Taxonomy of Some of the Salamandrids

 

Note: The following 6 articles were translated, with permission, from:
http://www.salamanderland.at/ [no longer available online]

All text © 2006 Günter Schultschik
Translation © 2007 Ralf Reinartz and Jennifer Macke

Topics:
Systematics of the genus Pleurodeles
Systematics of the genera Euproctus and Calotriton
Systematics of the genera Mertensiella and Lyciasalamandra
Systematics of the genus Salamandrina
Systematics of the genus Triturus
Triturus vulgaris and its subspecies
Triturus helveticus and its subspecies

 



 

Systematics of the genus Pleurodeles

The first animal from this genus to be described scientifically was a specimen that GERVAIS named Triton poireti in 1835. Incidentally, this newt came from Northern Africa from the peninsula of Edough in Algeria.

Between 1840 and 1842 GUICHENOT also received ribbed newts from Northern Africa. Believing he was looking at a new and previously unknown species, he described the animals as Pleurodeles nebulosus in 1850.

Using morphometric analysis as the only available method back then, WOLTERSTORFF went deeper into the matter and concluded in 1905 that the two different previously-described taxa had to be the same species. He recognized the genus Pleurodeles (GUICHENOT) but, on the basis of the priority rule, named the species poireti (GERVAIS) (resulting in the scientific name Pleurodeles poireti).

Unfortunately, WOLTERSTORFF wasn’t right either. Using a combination of molecular genetics and morphological analysis CARRANZA & WADE were able to prove in 2004 that these two taxa are distinct species. The ribbed newts from the peninsula of Edough in Algeria are indeed a different species. Thus, the original descriptions by GERVAIS and GUICHENOT are valid even today.

This separation is also supported by the fact that Edough was an island for a long period of time and thus separated zoogeographically from the rest of North Africa. (Because of the resulting genetic isolation, the ribbed newts on Edough were able to develop into a separate species.)

Molecular genetic analysis by CARRANZA & WADE also revealed that all (!) ribbed newts from the Iberian peninsula (i.e., Portugal and Spain) belong to the species Pleurodeles waltl, and that animals of this species found in North Africa were brought there by man.

All indigenous ribbed newts in North Africa therefore belong to the species Pleurodeles nebulosus, with the exception of animals from Edough, which form the third species within Pleurodeles and which were initially described as Triton poireti by GERVAIS in 1835.

References:

CARRANZA, S. & E. WADE (2004): Taxonomic Revision of Algero-Tunisian Pleurodeles (Caudata: Salamandridae) using molecular and morphological data. Revalidation of taxon Pleurodeles nebulosus (Guichenot, 1850). Zootoxa 488: 1-24.

GERVAIS, P. (1835): Communication sur les Reptiles de Barbarie. Bulletin de la Société de Sciences Naturelles de France, Séance du 23.12.1835: 112-114.

GUICHENOT, A. (1850) : Histoire naturelle des Reptiles et des Poissons. Exploration scientifique de l´Algérie pendant les annes 1840, 1841, 1842. Zoologie. Bibliothèque Français, Paris, 144 pp.

WOLTERSTORFF, W. (1905) : Zwergformen der Paläarktischen Urodelen. C. R. 6e Congrès International Zoologie Berne, 1904, 258-263.

 



 

Systematics of the genera Euproctus and Calotriton

For a long time the genus Euproctus consisted of three species:

  1. Euproctus platycephalus from Sardinia in Italy (described by GRAVENHORST 1829).
  2. Euproctus montanus from Corsica in France (initially described by SAVI 1838 as Megapterna montanus and later included in the genus Euproctus).
  3. Euproctus asper from the Pyrenees (described by DUGÉS 1852).

The zoogeographic origin of Euproctus/Calotriton: Twenty-nine million years ago, the conjoined island of Corsica/Sardinia (still a single land-mass at that time) separated from the Iberian mainland, splitting the ancestors of the current genera up into two isolated groups. Nineteen million years ago, Corsica and Sardinia separated into two different islands. This means that E. platycephalus and E. montanus formed a genetic unit for 10 million years.

