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A new species of Vitrina (Gastropoda, Pulmonata, Vitrinidae) from Kenya with a discussion of the genus in East Africa PDF

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Preview A new species of Vitrina (Gastropoda, Pulmonata, Vitrinidae) from Kenya with a discussion of the genus in East Africa

BASTERIA, 69:147-156,2005 Anewspecies ofVitrina(Gastropoda, Pulmonata, Vitrinidae) fromKenya witha discussion of thegenus inEast Africa B. Verdcourt Royal BotanicGardens,Kew, Richmond,Surrey,TW93AB,U.K. Vitrina chyuluensis spec. nov. is described from theChyulu Hills,Kenya.Ithas arather large shell and is characterised by a longspermathecalduct, very fine plicaebetween distal and proximalpartsofthepenis, and athree-ridgedjaw.The descriptionisfollowed byadiscussion onthe classification ofEastAfrican speciesofVitrina and onthevariation incharactersproba- bly oftaxonomic significanceatspecificlevel. The paperis concluded by somecomments on namingVitrinafromEastAfrica. Keywords: Gastropoda,Pulmonata,Vitrinidae,Vitrina,taxonomy,Kenya,EastAfrica. INTRODUCTION QuentinLuke during abotanicalcollecting expedition to theChyulu Hills, Kajiado District, Kenya (02°39' S, 37°52'E), collectedsome helicarionoidsnailsin montane forest onrecentvolcaniclava.Iatfirstassumedthattheseven specimensinthesamplebelonged to the same species, one more mature than the others. Dissection showed it tobe a Chlamydarion VanMol, 1970, whichIcouldnotmatchwithdescribed species. Inorderto verifycertainfeaturesI dissectedafurtherspecimen only tofindithadatotally different anatomyand wasinfactaVitrinaaswerefourotherspecimens. Oneotherwas ajuvenile Chlamydarion. Ihave for many yearsbeen attempting to make some sense of the East Africanspecies ofVitrinaDraparnaud, 1801,and dissectednumerous specimens fromthe East Africanhighlands. AlmostnomolluscshavebeencollectedintheChyulu Hills and Iwas not surprised tofindthe Vitrina differedin certain particulars fromany Ihad dis- sectedfromelsewhere. DESCRIPTIVEPART Vitrina(Calidivitrina) chyuluensis spec.nov. (figs 1-3) Materialexamined. - Kenya,KajiadoDistrict, ChyuluHillsmain forest,nearCamp 3,Mbirikani GroupRanch, alt. 1960 m, leg. QuentinLuke, 14.iii.2004 [RMNH 99586 (shell), (softparts)/holotype; RMNH99587/3paratypes]. Diagnosis. - AVitrinawitharather large shell characterisedby alongspermathecal duct, very fine plicae betweendistalandproximal partsof thepenis and a three-ridged jaw. Description. - Shell(fig. 1)typically vitrinoid, thin,very pale greenish brown,of3V2 whorls, withno apical puncturesbutfairlystrongirregularwrinkles. Animalwithmantlefaintly to distinctly spotted; right handmantleflap completely pale andunmarkedbutwithastripe onmantlejustanteriortoitparalleltotheshelledge andcrossingthebreathinghole;tailwithadistinctblackkeelstrip,notactuallykeeledbut 148 BASTERIA,Vol. 69, No. 4-6, 2005 Fig. 1. Vitrina chyuluensis spec. nov., shell ofholotype (RMNH 99586);measurements 13x 10x8 mm (Dmax x DminxH). Kenya,KajiadoDistrict,ChyuluHills,nearcamp 3, Mbirikani GroupRanch, alt. 1960m,Q.Luke leg.,March2004.PhotoJ.Goud,Leiden. in fact grooved.Onespecimen hadmuchpaler markings butofessentially thesamechar- acter. Footuniformly pale. Genital anatomy(fig. 3). Thepenial sheathconsistsof two parts, a muscularovoid distalpart slightly pointed andwiththepenial retractor attachedslightly belowtheapex; the proximal part is cylindrical, forming aright angle withthe upperpart with a deep groovebetween thematthe inner angle. Inother species ofVitrinaIhave dissectedthere is aseriesofstrongplicate foldsin this groovebut theseare muchreduced in the present species. The series of smalltubercles onthelower part nearly always presentinAfrican materialIhavedissected arepresent, therebeing nine. Thepenisitselfis clavate, follow- ing roughly the shape of the sheath but much more slender. The right ocular retractor passes betweenthemaleand thefemaleducts.Atriumwell developed. Vas deferensleav- ing thepenial sheathnearbaseofdistalovoid part. Spermatheca and ductmuchlonger than thepenial sheath, totalling ± 13 mm. Jaw (fig. 2) 1.25mm wide,oxygnathous but three-ridgedtransversely, a characternotobserved inany othervitrinid.Thinkingitmight be anabnormality I dissected asecond youngerspecimen which confirmedthe ridging andalso thelonger spermathecal duct.Radula formula40.14.C.14.40,the mainmarginals withlateralcusptowards thebaseof centralcusp,not pincer-likebut theoutermost mar- ginals havethecusps moreequal andpincer-shaped. Measurements.Shell:13xl0><8mm;foot:27x5 mm. Distribution.