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Synonymy and nomenclatural history of the Common or Viviparous Lizard, by this time: Zootaca vivipara (Lichtenstein, 1823) PDF

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1 Bonn zoological Bulletin 60 (2): 214-228 December 2011 Synonymy and nomenclatural history of the Common or Viviparous Lizard, by this time: Zootoca vivipara (Lichtenstein, 1823) JosefFriedrich Schmidtleri & Wolfgang Bohme^ ^OberfdhringerStrafie 35, D-81925 Milnchen, Germany; E-mail:[email protected] -Zoologisches Forschungsmuseum AlexanderKoenig, Adenauerallee 160. D-53113 Bonn, Germany; E-mail: [email protected] Abstract. We carefully reread and translatedthe Latin account by J.F. von Jacquin (1787) on his description ofa vivi- parous lizard ("Lacerta vivipara") in theAustrian Alps nearVienna. It turned out that - in contrast to common usage - this account cannot be regarded as the original description and scientific denomination ofthe taxon Zootoca (formerly Lacerta) vivipara. It is apparent that v. Jacquin did not at all intend to describe a new species, butjust wanted to point on his extraordinary observation that the lizard obsei-ved by him gave birth to young instead oflaying eggs (Latin: La- certavivipara=viviparous lizard). Forsecuringnomenclaturalstabilityofthiswell-knownandwidelydistributedspecies, we hadto search forthe next, subsequent authorusingv. Jacquin's name in the sense ofataxonomic denomination.Ac- cordingto ourextensive literaturereview, itwasLichtenstein (1823)who firstused"Lacertavivipara"as aspecies name (although he thought it tobe asynonymofLacertamuralis). In accordance with andto meetthe standards ofArticle 1 ofthe International Code ofZoological Nomenclature (ICZN 1999), the common lizard has now to be named Lacerta vivipara Lichtenstein, 1823, or, according to current concepts, Zootoca vivipara (Lichtenstein, 1823). Cocteau(1832)wastheonlyearlierauthorwhorecognizedthatvonJacquinhadnointentionofdescribinganewspecies. He therefore named his lizard from Schneeberg /Austria "Lacerta deJacquin' which has to be regarded as a replace- ment name (at least with respect to Lacerta vivipara Lichtenstein). Subsequently, Dumeril & Bibron (1839) discussed the biological details of"Lacerta vivipara" given by Cocteau (1832), but omitted his nomenclatural findings complete- ly, like all subsequent authors. Since then the common lizard was usually named "Lacerta vivipara de Jacquin, 1787", whereupon some oldernames before Lichtenstein's (1823) action were relegated into its synonymy. The viviparous lizard was for a long time the hidden member ofa voluminous collective species since early Linnean times (namedLacerta agilis by Linnaeus, 1758, orLacertus cinereus by Lacepede, 1788 ~andcomprising at least addi- tionally Lacerta agilis Linnaeus, 1768, Lacerta viridis Laurenti, 1768, and Podarcis muralis (Laurenti, 1768), in their current sense. Onthe otherhandWolfin Stunn (1805) and Mikan in Stunn (1805) had split thepresent-day Zootoca vi- vipara intothree species lateron .The discrepancy between several simultaneous lumping and splittingtrends immense- ly complicated the development ofnomenclature and systematics-not only in Zootoca vivipara (see Schmidtler, 2010) -up to the middle ofthe 19"^ century. In this paper, we provide a detailed list ofthe names erected anddescribedin connection withthe common lizardand discuss and evaluate their nomenclatural availability. Some ofthe old nomina were rediscoveredhere and are rathercu- rious examples from early post-Linnean times. Key words. Squamata: Lacertidae: Zootoca vivipara; nomenclature, authorships, history ofherpetology. INTRODUCTION The Common orviviparous lizard (Lacerta-nowZooto- subspecies, viz. sachalinensis Perelshin & Terentyev, & ca vivipara auct.) is the most widespread terrestrial rep- 1963, paniionica Lac Kluch, 1968, carniolica Mayer, tile in the world ranging from the Cantabrian mountains Bohme, Tiedemann & Bischoff, 2000, andlouislantziAi- and Ireland in the west through the entire Eurasian land- ribas, 2009 (see Dely & Bohme 1984, Kuprianova & mass to Sakhalin Id. (Russia) and Hokkaido Id. (Japan) Bohme 1997, Cabela et al. 2001, Mayer & Bohme 2000, in the east, and from the Po valley (Italy) and South Bul- Mayeret al. 2000, Odiema etal. 2000, 2004, Glandt2001, garia in the south to the Barents Sea in the noith. But de- Surget-Groba et al. 2001, Ambas 2009). Subsequently, it spite this huge range, this species was for long believed was even claimed that it would be possible (though still tobe monotypic (see e.g. Wemiuth 1951, Mertens &Wer- difficult) to define some of these taxa by external