Neurergus (Cope, 1862) Old
World / Middle Eastern Newts
Neurergus crocatus (Cope, 1862) Lake
Urmia Newt / Azerbaijan Newt
Neurergus kaiseri (Hallowell, 1860)
Luristan Newt / Kaiser's Spotted Newt
Neurergus microspilotus (Nesterov,
1916) Kurdistan Newt
Neurergus strauchii (Steindachner,
1887) Anatolia Newt / Stauch's Spotted Newt
The genus Neurergus is comprised of four species, N.
strauchii, N. crocatus, N. microspilotus, and N. kaiseri.
All four newts are found in the Middle East, in the countries of Iran,
Iraq, and Turkey, and occupy ranges of small proportion compared to many
other Salamandrid species.
The genus Neurergus is closely related to the Old World genus Triturus,
with which many similar characteristics are shared (Schmidtler, 1994).
N. kaiseri ins particular is especially similar to the Alpine Newt, Triturus
alpestris, in terms of morphology and habit.
N. strauchii, N. crocatus, and N. microspilotus are similar in general appearance, with dark colored bodies contrasting
bright yellow spots, and in fact, the genus was once considered a
monotypic one. N. kaiseri is perhaps the "outcast" of
the group, with its
black and white mottling, and orange dorsal stripe. N. kaiseri also
reproduce in still ponds, while the other three species are stream dwellers. N.
kaiseri's unique reproduction environment is thought to be a
representative example of genetic adaptation, as opposed to environmental
plasticity (Steinfartz et al., 2002). The larvae of N. kaiseri
are definitively pond type, with squat bodies, long gills, tall tail
fins, and a shorter larval period. There are also remarkable
osteological differences between N. kaiseri and N. strauchii
(Haller-Probst & Schleich, 1994).
Such genetic adaptations might
warrant separation at the genus level, or higher, in many cases, however, the
stream-dwelling characteristic that dominates in Neurergus is thought to be ancestral,
with the intrageneric conversion to pond environments in the case of N.
kaiseri occurring at some point in the
evolutionary process (Steinfartz et al., 2002), and subsequently leading
to genetic adaptations. This scenario is more
likely than a case of recurrent parallel evolution from a pond type
reproduction to a stream type in the case of Neurergus.
Neurergus crocatus (Cope, 1862)
Lake Urmia Newt, Azerbaijan Newt
Taxonomic Synonyms: Neurergus
crocatus (Cope, 1862), Molge crocata (Boulenger, 1890), Rhithrotriton
derjugini (Nesterov, 1916), Triturus crocatus (Fowler & Dunn, 1917),
Triton crocatus forma crocatus (Wolterstorff, 1926), Triton crocatus
forma derjugini (Wolterstorff, 1926), Triton (Mesotriton) crocatus (Bolkay,
1927), Neurergus derjugini (Terentjev, 1938), Triturus crocatus crocatus
(Bodenheimer, 1944), Triturus crocatus derjugini (Bodenheimer, 1944)
Vernacular Names: Lake Urmia Newt, Azerbaijan Newt
N. crocatus are similar to N. strauchii in size and
coloration; i.e. dark colored dorsa and contrasting yellow
spots, and lengths of up to 18 cm. Hoewever, the
spots of N. crocatus are larger than those of N. strauchii,
and the belly is solid or nearly solid red-orange, as opposed to the
thin orange ventral line of N. strauchii (Sparreboom et al.,
2000). N. crocatus males do not develop a bluish sheen along the
tail, as is observed in N. strauchii.
Neurergus kaiseri (Schmidt, 1952) Luristan Newt
Taxonomic Synonyms: Neurergus
crocatus kaiseri (Schmidt, 1952), Neurergus crocatus kaiseri (Thorn,
1968), Neurergus kaiseri (Schmidtler & Schmidtler, 1970)
Vernacular Names: Luristan Newt
As mentioned earlier, N. kaiseri is markedly different in
appearance and morphology than the other species. N. kaiseri
possess large, irregular black and white markings of various
size and shape about the dorsum. The percentage of black or white
coloration varies among individuals,
with some being mostly white, mostly black, or anywhere in between. N.
kaiseri also possess orange colored dorsal stripes, bellies,
undersides of limbs, and
orange coloration about the upper limbs and feet, adding to the striking color
and pattern of this species. N. kaiseri is the smallest species,
reaching only 10 cm in total length (Spareboom et al., 2000).
