MALACOLOGIA, 2006, 48(1-2): 77-132 REVISION OF THE GENUS ISLAMIA RADOMAN, 1973 (GASTROPODA, CAENOGASTROPODA, HYDROBIIDAE), ON THE IBERIAN PENINSULAAND DESCRIPTION OF TWO NEW GENERAAND THREE NEW SPECIES BeatrizArconada & Maria-Angeles Ramos Museo Nacionalde Ciencias Natural3es (1CSIC) José GutiérrezAbascal 2, 28006 - Madrid. Spain: [email protected] ABSTRACT The presence of the genus Islamia Radoman, 1973, on the Iberian Peninsula is con- firmed based on the detailed study ofa group ofspecies, ofwhich three were previously included in the genus Neohoratia Schutt, 1961. These species are most abundant in the south-southeastern Mediterranean region but also inhabitthe northern Mediterranean ar- eas of the peninsula, with scattered populations in central and western Spain. Iberian Islamia currently includes /. globulus (Bofill, 1909), /. lagan (Altimira, 1960), and /. ateni (Boeters, 1969), plus two new species, /. pallida and /. henrici, the latterwith two subspe- cies /. h. henriciand /. h. giennensis. Two new genera are also described, Milesiana and Josefus, each of which contains one species; M. schuelei (Boeters, 1981), which was previouslyassignedtoNeohoratia, andmostrecentlytoIslamia, andanewspecies,Josefus aitanica, respectively. Histological studyofthefemale genitalia confirmed the presenceof two seminal receptacles and the absence ofa bursa copulatrix in all species belonging to the three genera. In Islamia, the distal receptacle was once considered to be a reduced bursa copulatrix. We also confirm that there is no trace of glandular tissue on the penial lobe in any ofthe Islamia species forwhich histological evidence is available. Key words: Caenogastropoda, Hydrobiidae, Neohoratia, Islamia, Milesiana, Josefus, taxonomy, Spain, Iberian Peninsula. INTRODUCTION endemicity. Boeters (1988) recognized that species of two genera, Horatia Bourguignat, The European fauna of hydrobiids is par- 1887, and /\/eo/7orai/'aSchutt, 1961, inhabited ticularly rich in valvatiform species. However, thisgeographicalarea.An in-depthtaxonomic their morphological study is challenging be- review of the two genera is currently un- cause oftheirminute size. Many newgenera ravelling a very complex situation. Four new and species have been described on the ba- genera and several new species have been sis of shell features, which are known to be described in recent papers (Ramos et al., highlyconvergent. Sometimesotheranatomi- 2000;Arconada& Ramos, 2001 2002). Some , cal characters, which are frequently non-di- ofthe species in the new genera were previ- agnostic, are used inthesedescriptions. Data ously included in the above-mentioned gen- on character variability are absent or very era. Wecontinuethesestudiesbyrevisingthe rare. The result has been a much confused taxonomy of another group of species previ- taxonomic picturethatwas recently reviewed ously assigned to Neohoratia by Boeters and partially clarified by Bodon et al. (2001), (1988). who redescribed the type species of most of It has been difficult to distinguish the spe- the European valvatiform genera based on cies of the genera Neohoratia Schutt 1961, new anatomical studies and data in the lit- and Islamia Radoman, 1973, given their mor- erature. phological similarities (Bodon & Giovanelli, Preliminary studies on Iberian Peninsular 1994; Bodon et al., 1995; Manganelli et al., valvatiform hydrobiids (Ramos et al., 1992, 1998). The type species of Neohoratia is 1995;Arconadaetal., 1996)haveshowncon- Valvata (?) subpiscinalis Kuscer 1932 (Figs. siderable morphological diversity and high 1-5, paratypes from the Biological Institute, 77 78 ARCONADA & RAMOS Scientific Research Centre of Ljubljana. N Valvata coronado! Bourguignat, 1870; 1862, leg. Dr. J. Bole). This genus has under- Hauffenia {Neohoratia) coronadol schuelei gone several changes in itstaxonomicstatus. Boeters, 1981; \/a/uafa(rrop/d/>?a)fez/Altimira, has been regarded as a subgenus of 1960; Hauffenia {Neohoratia) gasulli Boeters It Hauffenia Pollonera, 1898, and of Horatia 1981; and Neohoratia azarum Boeters & Bourguignat, 1887 (Schutt, 1961; Boeters. Rolan, 1988). However, according to Boeters 1974: Bodon & Giovanelli. 