 - 90.","Baker R., Clarke K. & Howlett D. 1999. A survey of the broadland distribution of Pseudamnicola confusa (Frauenfeld). English Nature Research Reports 319: 1 - 21.","Abrehart T. & Forster S. 2012. Ecological Surveys of Saline Lagoons on the Benacre NNR and Suffolk Coast NNR February 2012. An ecological survey including floral and fauna observations undertaken for Natural England. Natural England Report.","Taybi A., Mabrouki Y., Ghamizi M. & Berrahou A. 2017. The freshwater malacological composition of Moulouya's watershed and Oriental Morocco. Journal of Materials and Environmental Science 8: 1401 - 1416.","Eschner A., Vinarski M. & Schnedl S. 2020. Addendum to the examination of the type material of freshwater mollusk species described by J. P. R. Draparnaud. Annalen des Naturhistorischen Museums in Wien B 122: 183 - 191.","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524","Verhaegen G., McElroy K. E., Bankers L., Neiman M. & Haase M. 2018. Adaptive phenotypic plasticity in a clonal invader. Ecology and Evolution 8: 4465 - 4483. https: // doi. org / 10.1002 / ece 3.4009","Holyoak D. T., Holyoak G. A. & da Costa Mendes R. M. 2017. Distribution and ecology of Mercuria tachoensis (Gastropoda: Hydrobiidae) in Portugal and evidence that M. edmundi is conspecific. Iberus 35: 1 - 8."]}

Mercuria maceana (Paladilhe, 1869) Figs 15–16 Amnicola maceana Paladilhe, 1869: 103. Mercuria maceana – Bank & Neubert 2017: 16. Revised diagnosis Shell ovate-conic; aperture obliquely broad ovate; periostracum yellowish to pale grey; protoconch microsculpture granulated; central radular tooth formula 3-C-3/1-1. Type material (not examined) Syntypes SPAIN • 1 spec. (sex unknown); Barcelona, C’an Tunis; MHNG-MOLL-105438/2 • 1 spec. (sex unknown); same collection data as for preceding; MHNL 45013950 • 1 spec. (sex unknown); same collection data as for preceding; UM.PDL.006 (Breure & Audibert 2017). Type locality C’an Tunis, Barcelona, Spain. The type locality of this species in the original description “récoltée à Antunez, près de Barcelone” refers to a series of local geographic names for a private property: C’an Tunis, Casa Antúnez, Camp de la Bota, Llano del Llobregat. These names have been written either conserving the original language, Catalan, sometimes using the Spanish form, or even using other local names for the same place. Material examined SPAIN • 30 specs; Barcelona, C’an Tunis; MNCN 15.05/7009 • 30 specs; same collection data as for preceding; MZB 77-8445, MZB 77-8107. Description SHELL. Ovate-conic, whorls 4–5, height 3.9–5.2 mm, width 3–3.8 mm (Fig. 15A–E); periostracum yellowish to pale grey; protoconch of 1.5 whorls, ca 350 µm wide, nucleus ca 200 µm wide (Fig. 16A); protoconch microsculpture granulated (Fig. 16B–C); teleoconch whorls convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin slightly reflexed; umbilicus narrow, not covered by the inner lip (Fig. 15A–E). RADULA. Length intermediate, ca 800 µm long (35% of total shell length), containing about 63 rows of teeth. Central tooth formula 3-C-3/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 16D). Lateral tooth formula (3)4-C-4(3), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 12–14 cusps; outer marginal teeth with 12–14 cusps (Fig. 16E–F). Radular data were compiled from a dry specimen (MZB 77-8445) collected from the type locality. Ecology and distribution This species is only known from the type locality, C’an Tunis (Casa Antúnez), Catalonia, Spain. The type locality has likely disappeared due to extensive alterations (e.g., construction, pollution) of the area as a result of port works conducted in the Barcelona harbour and the industrial zone in the 1980s and the construction work carried out for the 1992 Olympic Games (Jordi Corbella and Francesc Uribe pers. com.). Recent surveys conducted around Barcelona did not yield live specimens. Remarks Mercuria maceana is only known from its type locality, which has been destroyed over the past few decades. Specimens of this species have not been found in the nearby areas, suggesting that the species is extinct (IUCN category EX). The shells and dry specimens of M. maceana deposited in the Zoological Museum of Barcelona were examined to delineate the morphological variation presented by the species and to support its recognition. Our PCA based on shell shape differentiated this species (Fig. 3). Accordingly, M. maceana differs from M. egarensis sp. nov. by its shorter and more globose or subglobose shell and is more similar to the Moroccan species M. targouasensis. Based on the characters observed on the radulae, M. maceana cannot be distinguished from M. similis or M. egarensis. We were unable to study other anatomical features due to the lack of ethanol-preserved material., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 26-29, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Paladilhe A. 1869. Nouvelles miscellanees malacologiques. Chez Savy, libraire-editeur, Paris. https: // doi. org / 10.5962 / bhl. title. 12995","Bank R. & Neubert E. 2017. Checklist of the Land and Freshwater Gastropoda of Europe. Last update Jul. 16 th 2017. Molluscabase (Ed.).","Breure A. & Audibert C. 2017. Under the spell of Bourguignat: Alcide Paladilhe's malacological work, with notes on his collection. Folia Conchyliologica 41: 28 - 39."]}

Mercuria tachoensis (Frauenfeld, 1865) Figs 8–10; Supp. file 2: Tables S6–S9 Amnicola tachoensis Frauenfeld, 1865: 529. Mercuria edmundi Boeters, 1986: 126, fig 4–7. pl. 18a fig. 2. Mercuria edmundi – Boeters 1988: 209, figs 96–98. — Holyoak et al. 2017: 208, fig. 2. Mercuria tachoensis – Boeters 1988: 207, figs 99–103. non M. bayonnensis (Locard, 1894) – Boeters & Falkner 2017 [in part]: 230, fig. 6p. non M. anatina (Poiret, 1801) – Boeters & Falkner 2017 [in part]: 239, fig. 2. Revised diagnosis Shell ovate-conic; aperture obliquely and broadly ovate; protoconch microsculpture pitted; periostracum whitish to pale grey; central radular tooth formula (3)4-C-4(3)/1-1; female genitalia with bursa copulatrix elongate, ca 4 times as long as wide; seminal receptacle pyriform; penis darkly pigmented; penial appendix longer than the distal end of the penis, strongly pigmented at the junction with the penis, pigmentation gradually weakens from the junction until two-thirds of the appendix where it is weak; penial appendix base narrow, distally positioned on the inner edge of the penis; nervous system pigmented, elongate (mean RPG ratio = 0.61); cerebral ganglia approximately equal in size. Type material (not examined) Syntypes PORTUGAL • 2 specs (sex unknown); NHMW 113526 to 113531. Type locality According to the original description, the type locality is a spring close to the Tagus River [Quellen des Tajo bei Ajuda] near Ajuda, Lisbon, Portugal. Material examined PORTUGAL • 30 specs; Mataçaes, Fonte dos passarinhos Spring; MNCN 15.05/94807 • 30 specs; Coimbra, Fonte dos Amores Spring; MNCN 15.05/94809 • 30 specs; Coimbra, spring in Jardim da Sereia; MNCN 15.05/94810 • 30 specs; Leiria, Alpedriz Spring, Nascente do Senhor Jordão; MNCN 15.05/ 94811 • 30 specs; Leiria, Alpedriz, Das Mouras Spring; MNCN 15.05/94812 • 30 specs; Leiria, spring in Salir de Matos; MNCN 15.05/94813 • 30 specs; Leiria, São Gregório da Fanadia, Padre Antonio Spring; MNCN 15.05/94814 • 30 specs; Santarém, spring along road N114 near the crossing to Moçarria and Vila Nova da Babeca; MNCN 15.05/94815 • 30 specs; Santarém, Ereira, spring in Rua da Fonte; MNCN 15.05/94816 • 30 specs; Leiria, spring in São Mamede; MNCN 15.05/94817 • 30 specs; Lisbon, pond in Pragança; MNCN 15.05/94818 • 30 specs; Lisbon, Ericeira, Fonte do Cabo Spring; MNCN 15.05/94819 • 30 specs; Lisbon, Mafra, Fonte dos Tritôes Spring; MNCN 15.05/94820 • 30 specs; Lisbon, Carvoeira, spring in Valbom; MNCN 15.05/94821 • 30 specs; Mafra, Ribiera da Maciel Forro, Fonte da Ribera Spring; MNCN 15.05/94822 • 30 specs; Lisbon, Almargem do Bispo, spring in Vale de Lobos; MNCN 15.05/94823 • 30 specs; Setúbal, Oleiros, Fonte de Oleiros Spring; MNCN 15.05/94824 • 30 specs; Baixo Alentejo near Pomarão, Guadiana River; MNCN 15.05/94825. UNITED KINGDOM • 1 spec.; Arundel, stream outflowing from Swanbourne Lake; UGSB 14156 • 280 specs; Barking, Cuckold’s Haven Wetland, reed belt of Hand Trough Creek; UGSB 14157 • 30 specs; West Sussex, Burpham, Arun Banks; MNCN 15.05/94828. SPAIN • 30 specs; Asturias, Camoca, Fuente Tebia Spring; MNCN 15.05/94808. FRANCE • 30 specs; Mouguerre, stream near Chemin d’Elizaberry; MNCN 15.05/94826 • 30 specs; stream near the beach in Bidart; MNCN 15.05/94827. THE NETHERLANDS • 8 specs; Hoogvliet, small stream; UGSB 23178 • 8 specs; Hoogvliet, junction of the running surface in the tidal area; UGSB 23179 • 3 specs; Hoogvliet, Ruigeplaatbosch Park, Oude Maas River;; wet reed land; UGSB 23177. Additional locality information is provided in Supp. file 1: Table S1. Description SHELL. Ovate to ovate-conic, whorls 4–5, height 2–4 mm, width 2–3 mm (Fig. 8A–G; Supp. file 2: Table S6); periostracum yellowish to pale grey, occasionally dark brown; protoconch of 1.5 whorls, ca 350 µm wide, nucleus ca 180 µm wide (Fig. 9A–B); protoconch microsculpture pitted (Fig. 9C); teleoconch whorls convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin slightly reflexed; inner lip attached to the body whorl; umbilicus narrow, not covered by the inner lip (Fig. 8A, C–G). OPERCULUM. As for the genus, orange to brown, sometimes yellowish, about two whorls; muscle attachment oval, located near the nucleus (Fig. 8H–I). RADULA. Length intermediate, ca 800 µm long (35% of total shell length), containing about 65 rows of teeth. Central tooth formula (4)3-C-3(4)/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 9D). Lateral tooth formula (3)4-C-4(3), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 12–17 cusps; outer marginal teeth with 17–27 cusps (Fig. 9E–F). Radular data were collected from the following specimens: MNCN 15.05/94808 – Fuente Tebia Spring, Camoca, Asturias, Spain; MNCN 15.05/94820 – Dos Tritôes Spring, Mafra, Lisbon, Portugal; MNCN 15.05/94821 – spring in Valbom, Carvoeira, Lisbon, Portugal; MNCN 15.05/94824 – Fonte de Oleiros Spring, Oleiros, Setúbal, Portugal; MNCN 15.05/94826 – stream near Chemin d’Elizaberry, Mouguerre, France; MNCN 15.05/94827 – stream near the beach in Bidart, France; MNCN 15.05/94828 – Arun Banks, Burpham, West Sussex, United Kingdom. PIGMENTATION AND ANATOMY. Animal usually darkly pigmented, head and tentacles black to brown, pigmentation lighter on eye lobes and snout; snout about as long as wide, approximately parallel-sided, with medium distal lobation (Fig. 10B). Ctenidium occupying almost the total length of the pallial cavity; 20–26 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 10D). Pallial tentacle present. Osphradium elongate, more than 3 times as long as broad, positioned opposite middle of ctenidium. Stomach almost as long as wide with two chambers almost equal in size (Supp. file 2: Table S7); style sac longer than wide, with the unpigmented intestine surrounding its distal end before continuing on as a straight rectum (Fig. 10E). MALE GENITALIA. Prostate gland bean-shaped, about 2 times as long as wide (Supp. file 2: Table S9), connected by the posterior vas deferens to a convoluted seminal vesicle and the testis. Pallial vas deferens emerge near both the anterior end of the prostate gland and the external margin of the penis (Fig. 10C). Penis darkly pigmented, gradually tapering, attached to the neck behind the right eye; penial appendix longer than the distal end of the penis, strongly pigmented at the junction with the penis, pigmentation gradually weakens from the junction until two-thirds of the appendix where it is weak. Penial appendix base narrow, distally positioned on the inner edge of the penis (Fig. 10B). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide; albumen gland and capsule gland about the same length (Fig. 10A); bursa copulatrix elongate, ca 4 times as long as wide (Supp. file 2: Table S8); bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle pyriform, with a short duct, positioned on the distal part of the renal oviduct just above the junction with the bursal duct (Fig. 10A). NERVOUS SYSTEM. Slightly pigmented, elongate (mean RPG ratio = 0.61; see Supp. file 2: Table S15); cerebral ganglia approximately equal in size; pleuro-supraoesophageal connective ca 7 times as long as pleuro-suboesophageal one (Fig. 10F). Ecology and distribution Mercuria tachoensis has a lower conductivity tolerance than M. similis. The analysed populations of this species from Spain, Portugal and France inhabit springs, streams and ponds with conductivities from 359 to 1805µS/cm. Water parameters of the localities in the UK and the Netherlands were not available. Specimens were usually found inside the water. Amphibious behaviour has not been commonly observed in this species, except for populations located on the tide banks of the Guadiana River (David Holyoak pers. com.) and those of the Arun River (Martin Willing pers. com.). The species occurs sympatrically with the gastropod species Theodoxus aff. fluviatilis (Linnaeus, 1758), Belgrandia lusitanica (Paladilhe, 1867), and Potamopyrgus antipodarum. This species was previously recorded from springs and streams in Portugal. In this study, we report new records, extending its distribution to include northern Spain, northern France, the UK and the Netherlands (Fig. 1). At these localities, the species was found in the tidal mud of small rivers near the river mouth. According to this extended distribution, M. tachoensis inhabits water bodies that have an Atlantic drainage. The type locality of the species has likely disappeared due to the extensive growth of the city of Lisbon, intensive exploitation of local aquifers and pollution. Recent surveys around Ajuda, Lisbon, did not yield any new records (Rui da Costa Mendes pers. com.). Remarks Based on our morphological observations and previous phylogenetic results (Miller et al. 2022), we suggest several synonymization of several previously described species of Mercuria to M. tachoensis (see Discussion), which increases the previously known geographic range and intraspecific variation of this species. The species’ moderate shell variation can be observed across its geographic range (Figs 1, 8). Also, the PCA (Fig. 3) showed near the all populations identified by Miller et al. (2022) as M. tachoensis, (although it is closely related in shape to M. balearica), suggesting a low level of intraspecific variation in shell shape. This variability is due to differences in overall shell shape and colour pattern. Regarding shell shape, we found populations with only elongated ovate-conic shells [e.g., Fonte dos Amores Spring (MNCN 15.05/94809), spring in Jardim da Sereia (MNCN 15.05/94810), spring in Rua da Fonte, Ereira (MNCN 15.05/94816), Arun Banks (MNCN 15.05/94828)] or only ovate-conic shells [e.g., spring in Salir de Matos (MNCN 15.05/94813), Fonte dos Tritôes Spring (MNCN 15.05/94820)]. We also observed populations with both shell morphotypes [e.g., Fonte dos Passarinhos Spring (MNCN 15.05/94807), spring in Vale de Lobos (MNCN 15.05/94823), Fonte de Oleiros Spring (MNCN 15.05/94824)]. Colour pattern does not show high intraspecific variation: only the specimens from the spring in Vale de Lobos (MNCN 15.05/94823) are remarkable for being particularly darkly coloured with a dark brown thick periostracum (Fig. 8F) compared with the rest of the specimens, which have yellowish-brown periostracum. Anatomically, the bursa copulatrix is about the same size in all of the dissected specimens of the species, except for those from the spring in Jardim da Sereia (MNCN 15.05/94810) and Nascente do Senhor Jordão (MNCN 15.05/94811), which are considerably smaller (Supp. file 2: Table S8). Males from the stream near Chemin d’Elizaberry (MNCN 15.05/94826) and Nascente do Senhor Jordão, Alpedriz (MNCN 15.05/94811) have the smallest prostate glands and penises. Despite this, all males present the same penis characteristics: a rounded, pigmented penial appendix that is longer than the distal end of the penis. However, Boeters (1988) described the penis as longer than the penial appendix. Based on this discrepancy in the size and shape of the male genitalia, Holyoak et al. (2017) proposed M. edmundi as a junior synonym of M. tachoensis and described an allometric growth of the male genitalia related to the sexual maturity of the animals. Miller et al. (2022) confirmed with taxonomic multilocus data that only one species occurs in the localities where both species have been cited. Mercuria tachoensis differs from all the Iberian congeners by its pitted protoconch microsculpture (which is granulated in the rest of the species; visible only with electronic microscopy). In addition to this character, it differs from the phylogenetically closely related species M. similis and M. egarensis sp. nov. by often having one more cusp in the central radular teeth and a longer bursa copulatrix (ca 4 times as long as wide). Although our PCA (Fig. 3) was unable to differentiate M. tachoensis and M. carrillorum sp. nov. by shell shape, both species can be distinguished by differences in the radula (smaller in M. carrillorum and often with one cusp less in the lateral teeth), the bursa copulatrix (smaller in M. carrillorum) and the male genitalia (the distal end of the penis is longer than the penial appendix, which is flattened, ovate and less pigmented in M. carrillorum)., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 15-21, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Frauenfeld G. 1865. Verzeichniss der Namen der fossilen und lebenden Arten der Gattung Paludina Lam: nebst jenen der nachststehenden und Einreihung derselben in die verschiedenen neueren Gattungen. Verhandlungen der Kaiserlich-koniglichen Zoologisch-botanischen Gesellschaft in Wien 14: 561 - 672.","Boeters H. 1986. Unbekannte westeuropaische Prosobranchia, 7. Heldia 1 (4): 125 - 128.","Boeters H. 1988. Moitessieriidae und Hydrobiidae in Spanien und Portugal (Gastropoda: Prosobranchia). Archiv fur Molluskenkunde 118: 181 - 261.","Holyoak D. T., Holyoak G. A. & da Costa Mendes R. M. 2017. Distribution and ecology of Mercuria tachoensis (Gastropoda: Hydrobiidae) in Portugal and evidence that M. edmundi is conspecific. Iberus 35: 1 - 8.","Locard A. 1894. Les Bythinia du systeme europeen: revision des especes appartenant a ce genre d'apres la collection Bourguignat. Revue suisse de Zoologie 2: 65 - 134. https: // doi. org / 10.5962 / bhl. part. 75133","Boeters H. D. & Falkner G. 2017. The genus Mercuria Boeters, 1971 in France (Gastropoda: Caenogastropoda: Hydrobiidae). West-European Hydrobiidae, Part 13. Zoosystema 39: 227 - 261. https: // doi. org / 10.5252 / z 2017 n 2 a 4","Poiret J. 1801. Coquilles fluviatiles et terrestres observees dans le Departement de l'Aisne et aux Environs de Paris. Chez T. Barrois, Paris. https: // doi. org / 10.5962 / bhl. title. 35858","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

