Pseudamnicola (Corrosella) manueli, Delicado & Machordom & Ramos, 2011

Delicado, Diana, Machordom, Annie & Ramos, Marian A., 2011, Underestimated diversity of hydrobiid snails. The case of Pseudamnicola (Corrosella) (Mollusca: Caenogastropoda: Hydrobiidae), Journal of Natural History (J. Nat. Hist.) 46 (1 - 2), pp. 25-89 : 44-60

Pseudamnicola (Corrosella) manueli sp. nov

Type locality

La Garganta stream in Nava de San Pedro, Jaén, Spain, UTM: 30S 0509318/4194330.

Type material

Holotype MNCN 15.05 /49176a ( SEM preparation, Figure 9A,D ) and paratypes ( Figures 9E–G , 10 , 11 ) MNCN 15.05 /49176b ( SEM preparation, Figures 9E–G , 10 ,

and 70% ethanol, Figure 11 ) and MNCN / ADN 34899–34904 (frozen material), D.D., 12 October 2007 ; MNCN 15.05 /49175 (70% ethanol), 1 May 1990 , D.M.

Other populations studied

In addition to type specimens, we examined the following material collected in the province of Jaén: La Garganta spring in Nava de San Pedro, Jaén, UTM: 30S 0509318/4194330, D.D., 12 October 2007, MNCN 15.05 /49177 (70% ethanol); El Valle stream, La Iruela, Jaén ( Figure 9B ), UTM: 30S 0504010/4196581, D.D., 14 October 2007, MNCN 15.05 /49183 ( ESEM preparations and 70% ethanol) and MNCN / ADN 34908–34915 (frozen material); La Ponderosa spring in Hinojares, Jaén, UTM: 30S 0501944/4179988, D.M., 30 April 1990, MNCN 15.05 /49178 (70% ethanol); El Céfano spring in La Iruela, Jaén, UTM: 30S 0502253/4197687, B.A., 23 March 1998, MNCN 15.05 /49179 (70% ethanol); D.D., 14 October 2007, MNCN 15.05 /49180 (70% ethanol and ESEM preparations, Figure 9C ) and MNCN / ADN 34905–34907 (frozen material); Molino ditch in La Iruela, Jaén, UTM: 30S 0500538/4197129, D.M., 30 April 1990, MNCN 15.05 /49181 (70% ethanol); ditch in Finca Rechita, La Iruela, Jaén, UTM: 30 S 05028 /41981, B.A., 23 March 1998, MNCN 15.05 /49182 (70% ethanol); Sierra de Cazorla Hotel , La Iruela , Jaén , UTM: 30 S 05005 /41969, D.M., 30 April 1990, MNCN 15.05 /49184 (70% ethanol); spring in Prados de la Presa , Jaén, B.A., 24 March 1998, MNCN 15.05 /49185 (70% ethanol); La Mata spring in Mata Bejid , Jaén , UTM: 30 S 04553 /41721, B.A., 24 March 1998, MNCN 15.05 /49186 (70% ethanol); stream in La Toba , Jaén , UTM: 30S 0539412/4226322, B.A., 24 March 1998, MNCN 15.05 /49187 (70% ethanol); Arroyo del Museo de la Caza , Cazorla, Jaén, UTM: 30S 0500538/4197129, D.M., 1 May 1990, MNCN 15.05 /49188 (70% ethanol) .

Specimens examined for morphometry

Shell, anatomical, operculum and radular measurements (Appendix Tables 1–8) were made in male and female specimens collected at La Garganta stream in Nava de San Pedro (type locality), Jaén in March , April, May and October .

Etymology

Dedicated to Manuel Delicado, father of the first author of this paper, for his support and help with fieldwork.

Diagnosis

Shell with a bulging inflated body whorl relatively wider than the rest of whorls; oesophagus and intestine without pigmentation; anterior edge of style sac protrudes under the intestine; female genitalia with an elongated pyriform J-shaped bursa copulatrix, seminal receptacle slightly pigmented, above insertion of bursal duct; renal oviduct black pigmented until insertion point of seminal receptaculum; penis gradually tapering and pointed at its end, with a narrow distal pigment patch; nervous system black pigmented with supraoesophageal connective over four times longer than suboesophageal.

