Tylototriton species :
1- Tylototriton shanjing
2- Tylototriton verrucosus
1- Tylototriton shanjing :
courtesy to : www.caudata.org
Synonyms : Emperor Newt, Halloween Newt, Mandarin Salamander
Range : China: western, central and southern Yunnan
IUCN (Red Book) : Near Threatened
CITES : No listing
First described : Nussbaum, Brodie & Yang, 1995
A robust newt with granular skin and prominent boney ridges on the head and spine. A row of warts runs down each side. “The dorsolateral surfaces of these salamanders are marked vividly with orange, including the cranial crests, parotoid glands, middorsal crest, and lateral warts. In addition, the lower sides, limbs, tail, and venter are various hues of orange-yellow” (Nussbaum et. al., 1995). The rest of the animal is dark brown to black. The brightness of the orange markings varies between yellow-orange to almost rust colored. Although color varies between individuals, the patterns of the markings are relatively constant. Only slight variation is noted in the extent of orange coloration.
Natural Range and Habitat :
Found in the Yunnan province of China. It lives in the mountainous areas near the Nu, Yuan, and Lancang rivers. (Mahoney and Vredenburg, 1999)
A normally-colored individual, and an unusually light-brown individual.
This species requires only a small body of water. Although they rarely enter the water outside of breeding season, it’s always best to have it there in case the cage becomes too dry. Completely terrestrial, it should be given plenty of land area as well as places in which it can conceal itself. I have found that tightly packed soil approximately 2 to 3 inches deep works wonderfully. Use the substrate to create a moisture gradient so that one end of the enclosure is very wet and the other dry. Do this by adding water to only one end of the enclosure, it will slowly seep through the soil. This allows the animal to choose what it prefers. Do not allow the soil substrate to become muddy or too much moisture to build up, as they prefer fairly arid conditions.
If healthy, they feed well on a variety of foods. Common sources are wax worms, earthworms, chopped night crawlers, mealworms, and crickets. Feed a variety to assure balanced nutrition.
In the wild they typically breed May through August. They deposit eggs on rocks and plants in standing water bodies. (Mahoney and Vredenburg, 1999) In captivity, breeding can be initiated by mimicking their natural seasonal conditions. One way is to keep the animal in the lower 60s and on a fairly dry substrate during the winter. Then during summer, raise their temperature to the upper 70s and increase the humidity. If breeding is to occur, supply the captives with enough water for courtship and for depositing eggs, while still maintaining a nice sized land area.
A darker individual.
Cloaca of T. shanjing male.
Cloaca of T. shanjing female.
Mahoney, Meredith and Vance Vredenburg. Tylototriton shanjing: Yunnan Newt. University of California Berkeley. Nov. 18, 1999. .
Nussbaum, Ronald A., Edmund D. Brodie Jr., and Yang Datong. 1995. A taxonomic review of Tylototriton verrucocsusAnderson (Amphibia: Caudata: Salamandridae). Herpetologica, 51(3):257-268 .
2- Tylototriton verrucosus:
Synonyms :Crocodile Newt, Alligator Newt, Himalayan Salamander, Red Knobby Newt, Burmese Crocodile Newt
Origin : China, India, Myanmar, Nepal, Thailand and Viet Nam
IUCN : Red BookLeast Concern
CITES : No listing
First described : Anderson, 1871
n 1995, Nussbaum et al. published "A taxonomic review of Tylototriton verrucosus". Until 1995, most if not all authorities considered Tylototriton verrucosus and T. shanjing to be the same species. Comparing individuals of these species, the differences become quite apparent, and one may well wonder why the two were not considered separate species from the outset. A photo of a bright orange newt with the caption "Tylototriton verrucosus" is a common sight in books and literature. This situation is complicated by the fact that the species known as Tylototriton verrucosus occupies a large range (see below), and animals from different parts of the range may exhibit distinct morphological and behavioural differences, including large differences in adult size, colouration and body/appendage shape. These differences have led me to conclude that T. verrucosus is quite probably a complex of several subspecies or perhaps even full species. However, until phylogenetic and morphological studies are carried out on animals from known localities across the range, this group will remain one species. In my experience, there are at least two, possibly three commonly available variants within this species. For the present work these variants are:
1. T. shanjing-like" variant: This variant exhibits similarities in colouration and morphology to those of T. shanjing. The head is similarly shaped to that of T. shanjing, being less triangular and narrower than that of variants 2 and 3 described below. Maximum adult size: ~17 cm (~7 inches).
