A revision of the genus Alzoniella Giusti & Bodon, 1984 (Gastropoda, Caenogastropoda, Hydrobiidae) on the Iberian Peninsula and its implications for the systematics of the European hydrobiid fauna

2007·7.3 MB·English

by  Arconada, B

BASTERIA, 71: 113-156,2007
ArevisionofthegenusAlzoniellaGiusti &Bodon,1984(Gastropoda,
Caenogastropoda, Hydrobiidae) on theIberianPeninsulaandits implications
forthesystematics of theEuropean hydrobiid fauna
B.Arconada
MuseoNational deCienriasNaturales (CSIC),JoseGutierrez Abascal,2,E-28006Madrid,Spain
E. Rolán
MuseodeHistoriaNatural,CampusUniversitario Sur,E-15782SantiagodeCompostela,Spain
& H.D. Boeters
Karneidstr. 8,D-81545Miinchen,Germany
Thegenus Alzoniella Giusti&Bodon,1984,isreviewed forthe IberianPeninsula. Thisrevision
includes both thenominate subgenusAlzoniella (A.) and A.(Navarriella)Boeters, 2001,aswell
asthe genus Guadiella Boeters,2003,sinceits typespecies,G. andalucensis (Boeters, 1983),was
formerly regardedasbelongingtoBelgrandiellaA. J.Wagner,1928,togetherwith A.cantabrica
(Boeters 1983)and A.edmundi(Boeters, 1984),whicharenowclassified inAlzoniella. Most of
the newlyreportedlocalities enlargeknown distribution areas.Variousnewdata onanatomi-
cal aspects arepresented.Photographsofshells and theprotoconchs ofnearly all the species
discussed areincluded. Twonewspeciesaredescribed and three insufficientlyknown taxa of
Alzoniellaarementioned, but withoutassigningany specific name.
Key words:Gastropoda,Caenogastropoda,Hydrobiidae,Alzoniella,Navarriella,Guadiella,dis-
tribution,anatomy,specieskey, IberianPeninsula.
INTRODUCTION
Although many IberianPeninsular hydrobiids have been described in recent years
(Rolan, 1993;Arconada,2000; Ramosetalv2000; Arconada& Ramos,2001, 2006; Boeters,
2003),alarge numberof species stillneed tobe studiedin greaterdetail.The mostgeneral
review todatewaspublished byBoeters(1988).ThefirstspeciesofthegenusAlzoniellaGiusti
& Bodon, 1984,onthe IberianPeninsula,A.(Navarriella)elliptica, was describedin the 19th
century as Paludinellaelliptica (Paladilhe, 1874).Ittookover acentury foranother speciesof
thegenustobe discovered,viz.Alzoniella(A.) cantabrica(Boeters, 1983).Sincethen,andin a
shorttime,thenumberhasconsiderably increased, 10morespecies beingknownnowforthe
IberianPeninsulaandonefortheBalearicIslands(Boeters, 1983,1984,1986;Boeters&Rolan,
1988;Rolan, 1993). Itshouldbenotedthatupto 2001species ofAlzoniellahad always been
understoodasbelonging tothegenusBelgrandiellaA.J.Wagner, 1928.
Apart from papersonnewtaxa,therehavebeenvery few studies onthebiology of
thesespecies (Rolan, 1989).TheIberianspeciesofAlzoniellaarebasically distributedinthe
North of the IberianPeninsula(Rolan, 1993). In previous studies, SEMphotographs of
shells, protoconchs and radulaewere shown aswell as someanatomicalfeatures,partic-
ularlyof theloop of therectum in thepallialcavity, which canbe seenby transparency,
and thepenis (Boeters, 1983, 1986;Boeters & Rolan, 1988; Rolan, 1993). However, the
female genital system was seldom studied, except for some schematic drawings by
Boeters (1983, 1986, 1988) and Boeters& Rolan (1988). For the sake of completeness, it
114 BASTERIA,Vol. 71,No. 4-6,2007
shouldbementionedthatBoeters (2000,2001) revisedAlzoniellaforFrance.
Aimsofthisworkare (1)togive areview ofthespecies ofAlzoniellaandGuadiellathat
inhabitthe IberianPeninsula, (2) to present new informationonshell, protoconch and
radular characters, (3) to describeanatomicalfeatures that maybe relevantfor the sys-
tematicsofthetaxa, and(4)toestablishtheareas wherethesespecies occur.