- Kenya, Chyulu Hills. Ecology. - The montane forest is dominatedby Tabernaemontana (Apocynaceae), Neoboutonia(Euphorbiaceae) and Ficus thonningii Bl. (Moraceae); the snails were onthe leavesoftheunder-shrubPiper capenseL.f.(Piperaceae). TheChyulus areanareaofrecent Verdcourt:A newspecies ofVitrinafrom Kenya 149 Figs2-3. Vitrina chyuluensisspec. nov.(detailsas forfig. 1).2,jaw;3,genitalanatomyofholotype. Abbreviations: a, atrium;o, oviduct; p, penis (shownseparatelyontheleft);pp,penialplicae; pr,penialretractor;ps,penial sheath;pt,penial tubercules; s, spermatheca; sd, spermathecal duct;vd,vasdeferens. igneous activity withvastareasoflavaandhundredsofsmallconesratherthan largeiso- latedvolcanoes. Theblack markings ontheanimalsbrought tomind Vitrinanigrocincta vonMartens, 1897, describedfrom Kilimanjaro from materialcollectedby Volkensbetween 1900and 3800m. Itmightwell covermorethanonespecies. Itwas described fromthreesyntypes, thelargest being 8x6*5mm. Von Martens states thatthe animalis pale yellowish grey withsharply definedirregularblack marks, mantlewitha black stripe on theright side parallel to the peristome; the tail has a longitudinal medianblack stripe. Hubendick's description andillustrationsof the anatomyof thisspecies (Hubendick, 1953)arebased onSjostedt materialfrom3000-3500m onMt.Meru andmaterialfromKibonotoin Berlin. VonMartens doesnot mention this localityanditis notclearifitis typematerial; thereis noindicationwhocollectedit, buthemusthave hadaccess to preserved animalsto give such a detaileddescription. Hubendick'sillustrationsshow a spermatheca and ductno longer thanthepenis and materialI havedissectedfrom3100 m onMt.Kilimanjaro had themonly slightly longer. THE CLASSIFICATIONOF EASTAFRICAN VITRINA Thecommentarywhichfollowsformspartof apaperwhichwas neverpublished. A. Holmcollectedaspeciesof Vitrinafoundontheforestedrim ofNgorongoro Craterwhich Iconsideredwasnew.Unfortunately thematerialselectedasholotype andparatypewas 150 BASTERIA, Vol. 69,No. 4-6,2005 mislaid and neverrefound. No further material haseverbeen collected atNgorogoro so far as Iamaware.As IdissectedmorematerialfromEast Africafurthercharacters came to lightwhich Ihadnot consideredwhendealing withtheNgorongoro materialso until morematerialiscollectedthereitwouldbeunwise todescribeit.Sincethecrateris awell- known tourist attractiontheforestedcrater slopes probably remainundamaged and fur- thermaterialwillone daybecomeavailable. Von Martens (1895, 1897)wasthefirstpersonto describeVitrinidaefromEast Africa (as distinctfromEthiopia); hegaveinformationaboutthemantleflaps andcompared the radulaofVitrinaoleosavonMartens, 1895,withthatofVitrinadraparnaldi Cuvier, 1817[i.e. Phenacolimax major (Ferussac, 1807)]. Pollonera (1909) figured the genital anatomy of VitrinaibandensisPollonera, 1907.Pilsbry (1919:283)establishedthesubgenus Calidivitrina for Vitrinaoleosa largely based on thepremise that "the marginal teethall havesimple, thorn-like cusps,notbifid as partof themare intypical Vitrina”and "themantlehasno shell lobes or if present they are so small they disappear by contraction in alcohol". Anothercharacterwhichisalsomentionedisthelack ofpuncturation oftheshellso char- acteristic ofHolarctic species. The manydissections carried outontheextensivematerial already mentioneddemonstratethat either the description of Calidivitrinamust be con- siderably modifiedor more thanone groupof Vitrinawith unmodifiedvagina occur in EastAfrica. Many specimens havebeenexamined wherethemarginal teeth areobvious- ly bicuspid