On the distribution and chromatic differentiation of the … TREVISAN 1982). Afterwards only few data were published on the distribution of this new form (see: GROSSENBACHER 1994)

HERPETOZOA 13 (3/4): 171 -180 171Wien, 30. Dezember 2000

On the distribution and chromatic differentiation of theAlpine Salamander Salamandra atra LAURENTI, 1768,

between Val Lagarina and Val Sugana (VenetianPrealps): an updated review

(Urodela: Salamandridae)

Zur Verbreitung und Farbdifferenzierung des Alpensalamanders,Salamandra atra LAURENTI, 1768, zwischen Val Lagarina und Val Sugana

(Venetianische Voralpen): Ein aktualisierter Überblick(Urodela: Salamandridae)

Lucio BONATO & KURT GROSSENBACHER

KURZFASSUNG

Publizierte und unpublizierte Daten zur Detail-Verbreitung und zur Farb-Differenzierung des Alpensala-manders {Salamandra atra LAURENTI, 1768) in den südlichen Voralpen zwischen Val Lagarina und Val Suganawerden zusammengetragen und kritisch betrachtet. Bekannt sind Populationen vom nordwestlichen Teil der Hoch-ebene der Sette Comuni (Sieben Gemeinden) und von den Südhängen des Pasubio-Massivs, sehr wahrscheinlich gibtes Vorkommen aber auch im Gipfelbereich des letzteren sowie auf der Kette des Cornetto und im Carega-Massiv. We-nigstens zwei, wahrscheinlich allopatrische Typen von Populationen konnten aufgrund der Farbmerkmale unter-schieden werden: solche mit ausgedehnt gefleckten Individuen (beschrieben als S. atra aurorae) auf der Hochebeneder Sette Comuni, und solche mit schwach gefleckten, manchmal ganz schwarzen Individuen auf dem Pasubio-Massiv. Ein dritter Typ, bei dem völlige Schwarzfärbung häufiger auftritt, existiert möglicherweise auf Gebirgs-massiven im Südwesten.

ABSTRACT

Published and unpublished information was compiled and critically reviewed, in order to present an updatedsynthesis on the micro-distribution and the chromatic differentiation of the Alpine Salamander (Salamandra atraLAURENTI, 1768) in the Prealps between Val Lagarina and Val Sugana. Populations are known from the north-western part of the Sette Comuni plateau and the southern slopes of the Pasubio massif. The Alpine Salamander ismost likely present also at the top of the latter relief, on the Cornetto ridge and on the Carega massif. At least two typesof populations, probably allopatric, could be distinguished on the basis of chromatic features: those of extensivelypatched individuals (described as S. atra aurorae), on the Sette Comuni plateau and those of poorly patched individu-als, rarely completely black, on the Pasubio massif. A third type, in which entirely black individuals are more frequent,is supposed to exist on massifs in the south-west of the area.

KEY WORDS

Amphibia, Urodela, Salamandridae; Salamandra atra, S. atra aurorae, distribution, coloration, geographicdifferentiation,Venetian Prealps, Italy

INTRODUCTION

The occurrence of the Alpine Sala- S. H. I. sez. Lombardia 2000 - for Lom-mander Salamandra atra LAURENTI, 1768, bardia; CALDONAZZI et al. in press - for theis strictly limited to montane sites along Trento Province; Gruppo Nisoria in press -the Alpine and Dinaric chains (S. H. I. for the Vicenza Province; Gruppo per la re-1996; GROSSENBACHER 1997). The actual alizz. Progetto Atl. Erpet. Veneto 1996 -distribution is still only roughly known, for all of Veneto; LAPINI et al. 1996 - forparticularly in marginal regions such as the Friuli-Venezia Giulia).Southern Prealps. Here, recent investiga- Despite this situation, in the last twotions added new information but do not decades some findings and preliminaryseem to have yet reached a satisfactory studies brought the first insights into anview of the distribution of the species (see: interesting case of geographic colour differ-

©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

172 Lucio BONATO & KURT GROSSENBACHER

entiation within the species. This involvedpopulations of the Venetian Prealps, par-ticularly the Sette Comuni plateau and thePasubio relief. At the end of the Nineteen-seventies, an unusual population charac-terised by largely yellow-patched individu-als was discovered in a limited site on theSette Comuni plateau (TRE vis AN et al.1981; TREVISAN 1982). Afterwards onlyfew data were published on the distributionof this new form (see: GROSSENBACHER1994). As a result, studies on the geneticdifferentiation were carried out on animalsfrom a single site only until now (TREVISANet al. 1984; JOGER 1986; OLIVIERI 1991;STEINFARTZ et al. 2000). More recently,another interesting population, character-

ised by a less extensively patched colora-tion, was discovered on the Pasubio massif(BONATO 2000), suggesting somehow com-plex relationships among the populationsof Alpine Salamanders living in the regionbetween of Val Lagarina (Adige Valley) andVal Sugana (Brenta Valley).