In 2005 CARRANZA & AMAT concluded from their research that Euproctus newts from the Iberian mainland and from the Mediterranean islands of Corsica and Sardinia are paraphyletic, meaning that the mainland and the island populations had to have different ancestors on the Iberian mainland before the separation of the islands. Therefore, they revalidated the genus Calotriton (GRAY 1858) to include asper, as they found these animals to be more similar to the large Triturus species than to the Euproctus newts from Corsica and Sardinia.

In the course of their work CARRANZA & AMAT also stumbled on an isolated population of Calotriton newts in the El Montseny region south of the Pyrenees with some remarkably different traits than asper; they described these newts as a new species, Calotriton arnoldi.

Overview of systematic changes within the genera Euproctus / Calotriton:

Former  species name Current species name Distribution
Euproctus asper Calotriton asper Pyrenees
newly described Calotriton arnoldi El Montseny region
Euproctus asper castelmouliensis no longer a valid subspecies -
Euproctus platycephalus Euproctus platycephalus Sardinia
Euproctus montanus Euproctus montanus Corsica

 

References:

CARRANZA, S. & F. AMAT (2005) : Taxonomie, biogeography and evolution of Euproctus (Amphibia: Salamandridae), with the resurrection of the genus Calotriton and the description of a new endemic species of the Iberian Peninsula. Zoological Journal 145: 555-582.

DUGÉS, A. (1852): Recherches zoologiques sur les Urodelès de France. Annales des Sciences Naturelles París Zoologie 3: 253-272.

GRAVENHORST, J. L. K. (1829): Delicae musei zoologici vratislaviensis. 1. Chelonios et Batrachia. Leipzig : Sumptibus L. Vossii.: 84.

GRAY, J. E. (1858): Proposal to separate the family Salamandridae, Gray, into two families, according to the form of the skull. Proceedings of the Zoological Society of London 1858: 136-144.

SAVI, P. (1838): Descrizione della Salamandra corsica, e della Megapterna montana nuovi animali della famiglia dei Batrachii. Nuovo Giornale dei Letterati, Pisa 37: 208-217.

 



 

Systematics of the genera Mertensiella and Lyciasalamandra

In 1876 WAGA described a salamander of the Caucasia region. It was described not only as a new species, but as a new genus, on the basis of the prominent hedonic gland present at the base of the tail of this species’ males. He named it Exaeretus caucasicus. Unfortunately, WAGA didn’t have good luck in choosing this genus name, as Exaeretus was already in use at that time as a genus name for beetles (Coleoptera). Therefore, WAGA had to place this species within the genus Salamandra in 1878.

In 1891, STEINDACHNER, at a museum in Vienna, received a Lycian salamander from Professor Luschan. He recognized it as a new species and described it as Molge luschani, which BOULENGER later renamed Salamandra luschani in 1892. That same year another researcher named BEDRIAGA confirmed STEINDACHNER’s description as Molge luschani. From this it can be concluded that even back then BEDRIAGA did not consider the two species of Lycian and Caucasian salamanders to belong to the same genus. Some time later, in 1925, WOLTERSTORFF created the new genus Mertensiella for both species.  In his opinion, the characteristic trait of the males possessing a prominent hedonic gland at the base of the tail proved a mutual origin and ancestry of the animals.

However, looking at the morphology and life patterns of these two species one quickly finds reason to doubt their common ancestry. Whereas the Caucasian species lives in the vicinity of mountain brooks and lays eggs, the Lycian species is completely terrestrial and a live-bearer, giving birth to fully developed offspring. 

Research on the tissues of the hedonic gland by SEVER et al. in 1997 revealed that histologically these organs are totally different in the two species, raising the question of whether the two species are closely related at all. TITUS & LARSEN and VEITH et al. conducted molecular genetic research on the genus Mertensiella in 1995 and 1998, respectively. The results revealed that the Caucasian salamander is closely related to Chioglossa, whereas the Lycian species is closer to Salamandra. It also became obvious that both species have different ancestors and do not belong into the same genus, as WOLTERSTORFF had thought.  