andpincer-shaped andotherswheretheposition ofthesmallercuspbecomes higherand smallerandfinally isnomorethanasmallnick attheapexofthe maincusp. Whetherthesestatescorrespond to differentspecies oris sometimesmerevariation in one species is not yet clear. Shell lobes arevery well developed in mostspecies, sometimes almostenveloping the wholeshell. In fact someof the largest Ihave seenhavebeen in Holm61 fromthe Ruwenzori, MijusiValley, which is strongly atvariance withPilsbry's observations.Finally Vitrinavariopunctata Connolly, 1925,fromMtElgon (Connolly, 1925) haseither thewholeshell orallbuthalfofthebody whorl punctate. Itis cleartherefore thatnoneofthecharactersPilsbrymentionedto distinguishCalidivitrina, holdsfortherest ofEast Africa. Hubendick(1953) published areviewof theanatomyfortheeastern Africanspecies withdrawingsofthedissectionsofallthetypespecimens hecouldlocate.His maininter- estwasto determinetherelationships ofthesespecies totheEuropean species. Theresults didnotestablishhowmanyvalidspecies thereactually werenor howthey couldbe dis- tinguished anatomically. The drawingsofthegenitaliaarenotstrictly comparable. He did demonstratethattwo quitedifferentgroupsofvitrinidoccurred in eastern Africa. Some materialfrom Muhavura(on theborderof Uganda and Rwanda: 1°23'S, 29°41' E) col- lectedbyHedberg hadthespecialised vagina ofPhenacolimaxStabile, 1859,butnofurther materialhasbeendiscovered.Forcart(in litt.)diddeterminea green-shelled species with blackanimalsfromhighaltitudesonMtKenya asPhenacolimaxspec. nov. butlater(1978) madenoreferenceto thisandIthinkitwas in error. Forcart(inlitt.)statedthathehadseennomaterialfromEthiopia thatcouldbeattrib- uted to Calidivitrina.He had dissected a good dealof material collectedby Hugh Scott. Having dealtextensively withEuropean Vitrinidae(1944, 1949, 1952, 1954, 1955, 1956a, 1957, 1959)he turned to thoseof Africa (1956b, 1959, 1978). In his 1959 paperhe refers Ethiopian materialto PhenacolimaxHesse,subgenus ArabivitrinaThiele, 1931.1senthima considerableamountofmaterialbetween1957and 1960,buteventually hefoundhecould notprepareanaccount of the actual species involved;he still emphasized (in litt., 1978) thesimplemarginal cuspsand reducedmantleflaps asa distinctionofCalidivitrinawhich means, I think, thathecould nothaveexaminedall the materialIsent tohimmuch of whichhadclearlybicuspid marginals and veryextensivemantleflaps.Healsomentioned Verdcourt:A newspecies ofVitrinafrom Kenya 151 thelateralserrationbelowtheectocone insomeouter marginals intypical Vitrinawerenot foundinCalidivitrina, butIhavefound theoutermost marginals insome specimens are serrate withup to seven cusps(e.g. Mt.Kenya,Ragati Forest,2000m,Holm173). Forcart alsopointed out(inlitt., 10.iii.1958)thatEucobresiaBaker, 1929(SemilimaxHesse, 1923,non Gray, 1847) couldnotbe distinguished fromCalidivitrina, thelatter beingtheoldername. Hausdorf's(2002) recent cladistic analysis of theVitrinidaeis, however,strongly atvari- ance withthis.Calidivitrinais placedcloser toArabivitrinathantoEucobresiaandhepoints outthat theVitrininae auct.include thegroups inwhichthevaginal stimulatorhasbeen lost,namely Vitrinaand Calidivitrina, andispolyphyletic. Hesuggests thatsubfamiliesare notnecessary. Of theseventeen charactersusedby Hausdorfin his charactermatrixused forthe analysis, seven are not applicable to Calidivitrinasince the stimulatorand penial tunica areabsent; the seventeenthcharacter assumes that themarginal teethare never multicuspid but this is not always correct. He estimates the number of species of CalidivitrinatobeeightbutalsostatesfiveArabivitrinaoccur in EastAfricaso presumably he is including Ethiopia. I have seenno ArabivitrinafromEast Africa whichbearing in mindthecloseaffinityofEthiopian and Kenya highland plantsand animalsis surprising. He does not record Phenacolimaxfrom East Africa so presumably doubts Hubendick's record. VARIATION IN CHARACTERSATSPECIFICLEVEL 1.Colourofshellandanimal- The colouroftheshellvariesfromwhiteto darkolive green,the latter always associated withhigh altitudes on