Description

Shell ovate-conic, yellowish periostracum with 4.25–4.75 spire whorls, height 3.7– 3 mm ( Figure 9A–C ; Appendix Table 1); body whorl well-developed, about threequarters of shell length, and wider than the rest of whorls; deep suture and convex spire whorls; protoconch with approximately 1.8 whorls; protoconch width and width of nucleus around 420 µm and 180 µm, respectively ( Figure 9E,F ); protoconch microsculpture granulated ( Figure 9G ); oval aperture complete with thin outer lip and thicker inner lip; narrow umbilicus; edge of peristome straight ( Figure 9D ).

Operculum with around 2.5 spire whorls and an oval muscle attachment area near the nucleus ( Figure 10A, B , Appendix Table 2).

Radula medium size (18%) relative to maximum shell dimension ( Figure 10C , Appendix Table 4); with approximately 50 rows of teeth; central tooth with a large median cusp, sometimes slightly divided, and four or five lateral very small cusps of irregular shape decreasing in size ( Figure 10D,E ) giving the edge a serrated appearance; lateral teeth with three sharp lateral cusps; inner marginal teeth with approximately 18 tapered cusps and outer marginal teeth with around 22 tapered cusps smaller than inner marginal cusps ( Figure 10D,F ).

Pigmentation and anatomy. Head dark brown pigmented; internal side of the tentacle has a longitudinal streak without pigmentation ( Figure 11D ); foot intermediate in size and with dark brown pigment on its dorsal side. Ctenidium well-developed with 17–19 gill filaments taller than wide occupying two-thirds of pallial cavity; osphradium in opposite middle region of ctenidium ( Figure 11C , Appendix Table 3). Stomach almost as long as wide with both chambers equal in size (Appendix Table 5); style sac with protruding intestinal loop; oesophagus and intestine without pigmentation ( Figure 11F ); rectum S-shaped in pallial cavity containing orange faecal pellets.

Female genitalia with two glands in pallial oviduct: albumen gland, the most posterior, and capsule gland with two portions (the most proximal is whiter); albumen gland occupies more than one-third of pallial oviduct ( Figure 11G , Appendix Table 6); bursa copulatrix elongated J-shaped ( Figure 11H ); black pigmented renal oviduct fading from loop to insertion of seminal receptacle; renal oviduct lies over bursa copulatrix making two or three loops; elongated seminal receptacle without duct, situated on renal oviduct above the insertion of bursal duct.

Male genitalia with a bean-shaped prostate gland ( Figure 11E , Appendix Table 7) the seminal duct entering the posterior region and a pallial vas deferens emerging close to its anterior edge; penis simple, gradually tapered, with a distal patch of pigmentation and six or seven folds in its middle region ( Figure 11D ); the penis is attached to the central region of head, behind the eyes; penial duct straight or slightly undulating on the right/external side of the penis, running from base to penis tip.

Nervous system black pigmented, darker on ganglia than connectives and commissures; cerebral ganglia approximately same size; supraoesophageal connective is four times longer than suboesophageal connective ( Figure 11A,B , Appendix Table 8); RPG ratio is 0.44 (moderately concentrated); oesophagus running straight beneath cerebral commissure.

Remarks

Populations from Nava de San Pedro (type locality), la Mata and Padros de la Presa have shells with a wider body whorl and a more tapered penis with a small patch of pigmentation, whereas the shells of specimens from La Iruela and Cazorla village are more slender and the penis has a rounded tip and larger pigment patch. Furthermore, the pigmented area on the seminal receptacle is larger in females of the type locality than the rest. Molecular analyses (still in progress) also indicate certain genetic distance between the two groups of populations (3.9%), but this distance is insufficient to consider them two different species .

Pseudamnicola (C.) manueli sp. nov. differs from the rest of the species of Corrosella examined in having: an inflated body whorl of shells ( Figure 9A–C ), granulated protoconch microsculpture ( Figure 9G ), presence of four or five lateral very small cusps of the central radula tooth giving a serrated appearance ( Figure 10D,E , Appendix Table 4), long ctenidium (the longest being a mean of 1.43 mm, Appendix Table 3) and a tapered penis.

