Subspecies
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(1) Ichthyosaura
alpestris alpestris (Laurenti, 1768)
(2) Ichthyosaura alpestris apuana
(Bonaparte, 1839)
(3) Ichthyosaura alpestris cyreni
(Wolterstorff, 1932)
(4) Ichthyosaura alpestris lacusnigri
(Seliskar et Pehani, 1935)
(5) Ichthyosaura alpestris reiseri
(Werner, 1902)
(6) Ichthyosaura alpestris
veluchiensis (Wolterstorff, 1935)
There is significant disagreement
about the subspecies substructure of
this species. Dubois
and Raffaelli (2009) have
presented a list of all species and
subspecies of Salamandridae, also
including the present species.
However, they use unusual vocabulary
(e.g. "nucleospecies" which appears to
be similar but different from "type
species", because if the two terms
were identical, then the authors
surely would have used "type species"
and not invented a new term) and
unusual taxonomic units (e.g. "exerge"
which is probably similar to a
"species group", but is used
differently, because the authors seem
to interpolate the "exerge name"
between the species and the subspecies
epithet). Thus, it is unclear whether
the authors use a quadrinominal
name-form here (and these names are
excluded from the synonymy list
below). Because of this problematic
nature of the work, I do not follow Dubois
and Raffaelli (2009) here. I
follow here the analysis by Sotiropoulos
et al. 2007. They recognize 5
clades in Ichthyosaura alpestris, some
of them with further subgroupings and
not all of them yet with named taxa.
The five subspecies listed above
correspond to the clades B, C1, C3,
E1, E2 and the combined clade D.
Clades A and C2 do not yet have a
separate name. A full overview of the
correspondence of the clade structure
by Sotiropoulos
et al. (2007) and the 6
subspecies recognized above is given
below in the section "Synonyms". The
analysis by Sotiropoulos
et al. (2007) has largely been
confirmed by the analysis by Recuero
et al. (2014), but in addition,
this work provides evidence for a
separation of the northern Balcan and
the Carpathian populations into two
separate taxa (reiseri and carpathica,
respectively). I provisionally retain
these in a single subspecies until
more data on the nature of the two
groups are available. Of the six
subspecies recognized here, only the
nominate form occurs in Germany.
Note, however, that Vukov
et al. (2011) have shown that
the diversification of mitochondrial
DNA (mtDNA) discovered by Sotiropoulos
et al. (2007) is not accompanied
by significant morphological
diversification- at least in the
Balcan populations. Vukov
et al. (2011) suggest that the
evolution of the different populations
is non-adaptive. If this
interpretation should prove to be
correct, then this would support those
authors who argue that the Alpine Newt
does not have any subspecies.
Similarly, Speybroeck
et al. (2010) have argued that
the phylogenetic analysis of
mitochondrial DNA (as performed by Sotiropoulos
et al. (2007)) may be negatively
influenced by mtDNA introgression from
some subroups into other subgroups,
thus obscuring the true phylogenetic
relationships. Therefore Speybroeck
et al. (2010) argue that changes
in the intraspecific systematics of
Ichthyosaura alpestris are premature.
However, the analysis by Recuero
et al. (2014) also included
nuclear DNA and thus strongly suggests
that mtDNA introgression is not a
significant source of error in the
phylogenetic analysis by Sotiropoulos
et al. (2007).
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Synonyms
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I give here the
complete synonymy of all hitherto
described taxa and sort them according
to the clades in Sotiropoulos
et al. (2007).
(0) Clade A
(restricted to a small area in
Serbia) and Clade
C2
(north-eastern Italy) comprise
geographically restricted
populations that have not received a
scientific name yet.
(1) Clade C3 (Alps,
Central Europe, eastern France,
central portions of eastern Europe).
Ichthyosaura alpestris alpestris:
Triton alpestris Laurenti,
1768 (p. 38, p. 142)
Triton alpestris alpestris Laurenti,
1768
Salamandra alpestris (Laurenti,
1768)
Molge alpestris (Laurenti, 1768)
Hemitriton alpestris (Laurenti,
1768)
Triturus alpestris (Laurenti, 1768)
Triturus alpestris alpestris
(Laurenti, 1768)
Mesotriton alpestris (Laurenti,
1768)
Mesotriton alpestris alpestris
(Laurenti, 1768)
Ichthyosaura alpestris (Laurenti,
1768)
Ichthyosaura alpestris alpestris
(Laurenti, 1768)
Ichthyosaurus alpestris (Laurenti,
1768)
Type locality: "in Etschero
monte" (= Ötscher, Lower Austria).