Consequently, the new genus Lyciasalamandra was established by VEITH et al. for the Lycian salamander in 1998. Also, in the course of this research, the status of some former subspecies was raised to species level. A group of US scientists, however, did not follow this reasoning and placed the Lycian salamander into Salamandra, sticking to BEDRIAGA’s opinion from the 19th century (WEISROCK et al. 2001).

How could such a specific trait like the hedonic gland arise independently in two different genera? There are indications that this trait may have been present in a number of ancient genera and was simply lost in many species during the intervening course of evolution. Looking at the mating behaviour of modern fire salamanders and comparing it to that of Mertensiella caucasica one might even get the impression that the Salamandra male still pretends to have a hedonic gland.

Overview of changes within the genera Mertensiella and Lyciasalamandra:

Former species / subspecies name Current species / subspecies name
Mertensiella caucasica Mertensiella caucasica
Mertensiella luschani Lyciasalamandra luschani
Mertensiella luschani luschani Lyciasalamandra luschani luschani
Mertensiella luschani finikensis Lyciasalamandra luschani finikensis
Mertensiella luschani basoglui Lyciasalamandra luschani basoglui
Mertensiella luschani helverseni Lyciasalamandra helverseni
Mertensiella luschani atifi Lyciasalamandra atifi
Mertensiella luschani fazilae Lyciasalamandra fazilae
Mertensiella luschani antalyana Lyciasalamandra antalyana
Mertensiella luschani billae Lyciasalamandra billae
Mertensiella luschani flavimembris Lyciasalamandra flavimembris

References:

BOULENGER, G. (1892): Ann. Mag. Nat. Hist., Ser. 6, 9: 74.

BEDRIAGA, J. (1892): Congr. Internatl. Zool., Moscow, Part 1: 242.

SEVER, D. M., M. SPARREBOOM, G. SCHULTSCHIK (1997): The dorsal tail tubercle of Mertensiella caucasica and M. luschani (Amphibia: Salamandridae), J. Morphol., 232: 93-105.[PDF]

STEINDACHNER, F. (1891): Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 100: 306.

VEITH, M. & S. STEINFARTZ (2004): When non-monophyly results in taxonomic consequences - the case of Mertensiella within the salamandridae (Amphibia: Urodela), Salamandra, Rheinbach, 40(1): 67-80.

WOLTERSTORFF, W. (1925): Abh. Ber. Mus. Nat. Heimatkd. Magdeburg, 4: 244.

WEISROCK, D. W., J. R. MACEY, I. H. UGURTAS, A. LARSON, T. J. PAPENFUSS (2001): Mol. Phylogenet. Evol., 18: 434.

 



 

Systematics of the genus Salamandrina

This genus is endemic to the western Apennine Mountains. Like Euproctus, this genus originated from the Iberian peninsula and probably crossed the Mediterranean Sea to Italy on the ridges of the former Corsica and Sardinia islands. Unlike Euproctus, however, Salamandrina managed to reach the Italian mainland. On Sardinia today one can find fossil remains, which still await close examination.

The first historical record of this species comes from the region of the Mt. Vesuvius volcano. LACÉPÈDE described the animal in 1788 as Salamandra terdigidata. From the Tuscany region came another historical report in 1821 in which SAVI named his find as Salamandra perspicillata. In 1826 FITZINGER worked with the two taxa and placed them in the newly created monotypic genus Salamandrina. FITZINGER could not determine any differences in morphometrics or colouration between the southern and the northern populations.