2. "Light" variant: The largest variant, possessing a relatively slim profile, relatively narrow tail and head with faded/lighter brown colouration than variant 3. The head is wider than that of T. shanjing and it possesses some traits of variant 1 and some of variant 2. Maximum adult size: ~24 cm.
3. "Dark" variant: The least similar of the three variants to T. shanjing, possessing more rounded body features such as a wider head and body (less noticeable when comparing males), higher tail fin and dark brown to black base colouration. It has been my experience that the dark variant is more hydrophilic (i.e.the dark variant seems to like aquatic conditions more than the light variant). This variant possesses the most angular head of the three variants. Maximum adult size: ~17 cm but generally a little smaller.
Unfortunately locality data for the three variants is not known with certainty due to the acquisition of animals through the pet trade and the frequent "inaccurate" reporting of locality data by exporters. The three proposed variants are pictured below. Variants 1 and 2 appear to be quite similar but the head shape appears to be distinct and there is a large contrast between the adult body lengths of each.
All variants of the Himalayan Crocodile Newt are large and robust newts. The base colouration is chocolate brown-to-black, with areas of orange or brown on the tail, head, legs and tubercles of the rib peaks. The degree of orange varies between individuals and between “variants” and adults tend to be darker than juveniles. The head is flattened when viewed from above (less so in variants 1 and 2) and angular (in variants 1 and 2 this is less apparent and T. shanjing possesses a relatively straight head). The sides of the head have bony and glandular areas elevated to the level of the upper eyelids (Kuzmin). There are tubercles on the rib peaks, as in T. shanjing, but unlike T. shanjing the dorsal ridge of T. verrucosus variant 3 tends not to be bright orange. In variants 1 and 2 this ridge is generally a noticeably lighter brown than the base colouration or even slightly orange. When aquatic, the skin texture becomes less granular and the base colouration darkens, leading to more contrast with the lightly coloured or orange highlights.
The Himalayan Crocodile Newt is the most aquatic of the four species of crocodile newt (Tylototriton) that are likely to be encountered in captivity, the others being T. shanjing, T. kweichowensis and T. taliangensis. Depending on temperature, during the spring, summer and autumn months these newts will often remain completely aquatic and will rarely, if ever, emerge from the water. This is a very active and inquisitive newt when aquatic, and it can become reasonably tame in captivity, going so far as to swim actively at the glass for food and taking food from forceps and fingers.
These newts can live in excess of 10 years and probably a lot longer.
Natural Range and Habitat :
Tylototriton verrucosus has been recorded from Yunnan Province in China (Zhao and others), northern Thailand, northern Vietnam, northern Burma, northeastern India, Bhotan and eastern Nepal. It is debatable whether some of these records are of Tylototriton verrucosus and not T. shanjing. Its range seems to follow areas of reasonable elevation (foothills of mountains, etc) and it is said to occur in various habitats where mountain forests exist or previously existed, such as cultivated rice fields (Kuzmin and others). It is said to be unfussy when it comes to breeding pools, using anything from puddles to lakes.
These observations are primarily based on my experience with variant 3 and to a lesser extend on my experience with variant 2. Above about 18°C (64°F), the Himalayan Crocodile Newt can be maintained in completely aquatic conditions. An aquarium measuring 60 cm long x 30 cm deep x 37 cm tall (24 x 12 x 15 inches) is adequate for 2 adult pairs of variant 3, fewer in the case of the variant 2. Water depth is not important, but 12-15 cm (5-6 inches) is a good level. If maintained in an aquatic setup, a sizable island should be provided for the animals to emerge from the water should they choose to do so. An animal that spends most of its time out of the water (some young animals behave like this, particularly those of variant 2 and probably variant 1 as well) should be moved to a semi-aquatic setup.
In my experience, a good temperature range for these animals is 15-18°C (59-64°F) in winter with a rise in summer to about 25°C (77°F). Variant 3 can tolerate much higher (for periods of several days) and lower temperatures than these without ill effects. Filtration, if required, can be accomplished in any of the usual ways, and these newts are reasonably tolerant of flowing water, though as a general rule with pond-type newts this should be kept to a minimum. Like most newts, this species does well in a planted tank. A photoperiod of about 12 hours a day in summer is a nice touch, though not essential. Remember to ensure water quality is maintained by making regular water changes - 20-30% or so every two weeks is usually adequate unless there is a lot of waste food and detritus. Remember to treat tap water with a water conditioner prior to use as it may contain harmful substances (chlorine, chloramines and metals).
If you choose a semi-aquatic setup, such as a 50% land:50% water arrangement, be sure the animals can emerge from the water without difficulty.
