Salamanderland 

Tylototriton shanjing (Nussbaum, Brodie,Yang Datong, 1995)

Mandarin Salamander

First Author:
Nussbaum, R. A., E. D. Brodie, Jr., & Y. Datong (1995): A Taxonomic Review of Tylototriton verrucosus Anderson (Amphibia: Caudata: Salamandridae). Herpetologica 51 (3): 257-268. 

Diagnosis:
A large, massive newt of 17 cm / 6.7 inch overall length. The males are usually smaller. The flat, triangular head with large eyes is clearly set off from the trunk. On the head and along the back bone there are warty crests. The colour is dark-brown to black with characteristically orange marks. The orange marked warty row along the flanks is typical. They correspond with the ends of the ribs. A completely similar colouring can show Tylototriton verrucosus! The tail, the legs, parts of the head and the belly are likewise orange. The colouring reaches from bright orange over rusty up to a reddish brown. The design hardly varies from individual to individual. The skin is occupied with numerous large and small warts. Sexing: The sexes are difficult to determine, in particular out of the mating time. Frequently the male animals are somewhat smaller and slimmer. The most reliable characteristic of the distinction the expert can find in the different shape of the cloacas. In order to examine these, the outside lips of the cloaca has to be spread. While the male cloaca forms an oblong gap is the females rather round. During the reproduction time the males can be recognized by the prominent cloaca.

Differences to Tylototriton verrucosus:
From this taxon Tylototriton shanjing was systematically separated as an independent species (Nussbaum et al., 1995). The characteristic morphological demarcation to the species appears quite „soft” in the first description: It says falsely that Tylototriton verrucosus is an always uniformly black or brown coloured species. But it often has a similar design as Tylototriton shanjing. The marking is however frequently less strong in colour. Indeed it will be quite difficult to differentiate a strongly coloured individual of Tylototriton verrucosus from a paler Tylototriton shanjing. It does not easier that these animals come into our hands exclusively by the pet trade. This is the reason why collecting site data are completely unknown. The maintaining groups probably originate from different habitats. Many keepers of Tylototriton verrucosus try to select their breeding couples to get animals looking most like Tylototriton shanjing. So this two species (morphs?) hardly became distinguishable in the terrariums. It is an urgent necessity to refer these animals from reliable collectors and to isolate them from each other. One has to be careful also with photos from the Internet and the popular literature, since these do frequently not agree with the described species. Data in the primary literature before 1995 are also very difficult to assign because no differentiation between individual variants of Tylototriton verrucosus were made.
Recently (Zhang et al., 2007) strong doubts emerged whether the species status of Tylototriton shanjing versus Tylototriton verrucosus can be held. The authors react to these findings differentiated: Frost (2008) considers the argumentation sufficient to put Tylototriton shanjing into the synonymy of Tylototriton verrucosus, Mahoney & Vrendenburg (2009) wait for further findings. Ziegler et al. (2008) regard the species as valid without any comment. It should be noticed that the work of Zhang et al. (2007) seems to have lacking in proceedings.