1994), and as a (1988), these Iberian species, apartfrom hav- distinctgenus(Bole&Velkovrh, 1986; Boeters, ing a narrowing ('Einschnürung') of the outer 1988; Bole. 1993). /\/eo/?oraf/aischaracterised side of the female oviduct glands (capsule + by having a rather short, flat, blunt or slightly albumen glands), lacked a bursa copulatrix pointed peniswith 1-3 small, knob-like lateral and had a renal oviduct with two seminal re- lobes on its left side near the apex. The fe- ceptacles. This combination of characters, in male genitalia include a pin-like bursa copu- addition to a male genitalia with a penis usu- latrix and one proximal, small seminal ally having one glandular lobe on its left side, receptacle (Bole, 1993; Bodon et al., 2001). has been described as typical of the genus Boeters (1988) and Boeters & Rolan (1988) Islamla (Bodon et al., 1995; Bodon et al., overlooked these diagnostic characters while 2001). Islamla is attributed to a wide geo- including several species from the Iberian graphical distribution in the Mediterranean Peninsula in this genus {Amnícola globulus area [species are claimed to be from; Turkey Bofill, 1909; Microna ateni Boeters, 1969; (Schutt, 1964; Radoman, 1973b); the Balkanic FIGS. 1-5. Shell of Valvata subpisclnalis (IBCICL paratype n° 1862). REVISION OF THE GENUS ISLAMIA 79 Peninsula (Radoman, 1973a, b, 1978, 1983); MATERIALAND METHODS Italy (Giusti & Pezzoli, 1981; Bodon et al., 1995, 1996, 2001; Bodon &Cianfanelli, 2002); Field collections, anatomical studies, histo- Israel (Schutt, 1991; Bodon et al., 1995); logical protocols, and morphometricmeasure- Greece (Radoman, 1973b, 1978); and France ments are described in Ramos et al. (2000) (cited as Hauffenia Pollonera, 1898) (Ber- and Arconada & Ramos (2001). The number nasconi, 1984)]. of specimens studied for histology and mor- It was thus feasible that the species listed phometry, localities and sampling dates for above from the Iberian Peninsula could be each speciesare indicated inthecorrespond- attributed to the genus Islamia (type species ing section in the text. The morphological de- Hydrobia valvataeformisMöllendorff, 1873)or scriptions are based on terminology from even to new genera. In fact, two of them, Hershler& Ponder(1998). Scanning Electron Hauffenia {Neohoratia) gasulli [N. (?) gasulli, Microscope (SEM) photographs were made sensu Boeters, 1988] and Valvata {Tropidina) with a Philips XL20 following the methodol- fezi [N. (?) fezi, sensu Boeters, 1988] were ogy described in Ramos et al. (2000). Type recently allocated to two new genera, Tarra- materialofIslamiaglobuluswasphotographed conia Ramos &Arconada, 2000 (in Ramos et with a Environmental Scanning Electron Mi- a!.,2000),and Spai/?ogynaArconada&Ramos, croscope (ESEM) Philips Quanta 200 SEM at 2002, respectively. low vacuum mode, after being cleaned with Here we describe three new species and ultrasound (Figs. 18, 20, 23, 25, 27, 30. 31, redescribe the morphological characters (in- 33. 