MATERIALAND METHODS
In recent years, we investigated the malacological freshwater faunaof the Iberian
Peninsula, collecting samples from more than 1500 springs and water sources. Some
aspects of the collecting methodology and investigations have already been explained
(Rolan,1993).Thesamplingwascarried outby directobservation ofspecimens onstones,
by washing deadleaves, mossesand otherkindsofaquatic vegetation or by sieving mud
andvegetation. Livesnailswere drawntomakeapreliminaryclassificationbyexamining
shellcharacters andexternalmorphology such asthe penis andrectum shape. They were
later fixed in 70%ethanol.Dissection methodology is describedin Ramos et al.(2000).
Scanning electron microscope (SEM) photographs were madewith a PHILLIPS XL30.
Shellswere cleanedby removing theperiostracum throughimmersionin5% commercial
Chlorox, cleanedin anultrasonic50WRaypa apparatusandthencoatedwithagold layer
and fixedtothestubby meansofagraphite paste.
Sincenosignificant differenceswereobservedbetweentheopercula oftheAlzoniella
species investigated, only someof themhavebeenrepresented graphically. Spire whorls
arecountedfollowing Verduin's method(1977).
Locality names and UTM references are from the official map of the Army
Geographical Service, series 1:200.000to 1:25.000.Only new localities are listed in this
order: province, UTM number, locality. Distances in km are measuredalong the road.
Maps were donewithMapgrafix GIS for Macintosh. Only collecting localities are illus-
tratedonthemaps,eachpoint representing a 10km-square grid.
ABBREVIATIONS
Abbreviationsforanatomicalcharacters:Ag,albumengland;Be,bursacopulatrix; Cc,
cerebral commissure;Cg, capsule gland; Cgl, leftcerebral ganglion; Ct, ctenidium; Oes,
oesophagus; Ov, oviduct; P, penis; Pgl, leftpleural ganglion; PI, penial lobe;Po, pallial
oviduct; Pr, prostate; R, rectum; Sbo, suboesophageal ganglion; Spo, supraoesophageal
ganglion; SRI, seminalreceptaculum 1; SR2, seminalreceptaculum 2; Ss, style sac; St,
stomach;L, length; W, width.
For collections: BOE, H.D. Boeters, Munich, Germany; BOURG, J.R. Bourguignat,
Museum d'Histoire Naturelle, Geneve, Switzerland; CER, E. Rolan, Vigo, Spain; CZL,
Centro de Zoologia de Lisboa, Portugal; MHNS, Museo de Historia Natural of the
University of Santiago de Compostela, Spain; MNCN, Museo Nacional de Ciencias
Naturales, Madrid, Spain; PAL, A.Paladilhe, Faculte des Sciences, Montpellier, France;
SMF,Forschungsinstitut undNaturmuseumSenckenberg, FrankfurtamMain, Germany;
ZMA,Zoologisch MuseumAmsterdam,TheNetherlands.
Arconadaetah ArevisionofthegenusAlzoniellaontheIberianPeninsula 115
SYSTEMATICPART
Family Hydrobiidae Troschel, 1857
Key forthespecies
(notincludedareAlzoniella(A.) muritaandA.(A.) onatensis)
1 - Balearicspecies Alzoniella(A.)edmundi
- Iberianspecies 2
2 - Species fromtheSouthoftheIberianPeninsula Guadiellaandalucensis
- Species fromtheNorthof theIberianPeninsula 3
3 - Rectum loop not curved 4
- Rectumloop is marked 5
4 - Peniallobeis smallerthanthepenisand hasnopapillae Alzoniella(A.)rolani
- Penial lobeis as long asthepenis andhasseveral papillae 11
5 - Rectum loop is simple and its curved concavity is perpendicular or slightly points
towardstheanterior partof thebody Alzoniella(A.) lucensis
- Rectumloop ismarkedly curvedand itsconcavity points towardstheanteriorpartof
thebody 6
6 - Rectumloop is U-shaped 7
- Rectum loop is S-shaped 8
7 - Shellunder2mm, cylindrical;unpigmented orblack pigmentedpenis
Alzoniella(A.) cantabrica
- Shellover2 mm,almostovoid outline;pink pigmentation onthepenis
AAllzzoonniieellllaa((AA..))oovveetteennssiiss
8 - Shelllengthunder1 mm;shellovoid Alzoniella(A.)asturica
- Shell over 1 mm; shellovoid-elongated to cylindrical 9
9 - Shellwithalmostcylindrical outline Alzoniella(A.)galaica
- Shellwithdifferentoutline 10
10- Bodypigmented; shellovoidwitha deep suture Alzoniella(A.) marianae
- Unpigmented body; shellovoid-elongated withanotvery deep suture
AAllzzoonniieellllaa((AA..)) mmoonnttaannaa
11- RSI andRS2ofthesamelength Alzoniella(Navarriella) elliptica
- RS2 remarkably smallerthanRSI Alzoniella(Navarriella)pellitica
GenusAlzoniellaGiusti& Bodon, 1984
Typespecies:AlzoniellafinalinaGiusti & Bodon,1984.