Mts. Kenya, Kilimanjaro and Ruwenzori. Theanimalvariesfromvery pale whitish to jet black, thespecies withdark green shells always having black animals. On Mt. Kenya during the International Geophysical Year expedition in 1957 itwas notedby several collectorsthatVitrinavaried considerably incolour.Thereportatthetimementionedthecommon Vitrinaoccurring at 3000 m onLobeliamaybeonespecies butif so itis veryvariable.There mayofcoursebe several species livingtogether. Usually themantleand body are yellowish withblack or blackishmarkings andthehindendof thebody hasablackish dorsalstreak. The solehas numerous smallyellowish spotsandthetentaclesaredarkish.Somespecimens, however, areveryblackin appearanceandtheshellappearsdark;othersarebrownandyellowand some are entirely pale withvery pale markings and still othershave avery distinctred tinge. Someofthedifferencesremain noticeablein spirit material. There isevidence,however,that materialfromoneprecise locality canbe veryuni- form in colourand anatomy. Some 500 specimens collectednear thebase camp on the SirimonTrack by Wachtersduring theVanGoethemMt.Kenya Bio-Expedition weresam- pled andfoundtobe virtually identicalanduniformly pale in colour. Von Martens described Vitrina nigrocincta from Kilimanjaro with black markings around the mantle edge and much material fromthe mountain has such markings but some is totally devoidof them.It seems doubtfulthat species canbe distinguished on colour although the green-shelled black animals are not the same as the paler ones although littlematerial hasbeen availableforanatomicalwork. 2. Radula.-As hasbeenmentionedthereis considerablevariation in thecuspsof the marginal teeth varying from virtually thorn-like unicuspid teeth with a mere minute notchatthebaseofthelong cusps to distinctly bicuspid pincer-shaped teeth.My studies ofpopulations fromonearea indicatethattheshape isconstant butamore detailedstudy including possible changes duringthedevelopment oftheanimalis neededto checkthis. 152 BASTERIA, Vol. 69,No. 4-6,2005 The numberofteethin arow isknowntobehighlyvariableinmany groupsofsnailsbut itmaypossiblybeofsomeuse as acharacter. TheformulainVitrinaoleosafromHolm s.n. was givenbv Hubendick(1953)as 35.13.C.13.35and Ifound30.11.c.11.30and27.12.C.12.27 in Holm 61 and Holm47, all three from the same locality Bujuku Valley (Ruwenzori, Uganda). Widely differingnumbershavebeenfound inotherspecies. Ifound theformu- lafor V. variopunctata (Mt. Elgon, Holm76)tobe 15.9.C.9.15and Peile(in Connolly, 1931) gives 22.10.C.10.22.Up to 54 marginal teeth occur in material from the East Aberdares (Holm267).There is variation inthe cuspsof theoutermostmarginalteethwhichcanbe two-cusped liketherest or threeto five- oreven seven-cusped. Mordan& Marins (2001) show thatthevariation in themarginal teethofthe vitrinidgenusPlutonia Stabile, 1864, intheAzores is evenmore diverse. 3.Genital anatomy-Apart fromthePhenacolimaxfromMuhavura(Uganda) thevari- ationinthelower ductsof the East African materialdissected isrestricted. The narrow spermatheca gradually passing into a shortductfoundin Vitrinaoleosadiffersnoticeably fromtheoblong, ellipsoidor subglobose spermatheca moreclearly demarcatedfromthe ductfoundin allotherEast Africanspecies dissected.Theductvaries mostly fromunder 1 mm to 4mm butductsup to 8mmhavebeenfound in onespecimen fromMt. Kenya andprobably indicatesa distinctspecies. Thevaginavaries fromalmostobsoleteto3mm long. Theinternal structure of the penis, particularly of the folds andpenial gland, may provide the mostusefulcharacters for specific distinctions. Hubendick (1953) discusses thisandfiguressectionsbutIhavefoundthemvery confusing. Neubert(1998)gives more intelligible drawings of anArabianspecies. Thevariation and development withinone species needsinvestigationandalsotheeffectofdifferentmethodsofpreservation onthe appearanceof the organs. Penial glands which appear very differentin single prepara- tions mayowethedifferencestothesereasonsratherthantointrinsicspecific differences. 