34) or the periostracum removed by im- cluding previouslyunknowncharacters)ofthe mersion in 5% sodium hypochlorite (Figs. 19, above-mentioned species using a multi- 28). disciplinary approach based on type speci- Paratypes of Islamia cianensis Bodon, mens and a vastamountofrecently collected Manganelli, Sparacio& Giusti, 1995{n°6732), material. Additionally, histological studies of and /. gaiteri Bodon, Manganelli, Sparacio & these species provide evidence that the two Giusti, 1995 (n° 6733), from the Museo sac-like structures on the renal oviduct are Zoológico "La Specola" collection were used seminal receptaclesanddemonstratethenon- for comparisons. glandular nature ofthe penial lobe. Localities are listed according to the code: We conclude that two of the "Neohoratia" stream orspring, municipality, province, UTM species {sensu Boeters, 1988) from the Ibe- co-ordinates, samplingdate, collector'sinitials, rian Peninsula {Amnícola globulus and museum catalogue number and preservation Microna ateni) actually belong to the genus conditions (see abbreviations below). Local- Islamia, as hypothesized by Bodon et al. ity names and UTM co-ordinates were ob- (2001). Two other species, one of them with tained from the official Army Geographical two subspecies, are described as new and Service map (1;50.000 series). placed intoIslamia.Anotherspecies, Hauffenia {Neohoratia) coronadoischuelei, reported as StatisticalAnalyses N. schuelei{\r\ Boeters, 1988) and as Islamia schuelei(in Bodonetal., 2001), is redeschbed All statistics (mean value, standard devia- and placed intoa newgenus Milesiana, and a tion and coefficient of variation) were calcu- third new species is described and placed in lated using STATVIEW for Macintosh, and a newgenus, Josefus. Neohoratiaazarumhas standardized in ordertoavoid theeffectofthe not been included here because still unpub- measurement scale. lished data (Arconada, 2000) clearly demon- A discriminant función analysis (DFA) was strate that its anatomy is differs considerably performed on nine shell measurements (no from the genera and species described here. ratios) with STATISTICA v.6 for Windows in This paperincreasesthe numberofspecies orderto identifythe morphological characters and expands the distribution area of Islamia thatbestdifferentiatedspecieswhen noorfew (Schutt, 1961; Radoman, 1973a, b; Giusti et anatomical data were available. There were al., 1981; Bernasconi, 1984; Bodon et al., no missing data. The effects of violating as- 1995) in Europe and reinforces the hypoth- sumptions are minimized taking into account esis that the Iberian Peninsula is one of the the robustnessofthe Ftest (Lindman, 1974). richest hydrobioid {sensu Davis, 1979) diver- The significance ofthe overall discriminatory sity areas in the Mediterranean Basin (Arco- powerofthe analysis was tested using Wilk's nada & Ramos, 2003). Lambda. Canonical correlation was used to 80 ARCONADA & RAMOS measure interspecificvariation. Classification ture width: LBW. length of body whorl; NL: functions were computed for each group length of opercular nucleus: NW: width of (population) to determine, with the highest opercular nucleus; NSW: number of spire probability, which case belonged to which whorls: OL: operculum length: OW: opercu- population. Caseswereassigned tothegroup lum width: OLWL: length ofthe lastwhorl of with the highest classification score. theoperculum: OLWW:widthofthelastwhorl ofthe operculum: SL: shell length; SW: shell Abbreviations Used in the Text, Tables and width; WAW: width of the antepenultimate Figures whorl: WBW: width ofthe bodywhorl; WPW; widthofthepenultimatewhorl; CV: coefficient Shell and Operculum Characters: AH: aper- ofvariation: SD: standard deviation. ture height; AL: aperture length: AW: aper- AnatomicalCharacters:Ag:albumengland; Be: FIGS. 6-10. Histological sections ofthe anterior female genitalia of Milesians schueleishowing the positionofthespermatozoids insidethe seminal receptaculum. Notethe headsofthespermatozoids attached to the ciliated epithelial cells of the seminal receptacles. FIGS. 6, 7: Proximal seminal receptaculum; FIGS. 8, 9: Distal seminal receptaculum; FIG. 10: Inner epithelium of the widened renal oviduct. Abbreviations: c: cilia; sp: spermatozoids. REVISION OF THE GENUS ISLAMIA 81 bursa copulatrix; Cg: capsule gland; DBG: theepithelialcellsliningthelumenofthesemi- ductofthebursacopulatrix; Os: osphradium; nal receptacle (Figs. 