Subgenus Alzoniella(Alzoniella)Giusti&Bodon, 1984
For the characteristicsof this subgenus, see Giusti& Bodon(1984) who described
Alzoniellaand compared it with some other closely related European genera such as
AvenoniaNicolas, 1882,Belgrandia Bourguignat, 1869,BelgrandiellaA. J.Wagner, 1928(not
1927; see Gittenberger & Uit de Weerd, 2006: 131), Bythinella Moquin-Tandon, 1856,
Bythiospeum Bourguignat, 1882, Phraetica Velkovrh, 1970 and Pseudavenionia Bodon&
Giusti, 1982.
According to theoriginal description, theshell is small, whitish-transparent, conical
116 BASTERIA,Vol. 71, No.4-6, 2007
or cylindrical-conical, with 3-4 spire
whorls, quiteconvex, withanobtuseapex.
The aperture has a simple peristome, an
umbilicus that is slightly open or reduced
to a slit-like fissure. Operculum thin, cor-
neous, paucispiral, slightly yellowish atits
centre,withanexcentric nucleusandwith-
outinternaloutgrowths. The intestine sur-
roundsthestyle sac. Stomachlacking cae-
cum. Radula typically taeniaglossan, the
centralteeth arebutterfly-shaped, general-
ly withoneor twobasalcusps. Thelateral
teethhaveonebigdenticleand4-5moreon
eachside. Internalmarginal teethwith23-
28 smallanduniformdenticles.Ctenidium
not very developed, with 5-11 lamellae.
The nervous system (fig. 1) is typically
hydrobiid and has a long supraoesopha-
geal and a short suboesophageal connec-
tive. The oesophagus runs straight along
the cerebral ganglion. The right pleural
and the supraoesophageal ganglion are of
the same size. The leftpleural ganglion is
more rounded than the suboesophageal
ganglion. The penis has generally one
penial lobe located atitsconcavesidebut,
exceptionally, it can have more thantwo.
Theprostate is typically bean-shaped. The
anterior female genital system has an
Fig. 1. Partial nervous system of an Alzoniella ovoidbursacopulatrix varying frommedi-
species: A. (A.) lucensis (Rolán,1993), fromLugo, um to large, a long, unpigmented renal
Castillo de Doiras towards Ancares (PH63).Scale oviduct and two seminalreceptacles. The
bar: 250µm(forabbreviations,seethetext). renaloviductmakes an opencircularloop
that leans over the bursa and the pallial
oviduct.Thebursaprotrudes posteriorlybeyond theproximal end ofthealbumengland
Thecapsule gland is generally smallerthanthealbumengland.
Alzoniella(A.) cantabrica(Boeters, 1983)
(figs2, 15C,23-24,72-74, 93-94;map3)
Belgrandiellacantabrica Boeters,1983: 18.Typelocality: Santander,UN99: "Graben rechts derStrafie von
Treceno nachLa Revilla beiSan Vincente dela Barquera ,[UN99],
Typematerial:seeBoeters, 1983.
Localities (see Boeters, 1988;Rolan,1993).- New records (ito xxxii). Asturias: (i)UP70, 1km W.