Mercuria egarensis Miller, García-Guerrero & Ramos sp. nov. urn:lsid:zoobank.org:act: F4D85246-8580-48C8-8CFD-7C838D87B6B5 Figs 17–19; Supp. file 2: Tables S14–S18 Diagnosis Shell ovate-conic; aperture obliquely ovate; protoconch microsculpture granulated; periostracum whitish to pale grey; central radular tooth formula (2)3-C-3(2)/1-1; female genitalia with bursa copulatrix pyriform to elongate, ca 2.5–3 times as long as wide; seminal receptacle elongate with short peduncle; penis darkly pigmented, gradually tapering; penial appendix unpigmented, ovate, about equal in length or slightly shorter than the distal end of the penis and medially positioned on the inner edge of the penis; nervous system pigmented, elongate (mean RPG ratio = 0.57); cerebral ganglia approximately equal in size. Etymology The name ‘ egarensis ’ refers to the ancient Roman city of Egara, present-day Terrassa, Catalonia, where the species occurs. Type material Holotype SPAIN • sex unknown (dry preserved); Barcelona, Font de les Canyes; MNCN 15.05/200174H. Paratypes SPAIN • 38 specs (preserved in ethanol 80%, 10 dissected and 3 processed for DNA sequencing); same collection data as for holotype; MNCN 15.05/94752 • 13 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/94753 • 25 specs (preserved in ethanol 80%); same collection data as for holotype; MZB 2021-2798 • 1 spec. (dry preserved); same collection data as for holotype; NHMW 113529 • 2 specs (dry preserved); same collection data as for holotype; RMNH. MOL.507925 • 1 spec. (dry preserved); same collection data as for holotype; JPM-586 • 1 spec. (dry preserved); same collection data as for holotype; MCP • 1 spec. (dry preserved); same collection data as for holotype; MCP. Type locality Font de les Canyes, Catalonia, Spain 41.586217° N, 2.038500° E, 340 m a.s.l. Description MEASUREMENTS. Holotype: SL = 3.32 mm, SW = 2.53 mm, SL/SW = 1.31, AH = 1.51 mm, SL-LBW = 0.86 mm, WBW = 2.05 mm, AL = 1.61 mm, AW= 1.23 mm, WPW = 1.22 mm, WAW = 0.70 mm. SHELL. Ovate-conic, whorls 4–5, height 3–4.3 mm, width 1.7–2.8 mm (Fig. 17A–E; Supp. file 2: Table S14); periostracum whitish to pale grey; protoconch of 1.5 whorls, ca 350 µm wide, nucleus ca 150 µm wide (Fig. 18A); protoconch microsculpture granulated (Fig. 18B–C). Teleoconch whorls slightly convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin straight; umbilicus narrow, not covered by the inner lip (Fig. 17A–E). OPERCULUM. As for the genus, orange to brown, sometimes yellowish, about two whorls; muscle attachment oval, located near the nucleus (Fig. 17F–G). RADULA. Length intermediate, ca 800 µm long (35% of total shell length), containing about 55 rows of teeth. Central tooth formula (2)3-C-3(2)/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 18D). Lateral tooth formula (3)2-C-2(3), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 11–15 cusps; outer marginal teeth with 15–17 cusps (Fig. 18E–F). Radular data were obtained from specimens from the type locality. PIGMENTATION AND ANATOMY. Animal slightly brownish to darkly pigmented (Fig. 19C); head and tentacles dark brown, pigmentation lighter on eye lobes, snout and neck; snout about as long as wide, approximately parallel-sided, with medium distal lobation. Ctenidium occupying almost the total length of the pallial cavity; 19–23 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 19D). Osphradium elongate, more than 3 times as long as broad, positioned opposite middle of ctenidium. Stomach almost as long as wide with two chambers almost equal in size (Supp. file 2: Table S16); style sac longer than wide, protruding anteriorly into the intestinal loop (Fig. 19E). Intestine unpigmented, continuing on as a straight rectum. MALE GENITALIA. Prostate gland bean-shaped, about 2 times as long as wide (Supp. file 2: Table S18), connected by the posterior vas deferens to a convoluted seminal vesicle and the testis. Penis darkly pigmented, gradually tapering, attached to the neck behind the right eye; penial appendix unpigmented, ovate, about equal in length or slightly shorter than the distal end of the penis and medially positioned on the inner edge of the penis (Fig. 19F). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide; albumen gland longer than capsule gland (Fig. 19A); bursa copulatrix pyriform to elongate (Fig. 19A–B), ca 2.5–3 times as long as wide (Supp. file 2: Table S17); bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle elongate, with a short duct, positioned on the distal part of the renal oviduct just above the junction with the bursal duct (Fig. 19B). NERVOUS SYSTEM. Pigmented, elongate (mean RPG ratio = 0.57; see Supp. file 2: Table S15); cerebral ganglia approximately equal in size and have small black dots of pigment; pleuro-supraoesophageal connective ca 7 times as long as pleuro-suboesophageal one (Fig. 19G). Ecology and distribution The species is endemic to the Font de les Canyes Spring, alongside the Besós River. The conductivity of this spring is relatively high (1412 µS/cm). The species was collected from the rocky bottom of the spring at a low abundance. No other snail species were found in the spring. The water is used for human consumption, which may have had an impact on this and other snail populations. A series of sampling campaigns of this and nearby areas were carried out between 2016 and 2018 and the species was not found elsewhere. The spring, located in a small zone in the eastern part of the municipality of Terrassa, discharges to the Besós River basin. Watercourses from the western part of the municipality drain into the Llobregat River basin (Jordi Corbella pers. com.). Bofill & Haas (1920) cited M. similis from Terrassa; however, as the drainage of this spring is associated with another hydrological basin, it is unlikely that Bofill & Haas (1920) citation for M. similis corresponds to Font de les Canyes. Remarks Mercuria egarensis sp. nov. differs from the closely related species M. carrillorum sp. nov. by having a slightly shorter penis, a larger penial appendix, a longer radular ribbon, a larger number of cusps on the central radular tooth [(2)3–C–3(2)/1–1 vs (3)4–C–4(3)/1–1], a more concentrated perioesophageal ring, a pyriform SR1 positioned on the renal oviduct just above the junction with the bursal duct (in M. carrillorum, it is positioned at the junction) and the absence of a pallial tentacle. Regarding their ecology, the two species differ in their conductivity preferences, with M. egarensis occurring in waters with higher conductivity. Mercuria egarensis sp. nov. does not resemble any of the congeners found in the Catalonia region. However, according to our PCA results, M. egarensis and M. balearica have overlapping shell shape variation (Fig. 3), though the latter presents a more globose shell with a wider aperture (Fig. 11). Although M. egarensis also resembles M. tachoensis and M. similis in terms of shell shape, its granulated protoconch microsculpture discriminates it from the pitted one in M. tachoensis. The shape and colouration of the penial appendix differentiate M. egarensis from M. similis: it is highly ovate and unpigmented in the former species and triangular and pigmented in the latter. The sequence divergence of COI indicated that M. egarensis sp. nov. is closely related to M. carrillorum sp. nov. with a divergence of only 1.3%. Mercuria egarensis diverges 3.7% with M. similis and 8.1% with M. balearica. The species is most distantly related to M. tensiftensis and M. melitensis, diverging with both at around 9% (Miller et al. 2022)., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 29-33, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Bofill A. & Haas F. 1920. Estudi sobre la malacologia de les Valls Pirenaiques: Conca del Llobregat. Treballs del Museu de Ciencies Naturals de Barcelona 3: 380 - 830.","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

Mercuria lupiaensis Miller & Delicado sp. nov. urn:lsid:zoobank.org:act: A3B189DF-8A84-46B5-82BD-A083F46F8CF9 Figs 26–28; Supp. file 2: Tables S14–S18 Diagnosis Shell ovate-conic; aperture obliquely ovate; periostracum whitish to pale grey; central radular tooth formula 3-C-3/1-1; female genitalia with bursa copulatrix pyriform to elongate, ca 2 times as long as wide; seminal receptacle pyriform; penis darkly pigmented; penial appendix unpigmented, ovate, smaller than the distal end of the penis and medially positioned on the inner edge of the penis; nervous system pigmented, elongate (mean RPG ratio = 0.53); cerebral ganglia approximately equal in size, with small black spots. Etymology The specific epithet ‘ lupiaensis ’ refers to Lupia, the name of the city of Lecce during the time of the ancient Roman empire. Type material Holotype ITALY • sex unknown (preserved in ethanol 80%); Lecce, Frigole, Giammateo Creek; MNCN 15.05/200123H. Paratypes ITALY • 4 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/200123P • 13 specs (preserved in ethanol 80%, 7 dissected and 2 processed for DNA sequencing); same collection data as for holotype; UGSB 8015. Additional material examined ITALY • 9 specs; Apulia, Nardò, San Isidoro, Palude del Capitano Pond; UGSB 16236 • 7 specs; Apulia, Porto Cesareo, Ionian Coast, spring in Torre Castiglione; UGSB 16237. Additional locality information provided in Supp. file 1: Table S1. Type locality Giammateo Creek, Frigole, Lecce, Italy. 44.9436° N, 12.0213° E. Description MEASUREMENTS. Holotype: SL = 3.73 mm, SW = 2.95 mm, SL/SW = 1.26, AH = 1.92 mm, SL-LBW = 0.94 mm, WBW = 2.27 mm, AL = 1.96 mm, AW = 1.32 mm, WPW = 1.29 mm, WAW = 0.63 mm. SHELL. Ovate-conic, whorls 4–5, height 3.10–3.75 mm, width 2.50–3.00 mm (Fig. 26A–F; Supp. file 2: Table S14); periostracum whitish to pale grey; protoconch of 1.5 whorls, ca 300 µm wide, nucleus ca 150 µm wide; teleoconch whorls slightly convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin straight, inner lip attached to the body whorl; umbilicus narrow, not covered by the inner lip (Fig. 26A, C–F). OPERCULUM. As for the genus, orange to brown, about two whorls; muscle attachment oval, located near the nucleus (Fig. 26G–H). RADULA. Length intermediate, ca 800 µm long (35% of total shell length), containing about 60 rows of teeth (Fig 27A). Central tooth formula 3-C-3/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 27B). Lateral tooth formula 3-C-3, central cusp V shaped and slightly longer than the central tooth one (Fig. 27C). Inner marginal teeth with 14–17 cusps; outer marginal teeth with 22–25 cusps (Fig. 27D). Radular data were collected from specimens from Giammateo Creek, Frigole, Lecce (UGSB 8015); Palude del Capitano Pond, San Isidoro, Ionian coast, Nardó, Apulia (UGSB 16236) and a spring in Torre Castiglione, Ionian Coast, Porto Cesareo, Apulia (UGSB 16237). PIGMENTATION AND ANATOMY. Animal darkly pigmented (Fig. 28E, I); head and tentacles black pigmented, pigmentation lighter on eye lobes, snout and neck; snout about as long as wide, approximately parallel-sided, with medium distal lobation. Ctenidium occupying almost the total length of the pallial cavity; 23–27 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 28G). Stomach almost as long as wide with two chambers almost equal in size (Supp. file 2: Table S16); style sac longer than wide, with the unpigmented intestine surrounding its distal part and continuing on as a straight rectum (Fig. 28H). MALE GENITALIA. Prostate gland bean-shaped, about 2 times as long as wide (Supp. file 2: Table S18), connected by the posterior vas deferens to a convoluted seminal vesicle and the testis (Fig. 28F). Penis darkly pigmented, gradually tapering, attached to the central area of the neck; penial appendix unpigmented or slightly pigmented at the junction with the distal end of the penis. Penial appendix ovate, smaller than the penis, base narrow, medially positioned on the inner edge of the penis (Fig. 28C–E). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide; albumen gland and capsule gland about the same size (Fig. 28A–B); bursa copulatrix elongate, ca 2 times as long as wide (Supp. file 2: Table S17); bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle pyriform, with a short duct, positioned on the distal part of the renal oviduct just above the junction with the bursal duct (Fig. 28B). NERVOUS SYSTEM. Pigmented, elongate (mean RPG ratio = 0.53; see Supp. file 2: Table S15); cerebral ganglia approximately equal in size (Fig. 28I). Ecology and distribution The species is only known from the three nearby localities in the Salento Peninsula in the region of Apulia, Italy. The ecology of the species is not well known. The species was found in very clear running waters, close to the coastal margins. Co-occurring species are Pseudamnicola conovula (Frauenfeld, 1863), Eupaludestrina stagnorum (Gmelin, 1791) and Bithynia leachii (Sheppard, 1823). Remarks Mercuria lupiaensis sp. nov. now represents the easternmost species of the genus and potentially the second species reported in the Apennine Peninsula. The species M. similis has been documented from the Sale Basin (Bodon et al. 2005; Cianfanelli & Bodon 2017), the first species of Mercuria described from the Apennine Peninsula. Despite that the new species and M. similis have a similar shell shape; they can be easily distinguished by differences in the penial appendix: in M. similis, it is large, triangular and highly pigmented, whereas in M. lupiaensis, it is small, sometimes very small, ovate and slightly pigmented or unpigmented. In the new species, the albumen and capsule glands are about the same size, whereas, in M. similis, the albumen gland is longer than the capsule gland. They also present differences in the bursa copulatrix: elongate and about 2 times as long as wide in M. lupiaensis and pyriform to elongate and 3 times as long as wide in M. similis. Mercuria lupiaensis sp. nov. can be further distinguished from the phylogenetically closely related species M. veronicae sp. nov. by its smaller size, lighter dorsal pigmentation, similarly sized capsule and albumen glands (albumen gland longer than capsule gland in M. veronicae), a penial appendix that is shorter than the distal end of the penis and a smaller number of cusps on the central teeth. The sequence divergence estimates of COI showed that the species is most closely related to M. veronicae sp. nov. (mean divergence of 3.9%) and most distantly to M. midarensis (with a mean divergence of 9.3%) (Miller et al. 2022)., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 42-47, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Frauenfeld G. 1863. Vorlaufige Aufzahlung der Arten der Gattungen Hydrobia Htm. und Amnicola Gld. Hldm. in der kaiserlichen und in Cuming's Sammlung. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 13: 1017 - 1030.","Bodon M., Cianfanelli S., Manganelli C., Pezzoli E. & Giusti F. 2005. Checklist e distribuzione della fauna italiana. 10.000 specie terrestri e delle acque interne. In: Ruffo S. S. (ed.) Memorie del Museo Civico di Storia Naturale di Verona, 2 ° Serie, Sezione Scienze della Vita: 79 - 81. Ministerio dell´Ambiente, Verona.","Cianfanelli S. & Bodon M. 2017. Nuovi idrobiidi per il bacino del Fiume Sele, con una checklist dei molluschi dulciacquicoli della Campania (Gastropoda: Caenogastropoda: Hydrobiidae). Bollettino Malacologico 53: 79 - 120.","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