Pseudamnicola (C.) bareai sp. nov. is the genetically closest species to P. (C.) manueli sp. nov. (5.46% divergence) and both share characters such as: shell dimensions ( Figure 25 ), 18 cusps in inner marginal teeth, length of penis (1 mm approximately in both species) and prostate gland (1.6 mm average length). However, there are some characters that serve to differentiate both species: (1) central radular tooth with a wide, almost pentagonal, central cusp in P. (C.) manueli sp. nov., being narrower and more elongated in P. (C.) bareai sp. nov.; (2) three lateral cusps in lateral radular teeth in P. (C.) manueli sp. nov. and four in P. (C.) bareai sp. nov.; (3) different appearance and size of capsule gland ( Figure 11G ; Appendix Table 6), being longer in P. (C.) manueli sp. nov. and showing two regions of different opacity, whereas in P. (C.) bareai sp. nov. its appearance is more uniform ( Figure 14G ); (4) elongated bursa copulatrix folded into a J-shape in P. (C.) manueli sp. nov. without clear transition from bursal duct, which widens gradually ( Figure 11H ), and folded into a U-shape in P. (C.) bareai sp. nov. ( Figure 14H ) with clear transition from the bursal duct; (5) sharp penis with a long patch of pigmentation in distal region in P. (C.) manueli sp. nov. ( Figure 11D ), and round penis tip and smaller almost round patch of pigment in middle region of penis in P. (C.) bareai sp. nov. ( Figure 14D ).

Pseudamnicola (Corrosella) bareai sp. nov

Type locality

Spring at Ermita de las Santas, Collados de la Sagra, Granada, Spain, UTM: 30S 0542294/4201835.

Type material

Holotype MNCN 15.05 /49190a ( ESEM preparation, Figure 12A ), paratypes MNCN 15.05 /49190b ( ESEM preparation, Figures 12B,D–G , 13 , and 70% ethanol, Figure 14 ) and MNCN / ADN 34916–34930 (frozen material), D.D., 13 October 2007; JM.B., 6 June 2006, MNCN 15.05 /49189 (70% ethanol); JM.B., 21 May 2008, MNCN 15.05 /49191 (70% ethanol).

Other populations examined

All populations of the new species were found in the Castril mountains ( Granada province) with the exception on one population found at Siete Fuentes at the edge of the Cazorla mountains in Jaén province . Specimens were collected from: Fuente Nuevas, Castril, Granada, UTM: 30S 0512019/4181258, JM.B., 21 May 2008, MNCN 15.05 /49192 (96% ethanol) and MNCN /ADN: 34931–34932 (96% ethanol); El Laude, Castril, Granada, UTM: 30S 0517139/4187270, JM.B., 21 May 2008, MNCN 15.05 /49193 (96% ethanol) and MNCN /ADN: 34933 (96% ethanol); JM.B., 30 October 2008, MNCN 15.05 /49194 (70% ethanol); Agüerillo, Castril, Granada ( Figure 12C ), UTM: 30S 0517351/4186925, JM.B., 21 May 2008, MNCN 15.05 /49195 (96% ethanol) and MNCN /ADN: 34934 (96% ethanol); JM.B., 30 October 2008, MNCN 15.05 /49196 (70% ethanol); Siete Fuentes, Cuenca, Jaén, UTM: 30S 0502639/4176601, D.D., 12 October 2007, MNCN 15.05 /49197 (70% ethanol) and MNCN /ADN: 34935–34944 (frozen material) .

Etymology

Dedicated to José Miguel Barea, who besides being the discoverer of the type locality, kindly collaborated with us in sampling and protecting freshwater molluscs and their habitat in Andalusia.

Diagnosis

Shell with penultimate whorl tall in relation to previous ones; lateral radular tooth formula 4-C-4; intestine slightly pigmented; bursa copulatrix cylindrical U-shaped; renal oviduct darkly pigmented until insertion and seminal receptacle non-pigmented; penis gradually tapering with a clear narrow patch of pigment in the middle region; nervous system brown pigmented with supraoesophageal connective over four times longer than suboesophageal.