mor- muth 1960). However, more recent studies revealed con- phology (Clasen 2001, Guillaume et al., 2006, Arribas siderable differences in physiology, natural history, kary- 2009). ology, and - most important - molecular genetics which Because ofthis new andearlierunexpected diversityin resulted in the description and/or recognition of several the above-mentionedparameters ofthis species itbecame Received: 15.08.2011 Corresponding editor: P. Wagner Accepted: 10.10.2011 ) ) Synonymy and nomenclatural history ofZootoca rivi/xira (Lichtenstein, 1823) 215 important to define what Zootoca (orLaceiia respective- should be fticilitated when a sound nomenclatural basis ly) vivipara actually is. To this end, a re-analysis ofJ.F. is available. With this paper, we want to contribute to the V. Jacquin's (1787) tlrst description and a short biograph- creation of this basis by evaluating the complicated ical searchabouthimselfseemeduseful as a first step. This nomenclatural history ofZootoca vivipara which is more first description ofa viviparous lizard is a short account than 200 years old. ofone and a halfpages published in Latin language in the "Annales Helveticae". It deals with an observation the au- thor made when he was an eleven years old boy, accom- RESULTS panying his father, the famous botanistNikolaus Josefvon Jacquin, on a botanical excursion to the Schneeberg near L Synonymy list ofZootoca vivipara Vienna in 1778. During this walk, JosefFranz happened to catch a female lizard, put it into a small box, looked In the following we give an as complete as possible com- after it again two days later and found six juveniles but mented list of names applied to Z. vivipara since von no eggshells wherefore he drew the con-ect conclusion that Jacquin's (1787) account. The year ofdescription is fol- these juveniles should have come to light directly from lowed by the name in its original spelling and by the type their mother's body. A first important point in respect to localitiy- ifstated-likewise inthe original spelling. Our this article (written down nine years later, i.e. 1787) is that comments are referred to numbered notes in the second the term "Lacerta vivipara" (i.e. a viviparous lizard) is paragraph ofthis chapter. used only once, viz. in the title. The next mentioning of theanimal, the first inthe text, reads "Lacertapraegnans" 1830 Zootoca Wagler - Type species: ""Lacerta vivipara (i.e. apregnant lizard). But the statement given bythe au- Jacqu.". (Note 1) thor at the end ofhis article, that he would never dare to 1785 Seps? atra Schrank in Schrank & Moll - Berg state which species his lizard might belong to, makes def- Schwarzort bei Berchtesgaden. (Note 2) initely clear that "Lacerta vivipara" was not intended as 1787 "Lacerta vivipara"Jacquin-"Inmonte Schneeberg". a new scientific taxon name but only as a Latin term for (Note 3) the observed phenomenon ofviviparity. Ifhe would have 1788 Lacertiis cinereus Lacepede - (Note 4) written his account in German or any other language, the 1804Lacerta oedura Sheppard-"Atdifferenttimes found name Lacerta vivipara would never have been created at in vast abundance". (Note 5) that time and would never have been ascribed to J.F. von 1805 Lacerta montana Mikan in Sturm-"Riesengebirg". Jacquin (see Cocteau 1835, Bohme & Rodder 2006, (Note 6) Schmidtler & Bohme 2006). This means, thatLacerta vi- 1805 Lacerta nigra Wolfin Sturm-"Wenger-Alpe, Can- vipara Jacquin, 1787 does not at all meet the criteria of ton Bern". (Note 7) Art. 11.5 ofthe International Code ofZoologicalNomen- 1805 Lacerta crocea Wolfin Sturm- "Hiesige Gegend". clature (ICZN 1999) which states: "To be available, a (Note 8) name must be used as valid for a taxon when proposed". 1808 Lacerta fragilis Palmstruch & Swartz (Note 9) We were aware that this discovery is suited to endan- 1820 Lac. unicolor Kuhl - "Vaterland ?" (Note 10) gerthe stability ofthe nameZootoca vivipara because the 1820 Lac. ptychodes Kuhl - "Vaterland?". (Note 1 1 synonymy list is long and datesback to the early 19^^ cen- 1820LacertaptychodesMerrem-"Habitat...". (Note 11 tury. Changing the name ofthis famous reptile would be 1820 Lacerta pyrrhogaster Merrem - "Deutschland". catastrophic, not so much for herpeto-taxonomists (who (Note 12) are accustomed to such changes) but mainly for taxono- 1823 "'Lac. vivipara Jaquin" (sic!), Lichtenstein. (Note 3) my-users such as physiologists, ecologists, conservation- 1829 Lacerta schreibersiana Milne Edwards - "Envoyes ists etc., and the Code clearly states already in its pream- de Vienne". (Note 13) ble that stability anduniversaHtyofscientific names is its 1832Lacerta Schreibersiana