Mercuria veronicae Miller, Khalloufi & Delicado sp. nov. urn:lsid:zoobank.org:act: 986C0653-973B-4C3D-9A4A-C6A216321EBE Figs 29–31; Supp. file 2: Tables S14–S18 Diagnosis Shell ovate-conic; aperture obliquely ovate; periostracum whitish to pale grey; central radular tooth formula (5)4-C-4(5)/1-1; female genitalia with a bursa copulatrix pyriform to elongate, ca 2 times as long as wide; seminal receptacle elongate; penis darkly pigmented; penial appendix slightly pigmented, ovate, shorter than or equal in length to the distal end of the penis, medially positioned on the inner edge of the penis; nervous system pigmented, elongate (mean RPG ratio = 0.55); cerebral ganglia approximately equal in size; pleuro-supraoesophageal connective ca 11 times as long as pleuro-suboesophageal one. Etymology After Veronica Ruiz, the first author’s grandmother. Type material Holotype TUNISIA • sex unknown (preserved in ethanol 80%); Tozeur, Tamerza, Oasis Waterfall. El Waha Spring; MNCN 15.05/200124H. Paratypes TUNISIA • 3 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/200124P • 15 specs (preserved in ethanol 80%, 6 dissected and 1 processed for DNA sequencing); same collection data as for holotype; UGSB 17271. Additional material examined TUNISIA • 15 specs; Tozeur, Tamerza, Big Waterfall, Lekbir Spring; UGSB 17274 • 4 specs; Tozeur, Echbika, Echbika Waterfall, Echbicka Spring; UGSB 17276. Additional locality information provided in Supp. file 1: Table S1. Type locality El Waha Spring, Oasis Waterfall, Tamerza, Tozeur, Tunisia, 34.381933° N, 7.933450° E, 278.67 m a.s.l. Description MEASUREMENTS. Holotype: SL = 4.87 mm, SW = 3.56 mm, SL/SW = 1.36, AH = 2.03 mm, SL-LBW = 1.40 mm, WBW = 2.96 mm, AL = XX mm, AW = 2.06 mm, WPW = 1.67 mm, WAW = 0.85 mm. SHELL. Ovate-conic, whorls 4–5, height 3.25–5.3 mm, width 2.5–4 mm (Fig. 29A–F; Supp. file 2: Table S14); periostracum whitish to pale grey; protoconch of 1.5 whorls, ca 250 µm wide, nucleus ca 150 µm wide; teleoconch whorls slightly convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin straight; inner lip touching the body whorl; umbilicus narrow, not covered by the inner lip. OPERCULUM. As for the genus, orange to brown, sometimes yellowish, about two whorls; muscle attachment oval, located near the nucleus (Fig. 29G–H). RADULA. Length intermediate, ca 750 µm long (35% of total shell length), containing about 60 rows of teeth (Fig. 30A). Central tooth formula (5)4-C-4(5)/1-1, central cusp V shaped, cutting edge concave (Fig. 30B–C). Lateral tooth formula (3)4-C-4(3), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 11–15 cusps; outer marginal teeth with 21–25 cusps (Fig. 30D). Radular data were collected from the following specimens: UGSB 17271 – El Waha Spring, Oasis Waterfall, Tamerza, Tozeur, Tunisia; UGSB 17274 – Lekbir Spring, Big Waterfall, Tamerza, Tozeur, Tunisia; UGSB 17276 – Echbicka Spring, Echbika Waterfall, Echbika, Tozeur, Tunisia. PIGMENTATION AND ANATOMY. Animal darkly pigmented (Fig. 31A, D); head and tentacles dark brown, pigmentation lighter on eye lobes, snout and neck; snout about as long as wide, approximately parallel-sided, with medium distal lobation. Ctenidium occupying almost the total length of the pallial cavity; 28– 33 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 31B). Osphradium elongate, more than 3 times as long as wide, positioned opposite middle of ctenidium. Stomach almost as long as wide with two chambers almost equal in size (Supp. file 2: Table S16); style sac longer than wide, with the unpigmented intestine surrounding its distal part and continuing on as a straight rectum (Fig. 31C). MALE GENITALIA. Prostate gland bean-shaped, about 2 times as long as wide (Supp. file 2: Table S18). Pallial vas deferens emerge close to both the anterior edge of the prostate and the external margin of the penis (Fig. 31F). Penis darkly pigmented, gradually tapering, attached to the neck behind the right eye, distal end of the penis broad, triangular; penial appendix pyriform, shorter than or about to the same length as the distal end of the penis, slightly pigmented at the junction with the penis, medially positioned on the inner edge of the penis (Fig. 31D–E). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide, albumen gland longer than capsule gland; bursa copulatrix elongate, ca 2 times as long as wide (Supp. file 2: Table S17); bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle elongate, with a short duct, positioned on the distal part of the renal oviduct just above the junction with the bursal duct. NERVOUS SYSTEM. Pigmented, elongate (mean RPG ratio = 0.55; see Supp. file 2: Table S15); cerebral ganglia approximately equal in size; pleuro-supraoesophageal connective ca 11 times as long as pleuro-suboesophageal one (Fig. 31G–H). Ecology and distribution The studied samples of this species come from three localities in the Tozeur region, north of the Chotts, but its distribution extends to other regions around the Chotts el Gharsa and Djeride (Gafsa, Kebili, Nefzaoua and Gabes). It occurs in streams, springs and irrigation ditches of the oasis areas. This species is present at a high density (more than 3000 ind./m 2) on the edge of the streams, where the flow is low and the substrate muddy, and attached to aquatic plants. The main streams of Tamarza (Oued Frid) and Echbika (Oued Echbika) are highly saline (conductivity 8667 and 3945 µS/cm, respectively) and flow over an alluvial and clayey-sandy substrate of the Plio-Quaternary, with a predominance of sulphate-sodium, gypsum and phosphorites (Abidi 2007). Co-occurring species are Galba truncatula, Physella acuta (Draparnaud, 1805) (in low abundance), Eupaludestrina spp., Melanoides tuberculata (O.F. Müller, 1774), Melanopsis douvillei Pallary, 1916 and Melanopsis praemorsa (Linnaeus, 1758) (in high abundance). Remarks Intraspecies morphological differences were observed mainly in the penial features of males from the same population (Fig. 31D–E). Some individuals have a much-atrophied penis. Such variation may be parasite-induced, similar to the case reported in another species of Mercuria (Boulaassafer et al. 2018). Parasitized females also presented an atrophied pallial oviduct, making it difficult to conduct a complete anatomical study. The pigmentation of the penis is also very variable, ranging from very slight to pronounced dark pigmentation at the junction between the appendix and distal end of the penis. Mercuria veronicae sp. nov. differs from the phylogenetically closely related species M. lupiaensis sp. nov. and M. melitensis by having a high-spired shell and a more elongated bursa copulatrix compared with M. lupiaensis and thinner inner and outer shell lips compared with the characteristically thick ones of M. melitensis (Glöer et al. 2015). Mercuria veronicae also has a long, triangular and more pigmented penial appendix compared with the short (to very short), ovate and slightly pigmented or unpigmented one in M. lupiaensis. In M. melitensis, the penial appendix is only as long as the distal end of the penis. Compared with other geographically proximate species, M. veronicae resembles M. pycnocheilia (Bourguignat, 1862) in shell shape, although the latter presents a thicker inner shell lip (see Glöer et al. 2010) and the former, has smaller-sized shells. In addition, Glöer et al. (2010) and Glöer (2019) noted the acute apex of M. pycnocheilia (Bourguignat, 1862), which is not present in M. veronicae. Shell shape features also distinguish the new species from M. globulina (Letourneux & Bourguignat, 1887), which has a larger, more slender shell, and M. bourguignati Glöer, Bouzid & Boeters, 2010, which has a larger shell and smaller aperture (about half the shell length in M. bourguignati vs about three quarters the shell length in M. veronicae). The anatomy of these geographically proximate species is unknown. According to our sequence divergence estimates of COI, M. veronicae sp. nov. is most similar to M. lupiaensis sp. nov. (average divergence of 3.9%) and diverges most with M. tensiftensis (by 8.4%) (Miller et al. 2022)., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 47-52, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Abidi B. 2007. Caracterisation Hydrogeologique, Geochimique et Isotopique des Systemes Aquiferes du Synclinal de Tamerza et de la Plaine de Chott El Gharsa (Sud Ouest Tunisien). PhD thesis, University of Sfax, Tunisia.","Draparnaud J. P. R. 1805. Histoire naturelle des Mollusques terrestres et fluviatiles de la France. Chez Louis Colas, Paris. https: // doi. org / 10.5962 / bhl. title. 12856","Boulaassafer K., Ghamizi M. & Delicado D. 2018. The genus Mercuria Boeters, 1971 in Morocco: first molecular phylogeny of the genus and description of two new species (Caenogastropoda, Truncatelloidea, Hydrobiidae). ZooKeys 782: 95 - 128. https: // doi. org / 10.3897 / zookeys. 779.26797","Gloer P., Boeters H. D. & Walther F. 2015. Species of the genus Mercuria Boeters, 1971 (Caenogastropoda: Truncatelloidea: Hydrobiidae) from the European Mediterranean region, Morocco and Madeira, with descriptions of new species. Folia Malacologica 23: 279 - 291. https: // doi. org / 10.12657 / folmal. 023.024","Gloer P., Bouzid S. & Boeters H. D. 2010. Revision of the genera Pseudamnicola Paulucci 1878 and Mercuria Boeters 1971 from Algeria with particular emphasis on museum collections (Gastropoda: Prosobranchia: Hydrobiidae). Archiv fur Molluskenkunde 139: 1 - 22. https: // doi. org / 10.1127 / arch. moll / 1869 - 0963 / 139 / 001 - 022","Gloer P. 2019. The Freshwater Gastropods of the West-Paleartics, Vol. 1: Fresh- and Brackish Waters except Springs and Subterranean Snails. Biodiversity Research Lab, Hetlingen.","Letourneux A. & Bourguignat J. R. 1887. Prodrome de la Malacologie terrestre et fluviatile de la Tunisie. Imprimerie nationale, Paris. https: // doi. org / 10.5962 / bhl. title. 132280","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

Mercuria balearica (Paladilhe, 1869) Figs 11–14; Supp. file 2: Tables S10–S13 Amnicola balearica Paladilhe, 1869: 113. Mercuria balearica – Boeters 1988: 207, figs 106–107. Pseudamnicola balearica – Quintana 2019: 19, fig 1a. Revised diagnosis Shell ovate-conic; aperture obliquely broad ovate; protoconch microsculpture granulated; central radular tooth formula (4)3-C-3(4)/1-1; glandular oviduct 2.5 times as long as wide; albumen gland longer than capsule gland; bursa copulatrix pyriform to elongate; seminal receptacle elongate, with a short duct; penis darkly pigmented, as long as the penial appendix; penial appendix unpigmented or slightly pigmented at the junction with the penis, rounded, small, base narrow, medially positioned on the inner edge of the penis; nervous system pigmented, elongated (mean RPG ratio = 0.53). Type material (not examined) Syntype FRANCE • 1 spec. (sex unknown); UM.PLD.005 (Breure & Audibert 2017). Type locality According to the original description (Paladilhe 1869), the type locality is Port Mahon, Minorca, Balearic Islands [Port-Mahon (Illes Balears)]. Material examined SPAIN – Islas Baleares • 30 specs; Minorca, Colarsega River in Port Mahon; MNCN 15.05/94744 • 30 specs; Minorca, stream near Sant Joan de Carbonell; MNCN 15.05/94745 • 30 specs; Minorca, Cala en Porter; MNCN 15.05/94746 • 30 specs; Minorca, Son Bou, Barranc de Bec; MNCN 15.05/94747. – Málaga • 30 specs; Ardales, El Chorro Spring; MNCN 15.05/94732 • 30 specs; spring in Montecorto; MNCN 15.05/94731, MNCN 15.05/94737 • 30 specs; trough in Churriana; MNCN 15.05/94733 • 30 specs; Venta El Pilar Spring; MNCN 15.05/94734, MNCN 15.05/94738 • 30 specs; trough in Los Granados; MNCN 15.05/94735 • 30 specs; stream in Montecorto; MNCN 15.05/94736. – Cádiz • 30 specs; Alozaina, Valentín Spring; MNCN 15.05/94739. – Jaén • 30 specs; Fontanar, stream in Caño de la Rambla; MNCN 15.05/94740 • 30 specs; Peal de Becerro, Cinco Caños Spring; MNCN 15.05/94743. – Granada • 30 specs; Mojácar, Arabic Spring; MNCN 15.05/94741). – Murcia • 30 specs; spring in Sierra de la Muela; MNCN 15.05/94742. Additional locality information is provided in Supp. file 1: Table S1. Description SHELL. Ovate-conic, whorls 4–5, height 2–3.5 mm, width 1.4–2.6 mm (Fig. 11A–F; Supp. file 2: Table S10); periostracum yellowish to brown; protoconch with 1.5 whorls, ca 350 µm wide, nucleus less than 200 µm wide; protoconch microsculpture granulated (Fig. 12A–C); teleoconch whorls convex, separated by a deep suture; body whorl large, convex, occupying about two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; aperture margin straight, inner lip touching the shell wall; umbilicus narrow, not covered by the inner lip (Fig. 11A–F). OPERCULUM. As for the genus, orange to brown, sometimes yellowish, about two whorls; muscle attachment oval, located near the nucleus (Fig. 11G–H). RADULA. Length intermediate, ca 800 µm long (35% of total shell length), containing about 65 rows of teeth. Central tooth formula (4)3-C-3(4)/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 12D). Lateral tooth formula (3)4-C-4(3), central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 14–18 cusps; outer marginal teeth with 15–17 cusps (Fig. 12E–F). Radular data were collected from the following specimens: MNCN 15.05/94745 – stream near Sant Joan de Carbonell, Minorca; MNCN 15.05/94747 – Barranc de Bec, Son Bou, Minorca; MNCN 15.05/94738 – Venta El Pilar Spring, Málaga; MNCN 15.05/94741 – Arabic Spring, Mojácar, Granada; MNCN 15.05/94743 – Fuente de los Cinco Caños Spring, Peal de Becerro, Jaén. PIGMENTATION AND ANATOMY. Animal slightly darkly pigmented, although unpigmented specimens were also found (Fig. 11D); head and tentacles black, pigmentation lighter on eye lobes, snout and neck; snout about as long as wide, approximately parallel-sided, with medium distal lobation. Ctenidium occupying almost the total length of the pallial cavity; 21–27 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 13D). Osphradium elongate, more than 3 times as long as broad, positioned opposite to the middle of the ctenidium. Stomach almost as long as wide with two nearly equally sized chambers (Supp. file 2: Table S11); style sac longer than wide, with the unpigmented intestine surrounding its distal part before continuing on as a straight rectum (Fig. 13E). MALE GENITALIA. Prostate gland bean-shaped, about 2 times as long as wide (Supp. file 2: Table S13), connected by the posterior vas deferens to a convoluted seminal vesicle and the testis (Fig. 13C). Penis darkly pigmented, gradually tapering, attached to the neck behind the right eye; penial appendix and distal end of the penis nearly equal in size; penial appendix unpigmented or slightly pigmented at the junction with the penis, rounded, small, base narrow, medially positioned on the inner edge of the penis (Fig. 13B). FEMALE GENITALIA. Glandular oviduct 2.5 times as long as wide; albumen gland longer than capsule gland (Fig. 13A); bursa copulatrix pyriform (Fig. 14B–D) to elongate (Fig. 14A), ca 2 times as long as wide (Supp. file 2: Table S12), lying against the albumen gland; bursal duct shorter than bursa length; renal oviduct unpigmented, coiled, with three loops; coils can be wide open (Fig. 14E) or highly coiled (Fig. 14F–G); seminal receptacle elongate, with a short duct, positioned on the distal part of the renal oviduct just above the junction with the bursal duct (Fig. 14E–G). NERVOUS SYSTEM. Pigmented, elongated (mean RPG ratio = 0.53; see Supp. file 2: Table S15); cerebral ganglia approximately equal in size; pleuro-supraoesophageal connective ca 9 times as long as pleuro-suboesophageal one (Fig. 13F). Ecology and distribution Mercuria balearica presents broad ecological plasticity, inhabiting springs, streams and ponds with conductivities ranging from 179 to 2750 µS/cm. Population densities were low. During our field surveys, we found evidence of local extinctions in the regions of Andalusia and Minorca (e.g., extirpated populations in the towns of Zuheros, Arriate, Alozaina, Gaucín, Ferreries), suggesting that the species was more abundant in the late twentieth century. This species occurs sympatrically with Pseudamnicola meloussensis Altaba, 2007, Theodoxus aff. fluviatilis, Melanopsis spp. and Potamopyrgus antipodarum. The species was previously cited from springs, streams, small rivers and ditches in the southern Iberian Peninsula and on the Balearic Islands of Ibiza and Minorca (Boeters 1988; Glöer et al. 2015). We confirmed the species in Minorca (Ibiza was not included as a field site) and in several other springs and streams in south-eastern Iberia (Fig. 1). Remarks The taxonomy of the species has undergone several revisions, with the most recent change occurring in 2019, when Quintana (2019) placed the species in the genus Pseudamnicola Paulucci, 1878. He had found an abundant population living in Colarsega River at Port-Mahon (Minorca Island), which is the type locality of M. balearica. He sent a subsample to the MNCN in Madrid so that we could review the classification. We confirmed that all the specimens received belong to Pseudamnicola meloussensis and communicated to Quintana that more exhaustive sampling would be necessary to find specimens of M. balearica. However, he thought that if all the specimens found in this locality are Pseudamnicola, then M. balearica and P. meloussensis are probably the same species and they should be renamed as Pseudamnicola balearica, the species described by Paladilhe (1869) (as Amnicola balearica), with P. meloussensis consequently being a junior synonym of P. balearica. However, during a more detailed sampling of this population in 2018, we were able to collect a few specimens of M. balearica living together with P. meloussensis. The DNA-based sequences (Miller et al. 2022) and shell morphology (Fig. 11C) of these specimens correspond to those of a member of Mercuria, confirming their placement in this genus (and their continued presence at the type locality). Our morphological observations (this study) and DNA results (Miller et al. 2022) confirm the previous assignments of other Iberian populations to M. balearica (Boeters 1988; Glöer et al. 2015). Populations from the stream near Sant Joan de Carbonell, Minorca (MNCN 15.05/94745), Venta El Pilar Spring, Málaga (MNCN 15.05/94738) and Fuente Valentin Spring, Cádiz (MNCN 15.05/94739) accounted for most of the morphological variation observed between Minorcan and Iberian populations. Sequence divergence of COI among these populations was low (et al. 2022). Intraspecific morphological differences for this species are observed mainly in shell, penis and female genitalia features. Specimens from Minorca are generally larger in size and present globose to high-spired shells, whereas the Iberian specimens have only globose shells (Supp. file 2: Table S10). The specimens from Los Granados Trough in Málaga (MNCN 15.05/94735) have the smallest shells. All the specimens have a pigmented epithelium, though some specimens from Arabic Spring in Mojácar (MNCN 15.05/94741) are unpigmented. Variation in the male reproductive organs is as previously described for the genus: the penis is either slightly longer or shorter than the appendix, and the appendix is usually rounded and unpigmented (only the population in Peal del Becerro, Jaén presents a slightly pigmented appendix). The bursa copulatrix varies from pyriform in most of the populations to elongate in the specimens from Venta El Pilar Spring, Málaga (MNCN 15.05/94734). Populations from Minorca have both morphotypes. Females from Venta El Pilar Spring (MNCN 15.05/94734), the stream in Montecorto (MNCN 15.05/94736), Fuente Valentín Spring (MNCN 15.05/94739) and Arabic Spring (MNCN 15.05/94741) have a bursa copulatrix that surrounds the albumen gland, which is located beneath it (see Fig. 14C) rather than parallel to it (the typical orientation in other members of the genus). Mercuria balearica can be distinguished from M. similis by its smaller, less globose shell, less convex body whorl, shorter tapering triangular penis with a rounded unpigmented penial appendix that is slightly larger than or equal in size to the penis, pyriform bursa copulatrix and higher number of cusps on the central and lateral teeth of the radula. Mercuria balearica resembles M. carrillorum sp. nov. and M. felixi sp. nov. in shell shape (ovate-conic) and colour but differs from these congeners by having a shorter, more pigmented penis with a larger rounded penial appendix. Also, their distribution areas do not overlap. Sequence divergences for COI indicated that M. balearica is most closely related to M. tensiftensis and M. felixi (5.2% mean divergence with both species) and most distantly to M. midarensis and M. melitensis (Paladilhe, 1869) (8.6% mean divergence with both species) (Miller et al. 2022)., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 21-26, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Paladilhe A. 1869. Nouvelles miscellanees malacologiques. Chez Savy, libraire-editeur, Paris. https: // doi. org / 10.5962 / bhl. title. 12995","Boeters H. 1988. Moitessieriidae und Hydrobiidae in Spanien und Portugal (Gastropoda: Prosobranchia). Archiv fur Molluskenkunde 118: 181 - 261.","Quintana J. 2019. Los moluscos del nivel B de la Cova Murada (Barranc d'Algendar, Ciutadella de Menorca, Illes Balears). Implicaciones tafonomicas. Bolleti de la Societat d'Historia Natural de les Balears 62: 15 - 23.","Breure A. & Audibert C. 2017. Under the spell of Bourguignat: Alcide Paladilhe's malacological work, with notes on his collection. Folia Conchyliologica 41: 28 - 39.","Gloer P., Boeters H. D. & Walther F. 2015. Species of the genus Mercuria Boeters, 1971 (Caenogastropoda: Truncatelloidea: Hydrobiidae) from the European Mediterranean region, Morocco and Madeira, with descriptions of new species. Folia Malacologica 23: 279 - 291. https: // doi. org / 10.12657 / folmal. 023.024","Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