Noriega,springnearriver. Cantabria: (ii)UN79, 2km fromPineres justbeforeturn-offtoRoda,torrent;
(iii)VN29, Presillas,spring;(iv) VN29,Aes, springatchapel"Virgen de Gracia";(v) VN68,3km SSW.
exit from Ramales de la Victoria, old spring; (vi) VN68, at Valnera before the turn-off to Concha
(Karrantza),brook;(vii)VN59,4km S.Arredondo,Cueva de Coventora;(viii)VN49, 1.5km N.Linto at
Arconada etahArevisionofthegenusAlzoniellaontheIberianPeninsula 117
Fig. 2.Anatomical features ofAlzoniella (A.) cantabrica (Boeters, 1983);Santander,Cobrices (UP90). A,
digestive system; B, C. female genitalia; D. penis; E. rectum loop; F. osphradium and ctenidium.
Scalebar: 500µm(see abbreviations intext).
crossroadtoMiera,torrent;(ix) VN49,S.LaVega,several waterfalls.Burgos:(x) VN33, S.Escalada,near
Tubilla delAgua,brookonmountainside;(xi) VN33,N.ofCovanera,largespring;(xii)VN33,Covanera,
"PozoAzul";(xiii)VN32,S. exitofTubilla delAgua,springsinrocks;(xv) VN34,E entrancetoOrbaneja
delCastillo,spring; (xiv)VN45,between Areba and Hoz deAreba,brookalongsideriver;(xvi) VN45,1
kmNW.Incinillas,channel goingunder road;(xvii)VN57,N.exit oflocality"Rio deLasia";(xiii)VN47,
between locality "Rio deLasia" and Puerto dePas,between4and 6kmN.Rio deTrueba,springs; (xix)
VN57,3kmN. locality"Rio deLasia",spring;(xx) VN84,exitofSan Zadornil towards Villafria,spring;
(xxi)VN84,about 3km behind SanZadornil towards Villafria;(xii)VN84, 1.5kmbehind Villafria,"La
Fuentefria",spring. Alava:(xxiii) VN96,Aguiniga,spring; (xxiv) N94,Entrance toAstulez,brook;(xxv)
VN94, entrance toNograro, springwith pool; (xxvi) VN84, turn-offtoQuejo,springat side ofbrook.
Vizcaya: (xxii)VN78,Ambasaguas,nearturn-offtoConcha,spring.Palencia: (xxviii) UN75,W.exit of
Lebanza,springalongsideriver, (xxix) UN65,notfarW. ofSantibanez de Resoba,spring. Leon: (xxx)
UN45, about 5km W. ofValverde de la Sierra,spring; (xxxi) UN56. Llesca pass 5km W.border with
Cantabria,under bridge;(xxxii) UN57, downhill Puerto de San Glorio, between km 131 and 133, wet
rocks. Santander: (xxxiii)UP90,Cobrices [Cobreces],springLaMarrana.
Morphology. - Shell(cf.Boeters, 1983:figs37-41; 1988: figs 194-197;Rolan, 1993:pi. 1
fig. 2, pi. 2 figs 3-4). Some shells of thisspecies are illustratedby figs 23-24 and proto-
conchsby figs 72-74. Operculum notexamined.
Anatomy (after 5 femalesand 2 males from locality ii, 2 males from locality v, 3
femalesand2malesfromlocalityxxxiiiand2malesfromlocality xviii).- Thereisawell-
118 BASTERIA,Vol. 71, No.4-6, 1007
developed ctenidium(fig.2F) with 10-11 large lamellae.The intestine,whenleaving the
stomach forms the usualZ-shaped loop (fig. 2A). The rectum (figs 2E and 14E) is U-
shaped, usually pointing towardstheanteriorpartof thebody. (cf.Boeters, 1983:figs 7-8;
1988;Rolan, 1993:pi. 4fig.2).
Fig.3.Anatomicalfeatures ofAlzoniella(A.)edmundi(Boeters,1984);Majorca,Valldemossa(DD59).C,F,
penisfromalive specimen;A,B,D, E,G,H, femalegenitalia.Scalebars: 500µm
Arconadaetah A revisionofthegenusAlzoniellaon theIberianPeninsula 119
Theradulahasthegeneralcharacteristics ofthegenus(figs 93-94)although inoneof
the studiedpopulations (Cobrices), thecentralteethhave threebasal cusps oneachside
(fig.94), the thirdbeing very small. Inthis population, the upperborder of thecentral
teeth seemstobemoreexcavated, (cf.Rolan, 1993:pi. 6 figs 1-3).