var. a.fusca Gachet (p. 238) uppermostobjective, andthepriorityrule forexample has - "Environs de Bordeaux" (Note 13a) to serve stability and not vice versa (ICZN 1999). It was 1832 Lacerta Schreibersiana var. b. lutea Gachet (p. 239) therefore our first ambitious goal to solve the nomenclat- - "Environs de Bordeaux (dans une prairie humide)" ural problems arising fromthe insightthat Jacquin (1787) (Note 13a) is not the author ofthis taxon. Hence we provide here a 1832 Lacerta chysogastra Andrzejowski - "In vallibus commented list ofall synonyms or other names connect- circa Cremenecum". (Note 14) ed withZootoca vivipara. Such a list is certainly ofinter- 1835 Lacerta dejacquin Cocteau-"(novaactaHelvetica est because it might well be possible that further refine- 1787, page 33 Icon tab. 1)" (Note 15) ment ofthe infraspecific structure ofZ. vivipara and con- 1835 Lacerta guerin Cocteau - "La Foret d'Eu". (Note sequently further taxonomic splitting can happen. This 16) Bonn zoological Bulletin 60 (2): 214-228 ©ZFMK 216 JosefFriedrich Schmidtler & Wolfgang Bohme 1837 Zootoca montana Tschudi -"Wahrscheinlich das sel- (see text in fig. 1) on occasion ofSchrank's excursion to tenste schweizerische Reptil... Gebirgsbewohnerin". Mount Schwarzort near Berchtesgaden (then a clerical (Note 17) principality immediate to the so called Old German Em- 1837 Zootoca alpina Tschudi - "Schweiz". (Note 18) pire; now situated in southeastern Bavaria). The identifi- 1840 "Lacerta isidori Geoffi-r (sic!), Schinz (Note 19) cation as a viviparous lizard is nonetheless obvious: 1872 Lacerta vivipara var. pallida Fatio "Ca et la dans Schrankwas a goodadeptofthe local herpetofaunawhich nos Alpes et dans le Jura". (Note 20) he registered in the same work in his "20. Brief iiber 1897 Lacerta vivipara var. carnioUca Werner - Krain Berchtesgaden" (1785b). The simultaneous and progres- (Schneeberg). (Note 21) sive use of Laurend's (1768) new generic and specific 1898 Lacerta (Zootoca ) vivipara var. melanogastra Pra- names forthe species observedby Schrank himselfin this zak - "Leitomyschl, Carlsbad ", Bdhmen (Note 22) territory demonstrates well that he was able to differenti- 1902Lacerta vivipara var. barabensis Kashchenko~"Ba- atebetween lizards andsalamanders: Proteus tritonius (the rab", near Tomsk (Note 23) larvae oftheAlpinenewt, nowIchthyosaura alpestris; see 1923 Lacerta vivipara var. gedidyi Fejervaryi - "Felso- Schmidtler 2007), Salamandra atra, Salamandra macu- Babad, commune Ocsa, dep. Pest" (Note 24) losa, Seps viridis (here probably the male of the sand & A 1963 Lacerta vivipara sachalinensis Pereljeshin Ter- lizard,Lacerta agilis) beside ''Seps? atra''). certain un- entjev - Type locality not given, but by implication sureness concerningthe genus is mirroredby the question Sakhalin Island. (Note 25) markbehind Laurenti's new generic nameSeps forlizards. 1968 Lacerta vivipara pannonica Lac & Kluch - "Bo- In the case at handthe evidence ofquickness, colouration, t'any, Kapushansky Wald" (Note 26) the long tail andthe slenderbody are surelycrucial. There 2000 Zootoca vivipara carniolica Mayer, Bohme, Tiede- may be added the ecological description ofthe Schwar- mann & Bischoff- "Slovenia: Mt. Sneznik: 8 km SE zort mountain in the preceding "19. Brief (Schrank m Masun village, 1250 a.s.L". (Note 21) 1785a) which is pointing to a typical alpine montane for- 2009 Zootoca vivipara louislantzi Arribas - "Pla de Be- est. According to our present knowledge this altitudinal ret (Vail d'Aran, Lleida). Spain" (Note 27) belt in the Northern Calcareous Alps is characteristically inhabited by Zootoca vivipara. In the subsequent litera- ture this taxon is apparently alluded only once, by Jack- II. Notes el (1871: 85), who correctly related it to the melanistic Lacerta nigra Wolf(see below note 7). 1. Zootoca Wagler, 1830 Seps atra (grammatically and according to Art. 34.3 ICZN nomenclaturally correct, would be: ''Seps ater"; Lacerta vivipara is the type species ofZootoca Wagler, Seps has, contrary to Zootoca, a male gender; cf. Lauren- 1830 (see Stejneger 1907: 251); the later determination ti, final sentence ofthe "Errata": „Si Sepem inveneris in ofZootoca crocea by Fitzinger (1843: 20) is irrelevant, genere feminino lege masculinum") is therefore the old- since the type detennination had been carried out by Wa- est available synonym in the genus Zootoca. It is howev- gler(1830: 155) himself(by monotypy;Article 68.3 ofthe er a "nomen oblitum" (compared with vivipara; see be- Code). Wagler listed indeed Lacerta crocea beside other low note 3), but being available as a taxon ofthe species nomina, but they have been added in bracketsjust behind Lacerta vivipara. According to Wagler's handling, these nomina are deemed synonyms. Zootoca was re-elevated to full generic rank by Mayer & Bischoff(1996). 