4.Mantleflaps - Themantleflap variesfromanarrowarcto alarge elongate struc- ture whichalmostcoversthe shell reaching 9mmlong and 11mmwide withright hand lobe5 x 12mm.Thesize doubtlessvariesaccording tothe development oftheanimaland themodeof preservation mayalterthesize. Notesneedtobemadeonlivinganimals. NAMING VITRINAFROM EASTAFRICA Apart fromtypematerialmostsubsequent specimens havebeennamedVitrinasp.or given a namepurely on geographical grounds. The following comments may help to direct futurefieldwork undertakentotry andsort outthespecies. Vitrinaoleosa from Ruwenzori is apparently well characterised by its elongate or obovoid spermatheca gradually passing into the duct, reduced mantle lobes, aculeate marginal teethand yellow-green depressed shell.Pilsbry (1919) describedthe animalas dirtychamois with pale grey marbling but von Martens (1895) describes it asblackish. Von Martensstates thereis noneck flap and only asmalltongue shaped shell flapand Pilsbry statesthey aremore or lessabsent.My ownexaminationofHolm47and61 from Mijusi Valley at3950-4000magreeswiththeshapeof thespermatheca and ductandthe marginal teethbutthemantleflaps arevery well-developed inHolm65andperhaps sug- gestsanotherspecies maybepresent. Anything fromtheRuwenzorirangewitha green- ish ambershelland uniformlyblack animalcanbe assumedtobe V. ibandensis.Pollonera (1909)figures thegenitalanatomyofthisbutmakesnomentionofmantleflaps;using his Verdcourt:A newspecies ofVitrinafromKenya 153 informationthatthe drawing isenlarged by fourtimesthevaginais distinct, 2.5mm long andthelowerpartofthepenis elongated,5 mmlong. Thepenis certainlydiffersfromthe figuresgiven byHubendick(1953)of theothertwogreen-shelled Vitrinawhichhavebeen dissected - V. ericinellae d'Ailly, 1910, and V. viridisplendens d'Ailly, 1910,both from Kilimanjaro. Hubendick also states that the vagina is "very short" and "hardly any" respectively although thisis not clearfromhis figures. Polloneraalso describedV. cagnii Pollonera, 1906, fromRuwenzori butfigured only theshell. Pilsby (1919) reduces this withoutcommentto V.oleosaandthefigures oftheshellssuggestthisis correct, butifit iseventually proved thatseveralspecies occuronthemountain thisnamewillalsohave tobe taken into account. Attempts to find Pilsbry's original dissected material proved unsuccessful. Thiele (1911) also described V. bambuseti from Sabinjo (Sabinio) and V. tenuissimafromKarissimbi(Karisimbi),bothvolcanoesintheVirunga Mts.in easternD.R. Congo (formerly Zaire).The materialwas in spiritbut he didnot describe theanatomy although Hubendick(1953) laterdescribedthelatter. The Phenacolimaxknown fromMuhavura(Hubendick, 1953)hasnotbeennamedbut isunlikely tobeidenticalwiththeEthiopian species. Ihavenotseentheoriginal Hedberg materialand donot know howadequate itis. Nofurthermaterialhasbeen seen. Theonly species actually describedfrom Mt.Elgon is themost easily recognised of allEastAfricanspecies, viz. V. variopunctata withasmallvery compressed shell.Connolly pointed outthatitwas characterisedbythewholeshellbeingpunctured allover [not just theapical whorlasin Vitrinapellucida (Miiller, 1774)]andhealsofigures theradula(Peile, in Connolly, 1931). My observations on materialcollectedby Holm (Holm 76 from E. Elgon at2900 m) and Anderson(E. Elgon, 2490 m) indicatethat theextentof thepunc- turing canbe variableandoftenlacking onthemainpartofthebody whorl. Hubendick(1953) describesthe genitaliaof anunnamedVitrinacollected at 4250m on Koitoboss, Mt. Elgon. His figure does show the insideof the penis but he gives a description ofthefolds,whichis difficulttofollowbutsuggests itis a distinctspecies. Ihaveexaminedspecimens ofaVitrina(Holm 131)fromtheeastern sideofMt.Elgon at3150m withasmall ambershellbut verymarkedmantleflaps and avery characteris- ticpenial gland andpincer-shaped marginalteeth.Therearethereforeprobably threedis- tinctVitrinaonMt. Elgon. Material from the fairly close Cherangani Hills (Holm 55 and 65), 4 km N. of Kaisungor at2900m,has acombinationofdistinctivecharacters:a quitestrongly striate shell, along atrium, very distinctvagina, distinctivepenial