Revision of the genera Pseudamnicola PAULUCCI 1878 and

Transcription

Arch. Molluskenkunde
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95figures
| FrankfurtamMain,28.06.2010
Revision of the genera Pseudamnicola PAULUCCI 1878 and Mercuria BOETERS 1971
from Algeria with particular emphasis on museum collections
(Gastropoda: Prosobranchia: Hydrobiidae)
PETER GLÖER1, SLIMANE BOUZID2 & HANS D. BOETERS3
Abstract
Anextensiveinvestigationofnewlycollectedsamplesandmorethan140samplesofmuseum
collections (Museum Paris and Geneva) of Pseudamnicola spp. as well as of Mercuria spp. of
AlgeriaandTunisiahasbeencarriedout.NineoftheformerlydescribedPseudamnicolaspp,of
whichwebelievethatfourspeciesaresynonymsofPseudamnicola dupotetiana, couldbefound
inthecollections.Inaddition,ninePseudamnicolaspp.andtwoMercuriaspp.aredescribedhere
asbeingnew:Pseudamnicola boucheti n.sp., P. chabii n.sp., P. ghamizii n.sp., P. algeriensis n.
sp., P. gerhardfalkneri n.sp., P. calamensis n.sp., P. fineti n.sp., P. linae n.sp., P. rouagin.sp.,
Mercuria bourguignati n.sp., and M. gauthieri n.sp.
Key words:Gastropoda,Hydrobiidae,Pseudamnicola,Mercuria,Algeria,revision,newspecies.
Introduction
In the 19th century, the molluscan fauna ofAlgeria
was intensively studied. The most extensive malacological investigations inAlgeria have been carried out
by BOURGUIGNAT (1862, 1864) and LETOURNEUX (1870).
Inthecourseoftheseinvestigations,14Pseudamnicola
specieshavebeendescribedasbeingnew,inadditionto
the already known species Pseudamnicola dupotetiana
(FORBES1838),andP. luteola (KÜSTER1852).
Inthe20thCenturyPALLARY(1921:202)reportedon
Amnicola lanceolata PALADILHE1869,andA. similis DRAPARNAUD1805fromMorocco.GERMAIN(1908:269–273),
however,mentionedonlyBythinia numidica BOURGUIGNAT
1864,B. orsinii DE CHARPENTIER1852,andAmnicola dupoteti FORBES1838fromtheneighbouringcountryTunisia.
The species diversity in the genus Pseudamnicola
mentioned by B OURGUIGNAT (1864) and L ETOURNEUX
(1870)decreasedinthecourseoftimebymeansoftaxonomic lumping down to one species, P. dupotetiana,
mentionedbyVAN DAMME(1984:17),KRISTENSEN(1985:
7),andBROWN(1994:70).
When BOETERS(1971)designatedthenewgenusMercuria, which he separated from the genus Pseudamnicola, it was not clear which species of this new genus
occur inAlgeria. FromTunisia BOETERS (1976) mentionedMercuria confusa (FRAUENFELD1863)andM. punica (LETOURNEUX & BOURGUIGNAT 1887). Merely Mercuria conovulahasbeenmentionedfromAlgeriabyVAN
DAMME(1984:18).
Authors’addresses:
1
PETER GLÖER,Schulstr.3,D-25491Hetlingen,Germany;E-mail:gloeer@malaco.de
2
SLIMANE BOUZID,DepartmentofBiochemistry,AnnabaUniversity,BP12Annaba,Algeria,E-mail:sbouzid@univ-annaba.org
3
HANS D. BOETERS,Karneidstr.8,D-81545München,Germany;E-mail:boeters@online.de
eschweizerbartxxx_author
©E.Schweizerbart’scheVerlagsbuchhandlung(Nägeleu.Obermiller),2010,ISSN1869–0963
DOI10.1127/arch.moll/1869-0963/139/001-022
2
GLÖER, P., BOUZID, S. & BOETERS, H. D.:RevisionofthegeneraPseudamnicolaandMercuriafromAlgeria
GIUSTI et al. (1995: 132–138) comprehensively discussed the problems involved in species determination
within the genus Pseudamnicola and Mercuria. The
many species they dissected from Malta revealed only
slightdifferencesastotheirgenitalorganswhereasthere
isalargevariabilityoftheshells,sointheiropiniononly
onespeciesofeachgenusoccursinMalta.Incontrast,
BOETERS (in BECKMANN 1987: 7–9) found two distinct
Mercuriaspp.onMalta.
LETOURNEUX & BOURGUIGNAT (1887) stated that most
of the Pseudamnicola spp. were widely distributed in
Algeria. If this is the distribution pattern, it is in fact
distinct from those in other Mediterranean countries.
Pseudamnicolahasacircum-Mediterraneandistribution,
whiletheknownspeciesdiversityincreasesfromEastto
West.Inthosecountrieswherethisgenusiswellinvestigated(Tunisia:BOETERS(1976),Spain:BOETERS(1988),
Balearics: BECKMANN (2007)) the species richness is,
nevertheless,muchlowerthaninAlgeria,andincontrast
toAlgeria the species are more confined to smaller regionsconcerningtheirdistribution.
Thispaperisintendedtoelucidate(i)thespeciesrichnessinthegeneraPseudamnicolaandMercuria,(ii)the
distribution patterns of these Hydrobiidae inAlgeria,
and (iii) to describe the new species: Pseudamnicola
boucheti n.sp., P. chabii n.sp., P. ghamizii n.sp., P. algeriensis n.sp., P. gerhardfalkneri n.sp., P. calamensis
n.sp., P. fineti n.sp., P. linae n.sp., P. rouagin.sp., Mercuria bourguignati n.sp., and M. gauthieri n.sp.
Materialandmethods
The snails were collected by the second author with
asievefromanumberofspringsintheregionsofGuelmaandSoukAhras(northeasternAlgeria).Thesamples
were put into 75% ethanol.The dissections and measurements of the genital organs and shells were carried
out using a stereo microscope (Zeiss); the photographs
weremadewithadigitalcamerasystem(LeicaR8).
ToclarifythestatusofthePseudamnicola speciesdescribedsofar,weborrowedthesyntypesoftheMuséum
d’HistoireNaturelle,Geneva(MHNG,MHNG=collectionBourguignat),theMuséumNationald’HistoireNaturelle(MNHN),Paris,andtheNaturalHistoryMuseum
Goeteborg.TostudythespeciesrichnessandthedistributionofPseudamnicolaspp.andMercuriaspp.weexaminedinadditionmorethan120lotsfromtheMuséum
National d’Histoire Naturelle, Paris, and numerous lots
fromtheMuséumd’HistoireNaturelle,Geneva.
In doubtful cases, the genus group Pseudamnicola/
Mercuriacanonlybedeterminedbythemorphologyof
the penis.To make it possible to study the penis morphologyofdriedmaterialsfromoldsamples,too,weput
somesnailsforfiveminutesintohotwaterwithadrop
ofdetergenttore-hydratethedriedtissue.Thismethod
works,ifthespecimensformerlydriedrapidlyandwithoutanydecomposition.
To be sure that the species fromAlgeria are distinct
from those that have been described fromTunisia, we
borrowedthetypematerial,whichlookedsimilartosome
ofourspeciesthathavealreadybeenstudiedbyBOETERS
(1976).Wecomparedthesyntypesofthefollowingspecies with the material under discussion: Amnicola bythiniopsis(MHNG5274),A. barattei(MHNG5272),A.
saharica(MHNG5370),andA. sterea(MHNG5384).
Thestartingpointofourinvestigationswastolookat
thenewlycollectedmaterialfromtheregionsofGuelma
and SoukAhras, with regard to the variability of the
shellsaswellastotheanatomy.Followingthisapproach,
wecoulddistinguishfourPseudamnicolaspp.intheregionsofGuelmaandSoukAhras.Basedonthisconcept,
wecouldseparatethespeciesofthedriedmaterials:
Every species has at least one constant feature that allowsseparationfromotherspecies,
AtonesamplingsitemorethanonePseudamnicola/Mercuriaspeciescanoccur.
A constant distinguishing feature shows that speciation has taken place. But this concept only works with
samples that comprise a large number of individual
specimensinordertoaccountfortheconsiderablevariabilityconcerningthisspecies’shells.Inaddition,only
adultshellswithanidenticalnumberofwhorlsmaybe
compared, because the shells of younger specimens or
juveniles look distinct from the adults’ because of an
allometric growth. Especially those species of which
thejuveniles arespherical, while theadults areoval or
elongatedoval,canbeinterpretedasbeingtwodistinct
speciesorasbeingonespecieswithalargephenotypic
plasticity,iftheageofthesnails(numberofwhorls)is
not considered. Obviously aberrant shells are excluded
fromdetermination,becausee.g.verylargeshellscould
be the result of an parasitism (SZAROWSKA et al. 2006:
181).Inadditionzoogeographicalaspectsshouldbealso
takenintoconsideration(seee.g.Mercuria pycnocheilia
andM. gauthierin.sp.).
Abbreviationsofcollectionsused:
BOE privatecollectionBOETERS
GLÖ privatecollectionGLÖER.