This paper is an update of our knowl-edge on the distribution and differentiationof S. atra in the above region, integratingthe few published data and unpublishedinformation. Our aim is to present a back-ground (though a preliminary one) to fur-ther analysis of this interesting case of geo-graphical differentiation and to any actionplan for the conservation of these endemicforms.

STUDY AREA AND METHODS

Study area

Our survey was limited to the part ofthe Venetian Prealps between Val Lagarina(west) and Val Sugana (north and east).The region can be regarded as a naturalgeomorphological and biogeographical unit,being delimited by deep glacial valleys,that work as dispersal barriers for the Al-pine Salamander today. Moreover, all avail-able information suggests that an apprecia-ble differentiation of these animals can beobserved only inside this region.

The western part of the territory isoccupied by steep massifs, often more than2,000 m high, cut by several valleys andgradually degrading southwards to the Les-sinian Hills and the Venetian Plain. Fromsouth-west to north-east the following ma-jor mountainous groups can be recognised:Lessini, Carega, Cornetto, Pasubio, Cam-pomolon and Toraro, Becco di Filadonna.

The eastern part of the territory, in-stead, appears as a single large plateau(Sette Comuni plateau), about 1,000 m highbut more elevated on the northernmostpart; only few gorges open towards its core(Val d'Assa, Val Frenzela).

The western massifs and the easternplateau are separated by a fluvio-glacial in-cision, running about north-south, repre-sented by the deep Val d'Astico and thevalley of the Centa stream. A rather narrowridge, not more than 1,050 m high, seemsto be the only potential bridge present for

the dispersion of the Alpine Salamanders(BONATO 2000).

In the highest parts of the whole ter-ritory, the substrate is mainly dolomitic orcalcareous, consequently the landscape iskarstic; this feature characterises most ofthe Venetian Prealps and distinguishesthem from the true Alps. Average tem-peratures of +10/+20 °C in July, -5/+5 °Cin January, and precipitation intensity of1,500-2,500 mm/year are registered (FLIRI1975).

Methods

Information on the presence of Al-pine Salamanders was gathered by differentmethods: planned field researches, carriedout in the last years by the authors and oc-casionally by other persons; critical reviewof the available published data; requests forinformation directly addressed to personsoften visiting montane sites for differentpurposes and considered reliable in re-porting occasional encounters with theseanimals (e.g., botanists, entomologists andamateur naturalists).

Both published observations andother apparently reliable information wereevaluated according to the reliability de-gree as we could judge.

Each record was mapped with a pre-cision of one kilometre by assigning geo-graphical coordinates according to theUTM system (relative to the zone 32T).


Alpine Salamander (Salamandra atra) in the Venetian Prealps 173

Toponyms and UTM coordinates were de-rived from the last edition of the Cartad'Italia published by the Istituto GeograficoMilitare (Firenze).

The individuals observed were as-signed to the following chromatic types(fig. 1): body largely patched (P+) like inall S. atra aurorae TREVISAN, 1982 de-scribed from Bosco del Dosso (TREVISAN1982; KLEWEN 1988; BONATO 1998); bodypoorly patched (P-), like in most individu-

als described from the southern slopes ofMt. Pasubio (BONATO 2000); body com-pletely black (B), as is typical in S. atraatra according to all of the literature.

When permitted by the data available,we tried to assign each local population toone of the following chromatic types: (i) allindividuals largely patched; (ii) most in-dividuals poorly patched but some of themcompletely black; (iii) most or all individu-als completely black.

RESULTS

The collected data on distribution andchromatic differentiation of the AlpineSalamander between Val Lagarina and ValSugana are reported in table 1 and mappedin figure 2. They are summarised as fol-lows for each apparently suitable mountaingroup:

• Sette Comuni plateau. This wasthe most intensely investigated area: sincethe beginning of the Nineteen-eighties (af-ter the publication of the discovery of S.atra aurorae), many researchers visited theplateau, particularly north of Asiago. As aresult, a relatively rich series of observa-tions was gathered.