In 2005 NASCETTI et al. described the possible history of the colonization of the Italian peninsula and concluded that Salamandrina possibly reached the Italian mainland in two separate waves. This explains the presence of two genetically distinct groups of Salamandrina and calls for consequences in the systematic nomenclature of these groups. There are two species within Salamandrina: S. terdigidata in the south and S. perspicillata in the north; the authors correctly used the corresponding original names and revalidated them. Interestingly, this article was published even before the actual species description by CANESTRELLI et al. 2006. It is still difficult to distinguish the two species by appearance, just as it was for FITZINGER. Molecular genetic research, however, has allowed a distinct differentiation.

References:

CANESTRELLI, D., F. ZANGARI, G. NASCETTI (2006): Genetic evidence for two distinct species within the Italian endemic Salamandrina terdigidata (Lacépède, 1788) (Amphibia: Urodela: Salamandridae). Herp. J. 16(2): 21-227.

FITZINGER, L. J. (1826): Neue Classification der Reptilien nach ihrer natürlichen Verwandtschaften: Nebst einer Verwandtschafts-Tafel und einem Verzeichnisse der Reptilien-Sammlung des K.k. Zoologischen Museums zu Wien. J. G. Heubner, Wien.

LACÉPÈDE, B. G. E. (1788): Histoire naturelles des quadrupèdes ovipares et des serpentes. Paris, Imprimerie du Poi (Hôtel de Thou), vol I.

NASCETTI, G., F. ZANGARI, D. CANESTRELLI (2005): The spectacled salamanders Salamandrina terdigidata (Lacépède, 1788) and S. perspicillata (Savi, 1821): 1) Genetic differentiation and evolutionary history. Rend. Fis. Acc. Lincei, s.9, v.16: 159-169.

SAVI, P. (1821) : Descrizione (inedita) di una nuova specie di Salamandra terrestre, Salamanda perspicillata. Nob. Bibl. It. (Giornale di Letteratura, Scienze ed Arti), 22(2): 228-230.

 



 

Systematics of the genus Triturus

Without a doubt, this genus is one of the most complicated in herpeto-taxonomy, full of errors and mistakes.

Whenever the genus Triturus is mentioned in scientific literature, it is referenced to the very first author using this genus name: RAFINESQUE, 1815. However, when searching for this reference, which was probably a private publication by RAFINESQUE, it becomes evident that hardly anyone has ever seen this book. My own research failed to find a copy of this book anywhere in Europe. Only the Library of Congress in the USA lists this reference. Also the authors of the “Handbuch der Reptilien und Amphibien Europas” ( “Handbook of Reptiles and Amphibians of Europe ”) did not personally see this book, but adopted this information from another source (JEHLE, pers. comm.).

In his book, RAFINESQUE postulated the family Tritonia and listed the following genera, without any further descriptions: Triturus, Palmitus, Lophinus and Meinus. Thus, from a taxonomical viewpoint, these names are nomina nuda (Latin for “naked names”) meaning that these names are not available according to the rules of taxonomic nomenclature and that it shouldn’t have been permitted to use them. We conclude: Any text quoting RAFINESQUE, 1815 as source for the genus Triturus is most probably incorrect (more information on this in SCHMIDTLER, 2004, described below).

As late as 1820, five years after the initial publication, RAFINESQUE wrote a description of Triturus. From this description, the genus name Triturus becomes relevant to taxonomic nomenclature for the first time. Therefore all relevant text should cite RAFINESQUE, 1820 as the primary reference for the genus.

MERREM also published his concept of this newt genus in 1820, using the taxonomic name Molge resulting in the equally correct reference: Molge, MERREM, 1820. Both descriptions were published in 1820, and it would require investigative research to determine which one was published first and should therefore be given priority. Apart from this quaint situation in the issue of nomenclature, there were early doubts about whether Triturus/Molge is a systematically uniform genus. In 1928 BOLKAY suggested three subgenera on the basis of a comparative analysis of skull bone structures.

  1. Paleotriton (including vulgaris, montandoni, helveticus, italicus, boscai, vittatus)
  2. Neotriton (including cristatus [dobrogicus, carnifex, karelinii], marmoratus [pygmaeus])
  3. Mesotriton (including alpestris)

Recent genetic analyses by various researchers have all revealed that Triturus is indeed not a single genus. Also, the interrelations are probably rather complicated, leading authors to different conclusions and to proposals of different taxonomic relationships.