Mercuria felixi Miller, García-Guerrero & Ramos sp. nov. urn:lsid:zoobank.org:act: 798D1589-DD36-4893-9FEB-8BF7026FF008 Figs 23–25; Supp. file 2: Tables S14–18 Diagnosis Shell ovate-conic; aperture obliquely ovate; protoconch microsculpture granulated; periostracum dark brown to blackish; central radular tooth formula 3(4)-C-(4)3/1-1; female genitalia with bursa copulatrix pyriform to elongate, ca 3 times as long as wide; seminal receptacle elongate; penis darkly pigmented; penial appendix ovate, darkly pigmented at the junction with the penis, about the same length or slightly shorter than the distal end of the penis and medially positioned on the inner edge of the penis; nervous system pigmented, elongate (mean RPG ratio = 0.60). Cerebral ganglia are approximately equal in size. Etymology Named after Félix Ríos Jiménez, who provided us with valuable samples of the new species. Type material Holotype SPAIN • sex unknown (dry preserved); Cádiz, stream in Canuto de la Tala; MNCN 15.05/200176H. Paratypes SPAIN – Cádiz • 27 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/94754 • 22 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/94756 • 1 spec. (dry preserved); same collection data as for holotype; MZB 2021-2798 • 2 specs (dry preserved); same collection data as for holotype; NHMW 113528 • 1 spec. (dry preserved); same collection data as for holotype; RMNH.MOL.507926 • 2 specs (dry preserved); same collection data as for holotype; JPM-586 • 1 spec. (dry preserved); same collection data as for holotype; MCP. Additional material examined SPAIN – Cádiz • 30 specs; stream in Canuto del Zapato; MNCN 15.05/94755, MNCN 15.05/94757. Additional locality information provided in Supp. file 1: Table S1. Type locality Stream in Canuto de la Tala, Cádiz, Spain. 36.477025° N, 5.592611° W, 673.97 m a.s.l. Description MEASUREMENTS. Holotype: SL = 3.78 mm, SW = 2.87 mm, SL/SW = 1.31, AH = 1.77 mm, SL-LBW = 0.90 mm, WBW = 2.25 mm, AL = 1.75 mm, AW = 1.36 mm, WPW = 1.33 mm, WAW = 0.60 mm. SHELL. Ovate-conic, whorls 4–5, height 2.4–3.8 mm, width 2–2.9 mm (Fig. 23A–F; Supp. file 2: Table S14); periostracum dark brown to blackish; protoconch of 1.5 whorls, ca 270 µm wide, nucleus ca 125 µm wide (Fig. 24A); protoconch microsculpture granulated (Fig. 24B–C). Teleoconch whorls slightly convex, separated by a deep suture; body whorl large, convex, occupying more than two-thirds of the total shell length; aperture obliquely broad ovate, complete; inner lip thicker than outer lip; outer margin straight, inner lip touching the body whorl; umbilicus narrow, not covered by the inner lip (Fig. 23A, C–F). OPERCULUM. As for the genus, orange to brown, sometimes yellowish, with about two whorls; muscle attachment oval, located near the nucleus (Fig. 23G–H). RADULA. Length intermediate, ca 700 µm long (35% of total shell length), containing about 60 rows of teeth. Central tooth formula 3(4)-C-(4)3/1-1, central cusp V shaped, cutting edge slightly concave (Fig. 24D). Lateral tooth formula 3-C-3, central cusp V shaped and slightly longer than the central tooth one. Inner marginal teeth with 15–18 cusps; outer marginal teeth with 23–27 cusps (Fig. 24E–F). Radular data were collected from specimens from the type locality. PIGMENTATION AND ANATOMY. Animal weakly pigmented (Fig. 25E); head and tentacles unpigmented to weakly pigmented; eye lobe unpigmented; snout and neck weakly pigmented; snout longer than wide, approximately parallel-sided, with medium distal lobation. Ctenidium occupying almost the total length of the pallial cavity; 21–25 gill filaments; filaments broad, triangular, fused at the base by an epithelium (Fig. 25F). Osphradium elongate, more than 3 times as long as wide, positioned opposite middle of ctenidium. Stomach almost as long as wide with two chambers almost equal in size (Supp. file 2: Table S16); style sac longer than wide, with the unpigmented intestine surrounding its distal part and then continuing on as a straight rectum (Fig. 25G). MALE GENITALIA. Prostate gland bean-shaped, about 3 times as long as wide (Supp. file 2: Table S18), connected by the posterior vas deferens to a convoluted seminal vesicle and the testis (Fig. 25C). Penis darkly pigmented, gradually tapering, attached to the neck behind the right eye; penial appendix ovate, darkly pigmented at the junction with the penis, about the same length or slightly shorter than the distal end of the penis and medially positioned on the inner edge of the penis (Fig. 25D–E). FEMALE GENITALIA. Glandular oviduct 3 times as long as wide; albumen gland ca 2 times as long as capsule gland; bursa copulatrix pyriform to elongate, ca 3 times as long as wide (Fig. 25A–B; Supp. file 2: Table S17); bursal duct shorter than bursa copulatrix; renal oviduct unpigmented, highly coiled with three loops; seminal receptacle elongate, with a short duct, positioned on the distal part of the renal oviduct just above the junction with the bursal duct (position SR1). NERVOUS SYSTEM. Pigmented, elongate (mean RPG ratio = 0.60; see Supp. file 2: Table S15); cerebral ganglia approximately equal in size, pleuro-supraoesophageal connective ca 11 times as long as pleuro-suboesophageal one (Fig. 25H). Ecology and distribution This species was collected from two streams in southern Spain: the stream in Canuto de la Tala (type locality) and the stream in Canuto del Zapato. The ecology of the species is not well known. The species inhabits streams of very clear running waters with relatively low conductivities (75–292 mS/cm 2). The specimens collected from the stream in Canuto del Zapato were found in very low abundance. Co-occurring species are Ancylus fluviatilis and Galba truncatula in low abundance and Pisidium sp. in high abundance (Félix Ríos Jiménez pers. com.). Remarks Mercuria felixi sp. nov. differs from the phylogenetically closely related species M. tensiftensis and M. balearica by having a narrower aperture (wider in M. balearica); a more globose shell (high-spired in M. tensiftensis); a granulated protoconch microsculpture (grooved in M. tensiftensis); an ovate, pigmented penial appendix that is about the same length or slightly shorter than the distal end of the penis (rounded, unpigmented and shorter than the distal end of the penis in M. balearica and darkly pigmented and shorter than the distal end of the penis in M. tensiftensis); a thin, filiform, pigmented distal end of the penis (broad, triangular, and strongly and darkly pigmented in M. balearica, and triangular and pigmented in M. tensiftensis); a prostate gland that is 3 times as long as wide (2 times as long as wide in M. tensiftensis and M. balearica) and a smaller radular ribbon. The species also has one more cusp on the central teeth than in M. tensiftensis. The shell shape of M. felixi sp. nov. resembles that of M. carrillorum sp. nov. and M. tachoensis; however, our PCA (Fig. 3) showed that M. felixi can be differentiated from the rest of the Mercuria species. The penile morphology of M. felixi further differentiates it from the Iberian congeners: it has a very thin, pigmented penial appendix and a long, filiform and pigmented distal penis. According to DNA-sequence analyses (Miller et al. 2022), this species forms a clade with M. balearica (5.3% of mean COI divergence) and M. tensiftensis (5.1% of mean COI divergence). It is most distantly related to M. midarensis and M. saharica, diverging from these species by 8% and 8.6%, respectively., Published as part of Miller, Jonathan P., Delicado, Diana, García-Guerrero, Fernando, Khalloufi, Noureddine & Ramos, Marian A., 2023, Morphology and taxonomic assessment of eight genetic clades of Mercuria Boeters, 1971 (Caenogastropoda, Hydrobiidae), with the description of five new species, pp. 1-63 in European Journal of Taxonomy 866 (1) on pages 38-42, DOI: 10.5852/ejt.2023.866.2107, http://zenodo.org/record/7867353, {"references":["Miller J. P., Delicado D., Garcia-Guerrero F. & Ramos M. A. 2022. Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae). Molecular Phylogenetics and Evolution 173: 107524. https: // doi. org / 10.1016 / j. ympev. 2022.107524"]}

RADOMANIOLA PESICI SP. NOV. (FIGS 18, 19) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. o r g: a c t: 42A9C1DA-B6A9-40BE-9A96-4F381D133CA2 Etymology: Named after Professor Vladimir PeŠić, collector of this species, in recognition of his valuable contribution to the knowledge of the fauna of Montenegro. Type material: Holotype (MNCN 15.05 /200163), five paratypes (MNCN 15.05 /200164) in the MNCN collection and ~ 20 paratypes (UGSB 19048) in the UGSB collection. Type locality: Bar Spring, Donje Vrelo, Montenegro. M a t e r i a l s t u d i e d: B a r S p r i n g, D o n j e V r e l o, Montenegro, 42.86°N, 20.25°E, V.P., May 2005, MNCN 15.05 /200164 and UGSB 19048 (80% ethanol). Diagnosis: Protoconch microsculpture wrinkled; central radular tooth formula 7-C-7/1-1; bursa copulatrix globular, with a long duct; SR1 elongate, duct short; SR2 slightly shorter than SR1, elongate, duct short; penis unpigmented, gradually tapering, base narrow, approximately as long as head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.47). Description: Shell ovate-conic, 3.5–4.0 whorls, height 1.75–2.25 mm (Fig. 18A–D; Supporting Information, Table S6). Periostracum whitish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture pitted (Fig. 18G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 18B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 18E, F). Radular length intermediate, ~600 µm (~25% of total shell length), with ~60 rows of teeth (Fig. 18H). Central tooth formula 7-C-7/1-1 (Fig. 18I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 4-C-4. Inner marginal teeth having 30–32 tapered cusps, shortening toward the base. Outer marginal teeth with 30–35 sharp cusps (Fig. 18J). Animal darkly pigmented except for neck and tentacles (Fig. 19F). Ctenidium with ten or 11 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 19A). Stomach as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 19B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio= 0.47); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 19C). Female glandular oviduct approximately two times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix globular, slightly longer than wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 slightly shorter than SR1, elongate, with a short duct, located on renal oviduct near loop (Fig. 19D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 19H). Penis unpigmented, gradually tapering, approximately as long as head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 19F, G; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight. Remarks: This species resembles R. montana and R. filiola in bearing a more ovate shell than other congeners. Anatomically, R. pesici differs from the two species according to its larger bursa copulatrix, more slender penis, longer radula and larger central radular teeth. Average sequence divergence among the three species ranges between 5.3 and 7.9% for COI. Radomaniola pesici is also differentiated from the geographically proximate R. curta and R. wolffi in its shorter and more ovate shell, a larger number of cusps on the central and lateral teeth, smaller bursa copulatrix with a shorter duct, shorter penis and an average sequence divergence of 4.5–6.2% for COI (Supporting Information, Table S4)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 423-425, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584