Alpine Salamanders are found alonga stripe on the left of the Val d'Assa andnorth of the central basin, from Passo Vez-zena to Gallio; populations were detectedboth in woody slopes exposed southwardsand in shrubby elevations exploited in thepast as meadows, from 1,200 to 1,800 m.Despite the research efforts, no individualswere found in other apparently suitablesites, such as the northernmost ridges, theeastern part of the plateau and the rightside of Val d'Assa (just vis-à-vis of colo-nised slopes).

Only largely patched individualsseem to be present, their habitus agreeing

with that of the typical S. atra aurorae.Rather old findings of completely black in-dividuals reported in the literature (see:TREVISAN 1981; KLEWEN 1988) were notconfirmed by recent surveys and have to beregarded as unreliable.

* Becco di Filadonna. No data .* Campomolon and Toraro. No data.* Pasubio. A population of poorly

patched individuals, occasionally com-pletely black, is found at least on the south-ern slopes and gorges, between 1,500 and1,900 m.

Apparently black individuals wereseen on the upper plateau of this relief,from 1,800 to 2,000 m; these observations,however, are variable in reliability and un-til now our researches failed to confirmthem.

* Cornetto. An undocumented ob-servation of an apparently black individualwas reported from a steep slope, above1,500 m.

* Carega. A completely black indi-vidual was reported to have been collectedon the upper part of the massif, but unfor-tunately the preserved material cannot beidentified with certainty.

* Lessini. No data.

DISCUSSION

Distribution

As already suggested by GROSSEN-BACHER(1994), the range of the endemic S.atra aurorae turned out to be wider thanpreviously known from the literature.PoDulations chromaticallv referable to this

taxon appeared to be distributed in thenorth-western part of the Sette Comuniplateau, at least on a stripe about 15 kmlong and few km wide. However, whetherthe distribution is continuous or scattered ishard to argue. The geomorphological andenvironmental features of the neighbouring



P-

% *

Fig. 1 : Individual chromatic types recognised in the Alpine Salamander (Salamandra atra) in the area between ValLagarina and Val Sugana (Venetian Prealps): P+ = largely patched; P- = poorly patched; B = completely black.

Abb. 1 : Individuelle Zeichnungstypen des Alpensalamanders (Salamandra atra) im Gebiet zwischen Val Lagarinaund Val Sugana (Venetianische Voralpen). P+ = ausgedehnt gefleckt; P- = schwach gefleckt; B = völlig schwarz.

Table I (opposite page): Available data on the distribution and chromatic differentiation of the Alpine Sala-mander (Salamandra atra) in the area between Val Lagarina and Val Sugana (Venetian Prealps). For criteria adoptedand definition of chromatic types see text (Methods). Abbreviations in the column 'Habitat': A - Abies, F - Fagus,L - Larix, P - Picea.

Tab. 1 (gegenüberliegende Seite): Verfugbare Daten zur Verbreitung und Farbdifferenzierung des Alpensala-manders (Salamandra atra) im Gebiet zwischen Val Lagarina und Val Sugana (Venetianische Voralpen). Zu denangewandten Kriterien und zur Erklärung der Fârbungstypen siehe Text (Methods). Abkürzungen in der Spalte'Habitat': A - Abies, F - Fagus, L - Larix, P - Picea.


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176 Lucio BoNATO & KURT GROSSENBACHER

areas suggest that the actual range could beeven larger. Nevertheless, S. atra auroraeremains a very local endemic form, a vul-nerable one in a conservation perspective.

Further populations of Alpine Sala-manders were found to have colonisedother montane areas south-west of the SetteComuni plateau, but only within a distancenot exceeding 30 km. While all populationsknown from the Sette Comuni plateau canbe easily referred to S. atra aurorae, allthose discovered outside the plateau areclearly different in chromatic features fromtypical aurorae. However, insufficient in-formation does not permit a satisfactorycomprehension of their actual distribution,conservation status, phyletic relations andtaxonomic position (BONATO 2000).

Outside the region considered in thisreview, the Alpine Salamander appears tobe more widespread north of Val Suganathan in the other parts of the Venetian Pre-alps. Typically black populations are knownfrom the Orobian Alps (S.H.I, sez. Lom-bardia 2000) through Adamello-Presanella(BRUNO 1973; BENNATI 1988), Gruppo diBrenta (BRUNO 1973, but not confirmed inCALDONAZZI et al. in press), Lagorai ridgeand Cima d'Asta (CALDONAZZI et al. inpress), Vette Feltrine and Dolomiti Bellu-nesi (LAPINI et al. 1998; TORMEN et al.1998) to Friuli. A minimum distance ofabout 25 km can be estimated between S.atra aurorae and the nearest black popula-tion north of Val Sugana.