In the end, these conclusions and proposed taxonomies have been rearranged by third parties, adding to the confusion that has been accompanying Triturus for the past 200 years.

Currently there are 3 or 4 taxonomic approaches to these animals:

  1. The traditional taxonomy (e.g., “Handbuch der Reptilien und Amphibien Europas”) using only Triturus as the genus name.
  1. GARCIA-PARIS et al. published their findings in 2004 in Spanish. This group from Spain basically followed the arguments of BOLKAY, using, because of the priority rule, other and older names, but without studying the literature thoroughly enough to really use the oldest and therefore relevant references. They proposed the following taxonomy:
    • Lissotriton (BELL, 1839) (boscai, helveticus, italicus, montandoni, vulgaris)
    • Mesotriton (BOLKAY, 1928) (alpestris)
    • Triturus (RAFINESQUE, 1815?, 1820) (carnifex, cristatus, dobrogicus, karelinii, marmoratus, pygmaeus, vittatus)
  1. LITVINCHUK et al.  presented the following taxonomy in 2005:
    • Lophinus (Rafinesque, 1815) (boscai, helveticus, italicus, montandoni, vulgaris) (Using the nomen nudum from 1815(!). The correct reference therefore has to be: GRAY, 1850, since this author provides the first description.)
    • Mesotriton (Bolkay, 1928) (alpestris)
    • Triturus ( Rafinesque, 1815?, 1820) (carnifex, cristatus, dobrogicus, karelinii, marmoratus, pygmaeus).
    • Ommatotriton (Gray, 1850) (vittatus, ophryticus).
  1. SCHMIDTLER went even deeper into the matter in 2004. In an article he found, which had hardly been noticed by the scientific community until then, LATREILLE had described an odd caudate larva in 1802. It had been drawn previously by LAURENTI in 1768 and placed into the genus Proteus (Proteidae) as Proteus tritonius. In fact, the depicted animal was not an olm but, most likely, the larva of an alpine newt, and LATREILLE named the genus Ichthyosaura. Most probably this is the oldest genus name ever applied to European newts and should therefore definitely be used for alpine newts, in place of the genus name Mesotriton (BOLKAY, 1928).

SCHMIDTLER 2004 made some additional comments on the problem of genus names in European newts:

  • Molge (MERREM, 1820) was used as a substitute for the already-used genus name Triton by LAURENTI whose type-species therefore is Triton cristatus or Molge cristatus or Triturus cristatus.  (Problem: the simultaneous description of Molge and Triturus by RAFINESQUE, 1820, as described above, raises the inevitable question as to who came first by the priority rule!)
  • Geotriton (Bonaparte, 1832) was used before (!) Lissotriton, (Bell, 1839).
  • Palaeotriton (Fitzinger, 1843), for which THORN 1968 subsequently assigned Triturus vulgaris as type-species, wasn’t approved generally.

ITIS (Integrated Taxonomic Information System) currently does not see any need for action and sticks with the traditional (“all Triturus”) systematic nomenclature.

STEINFARTZ et al. also published on the matter in 2006, but did not deal with any taxonomic consequences.

LITVINCHUK et al. (2005) took it one step further and revalidated the genus Ommatotriton (GRAY, 1850) and further differentiated the species of vittatus. Interestingly, the authors initially mention Triturus vittatus. This is followed by a chapter in which three versions of taxonomy for the genus are discussed. The conclusion consists of the presentation of a systematic structure, including the genus taxon Ommatotriton (with the remarkable wording “we prefer”), and the description of a new subspecies: Triturus vittatus nesterovi.