5*3. ®d>ti>arjc &k(f>fc? (Seps? atra.) In addition, Wagler (1830) provided also the basis for the modem generic and specific systematic concept and ' fyAz bi'cfmf^ ?t>«*r ni(fet ^inJanglit^ gf. the current nomenclature ofEuropean lizards, next to Z. f«fi«n; ttidlcic^ft roar fc^irflr^ei; SDJoId?; okr €6 vivipara also fore.g. Lacertaagilis andPodarcismuralis, Wat bod) fcf)n<fl, unb bcr ©djtoonj fd)ifn mir Did although his generic concept was widely accepted much ti'mner unb \m^tx bep ben 9)^oId)en , unb bee later, since the early 1990s (see Schmidtler 2010: fig. 1). ^^orpcr frfilanfer. 5* ^ ' tSolbc imterm @^morj« & orfe (juf ftnem fflulen ^aumjlocf« gefe^wi, flHeln 2. Seps? atra Schrank in Schrank Moll, 1785 <if5 id) grclffn rooUle, roar n>eg, unb nic^t (Fig. 1) incf)c in finb^n. Even ifseen on the background offormer contemporary practices, the original description appears highly strange. Fig. 1. Text ofthe original description ofSeps atra Schrank, It corresponds only to a sight record of several seconds 1785. For translation see Note 2. Bonn zoological Bulletin 60 (2): 214-228 ®ZFMK Synonymy and nomenclatural history ofZootoca vivipara (Lichtenstein, 1823) 217 group, if the taxonomical distinctness of this Berchtes- fig. 3), which contains also manynew species descriptions. gaden taxon should emerge. (Art. 29 ICZN). From that time onwards (especially since Dumeril & The translation of Schrank's figure legend (Fig. I) is: Bibron, 1839) this name became more and more in gen- "53. Black lizard? (Seps? atra.). Ididnotsee this lizard eral use. The peculiarityhere is that inthis "Verzeichniss" sufficiently: perhaps it was a black newt; but it was by Lichtenstein 1823 the history ofdiscovery and the de- nonetheless quick, and the tail seemed to me much thin- scription of "L(7C. vivipara. Jaquin" (sic!) are treated in nerandlongerthan in newts, andthe bodyslimmer Ihave detail, but that this taxon was "only" ranked as a synonym seen it in theforest below of(Mt.) Schwarzort on a rot- under"14.Lacerta muralis Merr." (textunder fig. 3). But, ten tree trunk, however, when Iwanted to catch it, it dis- by virtue oftheArticles 11.5,11.6, 50.7 ICZN this fact is appeared, and was not to be traced again. " sufficientto make the nameLacerta vivipara from its type locality "Mons Schneeberg" (= Mt. Schneeberg, west of Vienna) available. 3. "Lacerta vivipara^'' Jaquin, 1787 and Lacerta vivipa- The oldersynonyms (between 1788 and 1820: atra, oe- ra Lichtenstein, 1823. (Fig. 2) dura, nigra, crocea, montana etc., see the synonymy list above) must not be considered as threatening the citation 3.1. Only a "viviparous lizard" but not an original de- ofviviparaby Lichtenstein(1823): None ofthemwasused scription. as a valid name after 1899 (Art. 23.9.1.1 ICZN), and it is no question here that Lacerta vivipara was validly used As mentioned already inthe introduction, J.F. de Jacquin in at least 25 publications by at least 10 authors in the last (1787) entitled a short note as follows: „Lacerta vivipa- 50 years (Art. 23.9.1.2 ICZN). ra, observatio Jos. Francisci de Jacquin".Afterhaving de- scribed the context ofhis discovery and the morphology ofa female with its hatchlings bom shortly after the cap- 3.3 The correct authorship ofLacerta vivipara ture, he stated that he would not dare to decide to which speciesthis specimen couldbe assigned, He even consult- "H. Lichtenstein" isheadedon thetitlepage ofthe"Verze- ed Linne's work in vain, who did also not seem to him to ichniss" (1823), but this fact does not exclude a possible know much in tenns ofmany other species of"ourAus- coauthorship ofotherpersons forparts ofthe publication tria". Thus the aforementionedcaption shouldcertainlynot (Art. 50.1 ICZN). So, among others, Tschudi (1837: 29) denote a new species but rather was intended as the sci- had nominated "Schulze" (sic!) without any substantia- entific communication ofa sensational biological anom- tion as a coauthorofthe mentioned citation. This was ap- aly, namely a "viviparous lizard" (Latin: "Lacerta vivip- parently based on Lichtenstein's own statement on his in- ara")within the class ofthe "oviparous quadrupeds" - the troductory page IX: ,J)amit ich mir nicht allein anzuma- present-day amphibians and reptiles (see Bohme & Rod- fien scheine, was Verdienstliches an dieserkleinenArbeit der2006; Schmidtler& Bohme 2006). Cocteau (1835; see sein mag. so darfich nicht unenvdhnt lassen, dass zwei note 14) was the first to spot these facts, but they were meinerwerthen