6-9). This is the typical P: penis; PI: penial lobe; Po: palliai oviduct; methodforspermstorageina molluscan semi- Pp: pseudopenis; Pr: prostate; Ro: renal ovi- nal receptacle (Thompson & Bebbington, duct; SR1: distal seminal receptacle; SR2: 1969; Giusti &Selmi, 1985; Fretter&Graham, proximal seminal receptacle; Ss: style sac; 1994: 303-306) and is morphologically re- St: stomach; Vc: ventral channel of capsule sponsible for the whitish-pearly réfringence gland; L: length; W: width.Theconcentration characteristic of this structure. On the other ofthenervoussystemwasdetermined bythe hand, the bursa copulatrix(gametolyticgland) "RPG" ratio (Davis et al., 1976): length of doesnotcontainspermatozoaorcontainsfew, pleuro-supraesophageal connective divided non-oriented spermatozoa (itscontent is cen- bythesumofthelengthsofrightpleuralgan- trally located and never réfringent) (see also glion, pleuro-supraesophageal connective Ramosetal., 2000; Bodónetal., 2001).There- and supraesophageal ganglion. Following fore, morphological réfringence can be used several studies, a synthesis of RPG ratios to distinguish bursa copulatrix from seminal from diverse hydrobioid taxa indicates: dor- receptacles or even to infer the possible role sal nerve ring concentrated (<0.29); moder- of sperm storage deposit in widened parts of ately concentrated (0.30-0.49); elongated the renal oviduct (Davis et al., 1992; Ramos (0.50-0.67); extremely elongated (> 0.68) et al., 2000, and papers cited therein) when (Davis et al., 1984, 1986, 1992). histological evidenceis notavailable.Thewid- Collections: MNCN: Museo Nacional deCien- ened portion ofthe renal oviduct has a thick, cias Naturales, Madrid, Spain; MZB: Museu moredeveloped innerepithelium in relationto de Zoología, Barcelona, Spain; NNM: Na- the portion between proximal and distal semi- tionaal Natuurhistorisch Museum, Leiden, nal receptacles, giving rise to a stretched lu- Naturalis,TheNetherlands; MHNG: Muséum menwherethespermatozoids move (Fig. 10). d'Histoire Naturelle, Genève, Switzerland; Histologicaldifferencesalongtherenaloviduct SMF: Forschungsinstitutund Natur-Museum epithelium are similar to those described for Senckenberg, Frankfurt, Germany; MZUF: Tarraconia gasulli (Ramos et al., 2000) and Museo Zoológico "La Specola", Universita suggest that the widened part of the oviduct di Firenze, Italy; IBCICL: SlovenianAcademy mayactasan additional sperm storage. How- of Sciences and Arts, Ljubljana, Slovenia; ever, we are not able to confirm this hypoth- NHMW: Naturhistorisches Museum, Wien, esis, because we have not had evidence of Austria. oriented spermatozoa in any of the species Collectors: R.A.: R.Araujo; B.A.: B.Arconada; studied. J.A.: J.Astigarraga;A. .:A. Bertrand; D. .: Careful analysis of serial sections of males D. Buckley; A. C.: A. Camacho; J. E.: J. belongingto /. globulus, I. pallida, M. schuelei, Escobar; S. J.: S. Jiménez; N. M.: N. Martín; and J. aitanica reveals that the penis and pe- . . D. M.: D. Moreno; N.: Noreña; J. P: J. I. nial lobe are made upofa thick layerofexter- Pino; J. M. R.: J.:M. Remón; J. R.: J. Roca; E. nal muscles beneath the outer epithelium R.: E. Rolan; G. G. Tapia. (Figs. 11-15). The inner structure consists of numerous vascular spaces of reticulated connectivetissue, denseralong the periphery GENITAL HISTOLOGY of the penis, with muscle fibres running be- tween them. There was no indication of any Histological studies of 4 pm serial sections glandular tissue either on the penial lobe or were conducted with special focus on female on any other part ofthe penis. This structure and male genital systems. For each species, is similar to that described for other molluscs thenumberand sexofspecimensinvestigated (Fretter & Graham, 1994: 302). The undulat- are indicated in the corresponding texts. ing penial ductcan also beobservedthrough- Considering