If we follow the authors’ reasoning, we would see the following situation within the revalidated genus Ommatotriton:

Old Name New Name
Triturus vittatus vittatus Ommatotriton vittatus vittatus
Triturus vittatus cilicensis Ommatotriton vittatus cilicensis
Triturus vittatus ophryticus Ommatotriton ophryticus with two subspecies:
    Ommatotriton ophryticus ophryticus in the East
    Ommatotriton ophryticus nesterovi in the West

Therefore, the name Triturus ophryticus nesterovi used for the first description of a new subspecies would instantly become a synonym of Ommatotriton ophryticus nesterovi in the same scientific article! This leaves one wondering about the authors' intent.

A thorough examination of the articles on the matter at hand by GARCIA-PARIS et al. and LITVINCHUK et al., reveals serious taxonomical mistakes in both publications, and they should therefore not be used for nomenclature. Unfortunately, this incorrect nomenclature, especially that of the Spanish group, has been distributed via the Internet and has been widely accepted by the public.

References:

BELL , T. (1839/1849): A history of British reptiles. Van Voorst, London, 142 pp.

BOLKAY S. J. (1928): Die Schädel der Salamandriden mit besonderer Rücksicht auf ihre systematische Bedeutung. Zeitschr. F. Anat. U. Entwicklungsgesch., München, Berlin, 86: 259-319.

BONAPARTE, C. L. (1832-1841): Iconografia della fauna Italica per le quattro classi degli animali vertebrati. Salviucci, Roma. Ministero dell´ Ambiente, Treviso, Reprint 2003, 3 Bände: 459, 1116.

FITZINGER, L. J. (1843): Sistema reptilium, fasc. Primus Amblyglossae. Braumüller et Seidel, Vindobonae, 106 pp.

GARCÍA-PARIS, M., A. MONTORI, and P. HERRERO (2004): Amphibia: Lissamphibia. Fauna Iberica Vol. 24. Madrid: Museo Nacional de Ciencias Naturales and Consejo Superior de Investigaciones Científicas.

GRAY, J. E. (1850): Catalogue of the specimens of amphibia in the collection of the British Museum. Part II, Batrachia Gradientia. Order of the Trustees, London, 72 pp.

LAURENTI, J. N. (1768): Specimen medium, exhibens Synopsin Reptilium. Trattner; Viennae, 214 pp.

LITVINCHUK, S. N., A. ZUIDERWIJK , L. J. BORKIN & J. M. ROSANOV (2005): Taxonomic status of Triturus vittatus (Amphibia: Salamandridae) in western Turkey : trunk vertebrae count, genome size and allozyme data. – Amphibia-Reptilia 26 (3):305-323.

MERREM, B. (1820): Versuch eines Systems der Amphibien (Tentamen systematics amphibiorum). Krieger, Marburg, 195 pp.

RAFINESQUE, C. S. (1815): Analyse de la nature ou tableau de l´univers et des corps organisés. Palerme, 224 pp.

RAFINESQUE, C. S. (1820) : III. Class. Erpetia. The Reptiles. Annals of Nature or annual synopsis of new genera and species of animals, plants etc. discovered in North America. 1820 (1): 4-6.

SCHMIDTLER, J. F. (2004): Der Teichmolch (Triturus vulgaris (L.)), ein Musterbeispiel für systematische Verwechslungen und eine Flut von Namen in der frühen Erforschungsgeschichte. Sekretär, Beiträge zur LGHT, Rheinbach, 4(2): 10-28.

SONNINI, C. S. & P. A. LATREILLE (1802) : Histoire Naturelle des Reptiles. Déterville, Paris, 4 Bände.

STEINFARTZ, S., S. VICARIO, J. W. ARNTZEN, A. CACCONE (2006): A Bayesian Approach on Molecules and Behavior: Reconsidering Phylogenetic and Evolutionary Patterns of the Salamandridae with Emphasis on Triturus Newts. J. Exp. Zool. (Mol. Dev. Evol.) 306(2): 139-162.

THORN, R. (1968): Les salamandres d`Europe, d`Asie et d`Afrique du Nord. Lechevalier, Paris, 376 pp.

GROSSENBACHER, K & B. THIESMEIER, Eds. (1999): Handbuch der Reptilien und Amphibien Europas, Schwanzlurche I, (pp 1-205), Aula, Wiebelsheim.