From the mountains just west of ValLagarina (Bondone-Stivo and Altissimo-Baldo ridges) no populations are known.Nevertheless, more research is needed, be-cause apparently suitable sites are availableand there is an unpublished, rather reliablebut not documented observation (M. Altis-simo: la Polsa, 1,200-1,250 m, UTM co-

ordinates 661-5050; May 1986; one indi-vidual found under stones, 14 cm long,completely black; P. LORENZI, pers. comm.).

East of Val Sugana, Alpine Salaman-ders seem to be absent from the Grappamassif, despite the fact that this area is en-vironmentally similar to the Sette Comuniplateau. Less surprising is the lack of datafrom the Cesen-Nevegal ridge where theelevation is rather low and the climate lesssuitable. Further east, the Cansiglio plateauand the M. Cavallo host well known popu-lations of completely black Alpine Sala-manders, although apparently limited tofew sites (DOLCE 1988).

Ecological notes

Between Val Lagarina and Val Su-gana, Alpine Salamanders live in a widehabitat range, from beech-woods to alpineprairies, from 1,200 to 2,000 m. Ecologicalconditions do not seem to differ from thosereported for the nearest colonised SouthernPrealps (DOLCE 1988; TORMEN et al. 1998;CALDONAZZI et al. in press).

In the considered region, the AlpineSalamander and the Fire Salamander S.salamandra (LINNAEUS, 1758), were neverfound syntopically. This may depend on theecological differences between the two spe-cies and the environmental configuration ofthe territory. While the viviparous and mi-crothermic Alpine Salamander can befound only in the karstic dry plateaus andridges, never below 1,200 m, the ovovi-viparous and mesothermic Fire Salamanderis widely spread on the marginal hills, veryrich in streams, mainly in the altitudinalrange of 100-600 m and only occasionallyup to 1,100 m (Gruppo Nisoria, in press).In the Sette Comuni plateau, Fire Sala-manders can reach the southern edge of

Fig. 2 (opposite page): Distribution and chromatic differentiation of the Alpine Salamander (Salamandra atra) in thearea between Val Lagarina and Val Sugana (Venetian Prealps), as suggested by the gathered data. For each square

kilometre of the UTM coordinate system, the following information was mapped:presence of the Alpine Salamander ( • A • = certain; A = suggested but to be confirmed);

chromatic type of local population ( • = most individuals with largely patched coloration; A A = most individualswith poorly patched coloration, yet some completely black; • = probably most individuals completely black).

Abb. 2 (gegenüberliegende Seite): Verbreitung und FarbdifiFerenzierung des Alpensalamanders (Salamandra atra) imGebiet zwischen Val Lagarina und Val Sugana (Venetianische Voralpen) auf Grundlage der gesammelten Daten.

Für jeden Quadratkilometer des UTM-Koordinatensystems wurden folgende Informationen dargestellt: Präsenz desAlpensalamanders ( • A • = sicherer Nachweis; A = vermutetes, zu bestätigendes Vorkommen); Färbungstyp in derLokalpopulation ( • = die meisten Individuen mit ausgedehnter heller Fleckung; A A = die meisten Individuen mit

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the plateau and the bottom of the deepervalleys (P. RIGONI, pers. comm.). In thecentral basin, however, a small stablepopulation of Fire Salamanders lives nearGallio, only few kilometres away from thenearest known population of Alpine Sala-manders (Gruppo Nisoria, in press).

A similar pattern of spatial vicariancebetween the two species is known for otherneighbouring prealpine regions, e.g., theVette Feltrine, the Dolomiti Bellunesi(LAPINI et al. 1998) and the Cansiglio pla-teau (POMINI 1936; DOLCE 1988). Cases ofsyntopy are however reported from inneralpine regions (e.g., KLEWEN 1988).

Chromatic differentiation

According to the chromatic featuresof the body surface, the populations clusterinto at least two groups, which are appar-ently allopatrically distributed:

• populations in which all individu-als show patches (no exceptions have beendocumented so far), the total extension ofwhich is - on the average - very large (P+).This type seems to be exclusive of the SetteComuni plateau.

* populations in which most indi-viduals show patches (some of them arecompletely black), the total extension ofwhich is - on the average - rather limited(P-). This type seems to be typical at leastin the Pasubio massif.

A third chromatic type, in which thepatches are even less frequent or com-pletely absent, is suggested but not provedto exist in the south-western mountains.The few reliable data reported from thosesites refer to individuals described as com-pletely black.