Description

Shell ovate-conic, yellowish periostracum, with 4–4.25 spire whorls, height 3.3– 2.5 mm ( Figure 12A–C , Appendix Table 1); body whorl occupying three-quarters of total shell length and penultimate whorl rather tall in relation to previous ones; whorls convex; protoconch net-shaped grooved ( Figure 12G ) with about 1.8 whorls; total width and nucleus width around 380 and 190 µm, respectively ( Figure 12E,F ); longitudinal ribs run parallel to protoconch suture; aperture complete oval; narrow outer lip in contact with last whorl practically hiding the umbilicus; inner lip wider than outer; in lateral view aperture straight and slightly backwards ( Figure 12D ).

Operculum with around three spire whorls ( Figure 13A , Appendix Table 2); muscle attachment area oval close to nucleus ( Figure 13B ).

Radula medium size (27%) relative to maximum shell dimension and nine times longer than wide ( Figure 13C , Appendix Table 4); around 61 rows of teeth; central tooth with a long tapered median cusp and four to six very small laterals, fused in some specimens ( Figure 13D,E ); basal tongue V-shaped; lateral teeth with four relatively tapered lateral cusps; inner marginal tooth contains approximately 18 cusps of decreasing size; outer marginal tooth with around 20 cusps smaller than inner marginal cusps ( Figure 13D,F ).

Pigmentation and anatomy. Head with uniform blackish pigment from snout to base of penis; pigmentation clearer on neck; ocular region without pigmentation; tentacles with a longitudinal band of pigmentation on the external side ( Figure 14D ); snout as long as wide with medium distal lobation; foot intermediate with dorsal pigmentation. Ctenidium with 16–18 well-developed gill filaments situated in the middle of the pallial cavity; osphradium less than 50% ctenidium length and located in the opposite middle of the ctenidium ( Figure 14C , Appendix Table 3). Stomach slightly wider than long with two similar-size cameras and medium-sized gastric caecum (Appendix Table 5); style sac approximately as long as stomach; intestine weakly pigmented ( Figure 14F ); rectum filled with orange faecal-pellets and S-shaped in pallial cavity.

Female genitalia with an albumen gland that occupies one-third pallial oviduct ( Figure 14G , Appendix Table 6); bursa copulatrix cylindrical U-shaped ( Figure 14H ) with a long duct; renal oviduct black pigmented until insertion of seminal receptacle, it makes a simple loop over the bursa copulatrix after two or three very dark loops; seminal receptacle elongated without pigment lies on renal oviduct slightly above the joining point of the bursal duct.

Male genitalia with a large bean-shaped prostate gland almost four times longer than wide and occupying a large section of pallial cavity ( Figure 14E , Appendix Table 7); seminal duct entering the medial-posterior region of prostate gland and a pallial vas deferens emerging close to its anterior edge; penis tubular with a narrow base, six or seven small folds in the middle region and clear blackish pigmentation, varying from a small patch in the middle region to a narrow long patch covering all the middle surface ( Figure 14D ); penis can appear coiled in live specimens; straight penial duct running on right side of the penis from base to the tip.

Nervous system with darker ganglia than connectives and commissures; cerebral ganglia approximately same size; right pleural ganglion smaller than left pleural ganglion; supraoesophageal connective four times longer than suboesophageal ( Figure 14B , Appendix Table 8); RPG ratio 0.42 (moderately concentrated); oesophagus runs underneath nervous system without loops or folds ( Figure 14A ).

Remarks

Pseudamnicola (C.) bareai sp. nov. can be distinguished from the other Pseudamnicola (Corrosella) species examined here by: lateral radular teeth with four lateral cusps on each side of the elongated central tooth, strong U-shaped bursa copulatrix and tubular penis with a narrow base and small pigment patch. Pseudamnicola (C.) manueli sp. nov. is morphologically and genetically the closest species (genetic divergence 5.46%, Appendix Table 9) but both species can be differentiated because: (1) P. (C.) bareai sp. nov. has a tubular penis with a narrow base, round tip and a small patch of pigment in middle-distal region, whereas the penis in P. (C.) manueli sp. nov. is sharp with a wider base and has a longer patch of pigment in its distal portion; (2) the bursa copulatrix is folded into a U-shape in P. (C.) bareai sp. nov. and is J-shaped in P. (C.) manueli sp. nov.; (3) lateral radular tooth formula is 4-C- 4 in P. (C.) bareai sp. nov. and 3-C- 3 in P. (C.) manueli sp. nov.; (4) protoconch microsculpture is granular in P. (C.) manueli sp. nov. and grooved in P. (C.) bareai sp. nov.

