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Neurergus strauchii, Neurergus crocatus, Neurergus kaiseri, Neurergus micropilotus
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


Neurerugs crocatus - photo courtesy of Henk Wallays, Henk.Wallays@pandora.beThe 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. 

Neurerugs strauchii strauchii - photo copyright Jessica J. MillerN. 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 crocatus. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus crocatus. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus crocatus. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus crocatus. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus crocatus. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus crocatus. Photo © Henk Wallays, Henk.Wallays@pandora.be 

For more photos, go to the Neurergus crocatus gallery



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 kaiseri. Photo © Christoph Bork, Christoph.Bork@t-online.de  


Neurergus kaiseri. Photo © Christoph Bork, Christoph.Bork@t-online.de  


Neurergus kaiseri. Photo © Christoph Bork, Christoph.Bork@t-online.de  




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 microspilotus. Photo © Henk Wallays, Henk.Wallays@pandora.be 




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.


Neurergus strauchii strauchii. Photo © Jessica J. Miller, jess@livingunderworld.org  


Neurergus strauchii barani. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus strauchii strauchii. Eggs. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus strauchii strauchii. Advanced larva. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus strauchii strauchii. "Golden" variant juvenile. Photo © Henk Wallays, Henk.Wallays@pandora.be 


Neurergus strauchii strauchii. Normal juvenile. Photo © Paolo Mazzei, Amphibians & Reptiles of Europe

For more photos, go to the Neurergus strauchii gallery



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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