MHNG Muséumd’HistoireNaturelleGenève,Switzerland
MNHN MuséumNationald’HistoireNaturelle,Paris,
France
SMF SenckenbergResearchInstitut,Frankfurtam
Main,Germany
ZMH ZoologicalMuseumHamburg,Germany
Samplingsites
Algeria can be subdivided from north to south into:
theTellianAtlas (or theTell), in the northern section
bordering the Mediterranean Sea, the Hauts Plateaux,
theSaharanAtlaswiththeAurèsMassif,andtheSahara
DesertwiththeSaharadepressionintheregionofTouggourt(Touggourtdepression).
NorthernAlgeria, theTellianAtlas, which includes
theDjurdjuraMassif,lieswithinthetemperatezone,and
itsclimateissimilartothatofotherMediterraneancountries.TheHautsPlateauxhasasemi-aridclimate,whose
basinprofileexplainsthepresenceofmanysaltmarshes
3
and dry or shallow lakes (sebkhas or chotts). South of
theSaharanAtlas,extendsahugedesertareaoftheAlgerianSaharawithanextremelyaridclimateaswellas
anirregularandunevenlydistributedrainfall.
Thesamplingsitesoftheoldmaterialscouldbefound
onoldmaps(PELET1902),andinadditionthepaperof
LETOURNEUX (1870) was very helpful, because he describedmanysamplingsitesveryprecisely.
Results
DeterminationoftaxathatbelongtothegenusPseudamnicolaandMercuriaisnoteasybecausetheyarenot
equippedwithawealthofdistinguishingfeatures.Neverthelessitwaspossibletodistinguishmanyspeciesof
thegenusgroupsunderdiscussion.Assomelotsofthe
museumcollectionsconsistedofonlyafewspecimens,
oras,inothercases,nosuitablefeaturesfordeterminationcouldbefound,notallspeciescouldbeidentifiedby
us.Manylotscontainedtwospecies,inmostcasesone
ofPseudamnicolaandoneofMercuria.
Toidentifyformerlydescribedspecies,itwasnotpossibletoworkwithtopotypes,becauseinsomeregionsup
tofivespeciesoccur,andthesamplingsiteshavenotin
everycasebeendefinitelydescribedinformertimes.
DissectionofthefemalesoftherecentlyfoundPseudamnicolaspeciesshowedusthatthebursacopulatrixis
notfoldedasinCorrosiella (see BOETERS1970),sothat
thesespeciesbelongtothegenusPseudamnicola.
The Pseudamnicola/Mercuria spp. ofTunisia, mentionedbyBOETERS(1976),couldnotbefoundinAlgeria,
exceptP. meluzziandMercuria globulina,thatoccurin
Algeria,too.
Onlyafewsyntypesandholotypesofthespeciesunderdiscussioncouldbefound(Figs2–5).Inmostcases
we had to determine the species by making use of the
original descriptions, and by those provided by BOURGUIGNAT(1862,1864)andLETOURNEUX(1870),whichare
suitableforspecies’identification.
However,somespeciescouldnotbefoundoridentified by us, because the description was vague or these
species did not occur in the examined samples.These
speciesarelistedbelow:
Pseudamnicola similis (DRAPARNAUD1805)(asCyclostoma, DRAPARNAUD1805:34,pl.1,Fig.15).TheidentityofCyclostoma similishasbeenclarifiedbyBOETERS
& FALKNER (2000).This species belongs to the genus
MercuriaandisdistributedintheMediterraneanregions
ofFrance.Thuswebelievethatthisspeciesdoesnotoccur inAlgeria and, considering BOURGUIGNAT’s description(1864:237),wecouldnotfindanyspeciessimilar
to this, either. In addition BOETERS & FALKNER (2000)
statedthatPseudamnicola confusa FRAUENFELD1863,is
a junior synonym of Mercuria similis. Pseudamnicola
confusahasbeenmentionedbyBOETERS(1976)tooccur
inTunisia.
Pseudamnicola seminium (MORELET 1857) (as Paludina,MORELET1857:376): Thetypelocalityis„…les
eauxthermales,auxalentoursdeConstantine“.InBOURGUIGNAT’scollectioninGenevaandParisthreelotscould
befoundwiththelabel„Pseudamnicola seminium“but
itwasinallcasesaMaresiasp.PossiblyP. seminiumis
anendemicspeciesthatlivesonlyinhotspringsatthe
typelocality.Syntypescouldnotbefound.
Pseudamnicola rouvieriana (LETOURNEUX 1870) [as
Amnicola,LETOURNEUX1870:316]:Thetypelocalityis
“HammamBou-Merzouk”(OuledRahmoun),butinthis
hotspringwecouldonlyfindPseudamnicola constantinae and Pseudamnicola chabii n.sp.MaybeP. rouvierianaisanaberrantformofthelatterspecies,thoughthe
originaldescriptionisverydistinctfromthetwospecies
thatwefoundthere.Syntypescouldnotbefound.
Pseudamnicola pomariensis (LETOURNEUX1870)[Amnicola, LETOURNEUX 1870: 317]: The type locality is
“Sefsef [=Es Safsaf] près deTlemcen” and we had no
lotsfromthissamplingsite.
Pseudamnicola costulata WESTERLUND 1886.WEST-
ERLUND (1886: 82) found this species together with P.
subrictainMarounania[=Marouania(OuedElAneb)]
nearAnnaba.ThisspeciescouldnotbefoundinWESTERLUND’scollection.
Pseudamnicola conovula (FRAUENFELD1863)[asAmnicola, FRAUENFELD 1863: 1026] has been mentioned
by VAN DAMME(1984)andotherauthorsforAlgeria,but
thetypelocalityofthisspeciesistheislandPag,Croatia
(FALKNER & BOETERS2003),thuswedonotbelievethat
this species occurs inAlgeria, and no species could be
foundofwhichtheshellshapecorrespondstotheshells
ofP. conovula.
Fig.1.Thesamplingsitesofthestudiedspecimens.
4
Table1.SamplingsitesofAlgeriaandtaxafound.
Samplingsite
AïnBendaoud(OuledDriss)
AïnFeïd-El-Bagrat(MedjezSfa)
AïnH’djar(MedjezSfa)
AïnKefKourrath(MedjezSfa)
AïnM’Daourouch
AïnM’lila
AïnOussera
AïnSfa(entreToukriaetSebaïnaioun)
AïnSultan
AïnZaaroura(MedjezSfa)
Region
SoukAhras
Guelma
Guelma
Guelma
SoukAhras
OumEl-Bouaghi
Djelfa
Tissemsilt
AïnDefla
Guelma
Algiers
Algiers
BahrdeTiguedidineprèsdeDjamaa
Boghar
Bône[=Annaba]
ElOued
Médéa
Annaba
Boufarik
Blida
Boughari[=KsarElBoukhari]
Bougia[=Bejaïa]
CapdeGarde
Champdemanoeuvre
Cherchell(fontaineprèsdelamer)
Constantine
Dellys
ElKhroub
Ferkane
Fontainechaude
FontainechaudeduruisseaudesZaatcha
FontaineentreDayaetTélag
FoukaprèsdeKolea
HammamSidiDjaballah(Cheffia)
HôpitalduDey
KsarChellala
Laghouat
M’Sila
Mefrouch
MoktaAl-Hadid
MoktaElOued
Negrine
Nemours[=Ghazaouet]
Médéa
Bejaïa
Annaba
Algiers
Tipaza
Constantine
Boumerdès
Constantine
Tébessa
Batna
Biskra
SidiBelAbbes
Tipaza
ElTarf
Algiers
Tiaret
Laghouat
M’Sila
Tlemcen
Annaba
entreDjelfaetBoughar
Tébessa
Tlemcen
entreBiskraet
Touggourt
Oran
Ouargla
Mila
Annaba
Touggourt(Ouargla)
ElOued
Touggourt(Ouargla)
Biskra
Mila
Oasisd’Ourir
Oran
Ouargla
OuedAthmenia
OuedElAneb
PuitsartésiendeSidiRached(OuedRir)
PuitsartésiensdeMeghaïer
PuitsartésiensdeSidiSlimane(Oued Rir)
Ruisseaudelafontainechaude
SaintDonat[=Tadjenanet]
Taxa
P. rouagi n.sp.
P. finetin.sp.
P. linae n.sp.
Pseudamnicolasp.