Pseudamnicola (Corrosella) marisolae sp. nov

Type locality

Pilar del Mono spring in Dúrcal, Granada, Spain, UTM: 30S 0449218/4095030.

Type material

Holotype MNCN 15.05 /49417a ( ESEM preparation, Figure 15A ), paratypes ( Figures 15D–G , 16 , 17 ) MNCN 15.05 /49417b ( ESEM preparation, Figures 15D– G , 16 , 70% ethanol, Figure 17 ), D.M and J.T., 17 October 1989; D.M. and E.R., 25 September 1989, MNCN 15.05 /49416 (70% ethanol); D.M., 15 October 1990, MNCN 15.05 /49418 (70% ethanol); D.M., 8 February 1992, MNCN 15.05 /49419 (70% ethanol); B.A., 27 March 1998, MNCN 15.05 /49420 (70% ethanol); JM.B., 22 September 2008, MNCN 15.05 /49421 (96% ethanol) and MNCN / AND 34945–34946 (96% ethanol).

Other populations examined

Specimens of this species were found in the centre and southern area of Granada as follows: Palmones spring, Padul, Granada ( Figure 15C ), UTM: 30S 0445770/4097570, JM.B., 28 February 2007, MNCN 15.05 /49422 (96% ethanol) and MNCN / ADN 34947–34951 (96% ethanol); Padul, Granada, UTM: 30SUG469/973, JM.B., 20 September 2006, MNCN 15.05 /49423 (96% ethanol) and MNCN / ADN 34952–34953 (96% ethanol); Fuente Grande, Alfácar, Granada ( Figure 15B ), UTM: 30S 0450991/4122546, JM.B., 18 October 2008, MNCN 15.05 /49424 (96% ethanol) and MNCN / ADN 34954–34955 (96% ethanol) .

Etymology

Dedicated to María Soledad Iglesias (Marisol), mother of the first author, for her help in collecting the material and her constant support.

Diagnosis

Shell slender of marked conic shape; central radular tooth with four lateral cusps; intestine and oesophagus without pigmentation; renal oviduct brown or black pigmented; bursal duct long and narrow except at the point joining renal oviduct where it has an expansion; penis long with a large patch of black pigment from middle region to tip and folds on the base and middle section; nervous system with a supraoesophageal connective around four times longer than suboesophageal.

Description

Shell yellowish periostracum with 4.5–5.5 spire whorls, height 4.60– 3.25 mm ( Figure 15A–C , Appendix Table 1); convex whorls with a very marked suture; body whorl occupies two-thirds of shell length; protoconch with around 1.6 spire whorls 500 µm wide and nucleus width approximately 150 µm ( Figure 15E,F ); protoconch microsculpture with grooves across entire surface ( Figure 15G ); peristome frontal, oval, complete, with a thin outer lip and thicker inner lip not in contact with body whorl; narrow umbilicus hidden behind inner lip of peristome; outer peristome simple and straight ( Figure 15D ).

Operculum translucent, with 3.5 spire whorls approximately ( Figure 16B , Appendix Table 2); internal side has a convex edge and oval muscle attachment near nucleus ( Figure 16A ).

Radula with around 50 rows of teeth of medium size (22% of total shell length); eight times longer than wide ( Figure 16C , Appendix Table 4); trapezoidal central tooth with a tongue-shaped median cusp and three or four pointed laterals ( Figure 16D,E ); lateral teeth longer than wide with two tapered lateral cusps and a rounded median cusp larger than laterals; inner marginal tooth with 11 cusps, approximately longer than those of outer marginal tooth ( Figure 16D,F ).