the female genitalia of Islamia outthedifferentsectionsofpenis until itenters globulus, I. henricihenrici, Milesianaschuelei, the nuchal area. Females of several species and Josefusaitanica, histological evidenceof havea nuchal node ora pseudopenis located "oriented sperm"in thetwosac-likestructures onthe rightsideofthehead, ina position simi- on the renal oviduct was obtained. The sper- lar to that of the male penis. These females matozoa are arranged with their heads an- have fullyfunctional genitalia with mature oo- chored to the cell surface among the cilia of cytes in the ovary (Fig. 16). . 82 ARCONADA & RAMOS y T* 11/ 12 f .-. . ^ sd /- f . /"sd '-> i'< 13 14 -- -i*jfc /- *»» *^ r. ifi i.\ REVISION OF THE GENUS ISLAMIA 83 lendorffaccordingto Radoman, 1983, and ac- Neohoratia globulus globulus (Bofill, 1909) - cepted by Bodon at al., 2001. Boeters, 1988: 214,figs. 137-144, 151-155, 163-170, pi. 2, fig. 22; Bech, 1990: 61. Diagnosis Islamiaglobulusglobulus(Bofill, 1909)-Bodon et al., 2001: 179, figs. 195-200; Bodon & Shell smallorverysmall,ovoidorplanispiral, Cianfanelli, 2002: 20. rarely ovate-conic; operculum without peg; central tooth with one or two basal cusps on Type Locality each side; penis with a well-developed non- glandularlobeon its leftside; femalegenitalia Fontdel Sotdel Pinell, close to Forteiletdel with two seminal receptacles, proximal (SR2) Montsech, Lérida, U.T.M.: GC16. larger and longer than distal (SRI); seminal receptacles located on opposite sides (or po- Material Examined sitions) on unpigmented renal oviduct; they can arise eitherclose to or ratherdistantfrom Type material: A lot containing 41 syntypes . eachother; proximal seminal receptacle(SR2) (dried) of globulus collected byArtur Bofill usually with evident duct and distal (SR1), at type locality were deposited in the MZB usuallywithoutaductevident; bursacopulathx (Bofill, 1917), cataloguenumber: 80-1589.The absent. specimen illustrated in Figs. 18, 23, 25, 27, 30, 33, is here designated lectotype (ICZN, Islamia glob,ulus (Bofill, 1909) 1999: Art. 74.7). The remaining syntypes are therefore paralectotypes. Lectotype (MZB80- Amnícolaglobulus 1909:205; 1915:57, 1589a)and 29 paralectotypesfromthis lotare 58, pi. 6, fig. 6; 1917: 35. in the MZB collections and 9 in the MNCN Amnícolaanatínaglobulus(Bofill, 1909)-Bofill collections with n° MNCN 15.05/46546. The & Haas, 1920: 50, 57, pi. Ill, figs. 19, 20. second lotwith around 1,000 syntypes (dried) Amnícola similis (Draparnaud) - Haas, 1929: is in the MZB collections (MZB 80-1628). 408, 409, fig. 163. Other populations examined: This species Pseudamnicola similis globulus (Bofill, 1909) iswidelydistributed in the provincesofLérida -Altimira, 1960: 10; 1963: 16. and Huesca (Fig. 17). Boeters (1988) also /. i>lohiilus 2- Í. Idi^ari 3- /. (itt'iii 4- /.pcillidd 5- /. Iienricilienrici 6-/. henrici^ieimensis 7- M. schuelei H- M. cf. schuelei 9-J. (litanica 200 km FIG. 17. Mapoflocalitiesofthegenera Islamia, Milesiana andJosefusinthe Iberian Peninsula. 84 ARCONADA & RAMOS cited itfrom Gerona, althoughwe cannotcon- May 1986, J. R., MNCN 15.05/46594; Laguar- firnn these data so far. One lot from Font La ta. Huesca, UTM; 30TYM374998, 12 April Figuereta (Lérida) population kept in the MZB 1995; B. A., MNCN 15.05/46542 (ethanol and (80-1629) was also examined and compared SEM preparation); 26 Oct. 1995, B.A. &E. R., with that from the same locality kept in the MNCN 15.05/46543 (ethanol, SEM prepara- MNCN (15.05/46540). Five specimens (etha- tionandfrozen material); Grimaspring, Gistain, nol) from Laguarta population were donated Huesca. UTM; 31TBH799184, 13 April 1995, to the MZB (n 2002-0537). B. A., MNCN 15.05/46544 (ethanol); Sopeira spring, Huesca, UTM; 31TCG1487, 24 July Localities (1e9t9h1an,oRl.Aa.n,dD.SME.,MJ.prMe.paRr.,atMioNnC).N 15.05/46545 Spring in Amargosa, Aristot, Lérida, UTM: 31TCG871948. 