Examinating the literature the following provisional suspicion can result: At least two forms seem to exist, which cannot be differentiated morphologically with the present knowledge. Both correspond to the appearance that at present Tylototriton shanjing (kursiv) senso stricto shows. Very clear differences result in the observations of the mating behaviour (Mudrack, 1969, 1972, 2005; Rehberg, 1986). One shows amplexus as Tylototriton verrucosus, which takes place obligatorily in water. The other one mates in a circular dancing on land without amplexus. Mudrack (1969, 1972) observed first the aplexus and some years later with other individuals (2005) the circular dance. Rehberg however (1986) only described the circular dance. Already Rehberg (1986) expressed the suspicion that one could have observed two different species, which would not be   distinguishable macroscopically.
With our present means it does not seem to be possible to differentiate both forms phenotypically. Another fact seems to be that the amplexus maybe can hybrid with Tylototriton verrucosus in the terrarium. Animals living now in our terrariums are under suspect to be hybrids of Tylototriton verrucosus senso stricto and Tylototriton shanjing senso lato. On the other hand Tylototriton shanjing of the circular dance form will not mate with Tylototriton shanjing of the amplexus form. We must search for the species border between these forms.
Mudrack (2005) gave very valuable material from an accurately well-known habitat at his disposal (Garfong village, Jingdong County, Yunnan Province) for his second investigation. This is the distribution area of Tylototriton shanjing senso stricto. These animals showed the circular dance. It is possible that Tylototriton verrucosus developed a cryptic form, which lives synoptically with Tylototriton shanjing. Both forms look alike to a large extent and  so far did not recognize the fact.
The hypothetical postulate could be: There is one „true Mandarin salamander“ looking alike Tylototriton shanjing senso stricto and one „false Mandarin salamander“ which is a colouring variant of Tylototriton verrucosus. Both forms are macroscopically not distinguishable.
Mayer  wrote in an additionally report: „If the story with the different mating behaviours is true, I am quite sure that there are actually 2 species. One of it will probably be verrucosus. But we need samples from both forms. Sounds very interesting! Without such an investigation a breeding program seems a pure plaything to me. It makes reliably little sense to breed a colour morph of verrucosus for a preservation programme…. “

Status of the subspecies:
So far no subspecies were described. Only  individuals deviating clearly in the morphology from the typical animals can be found in the international pet trade. Even the teeth exhibit partially differences. But without useful origin data a differentiation is impossible. The species has been seen as a colour morph of Tylototriton verrucosus up to its description

Distribution:
Terra typica: Dingpa, Jingdong County, Yunnan Province, China.
The species is limited on habitats of the province Yunnan. It settles areas at the rivers Nujiang, Mekong and Yuanjiang in 5 prefectures: Lijiang, Dali, Dehong, Kunming and Xishuangbanna. Tylototriton shanjing is limited in an area of 196073 km² / 121834 miles² in Yunnan on the northern, western and southwest portions of the province. At the Chinese west border to Myanmar the enormous distribution area of Tylototriton verrucosus begins, which is extended to northern India and Nepal. There is no known sympatric occurrences of the two species.

Habitat and ecology:
Little is known about the natural habitat. The animals live in rice fields in subtropical rainforest areas of the mountainous altitudes between 1000 and 2500 m / 3280 and 8202 feet asl. particularly at slowly flowing brooks, small ponds and irrigation canals. They seem to prefer moist and for local conditions cool habitats. The climatic course of the year in the area is clearly biphasic: A relative dry and cool time during the winter months and the monsoon time, which lasts from  the middle of May to October. The temperatures are highest during the monsoon, but vary strongly among different regions. The values measured by the meteorological stations can not be directly compared with the temperatures in the habitats. These might be during the warm season between 20° and 27°C / 68° and 80.6° F and decreases in the cool phases to lowest values of 8°C / 46.4°F. 

Food:
Nothing is known about nutrition in the natural habitat. The animals are greedy eaters in the terrarium. Middle and larger objects are preferred. Food, which is smaller than 4-5 mm / 0.15-0.19 inch, will not be recognized by the adults. All food organisms of suitable size used by salamander keepers are accepted.

Reproduction:
There is only little known about the reproduction in the natural habitat. The reproduction period partly covers with the monsoon time and lasts from May to August. The animals are spawning in smaller and larger standing and slowly flowing waters and set their eggs off separately or in groups at stones and plants. Outside of the reproduction time the species lives terrestrial.