foldsandclearlybicuspid pin- cer-shaped marginal radula teeth. Sample I.F. Thomas4, collectedby Pokot childrenon giantlobeliaat3150 m onCherangani,alsohadclearlybifidmarginal teethwiththemain cuspaboutthreetimesthesizeofthesmallerone; theinternalpenis structurewasnot con- formwiththatintheHolmmaterialandwas takenupentirelyby glandular material,the gland capis acuminate andbeneathitthereareaboutadozenannularflanges. Agreat dealof materialhas beencollectedin the Kenya Highlands andhas mainly been named Vitrina ugandensis Thiele, 1911, which was described from Kenya, Mau Escarpment fromshellsalone,or V.lacteaConnolly, 1925,describedfromMtKenya (mate- rial collectedby Kemp andby Gregory is cited). Connolly (1925)merely mentions thatV. lacteahasalessprominent apexbuthedoes,however,give radulainformationforV.lactea provided by Peile. This shows the marginals to be aculeate with a small entocone. Materialfromratherloweraltitudeshas usuallybeenattributedtoV. ugandensis andthat from over 3000 m to V. lactea. Smith (1903) gave the altitude of materialcollected by Doherty ontheMauEscarpment as 6500-9000ft(1950-2700m). Thelowestaltitudefor Vitrinain Kenya is materialcollected atThika, ChaniaFalls, 1500 m (Polhill 110). Holm267 from theE. Aberdares at2900 m is distinctivein having more marginal 154 BASTERIA, Vol. 69,No. 4-6,2005 teeth thanseen in any otherEast African Vitrina, the formulabeing 50-54.11.c.11.50-54; they are clearlytwo-cusped andtheouterones distinctly pincer-shaped. Thepenial gland is distinctivewith capitate apexwith abouta dozen flanges beneathand witha row of seven nodules arranged longitudinally and just overlapping the flanges. Whetherthese are homologous withthemuchmorespiniformpapillae describedby Mordan&Martins (2001)for oneof the vitrinidsfrom Santa Maria, Azores, is an interesting question. An immaturespecimen, Holm 168, just labelledAberdares, but fromthe samealtitude, had theformula29.9.C.9.29withsimilarteeth. There are at leastfourspecies of Vitrina onMt Kenya and itis necessary to decide whichshouldbe calledV.lactea.Hubendick(1953) gives adetaileddescriptionandfigures of the genitaliaof materialcollectedby Holm in theeastern partof the TelekiValley at 4150-4230m whichhe refersto V.lactea,butdoesnot mention thecolourof theshellsor animalsnor doestheraduladescriptionmentionthemarginal teethindetail.Othermate- rialcollectedbyHolmfromtheTyndall Glacierat4450misreferredtoV.baringoensis E.A. Smith, 1894,whichbothConnolly (1925)andI(Verdcourt, 1956,1983)havedemonstrated is notaVitrina.Theanatomyis saidtoscarcely differfromthematerialreferredto V.lactea andthecolouroftheshelland animaland detailsof themarginal radularteethare miss- ing. Vitrinabaringoensis wasbasedonmaterialfromtwo well-separated localities,Baringo and Mt. Kenya, and probably covers more than one species of shelled Urocyclidae (‘Helicarion’ofearlierauthors). Abundantmaterial(10 D), collectedby Wachters during theBelgian MtKenya Bio-Expedition at thebase camponthe SirimonTrack, has shells whichexactly matchthetypematerialofV.lactea and theradulaconforms. A.Andersoncollectedmanyspecimens neartheMeteorological Station ontheNaro- moruTrack at3080-3150m whichhave shellsmatching V. lactea and a radulaagreeing withthat depicted by Connolly. Otherspecimens examinedwithsimilar marginal teeth were collectedby Andersonin the TelekiValley at3715m. Elsewhereonthe mountain materialwithvery differenttwo-cusped pincer shaped marginals occur at varying alti- tudes,Holm 189fromRagati Forestat2000m,TattersfieldfromnearLakeEllisat3500m, andHolm s.n. at3280montheW.sideofthemountain. One specimen collected by Liliane (143D) near the base camp onthe Belgian Mt Kenya Bio-Expedition has the spermathecal duct 8 mm long, 2-3 times longer than in othermaterialexamined;itmaywell representa distincttaxon. Materialwithdark greenshellshas beencollected by Mrs Joy Bally (later Mrs Joy Adamson)at3150m intheMacKinderValley butnoanimalswerepreserved. M.Coecol- lectedmaterialwithvery soft deep shellsandintensely black animalsunderrocks with littlevegetation onthe Lewis