Proteus tritonius Laurenti,
1768 (p. 37, p. 140)
Ichthyosaura tritonius (Laurenti,
1768)
Type locality: "in lacu, qui est
in alpe Etscher", "in lacu
frigidissimo, qui est in alpe
Etscher" (= Ötscher, Lower
Austria).
Triton wurfbainii Laurenti,
1768 (p. 38)
Type locality: not given in
original description.
Triton salamandroides Laurenti,
1768 (p. 40)
Type locality: "in Germania", at
the time comprising most of
Central Europe.
Lacerta gyrinoides Merrem,
1789 (p. 194)
Gecko gyrinoides (Merrem, 1789)
Type locality: "auf dem Heimberge
in einem Sumpfe (das Lichtemeer
genannt)" (= now mostly dried-up
pond north of Kehr on Hainberg
hill near Herberhausen, Göttingen
(Niedersachsen)).
Salamandra ignea Bechstein, 1800
Molge ignea (Bechstein, 1800)
Triton igneus (Bechstein, 1800)
Type locality: Thüringen.
Salamandra cincta Latreille, 1800
Type locality: "France".
Salamandra rubriventris Daudin, 1803
Type locality: replacement
name for Triton alpestris, thus
same type locality.
Triton zonarius Millet de la
Turtaudiere, 1828
Type locality: replacement name
for Salamandra cincta, thus same
type locality.
Triturus alpestris winterli Sochurek,
1956
Type locality: Kärnten, Austria.
Triturus alpestris buekkiensis Dely,
1960
Type locality: Bükk mountains,
Hungary.
Triturus alpestris satoriensis Dely,
1960
Type locality: Istvan-Patak,
Hungary.
Salamandra wurfbaini auct.
(misspelling)
Molge wurfbaini auct. (misspelling)
Triton wurfbeinii auct. (misspelling)
Triton salamandroides wurfbani auct.
(misspelling)
(2) Clade B (most of
Italy). Ichthyosaura alpestris
apuana:
Triton apuanus Bonaparte, 1839
Triton alpestris apuanus (Bonaparte,
1839)
Triturus alpestris apuana
(Bonaparte, 1839)
Triturus alpestris apuanus
(Bonaparte, 1839)
Mesotriton alpestris apuanus
(Bonaparte, 1839)
Ichthyosaura alpestris apuana
(Bonaparte, 1839)
Type locality: Serravezza, Apuan
Alps, northwestern Italy
Triton neglectus Jan, 1857 (nomen
nudum)
Type locality: Appenine
mountains, Italy.
Triton alpestris immaculatus Duerigen,
1897
Type
locality: replacement name for
Triton apuanus, thus same type
locality.
Triton alpestris ocellata
Wolterstorff, 1934
Type locality: Genoa, Italy.
Triturus alpestris inexpectatus Dubois
et Breuil, 1983
Ichthyosaura alpestris inexpectata
(Dubois et Breuil, 1983)
Type locality: Calabria, Italy.
(3) Clade C1
(northern Spain). Ichthyosaura
alpestris cyreni:
Triturus alpestris cyreni
Wolterstorff, 1932
Mesotriton alpestris cyreni
(Wolterstorff, 1932)
Ichthyosaura alpestris cyreni
(Wolterstorff, 1932)
Type locality: Cantabrian Mountains,
northern Spain.
(4) Clade E1 (Montenegro).
Ichthyosaura alpestris lacusnigri:
Triturus alpestris lacustris Seliskar
et Pehani, 1935 (secondary junior
homonym of Lacerta lacustris
Blumenbach, 1821 (a junior synonym of
Triturus cristatus) when both were
considered congeneric in Triturus)
Type
locality: Crno Jezero, Montenegro.
Triton alpestris lacusnigri Seliskar
et Pehani, 1935
Triturus alpestris lacusnigri
(Seliskar et Pehani, 1935)
Ichthyosaura alpestris lacusnigri
(Seliskar et Pehani, 1935)
Type locality: valid replacement
name (Article 59.3) for Triturus
alpestris lacustris, thus same
type locality.
Triturus alpestris montenegrinus
Radovanovic, 1951
Ichthyosaura alpestris montenegrina
(Radovanovic, 1951)
Type locality: Montenegro, near
the Albanian border.