Keeping in the terrarium:
Due to the limited data of natural habitats the installation of the container is difficult and the phantasy of the keeper is demanded. The most favourable way is the aquaterrarium. This species prefers higher temperatures in the summer within the range of 20°-27°C / 68°-80,6°F. The animals usually look for the damper places in the terrarium and stay mainly on land. Outside of the reproduction time they can also be kept in a terrarium simply on land. Originally active during dawn and night the animals get accustomed to their artificial environment and can be observed in times of suitable air humidity also frequently outside of the hiding places during the day. The species can be kept in terrariums of different sizes. If no display is needed, the interior should be held reduced. The entire surface forms the water part with a level of 8 cm / 3.15 inch. Two pieces of bricks carry a glass plate, which is covered with foam material. It forms the land part. A mass of bark offer many hiding places. The water is filtered by an interior filter. The water should be ventilated. If situated outside no ventilation is needed.
Due to the fact that the animals are bad swimmers and stay hardly in water, a level of more than 10 cm / 3.9 inch is not useful. The animals should be able to crawl easily on land. They can be fed with rope worms, earthworms, waxworms, maggots, zophobas, crickets. Artificial food, dead food or frozen organisms are hardly accepted. The hardiness of the water should not be too low and the pH should be around 7. High hygiene standard is of great importance . Water should be as clean as possible. Stress will be badly tolerated . Inadequate interior, unsuitable food, not enough hiding places, environment change (transport!) etc. can cause rapidly dramatic losses, in particular with young animals.

Reproduction in the terrarium:
The animals mate in April/May after an appropriate cooler, drier winter time. The temperature does not have to be lowered more than around 15°C / 59°F. In addition, it is possible to hibernate the species in the same temperature regime (4°C / 39.2°F) like Triturus. For the introduction of the mating high air humidity (90%) and increased temperatures above 20°C / 68°F (monsoon!) is needed. Such conditions prevail in Europe occasionally in the summer months. If kept outside, mating will start  that days. The influence of the photo period on the reproduction is discussed frequently, but seems to play an insignificant role. A large problem in finding suitable reproduction partners is the usually unknown origin of the individuals. It might be difficult to synchronize animals from the subtropical lower regions with such from mountains. It would not be impossible to mate the amplexus type with the type of circular dance successfully. Maybe these circumstances are the reason for still rare breeding success so far. The mating takes place in the typical circular dance on land. During that the male sets off several spermatophores.  (Photos here and Mudrack, 2005) Tylototriton shanjing senso stricto never shows amplexus. This stands in contrastto Tylototriton verrucosus and speaks for the validity of the species. (Individuals of Tylototriton shanjing senso lato, which mate with amplexus, would be another form, standing near Tylototriton verrucosus) The animals of the author, they are kept for years, show the circular dance obligatorily on land and the eggs are laid on the shore to the majority. This will occur even if the land part is small (20 x 60 cm / 0.65 x 1.96 ft) and the water part is proportionally very large (approx. 0.8 m² / 2.62 ft²). The female starts to spawn a few days after mating. The eggs are large and there will be 300 and more. Oviposition will  mainly occur on and above the border between water and land, as well as on damp places ashore. The criterion for the choice of the substrate seems to be a certain firmness and humidity. Weak plants or other weak structures as well as completely dry places are avoided. Unfortunately frequently the fertilization rate of the eggs is low. The cause for this is unknown.
The diameter of the eggs is 6-7 mm / 0.23-0.27 inch with a diameter of the yolk of approx. 2 mm / 0.08 inch. The larvae hatch at temperatures between 20° and 24°C / 68° and 75.2°F after 3-4 weeks with a size of 13 to 15 mm / 0.5 to 0.6 inch. The raising of larvae prepares few problems at relatively high temperatures around 23°C / 73.4°F. But the larvae grow with different speed and if necessary must be sorted out according  sizes. The larvae are tendentious preying on each other. With sufficient hiding places (strands of synthetic wool tied together work satisfactorily) and capacity of the tanks it is not necessary to keep them single, if sufficiently fed. The raising in bald tanks (Ziegler et al., 2008) is not meaningful, since this can form an additional stress factor. (The animals try to escape and jump out of the container!) First the hatchlings are fed with Artemia after consuming the yolk. Feeding at least twice a day is indicated. Starting from a size of approx. 20 mm / 0.78 inch larger food organisms are needed, like Tubifex, Daphnia, red mosquito larvae etc. Attention to a maximum of hygiene should always be paid. Dead food leads to infections at legs and gills and will kill the larvae within hours. Sufficient filtering and the reduction of pollution in the water by UVC are recommended. The metamorphosis starts from the 4th month and lasts for a longer period. It will take again 4 months until all latecomers will be out.