P. numidica
P. bouchetin.sp.
P. dupotetiana
P. dupotetiana
P. gerhardfalkneri n.sp.
P. calamensis n.sp.
P. dupotetiana, P. boucheti n.sp., P. ghamiziin.sp.
M. bourguignati n.sp.,M. saharica, M. globulina
M. pycnocheilia
P. dupotetiana
P. letourneuxiana, P. luteola, P. meluzzi
M. globulina, M. bourguignati n.sp.,
P. boucheti n.sp., P. luteola, P. dupotetiana
M. bourguignati n.sp.
P. dupotetiana
P. ghamizii n.sp.
P. dupotetiana, P. ghamiziin.sp.
P. dupotetiana
P. meluzzi, P. numidica, P. boucheti n.sp.
P. numidica
M. pycnocheilia
P. boucheti n.sp.,P. chabii n.sp.
M. saharica
P. algeriensis n.sp.
P. letourneuxiana
P. dupotetiana
P. luteola
P. numidica
P. dupotetiana
M. saharica
M. pycnocheilia
M. gauthieri n.sp.
M. pycnocheilia
P. luteola
M. pycnocheilia
P. ghamiziin.sp.
P. boucheti n.sp.
M. saharica
M. sharica
M. saharica
M. saharica
P. bouchetin.sp.
5
6
Samplingsite
SidiM’Cid(thermalspring)
SidiMakhlouf
SourceduBoumerzoug(OuledRahmoun)
Sourcefaceàl’InstitutPasteur
ThenietEl-Had
Tlemcen
Zardezas
Region
Constantine
Laghouat
Constantine
Algiers
Tissemsilt
Tlemcen
Skikda
Taxa
P. meluzzi
M. saharica
P. chabii n.sp.,P. constantinae
P. ghamiziin.sp.
P. luteola
P. dupotetiana, P. algeriensisn.sp.
P. kuteola, M. globulina
Figs 2–5. Syntypes of Pseudamnicola spp. fromAlgeria from MHNG.—2: B. dersertorum (vicinity of Bône, synonym ofP.
letourneuxiana).3:B. pycnolena (Mascara,synonymofP. dupotetiana).4:B. perforata (vicinityofOran,synonymofP. dupotetiana).5:P. letourneuxiana (Djendell).
Pseudamnicola orientalis (FRAUENFELD1863)[asAmicola,FRAUENFELD1863:1027]couldnotbestudiedby
usbecausetheNaturalHistoryMuseumViennadoesnot
lend out type materials. InVienna exist 9syntypes and
inadditionalotwith2syntypesseparatedoftheoriginal
sample.The measurements, carried out by the curator
ANITA ESCHNER, are: shell height 1.7mm, shell width
1.25mm.Thisspeciescouldnotbefoundinthematerial
studiedbyus.
Identificationkeytothegenera
1.Freshshellswhitish,especiallytheapertureandumbilicus, opaque, umbilicus closed to slit-like,
peniswithalobe-likeappendix ......... Mercuria
1'. Shells not whitish, umbilicus opened, simple penis
withoutanappendix..................Pseudamnicola
Genus Pseudamnicola PAULUCCI1878
T y p e s p e c i e s :Bythinia lucensis ISSEL1866.
In Pseudamnicola species, the renal oviduct is pigmented in black and bears a bursa as well as a receptaculum(rs1).Theoviductenterswithonetothreespiral
loopsintothekidney,beforethepallialoviductloopsto
thewallofthestomach.
Identificationkey
Insomespeciesthejuvenileslookliketheadults,in
other cases the juveniles are of a more spherical shape
thantheadults.Thiskeyismadetoidentifyadultspecimenswithfourwhorlsandmore.
1. Shellspherical,shellheight2–3mm...................... 2
1'. Shellnotspherical................................................... 3
2. Shelllighthorn-coloured,glossy,2.1–2.9mm
.................................................... P. dupotetiana
2'. Shell dark-horn-coloured, surface lustreless, finely
striated,2.9–3.2mm ...................... P. numidica
3.Aperture triangular rounded, suture weak, umbilicus
closed .............................................................. 4
Figs6–20.Pseudamnicolaspp.ofAlgeria.6:P. dupotetiana(Algiers,champdemanoeuvre).7:P. numidica (Constantine).8:P.
meluzzi(Constantine).9:P. chabii n.sp. (Batna,Fontainechaude,holotype).10:P. constantinae(OuledRahmoun).11:P. gerhardfalkneri n.sp. (AïnSultan,holotype).12:P. calamensis n.sp. (AïnZaaroura,paratype).13: P. linae n.sp.(AïnH’Djar,holotype).14:P. algeriensis n.sp.(Tlemcen,holotype).15:P. luteola(ThenietEl-Had).16:P. rouagi n.sp.(AïnBendaoud,holotype).
17:P. fineti n.sp.(AïnFeïd-El-Bagrat,holotype).18:P. boucheti n.sp.(Batna,Fontainechaude,holotype).19:P. ghamizii n.sp.
(Algiers,champdemanoeuvre,holotype).20:P. letourneuxiana(Annaba).
7
8
3'.Apertureoval,suturecleartodeep........................... 6
4.Shellsmall,2.1–2.6mm ........................................... 5
4'. Shelllarger,3.4–4.1mm ....................P. constantinae
5.Shellopaque,silky,lighthorn-coloured ...... P. meluzzi
5'.Shellthickwalled,silky ......................P. chabii n.sp.
6.Shellelongateoval,heightsmallerthan3mm ......... 7
6'. Shellnotelongatedovalorheightlargerthan3mm 8
7.Sutureclear,whorlsconvex ...... P. gerhardfalkneri n.sp.
7'.Sutureflat,whorlsslightlyconvex. P. letourneuxiana
8.Shellelongatedoval,heightlargerthan4mm.......... 9
8'. Shell oval to elongated oval, height smaller than
4mm .............................................................. 10
9.Whorlsstepped,shellheight4.0–4.3mm
............................................ P. algeriensis n.sp.
9'.Whorlsrounded,shellheight4.5–5.0mm .. P. luteola
10.Surfacesilky................................... P. boucheti n.sp.
10'.Surfaceglossy ...................................................... 11
11.Shelllighthorn-coloured,3.9–4.3mm,shellheight:
width=0.68 ........................... P. ghamizii n.sp.
11'.Shelldarkhorn-coloured,height2.7–4.0mm....... 12
12. Shell small, height smaller than 2.7mm, oviduct
loop1smalland1largeloop.......P. linae n.sp.
12'.Shelllargerthan2.7mm,oviductloopwith1small
orthreesmallloops....................................... 13
13.Shelldarkhorn-coloured,2.7–3.0mm,oviductloop
withthreesmallloops........ P. calamensis n.sp.
13'.Oviductloopwithlessthanthreeloops ............... 14
14.Oviductloopwithonesmallloop.......P. fineti.n.sp.
14'.Oviductloopconsistsonlyofonelargeloop,tapered
attheproximalpart................... P. rouagi n.sp.
Pseudamnicola dupotetiana (FORBES1838)
Fig.6
1838 Paludina Dupotetiana FORBES,AnnalsofNaturalHistory,2ndseries:254,pl.12,Fig.3,loc.typ.:“Inmuddy
rivuletsneartheseaatAlgiersandBougia[=Bejaïa]”.
1862 Bythinia pycnolena BOURGUIGNAT,LesSpicilègesMalacologiques:120.
1862 Bythinia perforata BOURGUIGNAT,LesSpicilègesMalacologiques:118.
1870 Amnicola valvatidea LETOURNEUX, Annales de Malacologie,1[1870–1884]:319.
1887 Amnicola sterea LETOURNEUX et BOURGUIGNAT, Prodrome de la Malacologie terrestre et fluviatile de La
Tunisie:146.
1997 Pseudamnicola dupotetiana,–GHAMIZI&al.,Haliotis,
26:36(NeotypedesignationforPaludina Dupotetiana
FORBES1838).
M a t e r i a l e x a m i n e d : MNHN:Algiers (4samples:
2ex., 7ex., 10ex., champ de manoeuvre 9ex.); Cherchell
(fontaine près de la mer, 8ex.);Tlemcen (2samples: 13ex.,
14ex.);Boufarik(3samples:2ex.,8ex.,9ex.);Mefrouchprès
deTlemcen (2ex.);Tlemcen (numerous; SMF 334670/50);
Ksar Chellala (numerous; SMF 334669/42); Boghar, 2ex.;
Aïn Oussera, 6ex.; two unknown sampling sites (12ex.,
17ex.).—MHNG: MHNG 5270 (3ex., labelled as Amnicola barattei LETOURNEUX & BOURGUIGNAT)Aïn Kebiriti, entre le Chott Fedjedj et le Djebel Cherb,Tunisie. specimens
fromcoll.BOURGUIGNATinMHNG,withoriginallabel;MHNG
5281(numerous,labelledasAmnicola doumeti LETOURNEUX &
BOURGUIGNAT)Tala,Tunisiespecimensfromcoll.BOURGUIGNAT
withoriginallabel;MHNG5306(7ex.,labelledasAmnicola
globulina LETOURNEUX & BOURGUIGNAT) temple au Zaghouan,
Tunisiespecimensfromcoll.BOURGUIGNATwithoriginallabel;
MHNG 5308 (numerous, labelled as Amnicola globulina LETOURNEUX & BOURGUIGNAT)Tala,Tunisie,specimensfromcoll.
BOURGUIGNATwithoriginallabel;MHNG5383(5ex.,labelled
as Amnicola sterea BOURGUIGNAT)Aïn Sfa entre Toukria et
SebaïnAioun [ca. 7km east ofTissemsilt]), specimens from
coll. BOURGUIGNAT with original label; MHNG 5384 (numerous, labelled as Amnicola sterea BOURGUIGNAT) Limagues
(Nefzaouia),Tunisie,specimensfromcoll.BOURGUIGNATwith
originallabel;MHNG5385(1ex.,labelledasAmnicola sterea
BOURGUIGNAT)Limagues(Nefzaouia),Tunisie,withorig.label;
MHNG 5387 holotype of Amnicola valvatidea LETOURNEUX
1870 (1ex., labelled as Amnicola valvatidea,“d’Aïnser-Aoumeur,surlaroutedeSétifàTakitount“).Untilfurthernotice,
thisjuvenilespecimenishereconfinedasasynonymtoPseudamnicola dupotetiana.