Triturus alpestris serdarus
Radovanovic, 1961
Ichthyosaura alpestris serdara
(Radovanovic, 1961)
Type locality: Montenegro.
Triturus alpestris piperianus
Radovanovic, 1961
Ichthyosaura alpestris piperiana
(Radovanovic, 1961)
Type locality: Montenegro.
Triturus alpestris montenigrus auct.
(misspelling)
Triturus alpestris montenigrinus auct.
(misspelling)
(5) Clade E2
(northern Balcans, southern
Carpathians). Ichthyosaura alpestris
reiseri:
Molge alpestris reiseri Werner, 1902
Triton alpestris reiseri (Werner,
1902)
Molge reiseri (Werner, 1902)
Triturus alpestris reiseri (Werner,
1902)
Ichthyosaura alpestris reiseri
(Werner, 1902)
Type locality: Vranika mountains,
Bosnia.
Triturus alpestris carpathicus Dely,
1960
Ichthyosaura alpestris carpathica
(Dely, 1960)
Type locality: Sinaia, Romania.
This locality actually quite
exactly matches the locality of
Specimen 32 in Sotiropoulos
et al. (2007).
(6) Clade D (D1-D4;
Greece, Macedonia, southern Serbia).
Ichthyosaura alpestris veluchiensis:
Triturus alpestris graeca
Wolterstorff, 1934 (nec Wolterstorff,
1906; junior secondary homonym of
Triton vulgaris graeca Wolterstorff,
1906 (a good species in the genus
Lissotriton) when both were considered
as congeneric in Triturus)
Ichthyosaura alpestris graeca
(Wolterstorff, 1934)
Type locality: Mount Veluchi,
Greece.
Triturus alpestris veluchiensis
Wolterstorff, 1935
Ichthyosaura alpestris veluchiensis
(Wolterstorff, 1935)
Type locality: valid replacement
name (Article 59.3) for Triturus
alpestris graeca, thus same type
locality.
Mesotriton alpestris velouchiensis
auct. (misspelling)
The generic name of this species is
discussed controversially. First,
there is disagreement whether the
traditional genus Triturus is
monophyletic or not. Several recent
phylogenetic studies suggest that
Triturus in the traditional sense is
not a monophyletic group. Although
details of the phylogenies differ,
they all agree that the group
containing the type species of
Triturus, Triturus cristatus, is not
directly related to the other newt
species. I therefore adopt here the
notion that the traditional genus
Triturus has to be split into several
genera.
Second (if one accepts the splitting
of the traditional Triturus concept),
there is disagreement which genus name
is the valid name for the grouping
containing the nominal taxon Triton
alpestris. The name Ichthyosaura is
the oldest name used as a genus name
for a putative specimen of the present
species. This specimen has been
described as Proteus tritonius (Laurenti
1768) from a small lake in the
Ötscher mountain in Lower Austria and
later the genus Ichthyosaura has been
erected for it (Sonnini
and Latreille 1801 or 1802).
However, the identity of this specimen
is disputed and some authors regard it
as unidentifiable or as belonging to
other species of Salamandridae. In my
opinion there cannot be doubt about
the identity of the specimen described
and figured by Laurenti
(1768). It is a larva with four
fingers and five toes, relatively long
gills and a dorsal fin that almost
reaches the head. This combination of
characters and the given type locality
in Austria rules out that the specimen
is the larva of a species of Proteus,
Salamandra or Salamandrina. The figure
in Laurenti
(1768) clearly shows a tail fin
that does not taper towards the tail
tip and that has a rounded blunt end.
This is the typical fin and tail of
the nominal Triton alpestris and rules
out all other newts of the region.
Thus, the type species Proteus
tritonius of the genus Ichthyosaura is
clearly identical to Triton alpestris
and therefore the name Ichthyosaura is
the valid genus name for the genus
containing Triton alpestris. Please
note that the year of publication of
Sonnini and Latreille´s work is
unclear, because the title page only
gives "AN X", i.e. the year 10 of the
French Republican Calendar and there
is no other indication in the volume
that gives a more precise date of
publication. The year 10 of the French
Republican Calendar that corresponds
to the interval from September 23,
1801 to September 22, 1802, thus the
volume may have been published in 1801
or in 1802.
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Identification
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No
information has been entered yet.
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Distribution
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information has been entered yet.
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Biology
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No
information has been entered yet.
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Male during breeding season.
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