T. verrucosus pair in mating dance.
T. verrucosus pair in amplexus.
T. verrucosus are often very tame.
These newts are not reluctant feeders, and large females can consume an astonishing amount of food in a short time. They will eat earthworms (whole small-to-medium worms or chopped large worms), tubifex worms (live, frozen, or freeze-dried - some times incorrectly labelled as "bloodworms"), bloodworms (the larvae of Chironomus midges), blackworms (an aquatic relative of earthworms), crustaceans like shrimp, pieces of fish (avoid salted fish and marine fish), strips of beef heart or other lean red meat, etc. Like most newts, when aquatic they have a very keen sense of smell and they can find uneaten food, even at night in total darkness. As with most newts and salamanders, the warmer they are kept, the more regularly they should be fed. As much food as they will eat in 15 minutes is a good guide. They can be fed twice a week in summer, but I personally feed mine once every two days and less frequently in winter. If you feed a large amount of food at each sitting and then a few days later you observe your animals spending large periods of time on their island during warm periods, it is a good indication of high ammonia or nitrite levels present in the water from uneaten food, and one would be wise to feed more regularly with small amounts.
One final important note regarding feeding. Females tend to be more aggressive than males, and in a tank containing mixed sexes, the males frequently get less food than females if left to fend for themselves. It is important to monitor males to be sure they are eating well. Females tend to have a much higher capacity for food than males, so don't judge a male's eating habits by comparing its bulk to a female.
Female with eggs.
The following mating and courtship information is based on the author’s experience with variant 3. Size at maturity and courtship behaviour may vary for the other variants. These newts can reach sexual maturity in just a year, usually at a minimum length of 13 cm (5 in) in males and slightly larger in females. Males tend to be more brightly coloured than females, though this isn't a strict rule. The fin of the male may deepen during the warmer months of the year and the animal is aquatic. The cloaca of the sexually mature aquatic male is noticeably swollen when compared to that of the female. Animals not in breeding condition can be sexed by comparing the length of the cloacal opening - in males this is slightly longer than in females - and on average the male's tail fin height is greater than that of the female. One strange difference between the sexes of variant 3 is that the eyes of the sexually active male appear to be surrounded by a pale line which is not as highlighted in the female. The key to stimulating breeding in these newts seems to be a temperature change. In my experience, a temperature increase from 18°C (64°F) to 22-25°C (72-77°F) over a few hours or even a few days usually leads to breeding activity as will a large water change. This alone seems to be adequate, but photoperiod probably also plays a role. Males tail-fan females in the same manner as Triturus newts, but at a slower rate. I have observed that the male will clasp the female in aPleurodeles-like embrace on several occasions. Some sources describe spermatophore exchange taking place with the male using this embrace to guide the female onto a spermatophore, as is the case inPleurodeles. However I have observed spermatophore transfer take place many times during a circular courtship dance, similar to that employed by T. shanjing. The male deposits a number of spermatophores as he moves in the circle and the female picks up one or more in her cloaca. I have never observed spermatophore transfer via an embrace. In my experience males will only embrace females in the presence of other males, i.e. it is a method of monopolising a female. The male will frequently swim with the female in his grasp to an area of the tank that is free of other animals, and then a circular courtship will commence. Mating usually takes place at night. When the tank is allowed to undergo a yearly cycle of temperature change mating should occur naturally without any interference from the keeper.
Oviposition generally begins within 36 hours of mating, though sometimes it may take longer. In captivity, females can lay anything from 30 eggs to over 150 or more in one spawning, depending on the size and bulk of the female in question. The author once counted in excess of 300 eggs laid by one female in one spawning which led to over 280 larvae hatching (this was verified by taking a photograph of the bare tank and counting the larvae by hand). The eggs are relatively small for the genus at about 5 mm in diameter, including jelly. At 25°C (77°F) the eggs hatch in 10-12 days and are faster developing than many other species (e.g. Cynops ensicauda). Newly hatched larvae are about 8-9 mm long and only begin feeding a few days after hatching during which time they utilise the remainder of the yolk in their digestive tract (visible as a cream-coloured substance in the belly). The larvae do well at temperatures between 20 and 27°C (68-81°F), which is quite warm compared to most species. The initial few days can be a delicate time, but foods such as newly-hatched brine shrimp, young Daphnia and microworms are ideal first foods. Once the larvae reach 12 mm or so, young Daphnia can make up the bulk of their diet. When they reach 20-25 mm, they can begin feeding on foods such as frozen bloodworms and live blackworms (preferably minced/chopped as much as possible), in combination with adult Daphnia. Metamorphosis is reached at between 5 and 7 cm, though sometimes larvae will metamorphose at smaller or much larger sizes. Paedomorphic individuals occur in most clutches, and these sometimes don't finish metamorphosis until after sexual maturity is attained at 12 cm or more (Sparreboom; Wallays). They are quite capable of breeding in this state (Sparreboom). Unusual for most newts, after metamorphosis, Himalayan Crocodile Newts usually don't require a terrestrial phase, though a semi-aquatic setup is probably best until the animals are at least 10 cm (4 inches) long.