RADOMANIOLA CURTA SUBSP. OMBLENSIS SUBSP. NOV. (FIGS 8, 9) Radomaniola curta curta (Küster, 1842) – Hirschfelder, 2017 Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: 903123AE-A06C-4F45-8759-5F7E68F93BBF Etymology: Referring to Ombla Spring, in which this subspecies was originally found. Type material: Holotype (MNCN 15.05/200155), six paratypes (MNCN 15.05/200156) in the MNCN collection and ~ 70 paratypes (UGSB 18778) in the UGSB collection. Type locality: Ombla Spring, Rozat, Croatia. Material studied: Ombla Spring, Rozat, Croatia, 42.6749°N, 18.1374°E, T.H. and D.D., June 2016, MNCN 15.05/200156 and UGSB 18778 (80% ethanol). Diagnosis: Protoconch microsculpture wrinkled; central radular tooth formula 7-C-7/1-1; bursa copulatrix ovoid, with a long duct; SR1 elongate, duct short; SR2 small, pyriform to elongate, duct short; penis unpigmented, gradually tapering, smaller than head length and with one medial outgrowth on its left side; nervous system moderately concentrated (mean RPG ratio = 0.49). Description: Shell ovate-conic, 4.0–4.5 whorls, height 2.5–3.2 mm (Fig. 8A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture pitted (Fig. 8G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 8B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish to yellowish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 8E, F). Radular length intermediate, ~700 µm (~25% of total shell length), with ~50 rows of teeth (Fig. 8H). Central tooth formula 7-C-7/1-1 (Fig. 8I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 3-C-3. Inner marginal teeth having 20–25 tapered cusps, shortening toward the base. Outer marginal teeth with 18–26 sharp cusps (Fig. 8J). Animal darkly pigmented except for neck and tentacles (Fig. 9F). Ctenidium with 14–16 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 9A). Stomach approximately as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 9B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.49); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 9C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, about as long as wide. Bursal duct slightly longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, with a short duct, joining renal oviduct slightly above the insertion point with bursal duct. SR2 shorter than SR1, pyriform to elongate, with a short duct, located on renal oviduct near loop (Fig. 9D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 9I). Penis unpigmented, gradually tapering, smaller than head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 9F–H; Supporting Information, Table S9); penis attached to the area close to the right eye; penial duct narrow, near outer edge, almost straight. Habitat: Radomaniola curta omblensis is known only from its type locality, a large karstic spring fed by subterranean waters. Snails were deeply submerged (> 1 m) and attached to the walls, the aquatic vegetation and stones. A closely co-occurring species was Bithynia tentaculata (Linnaeus, 1758). Remarks: Hirschfelder (2017) found Radomaniola - like snails in Ombla Spring and assigned them to R. c. curta. Our morphological and genetic data from a Radomaniola population living in the same spring indicate that this population differs from R. c. curta and from any other subspecies. Specifically, this new subspecies differs from the conchologically similar R. c. curta and R. c. kicavica according to its more slender shell, slightly larger bursa copulatrix and shorter bursal duct. The COI sequence divergence with other subspecies ranges between 1.2 and 2.6%., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 410-413, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584, {"references":["Hirschfelder HJ. 2017. Neue Molluskennachweise aus der Ombla-Quelle bei Dubrovnik. Mitteilungen der Deutschen Malakozoologischen Gesellschaft 96: 33 - 38."]}

RADOMANIOLA VARIABILIS SP. NOV. (FIGS 22, 23) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: BACA04A4-0420-4FB3-B15B-A3CD6789A303 Etymology: From Latin variabilis, variable, referring to its high intraspecific variation in shell shape. Type material: Holotype (MNCN 15.05 /200169), three paratypes (MNCN 15.05 /200170) in the MNCN collection and ten paratypes in the UGSB collection (UGSB 23975). Type locality: Vellas Spring, Kalpaki, Greece. Material studied: Vellas Spring, Kalpaki, Greece, 39.8660°N, 20.6243°E, T.W. and C.A., May 2005, MNCN 15.05 /200170 and UGSB 23975 (80% ethanol); unnamed spring, 3 km west of Qafë Murrës, Albania, 41.63°N, 20.2°E, Z.F., June 2003, UGSB 14440 (80% ethanol); unnamed spring, 7 km south of Tepelenë, Albania, 40.27°N, 20.28°E, Z.F., October 2004, UGSB 14441 (80% ethanol); Syri i Kaltër Lake, 7 km west of Muzinë, Albania, 39.92°N, 20.18°E, Z.F., October 2004, UGSB 2294 (80% ethanol). Diagnosis: Protoconch microsculpture pitted; central radular tooth formula 7-C-7/1-1; bursa copulatrix ovoid, duct slightly longer than bursal length; SR1 pyriform, duct short; SR2 pyriform, duct short, slightly larger than SR1; penis unpigmented, gradually tapering, base narrow, approximately as long as head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.47). Description: Shell ovate-conic, 4.0–4.5 whorls, height 2.0– 2.6 mm (Fig. 22A–F; Supporting Information, Table S6). Periostracum yellowish, sometimes whitish. Protoconch ~425 µm wide, 1.5 whorls; nucleus ~175 µm wide; protoconch microsculpture pitted (Fig. 22I). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 22B, F). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 22G, H). Radular length intermediate, ~600 µm (~25% of total shell length), with ~60 rows of teeth. Central tooth formula 7-C-7/1-1 (Fig. 22J–L); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula (3)4-C-4(3). Inner marginal teeth having 25–32 tapered cusps, shortening toward the base. Outer marginal teeth with 27–34 sharp cusps (Fig. 22M, N). Animal darkly pigmented except for neck and tentacles (Fig. 23G, H). Ctenidium with 12 or 13 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 23A). Stomach as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 23B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio= 0.47); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 23C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, slightly longer than wide. Bursal duct slightly longer than bursal length. Renal oviduct unpigmented, coiled. SR1 pyriform, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 slightly larger than SR1, pyriform, with a short duct, located on renal oviduct near loop (Fig. 23D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 23F). Penis unpigmented, gradually tapering, approximately as long as head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 23G–K; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight. Habitat: Radomaniola variabilis lives in cold water springs and lacustrine habitats; it has been found on stones and aquatic vegetation. It co-occurs with Ancylus fluviatilis Müller, 1774, Radix cf. peregra (Müller, 1774) and Bithynia sp. in the type locality. Remarks: Populations of R. variabilis are particularly variable in the number of cusps on the lateral radular teeth, intensity of body pigmentation, penis width (Figs 22, 23) and shell morphology (e.g. shells of the Vella Spring population in Greece are slightly larger, more elongate than those of the Albanian populations and present a smaller aperture with a thin inner lip). Female genitalia could not be examined in detail because most of the dissected females from the type locality were parasitized and had stunted organs. Also, we found no females among the few specimens collected at the localities in Albania. Although the ABGD and GMYC methods subdivided this species into three entities, we consider this taxon as a single species based on the geographical proximity and phylogenetic clustering of the studied populations as a monophyletic group, a small intraspecific divergence COI and similar shape of the shell and penis. Fehér & Erőss (2009) assigned the three populations from Albania studied herein to R. curta based on shell morphology. However, penis features and genetic data suggest that these populations are members of R. variabilis and not of R. curta. The authors also reported the species R. curta and R. albanica co-occurring in the Syri i Kaltër Spring (Fehér & Erőss, 2009). Based on morphological and molecular results, we conclude that the snails from the same locality examined in the present study are likely to be conspecific with R. variabilis and R. albanica. Radomaniola variabilis is morphologically distinguishable from the geographically and phylogenetically proximate species R. dolens, R. albanica and R. haesitans according to its wider shell, wider shell umbilicus, longer penis, shorter bursal duct and SR1 and sequence divergence from these three species (5–6% average divergence for COI)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 427-430, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584, {"references":["Feher Z, Eross ZP. 2009. Contribution to the Mollusca fauna of Albania. Results of the field trips of the Hungarian Natural History Museum between 1992 and 2007. Schriften zur Malakozoologie 25: 3 - 21."]}

RADOMANIOLA WOLFFI SP. NOV. (FIGS 24, 25) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: 7373D604-929B-4E92-AF4E-11F19EB6DD96 Etymology: Named after Christian Wolff, collector of this and other gastropod species in the Balkan Peninsula. Type material: Holotype (MNCN 15.05 /200171), ten paratypes (MNCN 15.05 /200172) in the MNCN collection and ~ 50 paratypes in the UGSB collection (UGSB 19533). Type locality: Vitoja Spring, north of Lake Scutari, Montenegro. Material studied: Vitoja Spring, north of Lake Scutari, Montenegro, 42.3252°N, 19.3628°E, T.H. and C.W., 20 September 2009, MNCN 15.05 /200172 and UGSB 19533 (80% ethanol). Diagnosis: Protoconch microsculpture wrinkled; central radular tooth formula (3)4-C-4(3)/1-1; bursa copulatrix ovoid, with a duct as long as bursal length; SR1 pyriform, duct long; SR2 pyriform, duct short; penis unpigmented, gradually tapering, base narrow, longer than head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.45). Description: Shell ovate-conic, 4.0–4.5 whorls, height 2.3–3.0 mm, apex often eroded (Fig. 24A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~450 µm wide, 1.5 whorls; nucleus ~175 µm wide; protoconch microsculpture wrinkled (Fig. 24G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 24B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 24E, F). Radular length intermediate, ~600µm (~25% of total shell length), with ~45 rows of teeth (Fig. 24H). Central tooth formula (3)4-C-4(3)/1-1 (Fig. 24I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 3-C-3. Inner marginal teeth having 18–20 tapered cusps, shortening toward the base. Outer marginal teeth with 21–25 sharp cusps (Fig. 24J). Animal darkly pigmented except for neck and tentacles (Fig. 25F). Ctenidium with 14–17 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 25A). Stomach slightly longer than wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 25B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.45); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 25C). Female glandular oviduct approximately two times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, slightly longer than wide. Bursal duct as long as bursal length. Renal oviduct unpigmented, coiled. SR1 pyriform, duct long, joining renal oviduct slightly above the insertion point with bursal duct. SR2 shorter than SR1, pyriform, with a short duct, located on renal oviduct near loop (Fig. 25D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 25H). Penis unpigmented, gradually tapering, longer than head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 25F, G; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight. Habitat: Known only from a cold, large, karstic spring, Vitoja Spring, in Montenegro (for details of this locality, see Grego et al., 2019). The water conductivity was low (245 µS/cm) and vegetation sparse. The species was found mainly on stones, rocks, walls and detritus. Other gastropod species co-occurring in this spring are Pyrgula annulata (Linnaeus, 1758), Theodoxus sp., Bithynia sp., Planorbarius sp., Valvata sp. and Galba truncatula (Müller, 1774). Remarks: This species is conchologically similar and geographically proximate to Radomaniola elongata (Radoman, 1973), which was originally described based on the shell (Radoman, 1973), and only the anatomy of its penis is known (Falniowski et al., 2012). However, they can be distinguished by the narrower penis in R.wolffi (Fig. 25G) and an average sequence divergence of 9.1% for COI (Supporting Information, Table S4). Another Radomaniola species from the Lake Scutari region, Radomaniola lacustris (Radoman, 1983), for which only shell features are known (Radoman, 1983), differs from R. wolffi in having a more globose shell with a larger aperture., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 430-433, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584, {"references":["Grego J, Gloer P, Falniowski A, Hofman S, Osikowski A. 2019. New subterranean freshwater gastropod species from Montenegro (Mollusca, Gastropoda, Moitessieriidae, and Hydrobiidae). Ecologica Montenegrina 20: 71 - 90.","Radoman P. 1973. New classification of fresh and brackish water Prosobranchia from the Balkans and Asia Minor. Prirodnjacki Muzej Beogradu Posedna Izdanja 32: 1 - 30.","Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Srpska Akademija Nauka i Umetnosti, Posebna Izdanja (Odeljenje PrirodnoMatematitschkich Nauka) 57: i - ii + 1 - 256."]}

RADOMANIOLA JOVANOVSKAE SP. NOV. (FIGS 14, 15) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: 73C15FFA-001B-4C60-A237-C42DC17B52D8 Etymology: This species is named after Dr Elena Jovanovska, a dedicated researcher of the diatom flora in Lake Ohrid and Lake Prespa, who provided invaluable logistical support for this study. Type material: Holotype (MNCN 15.05 /200161), 12 paratypes (MNCN 15.05 /200162) in the MNCN collection and> 100 paratypes (UGSB 19517) in the UGSB collection. Type locality: Oteshevo Sirhan Spring, Sirhan, North Macedonia. Material studied: Oteshevo Sirhan Spring, Sirhan, North Macedonia, 41.0032°N, 20.9357°E, D.L. and S.N., May 2008, MNCN 15.05 /200162 and UGSB 19517 (80% ethanol). Diagnosis: Protoconch microsculpture wrinkled; shell aperture slightly oval; central radular tooth formula 7-C-7/1-1; bursa copulatrix ovoid, with a duct longer than bursal length; SR1 pyriform, duct short; SR2 pyriform, duct short; SR1 longer than SR2; penis unpigmented, gradually tapering, shorter than head length, base narrow; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.41). Description: Shell ovate-conic, 4.00–4.25 whorls, height 2.1–2.9 mm (Fig. 14A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture wrinkled (Fig. 14G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 14B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 14E, F). Radular length intermediate, ~650µm (~25% of total shell length), with ~55 rows of teeth (Fig. 14H). Central tooth formula 7-C- 7/1-1 (Fig. 14I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 4-C-4. Inner marginal teeth having 25 or 26 tapered cusps, shortening toward the base. Outer marginal teeth with 23 or 24 sharp cusps (Fig. 14J). Animal darkly pigmented except for neck and tentacles (Fig. 15F). Ctenidium with 11 or 12 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 15A). Stomach slightly wider than long, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 15B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.41); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 15C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, slightly longer than wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 pyriform, duct long, joining renal oviduct slightly above the insertion point with bursal duct. SR2 shorter than SR1, pyriform, with a short duct, located on renal oviduct near loop (Figs 15D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 15H). Penis unpigmented, gradually tapering, shorter than head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 15F, G; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, undulated. Habitat: This species has been collected from one spring in the watershed of Lake Prespa. Snails occur abundantly on stones together with Ancylus sp. Remarks: This species is conchologically similar to the geographically proximal R. c. kicavica (Supporting Information, Fig. S2). However, these two taxa can be distinguished anatomically according to a shorter bursal duct, more elongated bursa copulatrix and smaller seminal receptacles in R. jovanovskae, and larger and more tapering penis and larger prostate gland in R. c. kicavica (Supporting Information, Figs S6, S 8). The average sequence divergence between the two taxa was 4% for COI., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 418-419, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584

RADOMANIOLA NACHTIGALLAE SP. NOV. (FIGS 16, 17) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: 0139AB61-FB6A-42C4-814A-78E132AAEC1A Etymology: Named after Silvia Nachtigall, who helped us with much of the preparation of material and the laboratory work conducted for this and other studies. Type material: Holotype (MNCN 15.05 /200165), ten paratypes (MNCN 15.05 /200166) in the MNCN collection and> 100 paratypes (UGSB 18857) in the UGSB collection. Type locality: source of the Krupa River, Krupa na Vrbasu, Bosnia and Herzegovina. Material studied: Source of the Krupa River, Krupa na Vrbasu, Bosnia and Herzegovina, 44.6207°N, 1 7. 1 1 0 5°E, T.H. a n d D.D., Ju n e 2 0 1 6, M N C N 15.05/200166 and UGSB 18857 (80% ethanol). Diagnosis: Protoconch microsculpture slightly wrinkled; central radular tooth formula 7-C-7/1- 1; bursa copulatrix globular, with a long duct; SR1 elongate, duct short; SR2 elongate, duct short; penis slightly pigmented on the distal section, gradually tapering, shorter than head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.46). Description: Shell ovate-conic, four whorls, height 2.2–2.8 mm (Fig. 16A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~550 µm wide, 1.5 whorls; nucleus ~200 µm wide; protoconch microsculpture slightly wrinkled (Fig. 16G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 16B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 16E, F). Radular length intermediate, ~650 µm (~25% of total shell length), with ~60 rows of teeth (Fig. 16H). Central tooth formula 7-C-7/1-1 (Fig. 16I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula (5)4-C-4(5). Inner marginal teeth having 30–32 tapered cusps, shortening toward the base. Outer marginal teeth with 35–37 sharp cusps (Fig. 16J). Animal darkly pigmented except for neck and tentacles (Fig. 17F). Ctenidium with 13–15 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 17A). Stomach slightly longer than wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 17B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.46); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 17C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix globular, as long as wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 slightly shorter than SR1, elongate, with a short duct, located on renal oviduct near loop (Fig. 17D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 17H). Penis slightly pigmented on the distal section, gradually tapering, shorter than head length, base medium wide, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 17F, G; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight. Habitat: This species has been found only in a rheocrene spring of clear and shallow waters presenting low temperature (10.5 °C) and conductivity (465 µS/ cm). Aquatic plants are sparse. Snails are abundant on stones. Other co-occurring gastropods are Sadleriana sp. and Ancylus sp. Remarks: This species is closely related to the geographically proximate congeners R. bosniaca and R. szarowskae. The average COI sequence divergence between R. nachtigallae and these two species is 2% and 4%, respectively. This small genetic divergence contrasts with their considerable morphological differentiation in terms of shell size, shell shape and other characters of the reproductive system. Thus, R. nachtigallae differs from R. bosniaca in that it has a larger shell with taller spire whorls, wider base of penis, shorter penis compared with head length, more globular bursa copulatrix, longer bursal duct and more elongated SR1; and differs from R. szarowskae in its smaller shell, smaller penis and smaller bursa copulatrix (Supporting Information, Figs S2, S 6, S 8). These three species have a similar number of gill filaments and cusps on the radular teeth and RPG ratio., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 419-422, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584