D e s c r i p t i o n :Shellspherical,horn-coloured,surfaceglossy,apexblunt,4–5whorlswithaclearsuture,
umbilicus slit-like, spire small to medium sized, spire
height0.2ofshellheight,apertureovalandslightlyangled, peristome sharp, shell height 2.1–2.5mm, shell
width1.8–2.1mm.
D i s t r i b u t i o n :CentralandwesternTellandcentralHautsPlateaux.Surprisingly,thisspeciescouldnot
befoundintheeasternpartofAlgeria.
R e m a r k :BecauseofthemisinterpretationofP. dupotetianabyBOURGUIGNAT(seediscussion),hedescribed
Bithynia pycnolenain1862,whichisinouropinionconspecificwithP. dupotetiana.Onthetubeoftheholotype
ofB. pycnolenaalabelwrittenbyBOURGUIGNAT isfound,
and the sampling site corresponds to the locus typicus.
BOURGUIGNAT(1862)measuredashellheightof2.5mm
and a width of 2mm, our measurements of this single
shell (Fig. 3): shell height 2.1mm, width 1.8mm.The
original description of B. pycnolena corresponds well
with this shell. Concerning Bythinia perforata BOURGUIGNAT1862,BOURGUIGNATmentionedashellheightof
3mmandashellwidthof2.5mm.Wemeasuredashell
heightofthephotographedspecimen(Fig.4)of2.3mm
andawidthof1.9mm,theotheronehadashellheight
of 2.1mm and a width of 2.0mm. Both shells looked
uniform.ThisspeciescouldnotbefoundinthecollectionofMNHN,onlythesyntypesofB. perforataexist
in BOURGUIGNAT’s collection in Geneva. In our opinion,
thisspeciesisanaberrantformofP. dupotetianawitha
larger umbilicus.A specimen of MHNG 5385labelled
9
Figs21–25.Pseudamnicola luteola.21–23:Specimen(Zardezas),22:Penis,23:Mantle,24–25:Shells(ThenietEl-Had).-p=
penis,s=snout,t=tentacle.—Figs26–29.Pseudamnicola numidica. 26–27:Constantine,28–29:AïnM’Daourouch(afountain
inSoukAhrasregion,Bouzidleg.).
asAmnicola sterea BOURGUIGNAT1868wasisolatedand
labelled as "syntype" by BOETERS. Because this species
couldbeidentifiedbyusasP. dupotetiana, P. stereaisa
youngersynonymofP. dupotetiana.
Pseudamnicola luteola (KÜSTER1852)
Figs15,21–25
1852 Paludina luteola KÜSTER, In: MARTINI & CHEMNITZ:
SystematischesConchylien-Cabinet,(Ed. 2),I.21:44,
pl.9,Figs8–9,loc.typ.:“inderProvinzAlgier”.
1886 Pseudamnicola subricta WESTERLUND,Faunaderinder
paläarctischen Region lebenden Binnenconchylien. 6:
82),loc.typ.:“AlgerieninMarounania[=Marouania]
beiBona”.
M a t e r i a l e x a m i n e d :ThesyntypesofPaludina luteolacouldnotbefoundinKÜSTER’scollectioninSenckenberg
(R.JANSSENinlitt.).Thuswecouldindentifythisspeciesonlyby
utilisingtheoriginaldescription,andinadditionweconsidered
the drawing by KOBELT (1892: Orig. Icon. 826).—MNHN:
ThenietEl-Had(2samples:numerous;17ex.,SMF334674/36);
Boufarik(59ex.);Laghouat(2samples:8ex.,5ex.);Zardezas
(numerous);Oran(7ex.);Annaba(1ex.,coll.Westerlund);unknownsite(2samples:22ex.,numerous).
D e s c r i p t i o n :Shellelongatedoval,horn-coloured,
surface finely striated, glossy, apex blunt, 5whorls
roundedwithaclearsuture,umbilicusslit-liketoclosed,
spire height 0.3of shell height, aperture oval, slightly
oblique,peristomesharpbutflanged.Thelasttwowhorls
oftheadults(!)reachabout7/8oftheshellheight.Shell
height4.5–5mm,shellwidth2.5–3.1mm.
The mantle is brown coloured with a white border
(Fig.23).Thepenisisbroadandobtuseatthedistalend
(Fig.22).
D i s t r i b u t i o n : Eastern, central and westernTellian zone, central Hauts Plateaux and the northern edge
oftheSahara.
R e m a r k : In W ESTERLUND ’s collection (SMNH
StockholmType7480–syntypeofP. subricta)weidentifiedthespeciesasbeingasubadultspecimenofP. luteola. From Goeteborg we borrowed a sample of P. subricta, which contained one specimen of P. dupotetiana
andonejuvenileofP. luteola.Astheoriginaldescription
correspondstojuvenilesofP. luteola,webelievethatP.
subrictaisayoungersynonymofP. luteola.
Pseudamnicola numidica (CLESSIN1878)
Figs7,26–29
1878 Amnicola numidica CLESSIN,MalakozoologischeBlätter, 25: 118–119, Pl. 5, Fig. 4a–b), loc. typ.: “Hab.
Algeria(coll.WESTERLUND)”.
10
Figs30–31.Pseudamnicola letourneuxiana.Djendell,syntypes.—Figs32–33.Pseudamnicola constantinae.32:topotype,33:
SyntypeMHNG5278).—Figs34–37.Pseudamnicola boucheti n.sp. 34–35:paratypes,36:penis,37:headwithpenis.–o=operculum,s=snout.
M a t e r i a l e x a m i n e d :Thesyntypescouldnotbefound
in CLESSIN’s collection (Natural History Museum Stuttgart,
Niederhöfer in litt.), neither inWESTERLUND’s collection (VON
PROSCHWITZ, Goeteborg, in litt, and SINDEMARK KRONESTEDT,
Stockholm,inlitt.).Buttheoriginaldescriptionandthedrawings of the shells are suitable for determination.—MNHN:
Constantine(numerous;SMF334676/many);M’Sila(20ex.);
ElKhroub(numerous);AïnM’Daourouch[afountaininSouk
Ahrasregion]Bouzidleg.(numerous).
1862 Bythinia letourneuxiana BOURGUIGNAT, Les Spicilèges
Malacologiques:121,loc.typ.:“Habite,enAlgérie,les
sourcesthermales(42degrés)deDjenndel”.
1862 Bythinia desertorum BOURGUIGNAT,LesSpicilègesMalacologiques:118.
1870 Amnicola servainiana LETOURNEUX,Annales de Malacologie,1[1870–1884]:318.
M a t e r i a l e x a m i n e d :The syntypes exist in BOUR-
GUIGNAT ’s collection in Paris (MNHN 22977). The photo-
D e s c r i p t i o n : shell spherical to oval, dark horncoloured,surfacelustrelessandfinelystriated,apexobtuse, 4–4.5slightly convex whorls with a clear suture,
umbilicusclosed,spiresmall,apertureovalandnotangledwithasharpperiostome,shellheight2.9–3.2.mm,
shellwidth2.2–2.3mm.
P e n i s m o r p h o l o g y : penis triangularly, broad
attheproximalpartandslimroundedatthedistalpart
(Fig.29)
D i s t r i b u t i o n :EasternpartoftheAlgerianTell.
graphed syntype shell (Fig. 30) has a shell height of 2.2mm
and width of 1.7mm.The second syntype is perforated (Fig.
31) but both looked uniform.—MNHN: La Cheffia [Cheffia
near Bône] (2samples: 6ex., 17ex.); Bône (numerous; SMF
334673/27).—ThesyntypesofAmnicola servainianacouldbe
found in BOURGUIGNAT’s collection in Geneva: MHNG 10702
(lac desArbaouam (Senhadja), identified by BOETERS 1970,
2ex.,partofMHNG5373);MHNG5371(3ex.),AithOuaban
(Jurjura);MHNG5372(numerous),HammamSidiDjaballah;
MHNG 5373 (numerous), lac desArbaouam (Senhadja).The
lotsofA. servainiana,mentionedabove,havebeenidentified
byusasP. letourneuxiana.
Pseudamnicola letourneuxiana (BOURGUIGNAT1862)
D e s c r i p t i o n :shellconicallyoval,darkhorn-coloured,surfacesilky,apexobtuse,4–5regularlyincreas-
Figs20,30–31
ing whorls with a clear suture, umbilicus closed, spire
0.3of shell height, aperture oval, oblique, periostome
sharp,shellheight2.2–2.6mm,shellwidth1.4–1.5mm.
D i s t r i b u t i o n :Thisspeciesispossiblyconfinedto
thermalspringsaroundtheregionofAnnaba.
R e m a r k :The syntypes of P. desertorum exist in
BOURGUIGNAT’s collection in Geneva.The type locality
isthesameasforP. letourneuxiana,thusweinterpretP.
desertorumtobeanaberrantformoftheother.
Pseudamnicola constantinae (Letourneux1870)
Figs10,32–33
1870 Amnicola constantinae LETOURNEUX,Annales de Malacologie,1[1870–1884]:314,loc.typ.:“Habitedans
des Hammam à 32degrès, et dans le cours du BouMerzouk,prèsdeConstantine”.
11
Pseudamnicola boucheti n.sp.