Neurergus microspilotus
(Neserov, 1917) Kurdistan Newt
Taxonomic Synonyms: Rhithrotriton
derjugini var. microspilotus (Nesterov, 1916), Triton crocatus forma
microspilota (Wolterstorff, 1926), Triton crocatus microspilotus (Bokay,
1927), Neurergus crocatus microspilotus (Schmidt, 1939), Triturus
crocatus microspilotus (Bodenheimer, 1944), Neurergus microspilotus (Schmidtler
& Schmidtler, 1975)
Vernacular Names: Kurdistan Newt
N. microspilotus is also dark colored with bright yellow spots.
This species is rather small compared to N. strauchii and N.
crocatus, reaching lengths of 14-15 cm total. The ventral coloration
is similar to N. crocatus, consisting of solid or nearly solid
orangish-red. Like N. crocatus, N. microspilotus males do
not develop a bluish sheen along the tail during breeding season (Sparreboom
et al., 2000).
Neurergus
strauchii (Steindachner, 1887) Anatolia Newt, Strauch's Spotted Newt
Taxonomic Synonyms: Molge strauchii
(Steindachner, 1887), Triton crocatus forma strauchi (Wolterstorff,
1926), Neurergus crocatus strauchii (Schmidt, 1939), Triturus crocatus
strauchii (Bodenheimer, 1944), Neurergus strauchii (Schmidtler &
Schmidtler, 1970), Neurergus strauchii barani (Oz, 1994)
Vernacular Names: Anatolia Newt, Strauch's Spotted Newt, Baran's
Spotted Newt (spp. barani).
N. strauchii spp. are medium sized newts, reaching up to 18 cm in
total length. The dorsum is black or dark brown with contrasting yellow
spots. The spots are found from the head, to the tip of the tail,
including the limbs, head, and flanks. N. s. barani possess
spots of smaller circumference than those of N. s. strauchii. The
belly of both subspecies consists of an irregular orange line
extending from the chest to the tail tip, and surrounded in dark
coloration that lacks yellow spotting. When aquatic, the skin may take on a rather velvety texture,
while terrestrials usually have moderately rough skin. Males of both
subspecies develop laterally compressed tails, and a bluish-white sheen
along the lateral side. This dimorphic trait is not
observed in any other Neurergus species, and in fact is the only
sexually dimorphic color change observed in the genus (Sparreboom et
al., 2000). Over-wintered larvae develop adult coloration before
metamorphosis, and so juveniles typically possess adult coloration upon
metamorphosis. This species also produces a "golden" variant,
pictured below, whose yellow spots look as though they've been smeared
together.
Distribution & Habitat
All four species are found in the Middle East, in the countries of Turkey, Iraq,
and Iran. N. crocatus are found at the convergence point of the three countries,
within a range that includes southeast Turkey, northwestern Iran, and
northeastern Iraq (Steinfartz, 2002). Both subspecies of N. struachii are
found west of the Van Lake in Turkey, in two disconnected zones of the eastern
and central areas (Sparreboom, 2000). N. microspilotus is found
along the mid-border of Iran and Iraq, extending slightly into both countries. N.
kaiseri are found in western Iran, in the province of Luristan (Sparreboom,
2000.
 The ranges occupied by all species are rather small in comparison to
some other species, and all are disconnected. The map below, adapted from
Steinfartz, 2002, shows an estimation of distribution of all four species. As
mentioned previously, N. strauchii, N. crocatus, and N. microspilotus are
stream breeders, while N. kaiseri utilize still ponds for reproduction.
Outside of the breeding season, Neurergus typically remain terrestrial. The
terrestrial habitat is considered rather arid, and includes de-forested
grasslands, and semi-deserts near mountain streams at elevations of 500-1750 m.

Mountain Stream, Zagros Mountains, Iran, photo ©
Dr. Attila I. Gulyás, Institute of Experimental Medicine, Hungarian Academy of
Sciences, http://www.koki.hu/~gulyas.