LERNIANA GEN. NOV. Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: AAB227D3-CBD8-4E8A-A001-4AA29A9B4256 Type species: Hydrobia tritonum Bourguignat, 1852 (also known as Semisalsa tritonum; Schütt, 1980). Etymology: The genus name refers to the Greek name Λέρνη, Lerna, a spring region that was once a lake and today is home to a population of the type species; gender feminine. Diagnosis: Shell ovate-conic; 4.0–4.5 whorls; aperture oval with a straight narrow outer lip; umbilicus narrow. Operculum corneous, yellowish, thin, pliable, oval, without peg, and paucispiral with submarginal nucleus. One pair of basal cusps on radular central tooth. Ctenidium well developed, with 10–13 gill filaments. Osphradium positioned opposite approximate middle of ctenidium. Stomach without gastric caecum. Bursa copulatrix medium sized, pyriform, pedunculated and lying against the anterior section of the albumen gland; unpigmented renal oviduct; two seminal receptacles; SR2 smaller than SR1 and arising at the renal oviduct loop. Prostate gland bean shaped, about twice as long as wide. Penis strap-like, simple or with one small, medial outgrowth on its left side. Nervous system slightly pigmented, moderately concentrated. Remarks: Lerniana is distinguished here from Radomaniola according to its smaller shell with a shorter spire, smaller stomach, pyriform bursa copulatrix with a shorter duct (of globular shape and with a longer duct in Radomaniola), smaller SR2, strap-like penis [although sometimes also gradually tapering, see Falniowski etal. (2012: 29, fig.7G1, G5) and Georgiev (2013: 294, fig. 2.4)] with a small outgrowth on the left side (larger and gradually tapering with a more prominent outgrowth in Radomaniola) and a generally more concentrated dorsal nerve ring (RPG ratio Lerniana) (Supporting Information, Figs S3, S 7, S 9). Moreover, all phylogenetic analyses congruently depicted the three species assigned to this new genus (i.e. R. tritonum, R. feheri and R. seminula) as a well-supported clade outside Radomaniola (Fig. 2). Falsibelgrandiella Radoman, 1973 is remarkably different from Lerniana because it bears a tubular shell, distinct shape and size of the seminal receptacles, and a penis with a wider base (Radoman, 1983). Lerniana is more similar in shell shape to Turcorientalia Radoman, 1973 and Pseudorientalia Radoman, 1983 but differs from Graecorientalia Radoman, 1983 in having a larger and pyriform bursa copulatrix with a shorter duct, larger SR1 and narrower base of the penis; from Turcorientalia in terms of its smaller bursa copulatrix, larger seminal receptacles and longer and narrower penis; and from Pseudorientalia according to its narrower bursa copulatrix, larger seminal receptacles and narrower penis (Radoman, 1983)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on page 433, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584, {"references":["Schutt H. 1980. Zur Kenntnis griechischer Hydrobiiden. Archiv fur Molluskenkunde 110: 115 - 149.","Georgiev D. 2013. A new species of Radomaniola Szarowska, 2006 (Gastropoda: Hydrobiidae) from Peloponnese, Greece. Acta Zoologica Bulgarica 65: 293 - 295.","Radoman P. 1973. New classification of fresh and brackish water Prosobranchia from the Balkans and Asia Minor. Prirodnjacki Muzej Beogradu Posedna Izdanja 32: 1 - 30.","Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Srpska Akademija Nauka i Umetnosti, Posebna Izdanja (Odeljenje PrirodnoMatematitschkich Nauka) 57: i - ii + 1 - 256."]}

RADOMANIOLA CURTA SUBSP. MONTENEGRENSIS SUBSP. NOV. (FIGS 6, 7) Zoobank registration: zoobank.org:act: 6D859A59- 9357-4949-99F4-3960B1E5FBDD Etymology: The subspecies name refers to the occurrence of this snail in Montenegro. Type material: Holotype (MNCN 15.05/200157), five paratypes (MNCN 15.05/200158) in the MNCN collection and ~ 50 paratypes (UGSB 19515) in the UGSB collection. Type locality: Drobnica Spring, Reljići, Montenegro. Material studied: Drobnica Spring, Reljići, Montenegro, 42.2000°N, 19.0593°E, T.H. and C.W., September 2009, MNCN 15.05/200158 and UGSB 19515 (80% ethanol). Maricina Spring, Zabes, Montenegro, 42.2287°N, 19.0907°E, T.H. and C.W., September 2009, UGSB 19051 (80% ethanol). Outflow from the Velje Oko Spring, Gluhi do, Montenegro, 42.2142°N, 19.0512°E, T.H. and C.W., September 2009, UGSB 19043 (80% ethanol). Diagnosis: Protoconch microsculpture wrinkled; central radular tooth formula 7-C-7/1-1; bursa copulatrix ovoid, with a long duct; distal seminal receptacle (SR1) pyriform, duct medium length; proximal seminal receptacle (SR2) small, pyriform to elongate, duct short; penis unpigmented, gradually tapering, slightly longer than head length, attached to the central area of the head; nervous system moderately concentrated (mean RPG ratio = 0.47). Description: Shell ovate-conic, 4.0–4.5 whorls, height 2.5–3.0 mm (Fig. 6A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture wrinkled (Fig. 6G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 6B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 6E, F). Radular length intermediate, ~600 µm (~25% of total shell length), with ~60 rows of teeth (Fig. 6H). Central tooth formula 7-C-7/1-1 (Fig. 6I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 4-C-4. Inner marginal teeth having 24 or 25 tapered cusps, shortening toward the base. Outer marginal teeth with 23–26 sharp cusps (Fig. 6J). Animal darkly pigmented except for neck and tentacles (Fig. 7H). Ctenidium with 14 or 15 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 7A). Stomach approximately as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 7B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.47); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 7C). Female glandular oviduct approximately two times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, slightly longer than wide. Bursal duct slightly longer than bursal length. Renal oviduct unpigmented, coiled. SR1 pyriform, with duct of medium length, joining renal oviduct at the insertion point with the bursal duct. SR2 shorter than SR1, pyriform to elongate, with a short duct, located on renal oviduct near loop (Fig. 7D–G; Supporting Information, Table S8). Malegenitaliawithaprostateglandapproximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 7L). Penis unpigmented, gradually tapering, longer than head length, base narrow, weakly folded along inner edge and rarely with one medial outgrowth on its left side (Fig. 7H–K; Supporting Information, Table S9); penis attached to the central area of the head; penial duct narrow, near outer edge, almost straight. Habitat: Radomaniola curta montenegrensis has been collected from three closely proximal rheocrene springs located close to the Lake Scutari. These springs were of cold, low water flow. Snails were abundant on stones and in the mud at all localities. No direct co-occurring species was found. Remarks: All three studied populations of R. c. montenegrensis varied in shell size and shape and penis size (Figs 6, 7; Supporting Information, Tables S6 and S9). Despite their morphological differentiation, COI sequence divergence was low (0.1%). This subspecies differs from the geographically proximate R. c. curta, Radomaniola curta anagastica (Radoman, 1973) and Radomaniola curta maxima subsp. nov. in its shorter and more ovate shell (see Radoman, 1983; Supporting Information, Table S6), smaller bursa copulatrix (Supporting Information, Table S8), a shorter distance between seminal receptacles and an average sequence divergence with R. c. curta and R. c. maxima of 1.2–2.5% for COI. The COI sequence divergence between R. c. montenegrensis and one individual from the type locality area of R. c. anagastica, sequenced by Falniowski et al. (2012) (GenBank accession number: KC011773), was substantial (5.4%)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 407-410, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584, {"references":["Radoman P. 1973. New classification of fresh and brackish water Prosobranchia from the Balkans and Asia Minor. Prirodnjacki Muzej Beogradu Posedna Izdanja 32: 1 - 30.","Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Srpska Akademija Nauka i Umetnosti, Posebna Izdanja (Odeljenje PrirodnoMatematitschkich Nauka) 57: i - ii + 1 - 256."]}

RADOMANIOLA SZAROWSKAE SP. NOV. (FIGS 20, 21) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: 605481A3-A28F-4DC0-B532-9A1534B21887 Etymology: Named after Dr Magdalena Szarowska, in recognition of her many contributions to the evolution of Truncatelloidean gastropods of the Balkan Peninsula, including species of this genus. Type material: Holotype (MNCN 15.05 /200167), ten paratypes (MNCN 15.05 /200168) in the MNCN collection and 60 paratypes (UGSB 18566) in the UGSB collection. Type locality: source of the Cetina River, Cetina, Croatia. Material studied: Source of the Cetina River, Cetina, Croatia, 43.9766°N, 16.4301°E, T.H. and D.D., June 2016, MNCN 15.05 /200168 and UGSB 18566 (80% ethanol). Diagnosis: Shell ovate-conic, 3.5–4.0 whorls (apex often eroded); protoconch microsculpture wrinkled; central radular tooth formula 6-C-6/1-1; bursa copulatrix ovoid, with a duct as long as bursal length; SR1 elongate, duct short; SR2 elongate, same length as SR1, duct short; penis unpigmented, gradually tapering, base wide, shorter than head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.47). Description: Shell ovate-conic, 3.5–4.0 whorls, height 2.3–3.0 mm, apex often eroded (Fig. 20A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~500 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture pitted (Fig. 20G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 20B). Umbilicus narrow, not covered by the inner lip. Operculum oval, yellowish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 20E, F). Radular length intermediate, ~700 µm (~25% of total shell length), with ~50 rows of teeth (Fig. 20H). Central tooth formula 6-C-6/1-1 (Fig. 20I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 4-C-4. Inner marginal teeth having 20–25 tapered cusps, shortening toward the base. Outer marginal teeth with 20–24 sharp cusps (Fig. 20J). Animal darkly pigmented except for neck and tentacles (Fig. 21F). Ctenidium with 13–15 welldeveloped gill filaments, occupying ~50% of pallial cavity length and posteriorly positioned. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 21A). Stomach slightly longer than wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 21B; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.47); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 21C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, slightly longer than wide. Bursal duct as long as bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 as long as SR1, elongate, with a short duct, located on renal oviduct near loop (Fig. 21D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 21H). Penis unpigmented, gradually tapering, shorter than head length, base wide, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 21F, G); penis attached close to the right eye; penial duct narrow, near outer edge, almost straight. Habitat: The source of the Cetina River is a large, deep, karst spring located in the north-western slopes of the Dinara Alps. Snails are distributed on stones submerged at 0.5 m along the shore of the quiet pool formed by the spring waters. The temperature and conductivity of the water are low (9.6 °C and 350 µS/cm) and aquatic vegetation is sparse. Other co-occurring gastropods are Horatia klecakiana (Bourguignat, 1887) and Ancylus sp. Remarks: This is one of the largest species recorded so far for this genus. Conchologically, R. szarowskae resembles R. c. kicavica, but it differs anatomically from this subspecies according to its larger penis, larger and more globous bursa copulatrix, less concentrated nervous ring and a larger number of cusps on the lateral radular teeth. Phylogenetically, the two taxa are distantly positioned from one another (Fig. 2), and their COI average sequence divergence is 5.6%. Radomaniola szarowskae differs from the phylogenetically and geographically close species R. bosniaca and R. nachtigallae in that it has a more conic and larger shell, more globous and larger bursa copulatrix and in the shape and size of the penis (Supporting Information, Figs S2, S 6, S 8)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 425-427, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584

RADOMANIOLA CURTA SUBSP. MAXIMA SUBSP. NOV. (FIGS 10, 11) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: D86C5BA4-235B-410B-93B7-531D97B30A4D Etymology: From Latin maximus, very large, referring to its large body size. Type material: Holotype (MNCN 15.05 /200153), 20 paratypes (MNCN 15.05 /200154) in the MNCN collection and> 500 paratypes (UGSB 19540) in the UGSB collection. Type locality: Crnojevića River, Montenegro. Material studied: Crnojevića River, near the bridge, Montenegro, 42.3550°N, 19.0284°E, T.H., C.W. and D.E., September 2009, MNCN 15.05 /200154 and UGSB 19540 (80% ethanol). Diagnosis: Protoconch microsculpture pitted; central radular tooth formula 5-C-5/1-1; bursa copulatrix ovoid, with a long duct; SR1 elongate, duct short; SR2 small, pyriform, duct short; penis unpigmented, gradually tapering, shorter than head length, sometimes with one medial outgrowth on its left side; nervous system unpigmented, elongate (mean RPG ratio = 0.52). Description: Shell ovate-conic, 4.0–4.5 whorls, height 2.5–3.5 mm (Fig. 10A–D; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture pitted (Fig. 10G). Teleoconch whorls slightly convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 10B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 10E, F). Radular length intermediate, ~700 µm (~25% of total shell length), with ~50 rows of teeth (Fig. 10H). Central tooth formula 5-C- 5/1-1 (Fig. 10I); basal tongue V-shaped, length about equal to lateral margin. Lateral tooth formula (4)3-C- 3(4). Inner marginal teeth bearing 18–21 tapered cusps, shortening toward the base. Outer marginal teeth with 22–23 sharp cusps (Fig. 10J). Animal darkly pigmented except for neck and tentacles (Fig. 11F). Ctenidium with 13–16 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 11A). Stomach approximately as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 11B; Supporting Information, Table S7). Nervous system unpigmented, elongate (mean RPG ratio = 0.52); cerebral ganglia approximately equal in size (Fig. 11C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix ovoid, about as long as wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, with a short duct, joining renal oviduct slightly above the insertion point with bursal duct. SR2 shorter than SR1, pyriform, with a short duct, located on renal oviduct near loop (Fig. 11D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 11H). Penis unpigmented, gradually tapering, shorter than head length, weakly folded along inner edge and with one medial outgrowth (sometimes absent) on its left side (Fig. 11F, G; Supporting Information, Table S9); penis attached to the area close to the right eye; penial duct narrow, near outer edge, almost straight. Habitat: This is a riverine subspecies collected from the margins of the Crnojevića River when the water level was low. Snails were found in high abundance on stones and small rocks. Co-occurring species are Ancylus sp., Theodoxus sp., Bithynia sp. and Anagastina matjasici (Bole, 1961). Remarks: Although specimens of R. c. maxima are larger than those of other conspecific populations and present a more ovate shell shape, they are included in R. curta by the presence of a long SR1, an ovoid bursa copulatrix with a long duct, a gradually tapering penis with a medial outgrowth on the left edge and an RPG ratio from moderately concentrated to elongate. All these features have been described for R. c. curta by Radoman (1983). In this monograph, Radoman (1983) stated that Orientalina (= Radomaniola) curta subspecies differ from each other mainly in shell rather than in anatomical features, which applies to R. c. maxima. Other than by shell shape and size, this subspecies is diagnosable by its small number of cusps on the radular teeth, a high RPG ratio and a wide base of the penis. Our phylogenetic inferences also place the studied specimens of R. c. maxima unambiguously within the R. curta clade (Fig. 2)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 413-414, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584, {"references":["Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Srpska Akademija Nauka i Umetnosti, Posebna Izdanja (Odeljenje PrirodnoMatematitschkich Nauka) 57: i - ii + 1 - 256."]}