A series of photographs showing the early development of this species is available on this site atTylototriton verrucosus Developmental Photo Series.
Duellman, William E. and Trueb, Linda (1994). Biology of Amphibians. Johns Hopkins University Press.
Fleck, J. (1996). Neues über die Gattung Tylototriton [News about the Genus Tylototriton] elaphe 4(2): 79-80.
Fleck, J. (1999). Weitere Beobachtungen an den Gattungen Echinotriton/Tylototriton [Further observations of the genera Echinotriton/Tylototriton] elaphe 7(2): 66-67.
Kuzmin, Sergius L. Amphibia Web Data Sheet (http://elib.cs.berkeley.edu/aw).
Nussbaum, R. A., Brodie Jr., E. D., Datong Y. (1995). A taxonomic review of Tylototriton verrucosusAnderson (Amphibia: Caudata: Salamandridae). Herpetologica 51(3): 257-268.
Rehberg, F. (1986) Haltung und Zucht des Krokodilmolches Tylototriton verrucosus [Husbandry and Breeding of Tylototriton verrucosus] Anderson, 1871. Herpetofauna, Weinstadt 8(45): 11-17.
Sparreboom, M. (1999). Haltung und Nachzucht von Tylototriton verrucosus [Husbandry and Breeding ofTylototriton verrucosus] elaphe 7(2): 20-24.
Wallays, Henk (1999/2000/2001). Personal communication.
Zhao, E. and Adler, K. (1993). Herpetology of China. SSAR, Oxford (Ohio).
Juveniles, "light form".
Juvenile, "dark form".
Red and Mud Salamanders :
The mud salamander or mountain triton (Pseudotriton montanus) is a salamander in the family Plethodontidae. It is a red salamander with black spots that inhabitsswamps, bogs, and streams. It is often confused with the red salamander, but the mud salamander is distinguished by its dark eyes and short snout. It is indigenous to the eastern United States, but is currently an endangered species. Some states have programs to locate mud salamanders to try to keep the species alive.
The taxonomy of Pseudotriton montanus and its closest relatives is unsettled. Severalsubspecies have been described, and some of these might warrant full species rank. In particular, Pseudotriton diastictus maybe treated as a species or a subspecies (Pseudotriton montanus diastictus). The rest of this article is following the latter position.
The mud salamander is known for its reddish-brown color, brown eyes, stocky girth, and short tail. It also has between 30 and 40 distinct round black spots on its back by
the time it reaches adulthood. Younger mud salamanders are typically colored bright red, orangish-brown, or crimson, with unmarked stomachs and separated spots, while older mud salamanders’ colors darken with age and take on more of a brown coloring. They also acquire more spots, larger spots, and spotted stomachs. They can reach lengths of 3-8 in (7.6 to 20 cm) in adulthood, and are typically stocky. Red salamanders and mud salamanders are very difficult to distinguish from each other. The main differences are in their eyes and snouts. While the mud salamander has dark brown eyes and a short snout, the red salamander has bright yellow eyes and a long snout.
Mud salamanders breed during the warmer months of the year. Egg deposition commonly occurs during autumn and winter. The females reproduce at most once per year (usually once per two years), while males may breed several times a year. “When a mate is found, the male performs a tail undulation display. The female then straddles his tail, allowing glands on the male's tail to lubricate her. The male is then able to deposit his sperm into the female”. Females reach reproductive maturity around four to five years old, while males reach reproductive maturity around two to two and a half years old. “A female may stay with her eggs to aid the incubation process. Incubation typically lasts three or more months, with embryos hatching in the winter. Clutches range in size between 65 and 200.”