Pigmentation and anatomy. Head with dark brown pigment all over its surface except on the edge of snout and external edge tentacles ( Figure 17D ); dorsal region of foot also pigmented; pigment on neck clearer than on the head; snout as long as wide and tentacles shorter than snout; foot of intermediate size and anterior edge indented. Ctenidium with 20–24 well-developed gill filaments situated in the posterior section occupying most of the pallial cavity; osphradium located in opposite middle of ctenidium and two or three times longer than wide ( Figure 17C , Appendix Table 3). Stomach with a posterior chamber larger than anterior chamber, relatively long caecum in a ventral position of the posterior chamber (Appendix Table 5); style sac shorter than stomach, longer than wide; oesophagus and intestine without pigment ( Figure 17F ); rectum lightly S-shaped in pallial cavity.

Female genitalia with a capsule gland with two regions (the anterior region more whitish) and a smaller albumen gland ( Figure 17G , Appendix Table 6); bursa copulatrix pyriform J-shaped, relatively narrow after joining duct and later widely expanded; long and narrow bursal duct expanded near the point joining renal oviduct ( Figure 17H ); elongated seminal receptacle with short duct, slightly pigmented, attached close to base of renal oviduct; renal oviduct pigmented making one or two folds before the loop which is simple and long, later it continues straight from the loop until insertion of the bursal duct; pigmentation of renal oviduct fades strongly from loop to seminal receptacle ( Figure 17H ).

Male genitalia with a prostate gland four times longer than wide (Appendix Table 7); vas efferens entering the medial-posterior region and vas deferens exiting at the anterior ( Figure 17E ); long penis with a wide base, large pigment patch from middle section to tip and some folds in middle zone ( Figure 17D ); attached to central region of head, contains a wavy penial duct running on its right side.

Nervous system brown pigmented, darker on ganglia than on connectives and commissures; cerebral ganglia equal in size to pleural ganglia; supraoesophageal connective four times longer than suboesophageal ( Figure 17B , Appendix Table 8); RPG ratio 0.46 (moderately concentrated); straight oesophagus running beneath nervous system ( Figure 17A ).

Remarks

Two characters differentiate P. (C.) marisolae sp. nov. from the other Iberian Pseudamnicola (Corrosella) species: the shape of the bursa copulatrix and the expansion of the bursal duct terminus close to the pallial oviduct.

Morphological differentiation between this species and P. (C.) luisi , whose distribution areas are very close, is complex because both species have large conic shells and a pigmented long penis. However, shells of P. (C.) luisi are larger than P. (C.) marisolae sp. nov. and its peristome finishes in a thicker inner lip than in P. (C.) marisolae sp. nov. Furthermore, the penis of P. (C.) luisi is slender and around 0.30 mm longer than in P. (C.) marisolae sp. nov.; its base is also wider. The genetic distance between these two species is 6.0% (Appendix Table 9).

Despite P. (C.) marisolae sp. nov. and P. (C.) falkneri having conic shells, the two species differ according to a set of anatomical characters: the shell is more than 1.5 times taller in P. (C.) marisolae sp. nov.; radular formulae of teeth are very different in both species (Appendix Table 4); the bursa copulatrix is at least two times longer in P. (C.) falkneri ; the penis has a pointed tip, small patch of pale pigment and attachment area occupies a central position on the neck in P. (C.) falkneri , but in P. (C.) marisolae sp. nov., the penis has a rounded tip, a long patch of dark pigment and wider base located on the left side of the neck. The genetic distance between P. (C.) marisolae sp. nov. and P. (C.) falkneri is 9.19 % (Appendix Table 9).

With respect to other new species, their differentiation is clearer in terms of anatomy than shell shape. Pseudamnicola (C.) marisolae sp. nov. is the only new species whose lateral teeth formula is 2-C-2 (Appendix Table 4). The penis of P. (C.) marisolae sp. nov. has a wide base and a large dark patch of pigment in the distal region, whereas this pigmented area is smaller in P. (C.) iruritai sp. nov., P. (C.) manueli sp. nov. and P. (C.) bareai sp. nov. and clearer in P. (C.) andalusica sp. nov. and P. (C.) falkneri . The whole seminal receptacle is pigmented in P. (C.) marisolae sp. nov., whereas in P. (C.) manueli sp. nov., P. (C.) andalusica sp. nov. and P. (C.) iruritai sp. nov., pigmentation is restricted to the duct. The folded bursa copulatrix is J-shaped in P. (C.) marisolae sp. nov., P. (C.) manueli sp. nov., P. (C.) andalusica sp. nov. and P. (C.) falkneri , U-shaped in P. (C.) bareai sp. nov. and P. (C.) luisi and pyriform in P. (C.) iruritai sp. nov.