RADOMANIOLA DOLENS SP. NOV. (FIGS 12, 13) Z o o b a n k r e g i s t r a t i o n: z o o b a n k. org:act: 1DCAC494-B7E4-47B4-B0E7-7701B39DF9A9 Etymology: The name dolens means painful in Latin and is derived from Achérōn, the Greek name of the Acheron River, or ‘River of Woe’ in Ancient Greek mythology, where the type locality of this species is located. Type material: Holotype (MNCN 15.05 /200159), five paratypes (MNCN 15.05 /200160) in the MNCN collection and ~ 20 paratypes in the UGSB collection (UGSB 16932). Type locality: Springs of the Acheron River, Glyki, Greece. Material studied: Springs of the Acheron River, Glyki, Greece, 39.3280°N, 20.6235°E, M.B., August 2014, MNCN 15.05 /200160 and UGSB 16932 (80% ethanol). Diagnosis: Protoconch microsculpture wrinkled; central radular tooth formula 6-C-6/1-1; bursa copulatrix globular, duct longer than bursal length; SR1 elongate, duct short; SR2 elongate, smaller than SR1, duct short; penis unpigmented, gradually tapering, base narrow, shorter than head length; nervous system weakly pigmented, elongate (mean RPG ratio = 0.50). Description: Shell ovate-conic, 3.5–4.0 whorls, height 1.75–2.25 mm (Fig. 12A–C; Supporting Information, Table S6). Periostracum yellowish, sometimes whitish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture wrinkled (Fig. 12G). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (Fig. 12B). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (Fig. 12E, F). Radular length intermediate, ~600 µm (~25% of total shell length), with ~60 rows of teeth (Fig. 12H). Central tooth formula 6-C-6/1-1 (Fig. 12I); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 5-C-5. Inner marginal teeth having 28–30 tapered cusps, shortening toward the base. Outer marginal teeth with 25–27 sharp cusps (Fig. 12J). Animal darkly pigmented except for neck and tentacles (Fig. 13F). Ctenidium with 11–12 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (Fig. 13A). Stomach slightly longer than wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (Fig. 13B; Supporting Information, Table S7). Nervous system slightly pigmented, elongate (mean RPG ratio = 0.50); cerebral ganglia approximately equal in size, presenting small black granules (Fig. 13C). Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland. Bursa copulatrix globular, slightly longer than wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 smaller than SR1, elongate, with a short duct, located on renal oviduct near loop (Fig. 13D, E; Supporting Information, Table S8). Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (Fig. 13H). Penis unpigmented, gradually tapering, shorter than head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (Fig. 13F, G; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight. Habitat: Endemic to the large, cold spring of the Acheron River, in which the species lives in moderate abundance. Remarks: Genetic divergence in the small sample (N = 3) of R. dolens was relatively large (~2% for COI), as was the variation in shell shape and shell aperture observed in some individuals (Fig. 12A–D). Differences in shell shape might have been driven by shell erosion. We found no remarkable anatomical differences in this population. This species is distinguished from the geographically proximate and conchologically similar population of R. haesitans from the Louros River according to its wider shell umbilicus, smaller bursa copulatrix and SR1, more slender penis, lower RPG ratio and an average sequence divergence of 6.6% for COI (Supporting Information, Tables S4, S8 and S9)., Published as part of Delicado, Diana & Hauffe, Torsten, 2022, Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years, pp. 393-441 in Zoological Journal of the Linnean Society 196 on pages 415-418, DOI: 10.1093/zoolinnean/zlab121, http://zenodo.org/record/7035584

Details on the dataset, the included predictors, the diversification models, as well as tests for correlation among parameters and the influence of spatial heterogeneity

Genus Docleiana Delicado & Pe��ić gen. nov. urn:lsid:zoobank.org:act: 04639DB3-91D2-46B7-BFAB-FFC1EA3C27BD Type species Arganiella tabanensis Boeters, Gl��er & Pe��ić, 2014. Revised diagnosis Shell valvatiform; whorls ca 3; aperture complete, from rounded to ellipsoidal; outer lip narrow, straight in lateral profile; umbilicus wide. Operculum corneous, yellowish, thin, pliable, rounded, paucispiral with a central nucleus, without peg. Two pairs of basal cusps on each central radular tooth. Ctenidium well developed, with 10���11 gill filaments. Animal unpigmented. Osphradium positioned opposite approximately to the middle of the ctenidium. Stomach without gastric caecum; rectum forms a gentle V-shape in the mantle cavity. Bursa copulatrix small, globular, pedunculated and positioned beyond the posterior edge of the albumen gland; bursal duct longer than bursal length; unpigmented renal oviduct; one pyriform seminal receptacle arising at the insertion point with the bursal duct loop. Penis small and simple, gradually tapering. Etymology The new genus is named after Doclea, the name of the Roman city located on the site of modern Podgorica on whose municipal territory a new genus was found; gender feminine. Remarks Docleiana Delicado & Pe��ić gen. nov. can be distinguished from Arganiella as the former has a more ellipsoidal shell aperture, a narrower base of the penis, a smaller bursa copulatrix that is positioned beyond the posterior edge of the albumen gland, a larger seminal receptacle and more cusps on the lateral radular teeth (Fig. 2; for comparison, see Giusti & Pezzoli 1980; Boeters et al. 2014). The new genus differs from the closely related genus Kerkia by its smaller shell, absence of lobes on the inner edge of the penis, a smaller bursa copulatrix, a more globular seminal receptacle and V-shaped rectum (see Radoman 1978; Bodon et al. 2001). Mean COI sequence divergence for Docleiana Delicado & Pe��ić gen. nov. was 14.5% with Arganiella and 12.3% with Kerkia., Published as part of Delicado, Diana, Pe��i��, Vladimir & Ramos, Marian A., 2021, Arganiella Giusti & Pezzoli, 1980 (Caenogastropoda: Truncatelloidea: Hydrobiidae): a widespread genus or several narrow-range endemic genera?, pp. 140-155 in European Journal of Taxonomy 750 (1) on pages 146-147, DOI: 10.5852/ejt.2021.750.1369, http://zenodo.org/record/5451701, {"references":["Boeters H. D., Gloer P. & Pesic V. 2014. Arganiella tabanensis n. sp. from Montenegro (Mollusca: Gastropoda: Hydrobiidae). Ecologica Montenegrina 1 (3): 131 - 139. https: // doi. org / 10.37828 / em. 2014.1.20","Giusti F. & Pezzoli E. 1980. Hydrobioidea nuove o poco conosciute dell Italia appenninica (Gastropoda: Prosobranchia). Archiv fur Molluskenkunde 111: 207 - 222.","Radoman P. 1978. Neue Vertreter der Gruppe Hydrobioidea von der Balkanhalbinsel. Archiv fur Molluskenkunde 109 (1 - 3): 27 - 44.","Bodon M., Manganelli G. & Giusti F. 2001. A survey of the European valvatiform hydrobiid genera with special reference to Hauffenia Pollonera, 1898 (Gastropoda: Hydrobiidae). Malacologia 43: 103 - 215."]}

Genus Aretiana Delicado & Ramos gen. nov. urn:lsid:zoobank.org:act: 7B207B6D-47E8-4211-8139-0CF98DA353D7 Type species Boetersiella wolfi Boeters & Glöer, 2007. Revised diagnosis Shell trochiform; whorls 3.5–4.0; aperture complete, rounded; outer lip narrow, straight in lateral profile; umbilicus wide. Operculum corneous, yellowish, thin, pliable, oval to rounded, paucispiral with a central nucleus, without peg. Two pairs of basal cusps on each central radular tooth. Ctenidium well developed, with approximately 14 gill filaments. Osphradium positioned opposite to approximately the middle of the ctenidium. Stomach without gastric caecum; rectum forms a gentle U-shape in the mantle cavity. Bursa copulatrix medium-sized, pyriform, pedunculated and protruding beyond the posterior edge of the albumen gland; bursal duct shorter than bursal length; unpigmented renal oviduct that makes a complete loop over the pallial gland; one elongated or pyriform seminal receptacle arising from the renal oviduct, just above the insertion point with the bursal duct. Prostate gland bean-shaped, about twice as long as wide. Penis small and simple, gradually tapering. Nervous system unpigmented. Etymology The genus is named after Villa Aretiana, the Roman name of the town Aracena, which gives its name to the mountain range where the genus was found (i.e., Sierra de Aracena); gender feminine. Remarks Aretiana Delicado & Ramos gen. nov. can be distinguished from Arganiella as the former has a taller shell, a more oval operculum, pigmentation on the body and eyespots, fewer gill filaments, a narrower penis, a larger and pyriform bursa copulatrix located beyond the posterior edge of the albumen gland and fewer cusps on the lateral radular teeth (for comparison, see Giusti & Pezzoli 1980; Boeters & Glöer 2007; Arconada & Ramos 2007a). The new genus differs from the closely related genus Iberhoratia by its taller shell with a narrower umbilicus, the absence of lobes on the inner edge of the penis and of a proximal seminal receptacle (SR2) and presence of two pairs of basal cusps on each central radular tooth (see Arconada et al. 2007). Mean COI divergence for Aretiana Delicado & Ramos gen. nov. was 15.9% with Arganiella and 11.2% with Iberhoratia.

Biserta putealis gen. et sp. nov. urn:lsid:zoobank.org:act: AB67B9A3-8BA6-4AC2-97E3-97C61E6A4C32 Figs 7–8 Diagnosis Shell 1.6 to 2.0 mm high, elongate-conic, brown-yellowish; central radular tooth formula (5)4–C– 4(5)/1–1; bursa copulatrix pyriform, with a short duct; two seminal receptacles, SR1 more developed and pedunculated, SR2 small, globular, located near loop before the pouch; prostate gland bean shaped, ovate in section, folded; penis gradually tapering, unpigmented, with two sessile lobes closely positioned to one another. Distinguished from other closely related species by>8.0% for COI. Etymology The species epithet is from the Latin adjective ‘ putealis ’, meaning ‘from a well’. Type material Holotype TUNISIA • 1 spec. (sex unknown, preserved in ethanol 80%); Bizerte Province, 7 km south of Menzel Bourguiba, Soudene Well; 37.09046º N, 9.777997º E; 115 m a.s.l.; 15 Mar. 2015; N. Khalloufi leg.; MNCN 15.05/200099H. Paratypes TUNISIA • 5 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/200099P • 32 specs (preserved in ethanol 80%, 2 ♀♀, 2 ♂♂ dissected and 2 specs processed for DNA sequencing); same collection data as for holotype; UGSB 17664 • 70 specs (preserved in ethanol 95%, ca 40 individuals dissected); same collection data as for holotype; Khalloufi’s collection. Type locality TUNISIA: Bizerte Province, 7 km south of Menzel Bourguiba, Soudene Well; 37.09046º N, 9.777997º E; 115 m a.s.l. Description MEASUREMENTS. Holotype: SL = 2.00 mm, SW = 1.04 mm, SL/SW = 1.92, AH = 0.98 mm, SL-LBW = 0.70 mm, WBW = 0.97 mm, AL = 0.93 mm, AW = 0.70 mm, WPW = 0.78 mm, WAW = 0.17 mm. SHELL. Elongate-conic, brown to yellowish, with 4–4.5 whorls (Fig. 7 A–D; Table 2). Protoconch (Fig. 7G) obtuse, near planispiral, about 1.25 whorls, diameter about 300 µm; protoconch microsculpture granulated. Teleoconch whorls convex, separated by deep sutures; body whorl occupying ¾ of total shell length. Aperture about 50% of total shell length, pyriform, slightly angled on the top and narrowly disjunct from parietal wall of body whorl; inner lip thin but thicker than outer lip; peristome margin (Fig. 7B) straight. Umbilicus narrow and closed by inner lip. OPERCULUM. As for genus, thin, flat, whitish (nuclear region slightly yellowish), with ca 2 whorls (Fig. 7 E–F); attachment scar area oval and located near nucleus. RADULA. Length intermediate (25% total maximum shell length), having about 80 rows of teeth. Central tooth (Fig. 7 H–I) formula (5)4–C–4(5)/1–1, cutting edge strongly concave; basal tooth face M-shaped and relatively narrow wings; basal tongue broadly V-shaped, about equal to lateral margins; a single small pair of basal cusps, triangular. Lateral teeth (Fig. 7H) formula (4)3–C–5(6); central cusp large, narrow, pointed. Inner marginal teeth having 23–25 sharp cusps. Outer marginal teeth having 22–23 sharp cusps (Fig. 7J). PIGMENTATION AND ANATOMY. Visceral coil brown-yellowish, showing by transparency the gonad; pallial coil with uniform and dense pigmentation on the roof, anteriorly to edge of mantle, foot and area between tentacles (Fig. 8C). Black small eyes surrounded by brownish pigment. Snout with medium distal lobation. Tentacles parallel-side, distal end nonexpanded. Ctenidium filaments about 13, small, taller than wide. Osphradium whitish, ellipsoidal, positioned centrally along ctenidium (Fig. 8A). Stomach slightly longer than wide with a small gastric (posterior) caecum (Fig. 8B); rectum wide, folded and S-shaped. MALE GENITALIA. Penis gradually tapering (Fig. 8D), medium sized, unpigmented, slightly folded; filament short, tapering; basal portion intermediate in width, shorter than distal portion, attached centrally to the head well behind the eyes; penis having two developed lobes, very close to one another on opposite sides of penis; both lobes having distal papilla and showing by transparency tubular glands; ventral lobe large, pyriform, pedunculated; dorsal lobe small, pyriform, sessile. Prostate gland small, close, beanshaped, longer than wide (0.25 mm / 0.05 mm). Posterior vas deferens with several loops, yellowish and developed in its median portion (about 2 mm in length and 0.05 mm in section). Pallial vas deferens straight, narrow, whitish, unciliated, entering the posteroventral portion of gland and exiting from its anterior end. FEMALE GENITALIA. Ovary simple, with stalked lobes, occupying near 40% of visceral coil. Distal female genitalia occupying near 50% of pallial roof; albumen gland approximately as long as capsule gland (Fig. 8E). Bursa copulatrix pyriform, having a shorter duct than bursal length. Renal oviduct unpigmented, coiled, having a spherical pouch at the end of the coiled section. Two seminal receptacles; SR1 medium sized, elongate, pedunculate, joining renal oviduct just above the insertion point with bursal duct; SR2 small, globular, sessile, located on renal oviduct near loop before pouch (Fig. 8 F–G). Ecology and distribution This species was found only in a small well in southern Menzel Bourguiba city, northern Tunisia. The well is 1.5 m in diameter and 4 m deep. Specimens were very abundant on the rock walls and on the rocky and sandy bottom. Water salinity was about 1.5 ppt. Pseudamnicola sp. was the only co-occurring gastropod species. Remarks Some individuals of the collected population lack body pigmentation (Fig. 7C), which could indicate adaptation to subterranean conditions (Delicado 2018). Other anatomical structures resembled those of the pigmented individuals.