Unfortunately, an unresolved incon-gruency between different informationsources forbids us to rely on a completelyblack specimen preserved in the Museumof Natural History of Verona (cataloguenumber: CE 1009). It is labelled as follows:"Salamandra atra Laurenti - loc. Mte Pas-ubio - 14/8/64 - det. Maucci - ex coll. Mu-seo - sul sentiero che conduce al rifugio -leg. Raineri". Identical information is re-ported in the published catalogue of thecollection (MAUCCI 1971); the site reportedin BRUNO (1973) as "Col Santo presso ilRifugio V. Lancia, 1,800 m, sul M. Pas-

ubio", although more precise, is veryprobably derived from the same material.But the presumed collector recently deniedto have found such an animal (W. RAINERI,pers. comm.); moreover, a black individualcollected in the early Nineteen-eighties onM. Carega was indicated to have been de-posited in the same Museum (OSELLA1985; G. OSELLA, pers. comm.), but re-cently no material labelled from that sitecould be found in the collection. As a con-clusion, the present labelling is very prob-ably erroneous.

The chromatic variations observed inthe Alpine Salamanders between Val La-garina and Val Sugana seem to be exclu-sive of this prealpine region. An interestingcase of genetic differentiation underliesthese phenotypical differences, as somepreliminary studies already suggest (TRE-VISAN et al. 1984; JOGER 1986 - by meansof protein electrophoresis; OLIVIERI 1991 -by means of allozyme analysis; STEINFARTZet al. 2000 - by means of DNA sequenc-ing). Moreover, the updated available in-formation suggests that the phylogeo-graphical pattern occurring here may bemore complex than a simple atra-auroraedichotomy (as it was thought previously)and so stimulates further studies.

The evolutionary history of thesepopulations was obviously influenced bythe climatic and environmental changesthat occurred in the Venetian Prealpsmainly in the last million years. This ideawas already proposed on the basis of thetheory of the "refuge massifs" (GROSSEN-BACHER 1994; LAPINI et al. 1996; LAPINI etal. 1998; BONATO 2000) and supported byestimates of divergence times recently ob-tained by a "molecular clock" model(STEINFARTZ et al. 2000). The distributionof glaciers during the recent ice-ages in thepresent region was described with greatprecision already several decades ago(PENCK & BRUECKNER 1909; TRE vis AN1939; CASTIGLIONI 1940) and was recentlyconfirmed and given in more detail (CORÀ1998). These paleogeographical models areexpected to represent a good basis for thehistorical interpretation of the phylo-geographical pattern.

Although not extensively studied,several other cases of biological differen-tiation were reported for this region: en-


Alpine Salamander (Salamandra atra) in the Venetian Prealps 179

demie forms of montane and barely mobileorganisms, both plants (e.g., RAFFAELLI &BALDOIN 1997; PROSSER & SCORTEGAGNA1998) and animals (review in BAGNOLI etal. 1997), are known to live only in few

sites between Val Lagarina and Val Su-gana. Until now, however, the case of theAlpine Salamander is the only one knownamong vertebrates.

ACKNOWLEDGEMENTS

We thank all the persons which aided us in real-ising this review. M. BIONDO (Bern), P. D E FRANCESCHI(Verona), A. DE TOMASI (Vicenza), G. GUEX (Zurich),G. LAZZARO*! (Verona), P. LORENZI (Rovereto), M.MEINEGON (Trento), W. MISSLER (Plauen), A NOEL-LERT (Jena), B. G. OSELLA (L'Aquila), M. PAGANIN(Asiago), P. PEDRINI (Trento), F. PROSSER (Rovereto),

W. RAINER! (Genova), P. RlGONI (Asiago), S. RUFFO(Verona), S. SCORTEGAGNA (Schio), S. STEINFARTZ(Köln) and M. WlNISTOERFER (Bern) answered to ourrequest for information and furnished unpublished data.G. FRACASSO (Vicenza), G. MATESSI (Padova) and A.MlNELLJ (Padova) improved our draft both in contentsand in form.

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DATE OF SUBMISSION: March 28th, 2000 Corresponding editor: Heinz Grillitsch

AUTHORS: Dr. Lucio Bonato, Dipartimento di Biologia, Università di Padova, via U. Bassi 58b, 1-35131 Pa-dova, Italy [e-mail: [email protected]]; Dr. Kurt Grossenbacher, Naturhistorisches Museum, Bernastrasse 15,CH-3005 Bern, Switzerland [e-mail: [email protected]].


Documents

On the distribution and chromatic differentiation of the … TREVISAN 1982). Afterwards only few data were published on the distribution of this new form (see: GROSSENBACHER 1994)