Figs18,34–37
H o l o t y p e :Shell3.0mmhigh,2.4mmwide,Batna(Fontainechaude,25.III.1932),MNHN21440,Fig.18.—P a r a t y p e s :MNHN21441(numerous)fromtypelocality;MNHN
21910(37ex.),AïnM’lila;MNHN21915(St.Donat,18ex.,
Dr F.G. Marill 1953), Tadjenanet; MNHN 21914 (3ex.),
Boufarik; MNHN 21911 (3ex.), unknown Hammam; Batna
(4samples, Fontaine chaude, 3x numerous, M.A. Diezede
1932,SMF334666/54,MNHN21909,MNHN2191124ex.);
MNHN21912(numerous),OuedElAneb;Algiers(2samples:
MNHN 21916 (1ex.) Dr F.G. Marill 1956, MNHN 21917
(42ex.)coll.Jousseaume);MNHN21913(30ex.),mareroute
Kairouan à Enfida (Tunisia); MNHN 21918 (3ex.), Constantinecoll.Staadt1969;unknownsamplingsite(numerous).
E t y m o l o g y : Named in honour of PHILIPPE BOUCHET,
who borrowed us numerous samples from the Muséum Nationald’HistoireNaturelle,Paris.
D e s c r i p t i o n : shell elongated oval, yellowish
horn-coloured, surface silky and finely striated, apex
small,4.5–5slightlyconvexwhorlswithaweaksuture,
umbilicus closed, spire height 0.25of shell height, aperturetriangularrounded,periostomethick,shellheight
3.4–4.1mm,shellwidth2.4–2.6mm.
D i s t r i b u t i o n :Possiblyendemic.
D e s c r i p t i o n : shell conical oval, light horn-coloured, opaque, surface silky, apex small rounded, 4–
4.5slightlyconvexwhorlswithaclearsuture,umbilicus
slit-like, spire regularly increasing, spire height 0.25of
shell height, aperture oval and not angled, periostome
sharp,shellheight2.7–3.0mm,shellwidth2.3–2.4mm.
P e n i s m o r p h o l o g y :penissimplewith17folds
(Figs36–37).
D i f f e r e n t i a t i n g f e a t u r e s :The conical oval
shellis2.7–3.0mmhighandthesurfaceissilky.Other
Pseudamnicolaspp.whicharealsoconicaloval,andof
similarsize,haveaglossysurface.
D i s t r i b u t i o n : Eastern and central parts of the
Tell,easternHautsPlateaux.
Pseudamnicola meluzzi BOETERS1976
Pseudamnicola chabiin.sp.
M a t e r i a l e x a m i n e d : The syntypes exist in BOURGUIG-
NAT’s collection in Geneva: MHNG 5276 (19ex.), cours du
Bou-Merzouk;MHNG5277(=var.minor)(numerous),BouMerzouk; MHNG 5278 (3ex.), Hammam à 32 degrès.—
MNHN: Sources du Boumerzoug (Ouled Rahmoun) (topotypes)(numerous;SMF334668/16).
Fig.8
Fig.9
1976 Pseudamnicola meluzzi B OETERS , Archiv für Molluskenkunde, 107: 94, Figs 2–3and 12–13, loc. typ.:
“ineinemFluß,undzwarOuedMelahunweitderStationAonanaanderStraßezwischenMateurundSedjenane”.
M a t e r i a l e x a m i n e d :MNHN:Constantine(2series:
10ex., numerous); Sidi M’Cid, thermal spring (numerous,
SMF334675/35);Annaba(1ex.coll.WESTERLUND).
D e s c r i p t i o n :shellconicallyoval,lighthorn-coloured, thin, opaque, surface silky, apex rounded, 3.5–
4slightlyconvexwhorlswithaweaksuture,umbilicus
closed to slit like, spireheight 0.25of shell height, aperture triangular rounded, periostome sharp, operculum
horn-coloured, shell height 2.1–2.6mm, shell width
1.8–2.0mm.
D i s t r i b u t i o n :fromNEAlgeriatoNW-Tunisia.
R e m a r k : InWESTERLUND’s collection we found a
juvenilespecimenofthisspeciescollectedinAnnababut
determinedasP. subricta.
H o l o t y p e : Shell 2.4mm high, 1.8mm wide; Batna
(Fontaine chaude), MNHN 21442, Fig. 9. — P a r a t y p e s :
MNHN21443(7ex.)fromtypelocality;MNHN21896,SMF
334667/many, Source du Boumerzoug (= Ouled Rahmoun,
2lots,numerous).
E t y m o l o g y :This new species is named in honour of
YASSINE CHABI,adedicatedornithologistinAlgeria,whounfortunatelypassedawaysoearly.
D e s c r i p t i o n : shell conical oval, horn-coloured,
surfacesilky,apexsmall,4veryslightlyroundedwhorls,
suture flat, umbilicus closed, spire short, aperture triangularly rounded, periostome very thick, shell height
2.3–2.4mm,shellwidth1.8mm.
D i f f e r e n t i a t i n g f e a t u r e s :The conical shell
hasacharacteristicaperturewhichistriangularlyrounded.ItdiffersfromP. meluzzibyitsthickwalledshelland
theverythickperiostome.
A s s o c i a t e d s p e c i e s :Maresiasp.(numerousin
bothsamples).
D i s t r i b u t i o n :EasternHautsPlateaux.
12
Pseudamnicola ghamiziin.sp.
Figs19,38–39
H o l o t y p e : Shell 3.4mm high, 2.7mm wide, Cap de
Garde, MNHN 21444, Fig. 19.—P a r a t y p e s : MNHN
21445(25ex.)SMF334672/6,CapdeGarde;MNHN21900
(15ex.),OuedAthmenia;MNHN21901(7ex.),Alger;MNHN
21902 (6ex.),Algiers, source face à l'Institut Pasteur, champ
demoneuvre.
E t y m o l o g y : Named in honour of our colleague MOHAMED GHAMIZI,whocontributedgreatlytoourknowledgeof
Moroccanaquaticmalacofauna.
D e s c r i p t i o n :shellconicallyoval,lighthorn-coloured, surface glossy, apex small rounded, 4.5slightly
rounded and regularly increasing whorls, suture clear,
umbilicus opened to slit-like, spire conical, aperture
oval and not angled, periostome sharp and at the columella straight, shell height 2.7–3.4mm, shell width
2.4–2.7mm.
D i f f e r e n t i a t i n g f e a t u r e s :The conical oval
shell is about 4.0mm high and the surface is glossy.
Figs 38–39. Pseudamnicola ghamizii n. sp., paratypes, 38:
champdemanoeuvre,39:CapdeGarde.
OtherPseudamnicolaspp.whichhavealsoaglossysurfacearesmaller.
D i s t r i b u t i o n : Central and eastern parts of the
Tell.
Figs40–43.ThesamplingsitesofthePseudamnicolaspp.fromtheGuelmaregion.—40:AïnZaaroura(typelocalityofP. calamensis n.sp.).41:AïnFeïd-El-Bagrat(typelocalityofP. fineti n.sp.).42:AïnDamous(additionalsamplingsiteofP. fineti n.sp.),
43:AïnH’djar(typelocalityofP. linae n.sp.).
Pseudamnicola algeriensis n.sp.
Fig.14
H o l o t y p e : Shell 4.3mm high, 3.0mm wide,Tlemcen,
Algeria, MNHN 21446, Fig. 14.— P a r a t y p e s : MNHN
21447 (31ex.), SMF 334665/10, Tlemcen; MNHN 21898
(9ex.),“fontaineentreDayaetTélag“.
E t y m o l o g y :Namedafterthecountrywherethespecies
lives.
D e s c r i p t i o n :shelloval,darkhorn-coloured,surface
silky,apexsmall,4–5slightlyroundedandsteppedwhorls
thatincreaseregularly,umbilicusclosed,spireheight0.4of
shellheight,apertureovalandslightlyangled,periostome
thick,shellheight3.9–4.3mm,shellwidth2.8–3.3mm.
D i f f e r e n t i a t i n g f e a t u r e s :Thewhorlsofthe
shell are stepped and not rounded like inP. luteola. In
additiontheshellissmaller(about4mm)thantheshell
ofP. luteola(about5mm).
D i s t r i b u t i o n :westernpartoftheTellandwesternHautsPlateaux.
Pseudamnicola gerhardfalkneri n.sp.
Fig.11
H o l o t y p e : Shell 2.8mm high, 1.9mm wide,Aïn Sultan (coll. Denis), MNHN 21448, Fig. 11.—P a r a t y p e s :
MNHN 21449 (numerous), SMF 334671/39, Aïn Sultan;
MNHN21897(8ex.coll.H.Fischer),AïnSultan.
E t y m o l o g y : Named after G ERHARD F ALKNER who
foundthemanylotsinthecollectionoftheMuséumNational
d’HistoireNaturelle,Paris,forourstudies.
D e s c r i p t i o n : shell elongated oval, slim, colour
lightbrown,opaque,surfacefinelystriated,glossy,apex
small, 5–5.5convex whorls with a deep suture, umbilicus slit-like, spire conical, spire height 0.3of shell
height,apertureoval,slightlyoblique,periostomesharp,
shellheight2.7–2.8mm,shellwidth1.9mm.
D i f f e r e n t i a t i n g f e a t u r e s :Theshelliselongatedovalandslimvs.conicalorsphaerical.