THIESMEIER, B. & K. GROSSENBACHER, Eds. (2003): Handbuch der Reptilien und Amphibien Europas, Schwanzlurche IIA, (pp 206-758), Aula, Wiebelsheim.

THIESMEIER, B. & K. GROSSENBACHER, Eds. (2004): Handbuch der Reptilien und Amphibien Europas, Schwanzlurche IIB, (pp 759-1149) Aula, Wiebelsheim.

 



 

Triturus vulgaris and its subspecies

The following subspecies of Triturus vulgaris (Linnaeus, 1758) are considered valid at the time of this writing:

Subspecies Described by Location
T. vulg. ampelensis Fuhn, 1951 Valea Dosuluj, Romania
T. vulg. graecus Wolterstorff, 1905 Korfu, Greece
T. vulg. kosswigi Freytag, 1955 Abant Lake, West Anatolya, Turkey
T. vulg. lantzi Wolterstorff, 1914 Novorossijsk, Russia
T. vulg. meridionalis Boulenger, 1882 Turin, Italy
T. vulg. schmidtlerorum Raxworthy, 1988 Karacabey, West Anatolya, Turkey
T. vulg. schreiberi Wolterstorff, 1914 Lake Bokanjacko, Zadar, Croatia
T. vulg. vulgaris Linnaeus, 1758 Sweden

In the literature there is an interesting and recurring hint at another presumed subspecies in the lowlands of the Danube and Sava Rivers, distinguishable by their relatively small size. Interestingly, the smooth newts in the hills around Vienna, for example, also are 30 to 50% heavier than those from the Danube River floodplain.

The following are no longer considered to be valid subspecies:

Former subspecies Described by Location
T. vulg. borealis Kauri, 1959 Jämtland, Sweden
T. vulg. dalmatica Kolombatovic, 1907 Cetina Valley, Croatia
T. vulg. graecus
forma corcyrensis
Wolterstorff, 1908

Korfu, Greece

T. vulg. graecus
forma tomasinii
Wolterstorff, 1908 Tivat, Montenegro
T. vulg. intermedia Kolombatovic, 1907 Karakasika Bach, Croatia
T. vulg. kammereri Wolterstorff, 1907 Semmering, Austria
T. vulg. kapelana Méhely, 1905 Mrkopalj, Croatia
T. vulg. tataiensis Dely, 1967 Komarom, Hungary

 

Some characteristic traits of the valid subspecies:

  Dorsolateral zone in cross section Edge of crest Tail filament
presence, appearance
Hind feet webbing Belly spots
presence, size
Red neckstripe present
T. v. ampelensis edgy weakly undulated short light yes no
T. v. graecus bulky straight-edged long, thin strong, dark small yes
T. v. kosswigi bulky straight-edged long, thin strong, dark large yes
T. v. lantzi edgy finely serrated short, colorless dark yes no
T. v. meridionalis weakly pronounced straight-edged short, wide strong, dark small, few yes
T. v. schmidtlerorum edgy serrated no weak yes no
T. v. schreiberi ? serrated no ? no no
T. v. vulgaris rounded serrated hardly any light yes no

One curiosity is the occurrence of T. vulg. cf. graecus in the close vicinity of our facility at “Salamanderland” in Austria. Apparently, this is a released population with many individuals in an area of approximately 2 square kilometers (~1 sq. mile). This population has been documented since the 1980s, and may be showing a tendency to spread and out-compete the indigenous nominate form.

 



 

Triturus helveticus and its subspecies

This species was first described by RAZOUMOVSKY in 1789. He chose the name "Lacerta paradoxa seu Helvetica". Like the smooth newt, this species was then assigned a number of different names. Finally DUNN assigned the taxon Triturus helveticus in 1918, which remained in use for many years thereafter. Recent articles, however, doubt the affiliation of helveticus with the genus Triturus.