Gehiilfen, HerrDoctorRodig. (jetztPro- forgotten shortly after publication ofhis work. '"Lacerta fessorinMUnster) undHerrStud. Med. Ferd. Schultze von vivipara Jacquin, 1787" had therefore not become an Halle an dem Verzeichnifi derAmphibien undFische ei- available name (Art. 11.5 ICZN). nen wesentlichenAntheilhaben. Namentlich sinddie ein- gestreuten Bemerkungen iiberdie variablen EidechsenAr- ten unddieFassungdermehrsten Diagnosen in den letz- 3.2. The searchfor the correctspecies name. ten 4 Bogen ganz des letztern Werk, und von demselben wdhrendmeinerAbwesenheit von hier zum Druckbefor- According to theprinciple ofpriority (Art. 23.3.5 ICZN), dert' which reads in our translation: ,,In order not to as- thename vivipara would generally have to be replacedby cribe everythingto myselfwhatcouldbe meritious in this the next-oldest available synonym (see listing in chapter small work, I may not leave it unmentioned that two of I). However, the principle ofpriority is not a value in its my estimatedhelpers, Dr Rodig (nowprofessoratMUns- own, but it is a means to serve stability and universality ter) and Mr Stud . Med. Ferd(inand) Schultze ofHalle ofnomenclature (Preamble and Art. 23.2 ICZN). There- haveconsiderablyparticipatedin the list ("Verzeichniss") fore, names oflong acceptance should not be invalidated ofamphibians andfishes. Particularlythedispersedcom- butpreserved ifthis can anyhowbe achieved. This is par- ments on the variable lizard species and the version of ticularly true for the name of such a famous and wide- most ofthe diagnoses in the last 4 sheets are completely spread species as in this case! the latters work and were also sent by bim to theprint- , As far as noticeable for us, the name Lacerta vivipara er during my absence. " was first used as a "real" species name - and also by re- This highlyprecious andmannered information does not ferringto Jacquin-inLichtenstein's (1823) catalogue (see represent in itselfa proofthat F. Schultze really authored Bonn zoological Bulletin 60 (2): 214-228 ©ZFMK ; 218 JosefFriedrich Schmidtler & Wolfgang Bohme 4. Lacertus cinereiis Lacepede, 1788 (Figs 4, 5). 4.1. A confusingcollectivespeciesandan unavailablebi- nomen. This binominal taxon is mentioned by Lacepede (1788) only in his „Synopsis methodicaquadrupedum oviparum" being adherent to his herpetological encyclopedia ("L. cinereus^" underthe caption '"Lacertus''''). Apart from that, Lacepede was always speaking of the "Lezard gris" (= Grey lizard; cf. also "Lacertus cinereus'' and "Graue Ei- dechse" in Bechstein 1800: 537). However, Lacertus cinereus does not anymore figure in any contest with oth- er available taxa, since Lacepede"s new taxa in this first Fig. 2. "Lacerta vivipara" from de Jacquin (1787); adult fe- and not constantly binominal volume ofhis encyclopedia emnanlae.w-itNhotites 3n.ewborn offspring, from Schneeberg, west ofVi- were declared as not available („Lacepede, B.G.E. de la V. 1788. HistoireNaturellede Quadnipedes Ovipares, and all subsequent editions ofthis work ruled to be not avail- IIL Amx^hibien, 93 Thlr. Lac. vfvlpara. Jaqui'n. *) X^ae- collari integen'imo adnatO) squamis dorsi et latermn laevibu^ sexungularwiiSi squ> caudae annulatlm ahsclssis supra suhcai'i- nails. Os aciitius qnam in Lac- a g i 1 i. Linn. Lac. s1 1 rp iu m. Dand, series scutorum ..aodondnalium jx-sc portfcmorcdss 17- '^*) In me' *, . dio dorso Unea inbcrmpla pancLorum mgronml, sed sacpius deest. Fig. 3. Excerpt from the description ofLacerta vivipara in Lichtenstein (1823) as a synonym under "14. Merr. L. muralis. p. 67"(precedingpage 92). Belowpagenumber93 seetheprice columnwiththeabbreviation"Thlr" (=Thaler), atthattimethePruss- ian cuiTcncy. The publication ofLichtenstein (1823) belongs to an unusual and rare literary genre in which sales catalogue andzo- ological science are combined (see Junk, 1926-1936: 192, 198). Numerous animal species are described here under specification ofthe selling price per specimen. The explanations ofthe reptiles are at great length (see also Schmidtler 2004). -Note 3. the citations on Lacerta muralis or Lacerta vivipara, re- able for nomenclatural purposes, and no name to acquire spectively. Even the instance that F.S. Leuckart (see the status of availability by reason ofhaving been pub- Leuckart 1841: 14. footnote 64) had discussed with F. lished in any edition. Op. 2104"; see ICZN 1987, 2005). Schultze on the viviparity ofthe viviparous lizard ("1. cro- cea"), caimot be used as an argument. Therefore, Licht- enstein, authorofthe "Verzeichniss (1823)", shouldbe re- 4.2. Nonetheless: A key taxon important for the under- garded also as the author ofthe taxonomic