Development in the terrarium:
The raising of the young animals requires a high grade of hygiene and the continuous supplying of different food organisms of high quality. (Crickets gutloaded of suitable sizes, Drosophila, Enchytraeus, different species of insects etc.). The young animals react very sensitively on environment changes. For this reason the cleaning must be done carefully. Even necessary transport should be done carefully and fast. When the young animals reach the age of one year, they become clearly more stable and further growing up takes place without too many problems. This might be justified with a now sufficiently high production of skin poisons, which makes the animals recognizable more resistant against infections. Young animals aged less than one year seem to have insufficient skin poisons. At the age of 4-5 years the salamanders become adult. Their longevity will be more than 10 years.

Conservation Status:

Statements about the threats of this species according to the present data are purely speculatively. Even the systematic position of collected individuals can not be indicated for sure nowadays. Probably Tylototriton shanjing suffers under destroying of habitats and as a wanted species for pet trade there might be over-collection. However Rehberg (1986) already reports of regular imported individuals, which unabatedly takes place since then. In comparison with the high reproduction rate of the species collecting for pet trade might lead to a serious threat, only if this takes place in combination with destroying of habitats.

Literatur:
Frost, D. R. (2008): Amphibian Species of the World: an Online Reference. Version 5.2 (15 July, 2008). Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.php. American Museum of Natural History, New York, USA.

Mahoney, M. & V. Vredenburg (2008): Tylototriton shanjing in: AmphibiaWeb: Information on amphibian biology and conservation. [web application]. 2008. Berkeley, California: AmphibiaWeb. Available: http://amphibiaweb.org/. (Accessed: Dec 22, 2008).

Mudrack, W. (1969): Tylototriton verrucosus Anderson, 1871, ein seltener Molch aus Asien. Aqua Terra., 6, 134-136.

Mudrack, W. (1971): Tylototriton verrucosus Anderson, 1871, seine Pflege und Zucht. Aquarien- u. Terrarien Z. 24 (11), 388-390.

Mudrack, W. (1972): Ein seltener Krokodilmolch – Tylototriton verrucosus. Vom Ei zum Jungtier. Aquarien Mag., 6 (10), 406-409.

Mudrack, W. (2005). Nachzucht von Krokodilmolchen, Tylototriton shanjing. Amphibia, 4(1), 23–25.

Nussbaum, R. A., E. D. Brodie Jr., & Y. Datong (1995): A taxonomic review of Tylototriton verrucosus Anderson (Amphibia: Caudata: Salamandridae) Herpetologica, 51(3), 257-268.

Rehberg, F. (1986): Haltung und Zucht des Krokodilmolches, Tylototriton verrucosus. Herpetofauna, 8 (45): 11-17.  

Zhang, M., D. Rao, G. Yu & J. Yang (2007): The validity of Red Knobby Newt (Tylototriton shanjing) species status based on mitochondrial Cyt b gene. Zoological Research, 28(4), 430-436.  

Zhao, E. (1998): China Red Data Book of Endangered Animals: Amphibia and Reptilia. Science Press: Endangered Species Scientific Commission, P.R.C., Beijing.  

Ziegler, T., T. Hartmann, K. Van der Straeten, D. Karbe & W. Böhme (2008): Captive breeding and larval morphology of Tylototriton shanjing Nussbaum, Brodie and Yang, 1995, with an updated key of the genus Tylototriton (Amphibia: Salamandridae). Der Zoologische Garten, 77, 246-260.

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