Glaciermoraineat 4440 m. Forcart (in litt., 19.V.1968)sug- gested thismightbeanewPhenacolimaxbutnomentionwas madeof itin hispaper(1978) andI donotthinkhedissectedit. I hope totrace thismaterialandexamine it. Five species havebeendescribed fromKilimanjaro. Vitrina nigrocincta was described by vonMartens (1897) andhas beenmentionedpreviously. Ihaveexamined severallots ofmaterialwhichonewouldexpecttobethesameasVolkens'sloweraltitudespecimens. Holm218 from NW.of Bismarck (nowMandara) Hut at3100m had themarginal teeth withthelongcusps fivetimes aslong as theshortonesandHolm 203 fromneartheHut itselfat2480m hadsimilar cusps,but Holm208 fromneartheHut at2480m hadmar- ginals withpincer-like cusps.BothHolm203and208havetheanimalsmarkedwithblack orbrown.Holm266a fromKitoto onthenearby Mt.Meru atanunspecified altitudehas marginals similartothoseinhis203 and 218.D'Ailly (1910)wrotetheaccountof themol- luscs collected by Sjostedt's Kilimanjaro-Meru-Expedition; unfortunately he gave no anatomicaldetails.He records V. nigrocincta fromMeru andKilimanjaro, over 1600 speci- mens being collected oneachmountain.Hepoints out that vonMartens'smaterialwas Verdcourt:Anewspecies ofVitrinafromKenya 155 juvenile (presumably onevidenceof shelldimensions- thereisnoevidenceheactually saw the material). Materialwas recorded from 3000-4400m on Meru and2500 m on Kilimanjaro. D'Ailly (1910) also described three other species from Kilimanjaro: V. kiboschoensis isbased on46 specimens fromKiboscho at3000m in theheathzone found mostly betweentheleavesofLobeliadeckenii(Asch.) Hemsl.; V.ericinellaeisbased on 174 specimens fromKiboscho also at 3000 m altitudein theheathzonewitholive strigose shellswith a whiteapex and pale brown animals,and V. viridisplendens is basedon 64 specimens also fromKiboscho at3000-4000mnear theupperlimitof vegetation witha blackishanimaland yellow-greenor darkgreenishbrownshell.D'Ailly doesnot mention Dautzenberg's V. lobeliaecola(1908, based onmaterialcollectedby AlluaudonLobelia in "zone desprairies" at3200m) although hecites hispaperelsewhereso musthave been aware ofit. V. kiboschoensis couldbe synonymous;fromthe descriptions they are similar in colour,measurementsandhabitat.Hubendick(1953)has dissectedspecimens ofallthe species collectedby Sjostedt; it appearsthat V. nigrocincta hasa longer vagina andmore numerous marginal teeth than the otherthreebuthow far the species couldbe distin- guished by such detailsneeds are-examination of the abundantoriginal materialwith this view inmind.D'Ailly had akeen eye andhisspecies are probably valid.In 1955I examinedthe material in Stockholm and my notes give "dark greenish brown" for V. viridisplendens, "grey-black radially flammulateshellwhiteinside andwithwhitetip" for V.ericinellae and "yellow-brown" for V. kiboschoensis. I collected V. viridisplendens near Peter's (now Horombo) Hut at 3750m understones in January 1955but unfortunately sentallthematerialaway forexamination insteadofdoingit myself. Onespecimen ofgreatinterestremainstobementioned.Hubendick(1953)mentions aspecimen fromtheUsambaras,Tanzania,collectedin spiritandfigures itsanatomy.He seemstohavebeentotally unawarethatthisis anexceptional occurrence fromalow alti- tude(±1000m;Conradtcollectedonly in the East Usambaras) andis moreover themost southerly record of thegenus in Africa. Conradt'scollectionwas madelong beforevon Martens'sgreatworkonEastAfrican molluscs(1897)so itiscurioushemadenomention of thisspecimen. I sawno traceof aVitrina whilst Ilived in thearea in 1949-1950.Itis almost certain to be a distinct endemic species. Itis asurprisingly low altitudefor the genus. Describedby Thiele(1911) fromshellscollectedbyDoherty sentbyE.A.Smithtovon Martenswithouttheexactlocalitybeing given (seeConnolly, 1925). REFERENCES AILLY,A.d',1910. Mollusca.In:Y.SJOSTEDT,ed.,Wissenschaftliche Ergebnisseder schwedischen zoo- logischen Expeditionnach dem Kilimanjaro, dem Meru und dem umgebendenMassaisteppen Deutsch-Ostafrikas 1905-1906,unterLeitungvonProf.YngveSjostedt 1,6:1-32. CONNOLLY, M.,1925.NotesonAfricannon-marineMollusca withdescriptionsofmanynewspecies.