D i s t r i b u t i o n : This species is restricted to the
centralpartoftheTellianAtlas.
Pseudamnicola calamensis n. sp.
Figs12,44–47,52–54,60–61
H o l o t y p e : shell heigth 2.8mm, width 2.0mm, Aïn
Zaaroura [at Medjez Sfa, Guelma], 3625.906'N 0747.688'E,
237.74m alt., leg. S. BOUZID, 27.08.2006, ZMH 51388, Figs
12(typelocality),40.—P a r a t y p e s :ZMH51389(3ex.in
ethanol),SMF334682/68,GLÖ(18ex.),BOE3058(31ex.),
BOE3059(18ex.)fromtypelocality.
H a b i t a t :Naturalspring.
E t y m o l o g y : Calama was the name of Guelma during
theRomanera.
D e s c r i p t i o n :The brownish shell has 4.5–5convex whorls which increase regularly, the body whorl
13
dominates and is 3.5times higher than the spire; apex
blunt, suture deep, surface glossy; aperture oval at the
top moderately angled, broad at the basis, in adults reflected from the body whorl, peristome continuous;
horny operculum transparent colourless, thin; shell
height2.7–3.0mm,width2.0–2.1mm.
A n i m a l : mantle black, snout long, slim and crenated;penislongwithupto14penialfolds.
Anatomy: Bursa duct slightly curved, receptaculum
thumb-shaped.
D i f f e r e n t i a t i n g f e a t u r e s :Thespecies’shell
issimilartoP. linaen.sp.,P. finetin.sp.,andP. rouagi
n. sp.These species can be distinguished by the renal
oviduct:threesmallloopsinP. calamensis.
D i s t r i b u t i o n :Onlyknownfromthetypelocality.
R e m a r k s :The population contains also infected
♀♀,whichdevelopapseudopenis.
Pseudamnicola finetin.sp.
Figs17,48–51,55–56,62–63
H o l o t y p e :shellheigth3.2mm,width2.3mm,AïnFeïdEl-Bagrat[atMedjezSfa,Guelma],36°25.555'N07°51.386'E),
323.7malt.,leg.S. BOUZID,27.08.2006,ZMH51390,Figs17
(type locality).—P a r a t y p e s : ZMH 51391 (5ex. in ethanol),SMF334683/43,GLÖ(24ex.),BOE(xxex.)fromtype
locality; GLÖ (6ex.)Aïn Damous [at Medjez Sfa, Guelma],
36°25.350'N07°51.367'E,523.34malt.
H a b i t a t :Naturalsprings.
E t y m o l o g y :NamedafterYVES FINET,whoborrowedus
manymaterialofBOURGUIGNAT’scollection.
D e s c r i p t i o n :Thehorn-colouredshelliselongated
ovalandhas4.5–5convex,regularlyincreasingwhorls
with a deep suture, the last two whorls are prominent;
apexblunt,surfaceglossy;apertureovallyrounded,peristome continuous, umbilicus closed; horny operculum
transparent colourless, shell height 3.5–3.9mm, width
2.4–2.5mm.
A n i m a l :mantleblackwithanarrowwhiteborder,
penisbroadatthebasisandslimatthedistalend,upto
15penialfolds.
A n a t o m y :Bursaducthardtortuous,receptaculum
digitiform,which,however,doesnotfullyreachtheloop
oftherenaloviduct.
oviduct:onesmallloopinP. fineti.
D i s t r i b u t i o n :Onlyknownfromthetypelocality
andfromAïnDamous(atMedjezSfa,Guelma).
Pseudamnicola linae n.sp.
Figs13,67–70,57,64
H o l o t y p e : shell height 3.3mm, width 2.2mm,Aïn
H’djar [at Medjez Sfa, Guelma], 36°25.806'N 07°47.708'E,
256.03m alt., leg. S. BOUZID, 27.08.2006, ZMH 51394, Figs
14
Figs44–47.Pseudamnicola calamensisn.sp.44–45:shells(paratypes);46:renaloviduct;47:Headwithpenis.–e=eye,o=operculum,p=penis,s=snout.—Figs48–51.Pseudamnicola fineti n.sp.48–49:paratypes,50:renaloviduct,51:Headwithpenis(p).
13(typelocality),43.—P a r a t y p e s :ZMH51395(5ex.in
ethanol),SMF334684/142,GLÖ(156ex),BOE3062(32ex.)
fromtypelocality.
H a b i t a t :Naturalspring.
E t y m o l o g y :NamedafterLINA BOUZID,first-borndaughterofthesecondauthor.
D e s c r i p t i o n :Thebrownishshellisconicallyoval
andhas4.25–4.5slightlyconvexwhorlswhichincrease
regularly, the body whorl dominates and is 3.5times
higher than the spire; apex blunt, suture deep, surface
glossy;apertureovalatthetopmoderatelyangled,peristome continuous and sharp; horny operculum transparent colourless; shell height 3.3–3.5mm, width 2.2–
2.5mm.
A n i m a l : mantle black, snout broad and crenated;
penislongwithupto10penialfolds.
A n a t o m y :Bursaductslightlycurved,thestraight
finger-shapedreceptaculumisattachedbesidetherenal
oviductatthebursa.
oviduct:onesmallandonelargeloopinP. linae.
Pseudamnicola rouagin.sp.
Figs16,71–73
H o l o t y p e :shellheight3.1mm,width2.3mm,AïnBendaoud[atOuledDriss,SoukAhras],36°22.373'N08°04.402'E,
1118malt.,ZMH51392,Fig.16.—Paratypes:ZMH51393(5ex.
inethanol),SMF334681/5,GLÖ(58ex.)fromtypelocality.
H a b i t a t :Naturalfountain.
E t y m o l o g y : Named for RACHID ROUAG, promisingAlgerianherpetologist.
D e s c r i p t i o n :Thedarkhorn-colouredshellisconically oval and has 4.5slightly rounded whorls with a
cleartoweaksuture,thebodywhorlis3–4timeshigher
thanthespire;apexblunt,surfaceglossyandfinelystriated; aperture oval, peristome continuous and sharp;
horny operculum transparent colourless; shell height
2.7–3.1mm,width2.2–2.3mm.
A n i m a l : mantle greyish with a white broad border,snoutbroadandslightlycrenated;penislongwitha
roundedpenistip.
A n a t o m y :Oviductlooplarge,attheproximalpart
tapered.
issimilartoP. linaen.sp.,P. finetin.sp.,andP. rouagin.sp.
15
Figs52–66.52–58:ShellsofPseudamnicolaspp.fromtheregionofGuelma.52–54:P. calamensis;52:thesameex.asFig.58,
53:thesameex.asFig.59,54:thesameex.asFigs60–61.55–56:P. fineti n.sp.55:thesameex.asFig.62;56:thesameex.as
Fig.63.57:P. linaen.sp.,thesameex.asFig.64.58:Pseudamnicolasp.,thesameex.asFigs65–66,seediscussion.—58–66:
AnatomyofPseudamnicolaspp.fromtheregionofGuelma.58–59:P. calamensis.60–61:P. calamensisinfected♀;60:pseudopenis;61:mantlecavityandkidneycutopenandgillandrenaloviductuncovered.62–63:P. fineti n.sp.;62:AïnDamous;63:Aïn
Feid-El-Bagrat.64:P. linae n.sp.65–66:Pseudamnicola sp.66:infected♀withpseudopenis,seediscussion.—Abbreviations:
bc=bursacopulatrix,dm=wallofthemantlecavity,gi=gill,pp=pseudopenis,ro=renaloviduct,rs=receptaculumseminis.
(OriginaldrawingbyH.D.BOETERS).Allillustratedspecimensareparatypesfromtypelocality.
16
Figs67–70.Pseudamnicolalinaen.sp.67-68:paratypes,69:renaloviduct,70:Headwithpenis(p).—Figs71–73.Pseudamnicola rouagin.sp. 71:shell(paratype),72:renaloviduct,73:headwithpenis(p).
Thesespeciescanbedistinguishedbytherenaloviduct:one
largeloop,taperedattheproximalpartinP. rouagi n.sp.
Genus Mercuria BOETERS1971
T y p e s p e c i e s : Cyclostoma simile Draparnaud
1805.
The shell is whitish, especially in the region of the
umbilicus.The penis posses a large and flat triangular
appendix.
Identificationkey
1.Shelllargerthan4mm,elongatedconical................ 3
1'. Shellovaltospherical,smallerthan4mm .............. 2
2. Shell spherical, short spire 0.2–0.3of shell height,
shellheight2.4–3.0mm................M. globulina
2'.Shelloval,shellheightupto3mm
........................................M. bourguignati n.sp.
3. Shell whitish, solid, 6–6.5whorls, aperture slightly
angled,apertureheight:shellheightabout0.6
................................................. M. pycnocheilia
3'. Shell solid, yellowish to whitish, opaque, aperture
rounded ........................................................... 4
4. Shell large, > 5mm, aperture height: shell height
about0.5 ........................................M. saharica
4'. Shell smaller, < 5mm, aperture height: shell height
about0.4 ...............................M. gauthieri n.sp.