parts that deal standing ofthe history ofsystematics. with lacertids (Art. 50.1 ICZN). We conclude that the correct scientific denomination The story ofLacertus cinereus seems well appropriate to of the viviparous lizard is now Zootoca vivipara understand the hopeless confiisionwhich arose at the end (Lichtenstein, 1823). ofthe 18'h century in the systematics of lizards (and not Bonn zoological Bulletin 60 (2): 214-228 ©ZFMK Synonymy and nomenclatural history ofZootoca vivipara (Lichtenstein, 1823) 219 SCAJ.Y^ lilZAILD. Fig. 5. Excerpt with "Lezard gris (Lacerta agilis)" from Bonnaterre (1789: 44 and pi. 6, fig. 2). This copy ofPennant's Scaly lizard (=Zootoca vivipara; see fig. 4) was intended to represent a current French Podarcis muralis. See some single falsifications because ofthis cribbing: The more reduced pileus scalation, the reduced spotting upon the dorsum and the even scalation ofthe tail compared with the verticillate and rugose formadon ofthe original. -Note 4. only in their systematics!). Here, Linnaeus (1758) and Bibron 1839, Schmidtler2004, 2010, Kuzmin 2005: 242). Lacepede (1788) may be characterised as "lumpers". In It is noteworthy that Zootoca vivipara is not involved in theirspeciesLacerta agilisandLacertus cinereusILezard this splitting bacchanal. gris there are comprisedmore orless clearat leastthe sand As indicated,Lacertus cinereusLacepede (themale gen- & lizard(currently: Lacerta agilis Linnaeus), the green lizard der represents an unjustified emendation; cf. David al. (currently: Lacerta viridis Laurenti or L bilineata, 2002: 24) comprises also Zootoca vivipara. This fact is Daudin), the wall lizard(currently: Podarcismuralis (Lau- less identifiable from the ambiguous morphological de- renti)), the viviparous lizard (currently: Zootoca vivipara scriptions, but in fact from a publication on its vivipari- (Lichtenstein) see the comprehensive discussion in Ley- ty, being published covertly (Lacepede 1789: 491, "Ad- dig (1881: 161)). Contrary to them, Laurenti (1768) and ditions"), and forgotten later on. This citation worried Daudin (1802) took the line of an extreme "splitting". Lacepede. He only found a way by speculating that the They based new species uponjuvenile forms, colour va- grey lizards could not help but the salamanders in fact & rieties, males and females (for details see: Dumeril could (then also in the Linnean genus Lacertal), the fe- Bonn zoological Bulletin 60 (2): 214-228 ®ZFMK 220 JosefFriedrich Schmidtler & Wolfgang Bohme males ofwhich are sometimes egg-laying and sometimes By theway, Sheppard's article contains two furthernew giving birth to living young. Observations like these be- "Lacerta" species: Lacerta anguiformis ("*An Lacerta gan to shake systematics ofthat time. There intrinsically aquatica Linn.?" = Lissotriton vulgaris ?) and Lacerta oviparous quadrupeds and snakes (both in the class of maculata (a younger homonym ofbothLacerta macula- "amphibians") were facing the viviparous quadrupeds ta Shaw, 1802 (now: Ambystoma maculatum) \mALacer- (mammals). ta maculata Daudin, 1802); the latterdubious taxon is ap- parently not enclosed in later synonymy listings. 4.3. Problems ofspecific identification: Differentin every country. 6. Lacerta montana Mikan in Sturm, 1805 (Fig. 6) Ultimately Bell (1839: 22) hit the mark when discussing The „Riesengebirge": This mountain range forms today the use ofthe name ''Lacerta agilis'": „...thattheLinnean the border between Poland and the Czech Republic. See term agilis has been applied by zoologists ofdifferent also Zootoca montana Tschudi, 1837 (note 16). countries to thatspecies ofLizardwhich is bestknown or See in detail Leydig (1881 : 166) who showedthat Lin- mostcommon in theirown. Thus thePodarcis muralis, the naeus (1758: 203 and 1766: 363 ) had lumped together at common lizardofItalyandofFrance, has been so called leastthe current speciesLacerta agilis andZootoca vivip- by Italian or French writers; and our own little indige- ara underthe name Lacerta agilis. Wolfand Mikan were nous species, sofrequent in almost allparts ofEngland, the first who partitioned the Linnean collective speciesL. which Ishallpresently describe under itsproper appel- agilis in the fourth issue of Sturm's "Deutschlands Fau- lation ofZootoca vivipara. has hitherto receivedthesame na"(1805) The first importantprogress hadbeen madeby namefrom everyBritish naturalistwho has written on the Wolf in the second issue of Sturm's work when he de- subject''. scribed andmagnificently figuredthe male andthe female A good example is the unscrupulous cribbing ofPen- ofthe current species L. agilis under this name (see also nant's "Scaly lizard" (1776: fig. 4 this paper; definitely Schmidtler 2004). However, the current species Z. vivip- the female ofa British Zootoca vivipara by Bonnaterre ara was introduced by Mikan and Wolf as no less than ) (1789; fig. 5 this paper), who believed it a (current) French three different species see Note 7 & 8)! The mingling of Podarcis muralis but named it still Lacerta agilis the two species is still to be found as late as Brandt & ("Lezard gris")! Ratzeburg (1829) who reportedunderL. agilis also vivip- The factthattodaythebinomen ''Lacerta agilis'" is iden- arous individuals. - For more details about the complex tified with the sand lizard and not with the viviparous problems within collective species see note 4. lizard, depends lastly on Wolf in Sturm who had uncov- ered both taxa, living in Nuremberg, to be independent species and to describe and depict them elaborately (See 7. Lacerta nigra Wolf in Sturm, 1805 (Fig. 6) Wolfin Sturm 1799: some coloured images ofmales and females ofthe sand lizardLacerta agilis; depicted also by Thetype locality is "Schneegebirge, so genannteWenger- Schmidtler 2004. Wolf in Sturm 1805: below notes 6-8 Alpe", in the canton of Berne, Switzerland. See also and figs 6, 7 hoc loco). It was finally Wagler (1830; see Tschudi (1837: tab. 1), who regardedthis blackvariant as note 1 ) who leveraged Wolf's nomenclatural classifica- a variety ofhis Zootoca montana (note 16). tions togetherwith its basic systematical contents (see al- so Koch in Sturm 1828). 8. Lacerta crocea Wolfin Sturm, 1805 (Fig. 7) 5. Lacerta oedura Sheppard, 1804 Wolfwrote that he had receivedthis lizard from "hiesiger Gegend" (= "local region") whereby he meant his habi- "At different times found in vast abundance": Sheppard tation, the then Imperial City ofNuremberg (Niimberg). offers no exact information on the geographical origin of This taxonwas considered byWagler(1830) as a synonym his new species. However, it results from the caption and of Zootoca vivipara (cf note 1 and Dumeril & Bibron introduction ofhis article ("British lizards") that Britain 1839), but nevertheless persisted in the literature some has to be understood as type locality. decades as an independent species. This taxon is citedby Tschudi (1837: 3) underthe name The viviparous lizardwas firstdepictedbyNikolaus M. "Lacerta oedura Scheppard" (sic!) and undergoes a fur- Oppel (1782-1820) in aperfectmanner(Schmidtler2008: ther "mutation" in the paper by Schinz (1840: 19 as "La- 22, fig. 12), butremained subsequentlyunpublished. This certaaedura Scheppert(sic! See fig. 10), a spellingwhich figure stimulated Leydigto ahymn ofpraise inhis (1872: canbe found again in some laterreferences (Erber 1868). 224) historical chapter on "Lacerta vivipara". Bonn zoological Bulletin 60 (2): 214-228 ®ZFMK Synonymy and nomenclatural history ofZootoca vivipara (Lichtenstein, 1823) 221 Fig. 6. "Iconotypes" ofLacerta nigra Wolfin Sturm, 1805 (above) andLacerta montana Mikan in Sturni, 1805 (undesignated plates). - Notes 6 and 7. ®ZFMK Bonn zoological Bulletin 60 (2): 214-228 222 JosefFriedrich Schmidtler & Wolfgang Bohme Fig. 7. Different specimens in different positions ofLacerta crocea Wolfin Sturm, 1805 (Undesignated plates; "iconotypes"). -Note 8. iA & I 9. Lacertafragilis Palmstruch Swartz, 1808 (Fig. 8) The name Lacerta "fragilis " turns up only in the "Sys- tematiskt Register" at the end ofvolume I of"SvenskZo- ologi". There is a hint to no. 27 (bearing the caption : "Gra OdiaLacerta agilis... " in the first volume (part 5)) ofthis work. According to the main title page of« Sven- skZoologi" this work was founded in 1806 by C. Quensel andJ.W. Palmstruch.AfterQuensel's death in 1806 itwas continuedby J.W. Palmstruch and O. Swartz as displayed upon a new title page having been sent by the editor with part six in 1808 (R. Wahlgren and B. Dal in litt. 2011). According to the situation described above, this nam- ing of""Lacertafragilis" is therefore probably a lapsus. It refers indeed to the authors' ""Lacerta agilis'" (currently : Zootoca vivipara) then the collective species first de- , scribed by Linnaeus (1758; see also 1766 and Gmelin 1788: 1070; the latterreferencebeing explicitely citedby the authors). Concerning this Linnean collective species ""Lacerta agilis'", more details can be found under note 4 and 6. Fig. 8. Lacerta agilis resp. "'Lacertafragilis" from Palms- truch & Swartz (1808). -Note 9. Bonn zoological Bulletin 60 (2): 214-228 ©ZFMK Synonymy ;ind nomenclatural history ofZootoca vivipara (Lichtenstein, 1823) 223 10. Lacerta unicolor Kuhl, 1820 ed: Zootocapyrrhogastra seemedto be the most adequate name for this genus (meant is species) ofWagler's