14 March 1999, B. A., MNCN Material Examined for Morphometry and 15.05/46527 (ethanol and frozen material); Histology Blancaspring,VilanovadeMeya, Lérida, UTM; 31TCG371551, 25 Feb. 1986, J. R., MNCN Shellandanatomical measurements(Tables 15.05/46528 (ethanol, SEM preparation and 1, 3-7)correspond to populationsfrom Lérida frozen material); La Argentería spring, Baix and Huesca; Operculum and radularmeasure- Pallars, Lérida, UTM; 31TCG381842, 2 Oct. ments (Tables 2, 4)to Huesca (see table cap- 1986, J. R., MNCN 15.05/46529 (ethanol and tions). Male and females studied and SEM preparation); El Regué spring, Vilanova measured were collected in the following deMeya. Lérida, UTM; 31TCG304539, 27 Feb. months; Feb., March,April, May, July, and Oct. 1986, J. R., MNCN 15.05/46530 (ethanol); La For histology, four females and three males Fayeda spring, Abella de la Conca, Lérida, werestudiedfroma spring in Sopeira, Huesca UTM; 31TCG475668, 10 Oct. 1986, J. R., (July 1991), and one female from Laguarta, MNCN 15.05/46531 (ethanol); Fontanetspring, Huesca (Oct. 1995). Abelladela Conca, Lérida, UTM; 31TCG4269, 14 March, 1999, B.A.. MNCN 15.05/46593 Diagnosis (ethanol and frozen material); Les Greixes spring, SantEstevede La Sarga, Lérida, UTM; Shell ovate-conic, body whorl narrow; oper- 31TCG126635, 8 May 1986. J. R., MNCN culumovate;centraltoothofradulawithasingle 15.05/46532 (ethanol); Blanca spring, Gäbet basal cuspon each side; ctenidiumwell devel- de la Conca, Lérida. UTM; 31TCG301658, 13 oped; shortpleuro-subesophagealconnective; May 1986, J. R., MNCN 15.05/46533 (etha- esophagus running straight underneath cere- nol); D'Arcallo spring, Baix Pallars, Lérida, bralcommissure; bean-shaped prostategland; UTM; 31TCG482818, 29 Sept. 1986, J. R., big penis, usually black pigmented, with one MNCN 15.05/46534 (ethanol); La Sarga large, unpigmented non-glandular lobe, com- spring. Gäbet de La Conca, Lérida, UTM; monlyprotrudingfromthetipofpenis; pyriform 31TCG375567, 26 Feb. 1986, J. R., MNCN and pedunculated proximal seminal receptacle 15.05/46535(ethanol); Fredaspring,Abellade (SR2)andsmall,elongated, sessiledistalsemi- laConca, Lérida, UTM; 31TCG473677, 10May nal receptacle (SRI); receptacles emerge dis- 1986, J. R., MNCN 15.05/46536 (ethanol), 14 tinctly separated from each other. March 1999. B.A., MNCN 15.05/46616 (etha- nolandfrozen material); Freda springdeCasa Description Pallas,Aren, Lérida; UTM; 31TCG065908, 28 (Figs. 18-29, 30-35, 42-49; Tables 1-7; March 1987, J. R.. MNCN 15.05/46537 (etha- Bodon et al., 2001; figs. 195-200) nol); Bordons spring, Aren, Huesca, UTM; 31TCG085881, 31 March 1987, J. R., MNCN Shell: Shell ovate-conic, 4.1 whorls; sutures 15.05/46538 (ethanol); Adraén, Cadi moun- deep, aperture oval, slightly prosocline; tains, Lérida, UTM.; 31TCG767817, 15 Feb. peristome complete, slightly thickened at 1998, A. ., MNCN 15.05/46539 (ethanol and columelar margin, slightly reflected at lower SEM preparation); 15 March 1999, B. A., and columelar margin; body whorl very nar- MNCN 15.05/46541 (ethanol and frozen ma- row, over V; of the total shell length; proto- tehal); La Figuereta spring, AIós de Balaguer, conch consisting of 1.5 whorls; protoconch Lérida, UTM; 31TCG253439, 11 March 1986, width and width of the nucleus are 380 pm J. R., MNCN 15.05/46540(ethanol); LesBulles and 140 pm, respectively (Figs. 30-35); spring, Isona, Lérida, UTM; 31TCG371667, 8 protoconch pitted; umbilicus narrow, 130 pm REVISION OF THE GENUS ISLAMIA 85 FIGS. 18-29. Shells of Islamia globulus. FIGS. 18, 23, 25, 27: Lectotype (MZB 80-1589a); FIG. 19: Paralectotype (MZB80-1589b); FIG. 20: Paralectotype (MNCN 15.05/46546); FIG. 28: Paralectotype (MZB 80-1589c); FIGS. 21, 24,26, 29: Shellsfrom Laguarta; FIG. 22: Shell from Sopeira population. Scale bar = 1 mm (FIGS. 18-26); 500 pm (FIGS. 27-29). 86 ARCONADA & RAMOS Éé