Courtship
& Breeding
This section consists of a summary of results obtained by Sparreboom et al.,
2000, and should be viewed as a general guide to breeding behavior. The
courtship behavior of Neurergus is similar for all four species, and
resembles that of some Triturus species, namely T. alpestris. N.
crocatus, N. strauchii, and N. microspilotus will actively pursue
females and attempt to block their path, while pursuit is limited in N.
kaiseri. N. microspilotus males survey their surroundings from an
elevated spot, and return there after unsuccessful pursuits. Tail fanning is
observed in all four species, which consists of the male folding the tail along
the side of the body and undulating the the entire tail, or only the distal portion, usually while standing
perpendicular to the female. Males may fan from either the left or right side.
The amplitude of the fan is smallest in N. kaiseri. There is a noticeable
difference in the duration of fanning bouts between N. strauchii and N.
crocatus, with those of N. crocatus being longer in duration than N.
strauchii. After a period, the male will attempt to lead the female off by
creeping in front of her, while undulating a slightly raised tail. A responsive
female will follow, occasionally undulating her tail. Shortly thereafter, the
male deposits
a spermatophore and leads the female over it, at which point he pivots 90°
to a perpendicular position. This is commonly known as the "brake",
and effectively stops the females progression such that her cloaca is just over
the spermatophore. Competing males of N. strauchii may
interrupt a courtship display in progress, thus assuming the courting position. Females
of N. strauchii and N. crocatus deposit eggs on the undersides of
flat rocks, while N. microspilotus opt for crevices or hollows of stones.
N. kaiseri females deposit eggs on shaded, rough surfaces, including, but
not restricted to the undersides of stones.

N. s. strauchii eggs (left), N.
crocatus larvae (right), photos ©
Henk Wallays, Henk.Wallays@pandora.be
Proposed Monophyly
Molecular studies using N. strauchii as a
representative of the entire genus place Neurergus close to Triturus and
Euproctus asper (Steinfartz, 2002). As mentioned previously, there are
notable osteological differences between N. strauchii and N. kaiseri, while
N. kaiseri shares similar morphology and habit with Triturus alpestris
(Haller-Probst & Schleich, 1994). have shown that
Molecular studies have supported monophyly of Neurergus. Steinfartz et al., 2002, showed the existence of two further
clades within Neurergus; the crocatus-clade, consisting of N. crocatus, N.
microspilotus, and N. kaiseri, and the strauchii-clade,
comprised of only one species and two subspecies, N. s. strauchii and
N. s. barani. The two clades separated somewhere in the range of
8.5-13.9 mya, while speciation occurred within the crocatus-clade
at an estimated 5 mya, and finally subspecies differentiation within N.
strauchii occurred around 2.5-3 mya (Steinfartz et al.,
2002).

Journal References
Haller-Probst, M., H.H. Schleich. 1994. Vergleichende osteologische
Untersuchunen an einigen Urodelen Eurasiens. Courier Forsch, Inst.
Senckenberg 173: 23-77.
Schmidtler, J.J., Schmidtler, J.F. 1994. Eine Ubersicht neuerer
Untersuchungen und Beobachtungen an der vorderasiatischen Molchgattung Neurergus
Cope, 1862. Abh. Ver. Naturk., Magdeburg 17: 193-198.
Sparreboom, M., S. Steinfartz, G. Schultschik. 2000. Courtship behavior of Neurergus
(Caudata: Salamandridae). Amphibia-Reptilia 20: 1-11.
Steinfartz, S., U.W. Hwang, D. Tautz, M. Öz, M. Veith. 2002. Molecular
phylogeny of the salamandrid genus Neurergus: evidence for an intrageneric
switch of reproductive biology. Amphibia-Reptilia 23: 419-437.
Weitere, Markus, D. Tautz, D. Neuman, S. Steinfartz. 2004. Adaptive
divergence vs. environmental plasticity: tracing local genetic adaptation of
metamorphosis traits in salamanders. Molecular Ecology 13: 1665-1677.
Book References
Obst, Fritz Jugen, Udo Jacob, and K. Richter. Completely Illustrated Atlas
of Reptiles and Amphibians for the Terrarium.
Neptune City, NY: T.F.H. Publications, Inc., 1989.
Internet References
Rastegar-Pouyani, Nasrullah (2003). FrogLog 56: Ecology and Conservation of
the Genus Neurergus in the Zagros Mountains, Western Iran. Declining
Amphibian Populations Task Force.
http://www.open.ac.uk/daptf/froglog/FROGLOG-56-2.html
(Accessed: 2004)
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