Figures

Figure 9. Shells of Pseudamnicola (C.) manueli sp. nov. (A, D–G) Shells from La Garganta stream in Nava de San Pedro, Jaén (type locality); (B) shell from Arroyo del Valle, La Iruela, Jaén (MNCN 15.05/49183); (C) shell from El Céfano spring in La Iruela, Jaén (MNCN 15.05/49180). (A, D) Holotype (MNCN 15.05/49176a); (E–G) paratypes (MNCN 15.05/49176b), protoconch and microsculpture from the protoconch.

Figure 10. Operculum and radula of Pseudamnicola (C.) manueli sp. nov. from La Garganta stream in Nava de San Pedro, Jaén (type locality, MNCN 15.05/49176b). (A) Internal side of the operculum; (B) external side of the operculum; (C) radula; (D) rows of teeth of the radula; (E) detail of central tooth; (F) inner and outer marginal teeth.

Figure 11. Anatomy of Pseudamnicola (C.) manueli sp. nov. from type locality (MNCN 15.05/49176b). (A,B) Partial nervous system; (C) ctenidium and osphradium; (D) head of a male and penis; (E) prostate gland; (F) stomach; (G) female genitalia; (H) bursa copulatrix and seminal receptacle.

Figure 12. Shells of Pseudamnicola (C.) bareai sp. nov. (A, B, D–G) Shells from Spring in Ermita de las Santas, Collados de la Sagra, Granada (type locality); (C) shell from Agüerillo, Castril, Granada (MNCN 15.05/49195). (A, D) Holotype (MNCN 15.05/49190a); (E–G) paratypes (MNCN 15.05/49190b), protoconch and detail of its microsculpture.

Figure 13. Operculum and radula of Pseudamnicola (C.) bareai sp. nov. from Spring at Ermita de las Santas, Collados de la Sagra, Granada (type locality, MNCN 15.05/49190b). (A) Internal side of the operculum; (B) external side of the operculum; (C) radula; (D) rows of teeth of the radula; (E) central tooth; (F) outer marginal teeth.

Figure 14. Anatomy of Pseudamnicola (C.) bareai sp. nov. from type locality (MNCN 15.05/49190b). (A, B) Partial nervous system; (C) ctenidium and osphradium; (D) head of a male and penis; (E) prostate gland; (F) stomach; (G) female genitalia; (H) bursa copulatrix and seminal receptacle.

Figure 15. Shells of Pseudamnicola (C.) marisolae sp. nov. (A, D–G) Shells from Pilar del Mono spring, Dúrcal, Granada (type locality); (B) shell from Fuente Grande, Alfácar, Granada (MNCN 15.05/49424); (C) Palmones spring, Padul, Granada (MNCN 15.05/49422). (A) Holotype (MNCN 15.05/49417a); (E–G) paratypes (MNCN 15.05/49417b) protoconch and detail of the microsculpture of the protoconch.

Figure 16. Operculum and radula of Pseudamnicola (C.) marisolae sp. nov. from Pilar del Mono spring in Dúrcal, Granada (type locality, MNCN 15.05/49417b). (A) Internal side of the operculum; (B) external side of the operculum; (C) radula; (D) rows of teeth of the radula; (E) central teeth; (F) detail of an outer marginal tooth.

Figure 17. Anatomy of Pseudamnicola (C.) marisolae sp. nov. from type locality (MNCN 15.05/49417b). (A, B) Partial nervous system; (C) ctenidium and osphradium; (D) head of a male and penis; (E) prostate gland; (F) stomach; (G) female genitalia; (H) bursa copulatrix and seminal receptacle.

Figure 25. Plot of discriminant function analysis scores on the two canonical axes recorded for shell measurements made in seven Pseudamnicola Corrosella species.