- AnnalsandMagazineofNaturalHistory(9) 15:457-479. CONNOLLY, M., 1931.Descriptions ofnewnon-marine Mollusca fromnorth, southand centralAfrica withnotesonotherspecies. - Annals and MagazineofNatural History (10)8:305-321. FORCART, L., 1944. Monographieder schweizerischen Vitrinidae (Moll.-Pulm.). c Revue Suisse de Zoologie51:629-678. FORCART, L., 1949. DieErektion derKopulationsorganeund derKopulationmodusvon Phenacolimax major(Fer.).- Archiv furMolluskenkunde 77:115-119. FORCART, L., 1952. Neue FundevonVitrinidae. - Basteria 16:33-36. FORCART, L., 1954.RevisionvonVitrinakochiAndreae und Vitrinanivalis (Dumont&Mortillet)(Moll. 156 BASTERIA,Vol. 69, No. 4-6, 2005 Pulm.).- VerhandlungenderNaturforschenden GesellschaftinBasel65:264-269. FORCART, L., 1955.Dienordischen ArtenderGattungVitrina.Archiv furMolluskenkunde 84: 155-166. FORCART, L., 1956a.DieVitrinidae derOstalpen.- ArchivfiirMolluskenkunde 85:1-14. FORCART, L., 1956b. Journey to the High Simien (Northern Ethiopia), 1952-3. Three species of Phenacolimax (Gastropoda,Vitrinidae)with notes onthe taxonomyofthe genus.- Journalofthe Linnean SocietyofLondon,Zoology43:113-122. FORCART,L.,1957.DieVitrinidae vonBaden. (Moll.Styl.).- MitteilungendesBadischenLandesvereins fiirNaturkunde und Naturschutz,NeueFolge6,1:19-24. FORCART, L., 1959. Revision nordafrikanischer Vitrinidae. - Archiv. fiirMolluskenkunde 88:1-6. FORCART,L., 1978. Africandistribution ofgenera and subgeneraofthe familyVitrinidae(Pulmonata). - JournalofConchology29: 325-327. HAUSDORF, B., 2002.Phylogenyand biogeographyoftheVitrinidae (Gastropoda:Stylommatophora).- Zoological JournaloftheLinneanSociety14:347-358. HUBENDICK, B., 1953. The relationships ofthe EastAfrican Vitrinae with notesonthe taxonomyof Vitrina. - ArkivforZoologi(2)6: 83-96. MARTENS, E. VON, 1895.NeueArtenvonLandschnecken ausden GebirgenOst-Afrikas.- Sitzungs- Berichte derGesellschaftnaturforschender Freunde zuBerlin 18Juni1895: 120-129. MARTENS,E.VON, 1897. BeschalteWeichthiere Deutsch-Ost-Afrikas.In:K.MOBIUS,ed.,Deutsch-Ost- Afrika4:i-v+ 1-308.Berlin. MORDAN, P.B.,&A.M.F.MARTENS,2001. Asystematicrevision ofthevitrinidsemislugs ofthe Azores (Gastropoda:Pulmonata).- JournalofMolluscan Studies 67:343-368. NEUBERT, E., 1998. Annotated checklist of the terrestrial and freshwater molluscs of the Arabian Peninsula with descriptionsofnewspecies. - Fauna ofArabia 17:333-461. PILSBRY,H.A., 1919.Areview ofthelandmollusks oftheBelgianCongochieflybasedonthecollections ofthe AmericanMuseum CongoExpedition,1909-1915.- Bulletin ofthe American Museum of NaturalHistory40: 1-370. POLLONERA, C.,1906.SpedizionealRuwenzori diS.A.RLuigiAmedeo deSavoia Duca degliAbruzzi. Nuove specie di Molluschi terrestre (Diagnosipreventive).- Bolletino del Musei diZoologia ed Anatomia Comparatadella R.Universita diTorino 21,538: 1-4. POLLONERA, C.,1907.Spedizioneal RuwenzoridiS.A.RLuigiAmedeo deSavoia Duca degliAbruzzi. Nuovespecie diMolluschi (Diagnosipreventive). - Bolletino delMusei diZoologiaed Anatomia Comparatadella R.Universita diTorino 22,561:1-3. POLLONERA,C., 1909. Molluschi Stylommatophora.In:SpedizionealRuwenzori diS.A.R. ilPrincipe L.Amedeo diSavoia DucadegliAbruzzi,partescientifica 1:182-205. SMITH,E.A., 1894. Alistoftheland andfreshwatermolluscscollectedbyDr.J.W.GregoryinEastAfrica duringhisexpeditiontoMountKeniawithdescriptionsofafew newspecies.-Proceedings ofthe MalacologicalSociety ofLondon 1:163-168. THIELE, J., 1911. Mollusken der Deutschen Zentralafrika-expedition. In: H. SCHUBOTZ, ed., Wissenschaftliche Ergebnisse der deutschen Zentral-Afrika-Expedition1907-1908 unter Fiihrung AdolfFriedrich's,HerzogzuMecklenberg3(Zoologie1,6): 175-214. VERDCOURT,B., 1956. Notesonsomeland and freshwaterMollusca fromEthiopiaand Somalia (includ- ingsomerecords fromKenya). -Proceedings oftheMalacologicalSociety ofLondon 32:72-79. VERDCOURT, B., 1983. Alist ofthe non-marineMollusca ofEast Africa (Kenya, Uganda, Tanzania, excluding LakeMalawi).- Achatina 11:200-239.

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