genus Origin and fate of the holotype are unknown. Wagler Zootoca, because it characterizes exactly the coloration (1830: 155) quoted it under the comment "Mannchen (pyrrhogaster = "fire bellied"). Zootoca vivipara is non- adult, decolor." in the list ofhis synonyms ofZootoca vi- sense since both names are meaning the same"). See al- vipara. Cocteau (1835: 10) assimied that Wagler might so note 19. ha\e examined this specimen since the original descrip- tion was very enigmatic and highly insufficient ("a peu pres enigmatique et au dessous de toute critique"). Kuhl's 13. Lacerta schreibersiana Milne Edwards, 1829 consideration beside the vacuous description is remark- able: that the shape and sizeresemble entirelyL.ptychodes "Envoyes deVienne. parM. Schreibers" (see Milne Ed- .. (seethe followingnote 1 1). SubsequentlyL. unicolorwas wards 1829: pi. 5, fig. 5.): Carl Franz Anton Ritter von regardeded a dubious synonym ofZootoca respectively Schreibers (1775-1852) was then director ofthe Vienna Lacerta vivipara, cf Mertens & Wermuth 1960). Accord- „Naturalienkabinette". The specimens were „sent fromVi- ing to Hildenhagen (in litt. 201 1) the acronym 'icon O." enna"byhim. This wordingdoesnotnecessarily mean that applied here and in some other ofKuhl's species descrip- the specimens were also collected in or nearVienna. Ac- tions, means probablythatno figure was available ("Icon: cording to our present knowledge the viviparous lizard zero") does not occur today in the immediate vicinity ofVienna . but has few relict populations in the Viennese Basin (Ca- bela et al. 2001). 11. Lacertaptychodes Kuhl, 1820 („mihi") and Lacer- ta ptychodes Merrem, 1820 13a Lacerta Schreibersiana var. a.fusca Gachet, 1832 It seems to be intriguing that in 1820 the same specimen (p. 238) and Lacerta Schreibersiana var. b. lutea Ga- ofunknown origin was documented with the same name chet, 1832 (p. 239) by two different authors independently from each other. Indeed, Kuhl (1820: 121) mentioned an earlier contact Fiisca and hitea are nomina oblita, but surprisingly not with Merrem because ofthis specimen, but both articles comprised in Dumeril & Bibron (1839), and apparently reveal in each case that the original descriptions were au- forthe lasttime mentionedby Lataste (1876 : p. 82 + foot- thored without any reference whatsoever. Both descrip- note). Gachet's/i«C(3 represents also ayoungerhomonym tions are comparativelycomprehensive, inwhich Kuhl ac- of Lacertafusca Daudin 1802 (nomen substitutum pro centuated the colour pattern and Merrem the pholidosis Seps terrestris Laurenti 1768 =Lacerta viridis viridis Lau- (Merrem : die" runzelige Eidechse" = "rugose lizard"). In renti). On the other hand lutea is an available name en- both cases, Zootoca vivipara might have been hold very dangeringZootoca vivipara louislantziArribas, 2009. The well. In the later literature a lizard named Lacerta pty- description oflutea is very detailed but does not seem to chodes did never appear again, (see also Hildenhagen be absolutely certain in orderto ascertain synonymy (Ar- & 2010: 53 and Hildenhagen Hallermann 2010: 61) . ribas in litt. 2010). Further investigations are necessary. 12. Lacertapyrrhogaster Merrem, 1820 14. Lacerta chrysogastra Andrzejowski, 1832 Tschudi (1837: 27) apparently interpreted Merrem's "In vallibus circa Cremenecum" (Latin): "The valleys species name "'pyrrhogaster" as an adjective andtherefore around Cremenecum". Cremenecum is today Cremeniec, adapted the ending to feminine gender (Zootoca a city situated 100 km EastofLemberg(=Lwiw, Ukraine). pyrrhogastrd"). He interpreted Zootoca as a subgenus comprisingtwo species: Z.pyrrhogastra and"Z. nwntana nobis" (see note 16). 15. Lacerta dejacqiiin Cocteau, 1835 Tschudi's footnote on page 30 reveals that the princi- ples ofpriority and a common authoritative terminology Cocteau's quotation of de Jacquin's article suggests an were not yet comprised in the then usual understanding original description with a new name but is rather a re- ofnomenclature: ^Zootoca pyrrhogastra schien mir der placement name, at least with respect to Lacerta vivipa- passendste Name fur diese Gattung des Wagler'schen ra Lichtenstein, 1823 with the same type locality Genus Zootoca, da es die Farbung (= "feuerbauchig", "Schneeberg". From his point ofview Jacquin's publica- Anm. d. Verf.) genau kennzeichnet. Zootoca vivipara ist tion in 1787 didnot constitute avalid description ofa new Unsinn, dabeide Namen das Gleiche bedeuten" (translat- species (so explicitely upon his page (6; not paginated). ®ZFMK Bonn zoological Bulletin 60 (2): 214-228

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