Mercuria pycnocheilia (BOURGUIGNAT1862)
Figs74,79–82
1862 Bythinia pycnocheilia BOURGUIGNAT, Les Spicilèges
Malacologiques:117,loc.typ.:„EspècecommuneàTemascin[=Temacin],prèsdeTuggort[=Touggourt]“.
M a t e r i a l e x a m i n e d : The syntypes could be found
inBOURGUIGNAT’scollectioninGeneva:MHNG5351(40ex.),
Temascin,prèsdeTuggort.—MNHN:Ouargla(43ex.);Ferkane(34ex.);Oasisd’OurirentreBiskraetTouggourt(numerous; SMF 334680/23); Bahr deTiguedidine près de Djamaa
(22ex.);Negrine(18ex.).
D e s c r i p t i o n :Shellsolidandconical,colourwhitishopaque,surfacesmoothandglossy,Apex:smalland
acute,6–6.5slightlyroundedwhorlswithadeepsuture,
17
Figs74–78.Mercuriaspp.fromAlgeria.—74:Mercuria pycnocheilia,Temascin,prèsdeTuggort,syntype.75:Mercuria gauthieri
n.sp.,Nemours,holotype.76:Mercuria saharica,SidiRached.77:Mercuria globulina,Boufarik.78:Mercuria bourguignati n.
sp.,Boghari,holotype.
whorlsincreasewithconstantrates, umbilicussmallbut
openedanddeep,spire0.3ofshellheight,apertureoval
andangled,notcutbythebodywhorl,periostomethickened,shellheight3.8–4.9mm,shellwidth2.8–3.5mm
D i s t r i b u t i o n : Sahara depression and adjascent
areas.
R e m a r k :Theshellshapeisalittlesimilartothatof
M. gauthierin.sp.,butthespiresaredistinct:sliminM.
pycnocheiliaandbroadinM. gauthieri n.sp.Inaddition
bothspeciesliveindistinctzoogeographicalregionsof
Algeria.
label) El Kis, près de Feriana,Tunisie.—MNHN: Boufarik
(4samples: 26ex.,3xnumerous;SMF334679/many);Algiers
(numerous);Zardezas(35ex.);2unknownsamplingsites(2x
numerous).
D e s c r i p t i o n : Shell conically oval with a prominentbodywhorl,colourwhitishopaque,surfacesmooth
and glossy, apex small, 4.5–5whorls rapidly increasingwithadeepsuture,umbilicusopenbutsmall,spire
height 0.2–0.3of shell height, aperture oval rounded,
periostome thick, shell height 2.5–3.0mm, shell width
2.3–2.5mm.
D i s t r i b u t i o n : Central and eastern parts of the
Tell.
Mercuria globulina (LETOURNEUX & BOURGUIGNAT1887)
Figs77,83–84
1887 Amnicola globulina LETOURNEUX & BOURGUIGNAT,Prodrome de la Malacologie terrestre et fluviatile de La
Tunisie:147,loc.typ.:„Tala,El-Kis,prèsdeFeriana,
ainsi que dans le bassin du temple du Djebel, Zaghouan.“
M a t e r i a l e x a m i n e d :syntypesMHNG5307(numerousspecimensfromcoll.BourguignatinMHNG,withoriginal
Mercuria saharica (BOURGUIGNAT1887)
Figs76,85–88
1887 Amnicola saharica BOURGUIGNAT, In: LETOURNEUX &
BOURGUIGNAT,ProdromedelaMalacologieterrestreet
fluviatile de LaTunisie: 144, loc. typ.: “Sidi Sliman;
etc.”.
1887 Amnicola subscalaris LETOURNEUX & BOURGUIGNAT,
Prodrome de la Malacologie terrestre et fluviatile de
18
Figs79–82.Mercuria pycnocheilia.79–81:shellsofOuargla,82:spireof79.—Figs83–84.Mercuria globulina.83:Boufarik
(aberrantshell);84:syntype(MHNG5307).—Figs85–88.1-2:Mercuria saharica.85:SidiSlimane(syntypeMHNG5370);86:
marekm139routeKairouanàEnfida;87:OuedGabès(MHNG5275);88:SidiMakhlouf.
LaTunisie:144,loc.typ.:“Algérie,àMokta-el-Oued,
entreDjelfaetBoghar;Tunisie,OuedGabèsàGabès”.
1887 Amnicola bythinopsis LETOURNEUX & BOURGUIGNAT,
ProdromedelaMalacologieterrestreetfluviatiledeLa
Tunisie:144,loc.typ.:“dansleseauxdesenvironsde
Gabès”.
M a t e r i a l e x a m i n e d : syntypes saharica: MHNG
5365 (numerous) Ruisseau de la fontaine chaude de Biskra,
Algérie, with original label; MHNG 5366 (numerous) Puits
artésiendeM'Zaiez(=Meghaïer),entreBiskraetTuggurt,Algérie,withoriginallabel;MHNG5367(3ex.)Fontainechaude
duruisseaudesZaatcha,dansleZaboccidental,Algérie,with
originallabel;MHNG5368(7ex.)OuedGabèsàGabès,Tunisie,withoriginallabel;MHNG5369(numerous)puitsartésien
deSidi-Rached(OuedPriz[=OuedRir]),withoriginallabel,
MHNG 5370 (numerous, with original label), puits artesiens
de Sidi Sliman (Oued Rir = Sidi Rached).—The following
samples are labelled as A. saharica (see remark): MHNG
5455(2ex.)environsdePalerme,Sicile,specimensfromcoll.
BOURGUIGNATinMHNG,withoriginallabel.[=Mercuriasp.];
MHNG5456(2ex.)fleuveOreto,prèsdePalerme,Sicilespecimensfromcoll.BOURGUIGNATwithoriginallabel[=Mercuria
sp.].—syntypesubscalaris:MHNG5386(1ex.)OuedGabès
19
Figs89–90.Mercuria bourguignatin.sp.89–90:shells,paratypes.—Figs91–94.Mercuria gauthierin.sp.91:holotype;92–93:
shellsofparatypes,Nemours;94:spireof91.
à Gabès,Tunisie, with original label [theAlgerian specimens
are not present in MHNG].—syntypes bythinopsis: MHNG
5275 (2ex.) Oued Gabès.—MHNG 5274 (1ex.) Mokta El
Oued, entre Djelfa et Boughar [this lot is named bythinopsis
, however, this is the “type” locality of subscalaris and very
probablyrepresentsthemissinglot;thisisprobablyanoriginal
confusionbyBOURGUIGNAThimself].—allfollowinglotsfrom
MNHN:SidiMakhlouf(numerous);marerouteKairouanàEnfida(Tunisia)(8ex.);unknownpuitsartesiens(16ex.);Algiers
(numerous).
D e s c r i p t i o n :shellelongatedconical,surfacefinelystriatedandglossy,apexacute,5.5–6whorlsslightly
roundedandrapidlyincreasingwithadeepsuture,whitishopaque,umbilicusclosedtoslit-like,apertureheight
about 0.4–0.45of shell height, aperture oval rounded,
periostomethick,shellheight5.5–6.2mm,shellwidth=
3.0–3.6mm.
D i s t r i b u t i o n : Sahara depression and adjacent
areas,centralHautsPlateauxandcoastalregionsofAlgeriaandTunisia..
R e m a r k :GAUTHIER(1928:66)mentionedthisspeciesfrom“l’OuedGabès,àRas-el-Oued...dansl’Oued
Béchar”,too.NexttothelocalityinAlgeria(originally
consideredas“typelocality”),LETOURNEUX & BOURGUIGNAT(1887:144)mentionedA. subscalarisalsofromTunisia“OuedGabès,àGabès”;inparallel,theydescribed
A. bythinopsis from the same are inTunisia.The only
difference between the shells of both species is in the
shapeoftheaperture.BecauseaberrantaperturesinMercuria spp. are not unusual (see Mercuria bourguignati
n.sp.,Figs16.5and20.1),webelievethatbothspecies
areconspecific.LETOURNEUX & BOURGUIGNAT(1887:145)
mentioned M. saharica species also from l’Oreto near
Palermo (Sicily). GIUSTI & PEZZOLI (1980) did not list
this species from Italy, but the shells of BOURGUIGNAT’s
collection (MHNG 5455 and MHNG 5456), both samples collected near Palermo, are indeed very similar to
M. saharica.
Mercuria bourguignati n.sp.
Figs78,89–90
H o l o t y p e :shellheight2.9mm,width2.0mm,Ksar el
Boukhari, MNHN 21450, Fig. 78.—P a r a t y p e s :MNHN
21451(2lots,numerous),SMF334677/55,KsarElBoukhari(=
Boghari);MNHN21907(38ex.),FoukaprèsdeKolea;MNHN
21905(3samples:19ex.,2xnumerous);MNHN21903(18ex.),
Boufarik,Algiers(HôpitalduDey);MNHN21904(numerous)
MoktaAl-Hadid(coll.Jousseaume),MNHN21906(numerous),
Dellys;MNHN21908(3ex.),unknownsamplingsite.
E t y m o l o g y :NamedafterJULES-RENÉ BOURGUIGNATinappreciationofhisworkonthecontinentalmolluscfaunaofAlgeria.
D e s c r i p t i o n :shellelongatedoval,colourwhitish
opaque, the juveniles are spherical, surface finely striatedandglossy,apexblunt,whorlsslightlyroundedwith
aflattoclearsuture,umbilicusopenedbutsmall,spire
20
height0.3ofshellheight,apertureoval,peristomesharp
andflangedatthecolumella,shellheight2.9mm,shell
width2.0mm.