Genitalsystem (table3). Theovoidbursacopulatrix iswell-developed andhasalong
and curved duct(fig.2B-C). TheSRI is longer thantheSR2which,inanaturalposition,
leansover the bursa copulatrix (fig. 2B). The albumengland is larger than thecapsule
gland (fig.2B) (cf. Boeters, 1983: figs 28-30, 1988:figs 210, 224, 227).Thepenis (figs 2D,
15C)is long and wideand has arounded lobelocated in a medialposition. Sometimes
thereis apigmentedarea distallybehindthis lobe (cf.Boeters, 1983:figs15-22; 1988:208-
227;Rolan, 1993:pi. 4figs 10-16)
Distribution (map 2). - This species has a wide range comprising some northern
Spanish provinces suchas theeastern partofAsturias,Cantabria, theBasque Country, the
NorthofLeon,Burgos andPalencia.
Remarks. - The number of new localities greatly increases the known range of
Alzoniella(A.) cantabrica,whichcanbefoundsometimes sympatric with A.(A.)ovetensis
(Rolan, 1993) (UP70) or A. (A.) montana (Rolan, 1993) (UN57, VN68). Female genitalia
show littlevariability. On the contrary, males of this species show greatervariability as
regardspenis shape andpigmentation (Boeters, 1983;Rolan, 1993).
Alzoniella(A.)edmundi(Boeters, 1984)
(figs3, 25-28,76-77, 95;map1)
Micronasaxatilis (Reynies, 1843);Moolenbeek,1980:101.
Belgrandiellaedmundi Boeters, 1984: 10,fig. 3, pi. 1 fig. 9a-b. Type locality: Baleares, DD69:Majorca,
Valldemossa,FontdeS'Aiqueta
Typematerial: seeBoeters, 1984.
Localities.- Two new localities increase the number of known localities: Majorca: (i) DD59,
Estellenchs,spring; (ii)DD59,S. exitofValldemossa,FontS'Aigueta.
Morphology. - Shells(cf. Moolenbeek, 1980;Boeters, 1984:10)are illustratedby figs
25-28 andprotoconchs by figs76-77. Operculumnotexamined.
Anatomy (after2 femalesand2 malesfromlocality i).- Livespecimens (fig. 3C) are
completely unpigmented so that the pink radular sac can be seenby transparency. The
eyes areblack spotsandthereisaposterior areawithwhitespots. The ctenidium(fig.3E)
is barely developed andhas about 3-5lamellaesituated nearthe S-shaped rectum loop
(fig.3H), whichpoints towardstheanteriorpartof thebody.
Theradula(fig.95) hastheusualcharacteristicsofthis genus.Thecentralteethhave
twobasalcusps oneach sideandtapered denticles.
Genitalsystem (table 4).Thefemaleshaveasmallpinkpseudopenis originating inthe
right partofthehead,posterior totheright cephalic tentacle,withasimilarshapeto that
of themales(fig.3D). Thebursacopulatrix (fig.3A) issacciformandhasamediumsized
straight duct.The SRI (fig. 3B) isshorter andwiderthan theelongated SR2which,in its
naturalposition, leansover thebursa copulatrix (fig.3A). Therenal oviductposteriorto
theRS2is slightly widenedand is pearl-coloured duetothe accumulatedsperm inside.
Thepenis (figs 3C, 3F, 3G) is very large,broad and longer than the head. Its roundish
penial lobe is locatednear the pointed tip. In fixed specimens, the penis is completely
unpigmented,but thepenises oflivespecimens presentseveralwhitespots.
120 BASTERIA,Vol. 71, No.4-6,2007
Fig.4.Anatomicalfeatures ofAlzoniella (A.)rolani (Boeters,1986);Pontevedra,Rasas,Gondomar (NG25).
Drawings afterlive specimens.A-B, distal female genitalia; C,penis; D, osphradium and ctenidium.
Scalebar: 500µm.
Fig.5.Anatomical featuresofAlzoniella (A.)asturica(Boeters&Rolán, 1988);Asturias, Grado,La Fontona
(QJ30).Drawingsfrom fixed material. A,B, distal femalegenitalia;C,osphradiumand ctenidium;D,F,
penis;E,rectumloop.Scale bars:500µm.
Arconadaetat ArevisionofthegenusAlzoniellaon theIberianPeninsula 121
Distribution (map 1). - This species is only known from the islandof Majorca
(Balearic Islands).