Also quite unclear and complicated is the status of the subspecies within Triturus helveticus.
It is an interesting fact that, despite having a large geographic distribution, variation within this species is remarkably low.

The first taxon on the subspecies level from the terra typica Porto was named by WOLTERSTORFF in 1905 as Triton palmatus sequeirai. This species was later renamed Triturus helveticus sequeirai by MERTENS & MÜLLER in 1928.

Already in 1884, SEOANE had described a newt from another location on the Iberian peninsula (terra typica Cabanas en La Coruna ) as Triton alonsoi. Nevertheless, in his 1974 article, SALVADOR reaches the conclusion that T. h. sequeirai is a synonym of Triton alonsoi and, because of the priority rule, the latter name should be used. The distribution area of this Triturus helveticus alonsoi would be northern Portugal and northwestern Spain and would therefore include the habitat of T. h. sequeirai.

In 1969 SCHMIDTLER described a conspicuous colour variant from the karst lake of Pozo Negro (=terra typica) at 1770 m above sea level and called it Triturus helveticus punctillatus. The Latin name reveals the main characteristic trait of these animals: numerous dark dots on the back, sides, and tail. SALVADOR (1974), however, doesn’t recognize this subspecies and regards T. h. punctillatus to also be a synonym of T. h. alonsoi.

As recently as 2005, this problem was again addressed by GARCIA-PARIS et al. taking this matter one step further and recognizing the nominate form L. h. helveticus and two subspecies, L. h. punctillatus and L. h. alonsoi, but not T. h. sequeirai. In this respect, they conform with SALVADOR (1974).

We will have to wait for further developments in this matter. At the moment, the following three or four taxa are in use:

  • Triturus helveticus helveticus
  • Triturus helveticus sequeirai
  • Triturus helveticus punctillatus
  • (Triturus helveticus alonsoi)

The genus name Triturus is used for those who decide not to follow GARCIA-PARIS et al. and their genus Lissotriton nor LITVINCHUK et al. and their genus Lophinus. (Both are most probably incorrect.)

References:

DUNN, E. R. (1918): Bull. Mus. Comp. Zool., 62: 452.

GARCÍA-PARIS, M., A. MONTORI, and P. HERRERO (2004): Amphibia: Lissamphibia. Fauna Iberica Vol. 24. Madrid : Museo Nacional de Ciencias Naturales and Consejo Superior de Investigaciones Científicas.

LITVINCHUK, S. N., A. ZUIDERWIJK , L. J. BORKIN & J. M. ROSANOV (2005a): Taxonomic status of Triturus vittatus (Amphibia: Salamandridae) in western Turkey : trunk vertebrae count, genome size and allozyme data. – Amphibia-Reptilia 26 (3):305-323.

MERTENS, R. & P. MÜLLER (1928): Abh. Senckenb. Naturforsch. Ges., 41: 12.

RAZOUMOVSKY, G. (1789): Hist. Nat. Jorat, 1: 111.

SALVADOR , A. (1974): Guia de campo de los Anfibios Espanoles. Identificacion, historia natural y distribucion. Talavera de la Reina (Canseco Editores), 269 pp.

SEOANE, V. L. (1885): Identidad de Lacerta schreiberi (Bedriaga) y Lacerta viridis, var. gadowii (Boulenger) é investigaciones herpetológicas de Galicia . La Coruña (Imp. y est de Vicente Abad), 19 pp.

SCHMIDTLER, J. F. (1969): Herpetologische Beobachtungen in den Iberischen Randgebirgen, mit Beschreibung einer neuen Unterart von Triturus helveticus (Salamandridae, Amphibia). Abh. Ber. Naturk. Vorgesch. Magdeburg, 11: 219-231.

WOLTERSTORFF, W. (1905): Zwergformen der Paläarktischen Urodelen. Comptes rendus du 6e Congrès international de Zoologie. Session de Berne 1904: 258-263.


All text © 2006 Günter Schultschik. Translation © 2007 Ralf Reinartz and Jennifer Macke.

 

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