D i f f e r e n t i a t i n g f e a t u r e s : Shell elongated
oval and not spherical like M. globulina.The shell is
smaller(about3mm)thantheotherMercuriaspp.(about
4–6mm).
D i s t r i b u t i o n :EasternandcentralTellandcentral
HautsPlateaux.
Mercuria gauthieri n.sp.
Figs75,91–94
H o l o t y p e : shell height 4.6mm, width 3.2mm,prèsde
Nemours [= Ghazaouet], MNHN 21452, Fig. 91.—P a r a t y p e s :MNHN21453(numerous),SMF334678/37fromthe
typelocality;MNHN21899(40ex.),Nemours.
E t h y m o l o g y :Named,inmemoriam,forHENRI GAUTH-
IER, in recognition of his extensive work on the continental
aquaticfaunaofAlgeriaandTunisia.
D e s c r i p t i o n :shellsolidandconical,colourwhitish opaque, surface smooth and glossy, apex rounded,
4.5–5slightlyroundedwhorlswithaclearsuture,whorls
regularly increasing, umbilicus small but opened, spire
0.2–0.3ofshellheight,apertureovalandangled,notcut
by the body whorl, periostome thickened, shell height
3.5–4.7mm,shellwidth2.5–3.1mm.
D i f f e r e n t i a t i n g f e a t u r e s :Theshellshapeis
alittlesimilartothatofM. pycnocheilia,butthespires
are distinct: slim in M. pycnocheilia and broad in M.
gauthieri n.sp.Inadditionbothspeciesliveindistinct
zoogeographicalregionsofAlgeria.
D i s t r i b u t i o n :Thisspeciesisapparentlyconfined
tothewesternmostpartoftheAlgeriancoast.
Discussion
Itisnotpossibletodecideonlybymeansofcontemplatingwhethertheshellsofaparticulartaxonrepresents
aspeciesorasubspecies,butasmostofthetaxathatwe
foundwerenotrestrictedintheirdistribution,wecanexcludegeographicalraces.Forthisreasonweconsiderall
thesetaxafoundbyusasbeingtaxaofspeciesrank.
However, identifying the oldest name for a given
species may cause some problems, because in former
timesmanyspeciesofthegenusunderdiscussionhave
beendescribedasAmnicola,Hydrobia,Paludinella,and
Bithynia,andonlyafewoftheoldnameshavealready
beenidentifieduntilnow.
BOURGUIGNAT (1864: 245) misinterpreted the species
descriptionofPseudamnicola dupotetiana (FORBES1839)
and recognised a high degree of variability concerning
the shell size (3–5mm).The drawing he provided (his
pl.14,Fig.31)hasnothingincommonwiththeoriginal
drawing by FORBES (1839) (Fig. 95).All other authors
uptonow,e.g.WESTERLUND(1886:79),PALLARY(1898:
133), and GHAMIZI et al. (1997) followed him. GHAMIZI
et al. (1997) designated a neotype for P. dupotetiana,
butusedthisnamealsoinawidesenseconcerningthe
shellshape(theirFigs5–7,p.43)andpenismorphology
(theirFigs20–28,p.45).Theneotypecorrespondstothe
originaldescriptionaswellastoourspeciesdefinition,
buttheadditionallyillustratedspecimens(theirFigs5–7,
p.43)are,inouropinion,distinctfromP. dupotetiana.
The misidentification of Pseudamnicola dupotetiana
by BOURGUIGNAT (1862, 1864) and other malacologists
possiblyhappenedbecauseMercuria globulinainhabits
thesameregion,whichhasasimilarshellshapeifcomparedtoP. dupotetiana,butdiffersbyahigherdegreeof
variability,anduntilthedescriptionofthegenusMercu-
ria,bothspeciescouldnotbedistinguished.Inaddition,
juvenilesofmanyPseudamnicolaspp.aresimilartoP.
dupotentiana, because their shells are more spherical
than the shells of the adults.This means, if two Pseudamnicola/Mercuriaspp.occurinsympatricconditions,
parallel existence of differing species does not become
obviousatthefirstglancewithoutcountingthenumber
ofwhorls.
BOETERS(1988)alreadystatedthatinrepresentatives
ofPseudamnicola,theductofthebursa,i.e.thepedunculus,canbecurvedorevenbetortuous.Thus,BOETERS
(1988)differentiatedforexampleP. gasulli BOETERS,1981
(BOETERS1988:204figs78–80)bymeansofitscurved
ductfromP. spirata (PALADILHE1869)(loc.cit.:204figs
81–84)withitstortuousduct.Bothspeciesoriginatefrom
the Iberian peninsula. Further, for P. meluzzi BOETERS,
1976(BOETERS1976:96Fig.3)hedescribedtheductto
becurved,yetfor P. conovula (FRAUENFELD1863)(loc.
cit.:96Fig.6andfigs5and7,resp.),bothfromTunisia.
However,accordingtoourpresentunderstandingitcannotbeexcludedthatthematerialasidentifiedbyBOETERS
(1976)asP. conovulamighthavecomprisedmorethan
onesinglespecies;asregardsarevisedinterpretationofP.
conovulaseeforexampleFALKNER & BOETERS(2003:23).
Itmaybeargued,ofcourse,thatthetorsionoftheoviduct
isduetotheextremeretreatoftheanimalduringpreservationinethanol.However,evenunderthisassumption
thefactremainsthatonewayofkillingwillcauseatorsionoftheoviductwhileanotherwillnot.Inadditiona
tortuous duct resulted by fixation cannot be a constant
characterforaspecies,aswefounditinoursamples.
The three species from the Guelma region (P. calamensis n. sp., P. fineti n. sp., and P. linae n. sp.) differ
21
Fig.95.FaksimileoftheoriginaldescriptionofPaludina dupotetiana FORBES1839,andFig.3ofplate12.
onlyslightlyintheirshellshape,theirpenismorphology,
theirbursaduct,andtheirrenaloviduct.Butthedifferencesintherenaloviductareconstant,sothesespecies
can be interpreted as being distinct. Many species describedhererevealmoredifferencesintheirshellshape
thanthosefromGuelmaregion.
One species of Pseudamnicola (sampling site:Aïn
Kef Kourrath, at Medjez Sfa, Guelma, 36°26.395'N
07°51.996'E, 384.05m alt., Figs 58, 65–66) could not
be identified because all specimens were infected, and
allfemalesbearedapseudopenis.Wedonotknowifthe
Oviductloop,whichdifferedfromthatoftheotherspeciesinthisregion,hasbecomeaberrantbytheparasites,
but the shell sizes seems not to be influenced, as suggestedbySZAROWSKAetal.(2006).
Someofthespeciesmentionedinthispaperarewidely distributed inAlgeria, while others, especially those
that live in thermal springs, are obviously restricted in
theirdistributionrange.InothercountriessuchasSpain
for example (see BOETERS 1988), the distribution areas
of Pseudamnicola spp. are smaller.Additionally, speciesrichnessofPseudamnicolaspp.andMercuriaspp.
inAlgeria is much larger. Because the number of specieslistedinachecklistdependsnotonlyontheactual
diversity, but also on the thoroughness of investigation
andonthetaxonomicalexpertise.Inourcase,thegreat
numberofsamplesstudiedbyusmayalsohavecontributedtothespeciesdiversityreportedhere.Itshouldbe
stressedthatAlgeria,especiallyEasternAlgeria,isahot
spotofPseudamnicolaspeciation.
Acknowledgements
WewouldliketoexpressourthankstoYvesFinet(Muséum
d’Histoire Naturelle, Geneva) for the loan of the syntypes
and many other lots of BOURGUIGNAT’s collection, toTED VON
PROSCHWITZ (Natural History Museum Goeteborg), for sendingspecimensofPseudamnicola subrictafromWESTERLUND’s
collection, and toANITA ESCHNER (Naturhistorisches Museum
Wien),whomeasuredthesyntypesofPseudamnicola orientalis,toPHILIPPE BOUCHET(MuséumNationald’HistoireNaturelle,
Paris), for providing the numerous Pseudamnicola/Mercuria
samplesofAlgeria,toGERHARD FALKNER(Hörlkofen)aswellas
VIRGINIE HÉROS(MuséumNationald’HistoireNaturelle,Paris),
whobothfoundthesamplesinthecollectioninParis,andto
EIKE NEUBERT, who found some important lots in Bourguignat’scollectioninGeneva..WeareespeciallyindebtedtoNICK
DRAKE(DepartmentofGeography,KingsCollege,London)for
providing useful information regarding locations in southern
Tunisia and we are also so thankful to DAVID WALKER (Hamburg) for checking the English as well as to two anonymous
refereesfortheirhelpfulcomments.
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GIUSTI, F. & PEZZOLI, E. (1980): Guido per il riconoscimento
delle specie animali delle acque interne Italiane B.
Gasteropodi,2.—ConsiglioNazionaledelleRicerche:1–66;Siena.
Manuscriptsubmitted:05.05.2009
Revisedmanuscriptaccepted:06.04.2010

Revision of the genera Pseudamnicola PAULUCCI 1878 and

Transcription

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