Belgrandiellopsis chorfensis gen. et sp. nov. urn:lsid:zoobank.org:act: 2C5704A1-BDFC-41BA-86C6-CF17A999BBC8 Figs 3���4 Diagnosis Shell 1.45 to 1.65 mm high, elongate-conic, brown-yellowish; central radular tooth formula (5)4���C��� 4(5)/1���1; bursa copulatrix ovoid, with the same length as the duct; two seminal receptacles, SR1 more developed and pedunculated, SR2 small, globular, located near loop before the pouch; prostate gland bean shaped, ovate in section, folded; penis cylindrical, slender, distal end tapered, unpigmented, with two lobes distantly positioned; basal lobe more developed. Distinguished from other closely related species by> 8.0% for COI. Etymology A geographic epithet referring to Chorfa Spring, where the species was found. Type material Holotype TUNISIA ��� 1 spec. (sex unknown, preserved in ethanol 80%); B��ja Province, ca 15 km west of Joumine, Chorfa Spring; 36.865918�� N, 9.363313�� E; 430 m a.s.l.; 15 Mar. 2015; N. Khalloufi leg.; MNCN 15.05/200097H Paratypes TUNISIA ��� 5 specs (preserved in ethanol 80%); same collection data as for holotype; MNCN 15.05/200097P ��� 40 specs (preserved in ethanol 80%, 2 ♀♀, 2 ♂♂ dissected and 2 specs processed for DNA sequencing); same collection data as for holotype; UGSB 17663 ��� 150 specs (preserved in ethanol 95%, ca 70 individuals dissected); same collection data as for holotype; Khalloufi���s collection. Type locality TUNISIA: B��ja Province, ca 15 km west of Joumine, Chorfa Spring, 36.865918�� N, 9.363313�� E, 430 m a.s.l. Description MEASUREMENTS. Holotype: SL = 1.61 mm, SW = 0.88 mm, SL/SW = 1.83, AH = 0.66 mm, SL-LBW = 0.49 mm, WBW = 0.84 mm, AL = 0.78 mm, AW = 0.53 mm, WPW = 0.64 mm, WAW = 0.10 mm. SHELL. Elongate-conic, brown to yellowish, with 4���4.5 whorls (Fig. 3 A���D; Table 2). Protoconch (Fig. 3G) obtuse, near planispiral, about 1.2 whorls, diameter about 350 ��m; protoconch microsculpture granulated. Teleoconch whorls weakly convex, separated by deep sutures; body whorl occupying �� of total shell length. Aperture about 40% of total shell length, pyriform, strongly angled on the top and narrowly disjunct from parietal wall of body whorl; inner lip thin but thicker than outer lip; peristome margin (Fig. 3B) straight. Umbilicus narrow and closed by inner lip. OPERCULUM. As for genus, thin, flat, whitish (nuclear region slightly yellowish), with ca 2 whorls (Fig. 3 E���F); attachment scar area, oval and located near nucleus. RADULA. Length intermediate (20% total maximum shell length), having about 80 rows of teeth. Central tooth (Fig. 3 H���I) formula (5)4���C���4(5)/1���1, cutting edge concave; basal tooth face M-shaped and relatively narrow wings; basal tongue broadly V-shaped, about equal to lateral margins; a single small pair of basal cusps, triangular. Lateral teeth (Fig. 3J) formula (4)3���C���5; central cusp large, narrow, pointed. Inner marginal teeth (Fig. 3H) having 25���27 sharp cusps. Outer marginal teeth having 20���22 sharp cusps. PIGMENTATION AND ANATOMY. Visceral coil brown-yellowish, showing by transparency the gonad; pallial coil nearly pale with traces of pigment on the roof, anteriorly to edge of mantle; foot and area between tentacles slightly pigmented (Fig. 4C). Black small eyes surrounded by brownish pigment. Snout with medium distal lobation. Tentacles parallel-side, distal end nonexpanded. Ctenidium filaments about 12, small, taller than wide. Osphradium whitish, ellipsoidal, positioned centrally along ctenidium (Fig. 4A). Stomach slightly longer than wide with a small gastric (posterior) caecum (Fig. 4B); rectum wide, folded and S-shaped. MALE GENITALIA. Penis cylindrical (Fig. 4 D���E), medium sized, unpigmented, slender, slightly folded; filament short, tapering; basal portion intermediate in width, shorter than distal portion, attached centrally to the head well behind the eyes; penis having two developed lobes located relatively far from one another on opposite sides of penis; both lobes with distal papilla, showing by transparency tubular glands distal; basal lobe larger, thumb-like, not pedunculated; distal lobe smaller, pyriform, pedunculated. Prostate gland small, close, bean-shaped, longer than wide (0.2 mm / 0.10 mm). Posterior vas deferens developed in its median portion with several loops (about 4 mm in length and 0.01 mm in section); seminal vesicle yellowish. Pallial vas deferens straight, narrow, whitish, unciliated, entering the posteroventral portion of gland and exiting from its anterior end. FEMALE GENITALIA. Ovary simple, with stalked lobes, occupying near 40% of visceral coil. Distal female genitalia occupying near 70% of pallial roof; albumen gland approximately as long as capsule gland (Fig. 4F). Bursa copulatrix ovoid to pyriform, having a duct as long as bursal length. Renal oviduct unpigmented, coiled, having a spherical pouch at the end of the coiled section. Two seminal receptacles; SR1 medium sized, elongate, pedunculate, joining renal oviduct just above the insertion point with bursal duct; SR2 small, globular, sessile, located on renal oviduct near loop before pouch (Fig. 4 G���H). Ecology and distribution Belgrandiellopsis chorfensis gen. et sp. nov. was found in a small spring near a rural settlement called Chorfa between the cities of Mateur and Beja, in northern Tunisia. It is known only from the type locality and some springs in this region (e.g., Ain Elgaid Spring). The specimens were found on the bottom and attached to the underside (algae-side) of gravel and stones. Density decreased drastically downstream from the water resurgence. Pseudamnicola sp. was the only co-occurring gastropod species. Remarks Differentiated from its congener according to its smaller shell size (Table 2), smaller number of lateral cusps on the central radular teeth, larger penis and slightly smaller bursa copulatrix. Belgrandiellopsis chorfensis gen. et sp. nov. also differs in its mean COI divergence by 8.4% from B. secunda gen. et sp. nov., Published as part of Khalloufi, Noureddine, B��jaoui, Mustapha & Delicado, Diana, 2020, Two new genera and three new subterranean species of Hydrobiidae (Caenogastropoda: Truncatelloidea) from Tunisia, pp. 1-27 in European Journal of Taxonomy 648 (648) on pages 10-14, DOI: 10.5852/ejt.2020.648, http://zenodo.org/record/3836080

Genus Belgrandiellopsis gen. nov. urn:lsid:zoobank.org:act: 6073 FD 1 F- 91C0-4C93-B1 CB- 09 AA 54B1 FABD Type species Belgrandiellopsis chorfensis sp. nov., by present designation. Diagnosis Shell small (maximum length slightly exceeding 2 mm), ovate-conic to elongate-conic, with obtuse and planispiral protoconch; whorls slightly convex, with weak sutures. Penultimate whorl tall relative to remaining whorls. Aperture pyriform to ovate, complete, angled on top and often fused to the body whorl. Umbilicus closed by inner lip. Operculum corneous, whitish to yellowish, thin, pliable, elongateellipsoidal, spiral, paucispiral, with submarginal nucleus. One pair of small triangular basal cusps on radular central tooth. Ctenidium well developed. Stomach with a small gastric (posterior) caecum. Presence of bursa copulatrix; unpigmented renal oviduct with a spherical pouch at the end of the coiled section; two seminal receptacles. Penis attached to the neck behind the right eye with two glandular lobes distantly positioned, each of them bearing a terminal papilla. Etymology Refers to the shell similarities with the hydrobiid genus Belgrandiella; gender feminine. Remarks Belgrandiellopsis gen. nov. differs from the conchologically similar genus Belgrandiella in having a strap-like penis with two glandular lobes, two seminal receptacles and one pair of basal cusps on the central radular teeth (for comparison see Szarowska 2006); from the closely related genus Bullaregia according to the presence of two lobes on the inner edge of the penis, smaller gastric caecum, shorter bursal duct, larger bursa copulatrix and in having generally a smaller number of cusps on radular teeth (see Khalloufi et al. 2017); and from Biserta gen. nov. according to its smaller shell umbilicus, longer penis with a larger distance between penial lobes, slightly smaller bursa copulatrix and longer bursal duct (Figs 4, 6, 8). Belgrandiellopsis gen. nov. differs in its mean COI divergence by 9.3% and 8.4% from the closely related Bullaregia and Biserta gen. nov., respectively, and by 17.6% from Belgrandiella., Published as part of Khalloufi, Noureddine, B��jaoui, Mustapha & Delicado, Diana, 2020, Two new genera and three new subterranean species of Hydrobiidae (Caenogastropoda: Truncatelloidea) from Tunisia, pp. 1-27 in European Journal of Taxonomy 648 (648) on pages 8-9, DOI: 10.5852/ejt.2020.648, http://zenodo.org/record/3836080, {"references":["Szarowska M. 2006. Molecular phylogeny, systematics and morphological character evolution in the Balkan Rissooidea (Caenogastropoda). Folia Malacologica 14 (3): 99 - 168. https: // doi. org / 10.12657 / folmal. 014.014","Khalloufi N., Bejaoui M. & Delicado D. 2017. A new genus and species of uncertain phylogenetic position within the family Hydrobiidae (Caenogastropoda: Truncatelloidea) discovered in Tunisian springs. European Journal of Taxonomy 328: 1 - 15. https: // doi. org / 10.5852 / ejt. 2017.328"]}

Belgrandiellopsis secunda gen. et sp. nov. urn:lsid:zoobank.org:act: D3A45301-6EBE-4086-A740-FEFBE9328FB5 Figs 5–6 Diagnosis Shell 1.7 to 2.1 mm high, elongate-conic, brown-yellowish; central radular tooth formula (6)5–C– 5(6)/1–1; bursa copulatrix ovoid with short duct; two seminal receptacles, SR1 more developed and pedunculated, SR2 very small, globular, located near loop before the pouch; prostate gland bean shaped, ovate in section, folded; penis cylindrical, slender, distal end tapered, unpigmented, with two lobes relatively distant positioned; basal lobe more developed and middle positioned. Distinguished from other closely related species by>8.4% for COI. Etymology Referring to the second species of the genus Belgrandiellopsis gen. nov. discovered so far. Type material Holotype TUNISIA • 1 spec. (sex unknown, preserved in ethanol 80%); Bizerte Province, 40 km south of Mateur, Ettoute Spring; 36.883898º N, 9.505894º E; 415 m a.s.l.; 15 Mar. 2015; N. Khalloufi leg.; MNCN 15.05/200098H. Paratypes TUNISIA • 5specs (preserved in ethanol80%);same collection data as for holotype; MNCN15.05/200098P • 26 specs (preserved in ethanol 80%, 2 ♀♀, 2 ♂♂ dissected and 2 specs processed for DNA sequencing); same collection data as for holotype; UGSB 17666 • 250 specs (preserved in ethanol 95%, ca 70 individuals dissected); same collection data as for holotype; Khalloufi’s collection. Type locality TUNISIA: Bizerte Province, 40 km south of Mateur, Ettoute Spring; 36.883898º N, 9.505894º E; 415 m a.s.l. Description MEASUREMENTS. Holotype: SL = 2.09 mm, SW = 1.16 mm, SL/SW = 1.80, AH = 0.90 mm, SL-LBW = 0.59 mm, WBW = 1.09 mm, AL = 0.99 mm, AW = 0.70 mm, WPW = 0.78 mm, WAW = 0.09 mm. SHELL. Elongate-conic, brown to yellowish, with 4–4.5 whorls (Fig. 5 A–D; Table 2). Protoconch (Fig. 5G) obtuse, near planispiral, about 1.3 whorls, diameter about 350 µm; protoconch microsculpture granulated. Teleoconch whorls weakly convex, separated by deep sutures; body whorl occupying ¾ of total shell length. Aperture about 50% of total shell length, pyriform, strongly angled on the top and narrowly disjunct from parietal wall of body whorl; inner lip thin but thicker than outer lip; peristome margin (Fig. 5B) straight. Umbilicus narrow and closed by inner lip. OPERCULUM. As for genus, thin, flat, whitish (nuclear region slightly yellowish), with ca 2 whorls (Fig. 5 E–F); attachment scar area oval and located near nucleus. RADULA. Length intermediate (25% total maximum shell length), having about 80 rows of teeth. Central tooth (Fig. 5 H–I) formula (6)5–C–5(6)/1–1, cutting edge slightly concave; basal tooth face M-shaped and relatively narrow wings; basal tongue broadly V-shaped, about equal to lateral margins; a single small pair of basal cusps, triangular. Lateral teeth (Fig. 5H) formula 5–C–3(4); central cusp large, narrow, pointed. Inner marginal teeth having 32–35 sharp cusps. Outer marginal teeth having 30–33 sharp cusps (Fig. 5J). PIGMENTATION AND ANATOMY. Visceral coil pale-yellowish, showing by transparency the gonad. Pallial coil with uniform and dense brown pigmentation on roof (left side), anterior edge of mantle, foot and between tentacles (Fig. 6C). Black small eyes surrounded by brownish pigment. Snout with medium distal lobation. Tentacles parallel-side, distal end nonexpanded. Ctenidium filaments about 12, small, taller than wide. Osphradium whitish, ellipsoidal, positioned centrally along ctenidium (Fig. 6A). Stomach slightly longer than wide with a small gastric (posterior) caecum (Fig. 6B); rectum wide, folded and S-shaped. MALE GENITALIA. Penis cylindrical (Fig. 6D), medium sized, unpigmented, slender, slightly folded; filament short, tapering; basal portion intermediate in width, longer than distal portion, attached centrally to the head well behind the eyes; penis having two developed lobes, both with distal papilla, showing by transparency tubular glands; lobes located relatively distant from one another on opposite sides of penis; basal lobe large, conic, sessile, positioned centrally along penis; distal lobe smaller, pyriform, pedunculated. Prostate gland small, close, bean-shaped, longer than wide (0.2 mm / 0.10 mm). Posterior vas deferens whitish, forming the seminal vesicle and developed in its median portion with several loops (about 3.5 mm in length and 0.05 mm in section). Pallial vas deferens straight, narrow, whitish, unciliated, entering the posteroventral portion of gland and exiting from its anterior end. FEMALE GENITALIA. Ovary simple, with stalked lobes, occupying near 40% of visceral coil. Distal female genitalia occupying near 50% of pallial roof; albumen gland less developed than capsule gland (Fig. 6E). Bursa copulatrix ovoid to pyriform, having a duct slightly shorter than bursal length. Renal oviduct unpigmented, coiled, having a very small, ovoid to spherical pouch at the end of the coiled section. Two seminal receptacles; SR1 medium sized, elongate, pedunculate, joining renal oviduct just above the insertion point with bursal duct; SR2 small, globular, sessile, located on renal oviduct near loop before pouch (Fig. 6 F–G). Ecology and distribution This species was found in a spring and its outflow in the rural village Ettoute between the cities of Mateur and Beja, in northern Tunisia. Specimens were attached to the underside of gravel and stones covered by algae. In the small stream flowing out of the spring, the snails were found in the central section featuring clean and oxygenated waters and a silty substrate. The density of this species decreased drastically downstream from the water resurgence. Co-occurring gastropod species were Pseudamnicola sp., Galba truncatula (O.F. Müller, 1774), Bulinus truncatus (Audouin, 1827) and Physella acuta Draparnaud, 1805. Remarks Despite similarities in mean COI divergence between this species and the closely related species Bullaregia tunisiensis Khalloufi, Béjaoui & Delicado, 2017, Biserta putealis gen. et sp. nov. and Belgrandiellopsis chorfensis gen. et sp. nov. (9.1%, 8.9% and 8.4%, respectively), B. secunda gen. et sp. nov. was assigned to Belgrandiellopsis gen. nov. based on the combination of two glandular lobes located relatively distant from one another on opposite sides of the penis [one lobe in Bullaregia (Khalloufi et al. 2017), two lobes closely positioned to one another in Biserta gen. nov.], small bursa copulatrix with a long duct (larger bursa copulatrix with a shorter duct in Biserta gen. nov.), small gastric caecum (larger in Bullaregia) and small shell umbilicus (larger in Biserta gen. nov.).

The western Palearctic freshwater snail genus Mercuria (Caenogastropoda: Hydrobiidae) comprises 26 species primarily distributed in lowland localities of Western Europe and North Africa. Although this genus in North Africa has received considerable attention in terms of species discoveries through morphological descriptions, its distribution and phylogenetic patterns remain unknown. Based on morphological and mitochondrial DNA (mtCOI) evidence, this study examines the three Mercuria species (M. bakeri, M. tingitana, and M. targouasensis) from Morocco identified so far. Besides expanding on information regarding the anatomy of these species, two new species (M. midarensis sp. n. and M. tensiftensis sp. n.) are described for this region and phylogenetic relationships inferred between these species and the European M. emiliana and M. similis. All Moroccan and European species were recovered as independent entities according to these phylogenetic inferences (uncorrected p-distances 2.8â 8.5%) and DNA barcode data. Moroccan Mercuria species clustered with M. emiliana from Spain, although basal relationships within this clade were not well supported. Given that factors such as the season when specimens are collected, habitat type, and parasites could be responsible for the remarkable intraspecific variation observed in shell and penis morphology, it is proposed that the most efficient approach to delimit and identify Mercuria species is to combine morphological descriptions with genetic data.

Table S1. Species names, locality data, locality codes and GenBank accession numbers for the taxa studied. Table S3. Abbreviations of the morphological characters depicted in Additional file 3: Table S2. Figure S2. Comparison of macroevolutionary models. Boxplots showing the relative fit (ΔAIC) of quantitative state speciation and extinction (QuaSSE) models estimating the evolution of temperature preference and its potential influence on speciation rate along 100 random post-burn-in trees of Pseudamnicolinae for (a) mean annual temperature, (b) mean temperature of the warmest season, and (c) mean temperature of the coldest season. Each set of temperature included a standard deviation (SD) of 10, 20, 30%. Dots display the relative fit for the MCC tree. Tables S4, S5. Models testing the relationship of speciation rates with both environmental temperature and elevation, respectively. We fitted quantitative state speciation and extinction (QuaSSE) models with genus specific or independent coefficients for rates of speciation (λ), change of speciation rate with elevation (λElv) and environmental temperature (λTem), and rates of elevational and environmental temperature evolution (σ2). Subscripts P and C denote the two genera Pseudamnicola and Corrosella, respectively. Empty cells indicate identical coefficients of the respective parameter for both genera. Not available (NA) parameters were not included in the respective model. For each model, the first row indicates the coefficients and model fit of the MCC tree. Brackets in the second row include the average and standard deviation of these coefficients as well as the model fit based on 100 random post-burn-in trees. Table S6. Testing mode and rates of morphological divergence of the two sister-genera Pseudamnicola and Corrosella along the MCC. The Brownian Motion (BM) model includes the rate of morphological divergence (σ2) and the morphological optimum (θ). We used AIC-based model comparison to test additional variants of this model that allowed for genus and/or trait specific coefficients. Empty cells indicate identical coefficients for both trait axes or both genera, respectively. For each tested model, the first row indicates the coefficients and model fit of the MCC tree and the brackets in the second row include the average and standard deviation of these coefficients as well as the model fit based on 100 random post-burn-in trees. Because of the relatively small number of species, we could not fit more complex models with genus- and trait specific parameters or selection. (PDF 625 kb)

Figure S1. Phylogenetic relationships of Corrosella and Pseudamnicola species based on a Bayesian inference of the combined COI, 16S, and 28S datasets. Bayesian posterior probabilities are indicated with black dots when