Remarks. - Anatomicaldatawere unknown until now and only shells had been
described.Thisspecies canbefoundsometimestogetherwithPseudamnicolaspec. (DD59).
Alzoniella(A.) rolani(Boeters, 1986)
(figs4, 14A-B,29-32,58-60, 78-79,96-97;map 1)
Bythinellabrevis; Rolan,1983:120.
BelgrandiellarolaniBoeters, 1986:127,pi. 18afig. 3.Typelocality: Pontevedra,NG07: Islas Cies,S.Martin.
Typematerial:seeBoeters,1986.
Localities. - Localities (i),(iii)and (v) can beadded asnew (cf. Rolan, 1989,1993). ACoruna: (i)
GA98,atFeiradoTres,springknown asOPlumar.Pontevedra:(ii)NG25,Mill 2kmS. ofGondomar.(iii)
NG29, Carballeira deSan Xusto,springsnearold sulphurbaths.Portugal: (iv) NE25,BeiraBaixa,near
Rebordosa;(v)NF62,DouroLiteral,highwayfromFafe toPovoa doLanhosoenteringviaturn-off5km
from SerafaotoVilarielho,about2km fromLugardela Igreja,spring; (vi)NF95,DouroLiteral,Serra do
Marao.
Morphology. - Shell.Severalshellsare illustratedby figs 29-32.Twoofthese(figs 29-
30) arecharacteristicforthisspecies, whiletheshellsfromtheSerradoMarao(locality vi;
fig. 31) and Rebordosa(locality iv; fig. 32) are atypical (cf. Boeters, 1986;Rolan, 1989,
1993).Theoperculum(figs 58-60)is characteristicforthis genus.
Anatomy(after 5femalesand 2malesfromlocality ii and 2malesfrom locality vi).-
SeealsoBoeters(1986). Thectenidium(fig.4D) iswell-developed and hasapproximately
8to 13lamellae.The rectum describesasmoothcurve inthepallial cavity(figs 4A,14A)
and theanus doesnotend close totheedge of thiscavity(cf. Boeters, 1986;Rolan, 1989,
1993).
Theradula(figs 96-97) hastheoverall shape of theotherspeciesof this genus.Ithas
awidesinglecusp ateach sideof thecentral teeth.
Genital system (table 5). The capsule gland (fig. 4A) constitutes almost 50% of the
length of thepallialoviduct, which completely covers therectum loop. In somefemales,
the pallial oviduct is less developed. The bursa copulatrix (fig. 4B) is sacciform, well-
developed andhas amedium-sizedstraight duct. TheSRI is long andwide; in natural
position itis hiddenbytheloop oftherenaloviduct.TheSR2isroundishandvery small.
Thepenis (fig. 4C) is smalland has abluntedtip. Thepenial lobe is located in amedial
position; it is very long and roundish, sometimes longer than total penial length (cf.
Boeters, 1986;Rolan, 1993).
Distributon(map 1).- This species is known fromthefour provinces ofGalicia and
fromnorthernPortugal southwardtoCoimbra.
Remarks. - Thenew records do not modify the distribution areaknown for this
species (Rolan, 1989, 1993). Theshells (figs 31-32) fromthe marginal populations of the
southern part of the range differconspicuously from those of the other populations,
which have a consistently uniformshell. Nevertheless, noanatomicaldifferenceshave
been found for those conchologically deviating populations. This widely distributed
species shows acertainamountofvariability insomeanatomicalfeatures.Boeters (1986)
describes thepenis as "intensely black pigmented". There arealsospecimens withcom-
pletely unpigmented penises however, indicating thatthis pigmentation is variable.As
regardsthebursacopulatrix, Boetersdid notmentionany seminalreceptaculum. Therec-
122 BASTERIA,Vol. 71, No. 4-6, 2007
Fig. 6.Anatomical features ofAlzoniella (A.) galaica (Boeters & Rolán, 1988);Lugo, Peares dam (PH00).
Drawingsfromfixedmaterial. A,B,femalegenitalia;C,penis;D,rectumloop;E,osphradiumandctenidium.
Scalebars: 500µm.
Fig. 7.AnatomicalfeaturesofAlzoniella (A.)lucensis(Rolán,1993);Lugo,Castillo deDoiras(PH63).A,B,E,
female genitalia;C,F,penis; D, osphradiumand ctenidium;G,prostate and rectum. Scalebars:500 µm.