The mud salamander inhabits swamps in low elevations, bogs, seeps, springs, and streams that not only provide a muddy bottom, but also clean and clear water. The mud salamander, a burrowing species, seeks shelter in burrows beneath leaf litter, logs, stones, or bark. The mud salamander may also build tunnels in creek banks, as well. These amphibians spend most of their lives in close proximity to water, but also burrow into the soil of the surrounding area. Larvae are usually underground in muddy springs; they are often found in leaf litter, debris, and muck of muddy springs, seeps, and streams. After they lose their gills and become adults, they make burrows in muddy areas. They often use burrows of crayfish and will sit with their heads sticking out of these burrows waiting for prey to pass by. They come out of these burrows at night and forage in the surrounding area. Generally, mud salamanders do not wander as far from their main habitat as their close relatives, the red salamanders. Mud salamanders seem to favor small, muddy seeps and springs that dry up in the summer. Both larval and adult mud salamanders go deep underground during the hottest months of the summer, especially in the small springs and seeps that dry up. The gilled larvae go deep in the mud where the water is underground and the adults remain deep in burrows. During the hottest times of the year, they are usually only found at night or during rains foraging for a short time before they return to their burrows in the mud. Dusky salamanders are often found in the same habitat as mud salamanders and are much easier to find than the mud salamanders. When this is the case, the more abundant dusky salamanders often serve as a food source for the mud salamanders. There are many scenarios where small muddy springs where mud salamanders live feed into larger streams that have more common species such as dusky and two-lined salamanders. In this case, the muds venture into the main stream and can often be found in it because they are looking for more food outside of their smaller more primary habitat. Some reasons for this is less competition outside of a smaller habitat full of mud salamanders, another reason is the abundant two lined and dusky salamanders that are food for the larger muds. They can often be found in creeks that do not seem like ideal habitat for them because they have ventured out of their primary habitat for food. Chances are an ideal muddy habitat is within walking distance from where the mud salamander was found in the stream.
A mud salamander's diet varies with age. In the larval stage, the small creatures tend to feed on equal-sized or smaller, aquatic invertebrates. The salamander larvae are also said to consume other salamander larvae. As an adult, though, the salamander's diet increases in variety, but it still eats smaller prey. Though not much is known about an adult salamander’s eating habits, it is known that they are likely to feed on earthworms, beetles, spiders, and even smaller kinds of salamanders. Mud salamanders also can eat insects as small as mites. What the mud salamander tends to eat however, mainly lies in the habitat in which it lives.
Conservation status :
Because of the mud salamander's extreme rarity in Virginia, it was put on the threatened species list in 1979. The VirginiaHerpetological Society regards this species to be secure globally, but in danger in Virginia because of its extreme rarity there. Many surveys and searches were run in the 1980s to locate the populations of the mud salamander in western Virginia. Although efforts were great, few sightings of this species were made. Because little information about the species is known, it is difficult to find possible threats, but threats to other types of salamanders probably affect mud salamanders. Tristan Clark has found multiple populations in East Tennessee in Sullivan and Hawkins county. Because this species is so hard to find, it is important to be treated as an endangered species in any state because enough specimens cannot be found to have an idea of their true abundance. (Clark, 2014) UPDATE several located in Macon County North Carolina (12/15/2015) .
Salamandra Species & Subspecies Guide
courtesy to : www.caudata.org By William Jones
The genus Salamandra consists of 6 species, 4 of which have defined subspecies. Additional information about these salamanders and their care in captivity can be found in the Salamandra caresheet. This guide has been created to show the wide range of appearance and geographical distribution among members of the genus. This document should not be used as an identification guide, because many subspecies and species have high levels of variation between populations and even among different animals within the same population (Salamandra salamandra bernardezi is a textbook example of a subspecies with high variability among individual animals). The range maps give a rough indication of where each species/subspecies lives, but should not be used as an exact guide. Some subspecies have links to photos of "other forms", but many additional subspecies have other forms not shown. If there are any errors, please contact the author using the Contact page for this website.
1- Salamandra salamandra salamandra
Balkan penninsula, the Carpathians, eastern Germany, northern Italy, and southeastern France.
- S. s. alfredschmidti
Spain: Tendi Valley.
- S. s. almanzoris
(Müller and Hellmich, 1935)
Range : Former glaciers of Gredos. (Laguna Grande de Gredos, Sierra de Gredos). The salamanders of the Penalara lagoon in Madrid show similar charactaristics.
- S. s. bejarae
(Mertens and Müller, 1940)
Range : Central mountain ranges of the Spanish peninsula, (except in the highest points of the Sierra de Gredos, and in the mountain ranges of Toledo).
-S. s. bernardezi
Other forms: Lined,yellow
Range : Asturias, northern and eastern Galicia.
-S. s. beschkovi
Range : Pirin mountains, Bulgaria.
- S. s. crespoi
Range : Portugese Algarve.
-S. s. fastuosa
S. s. "bonnali". Now considered to be synonymous with S. s. fastuosa.
Range : Cantabria, the Basque region, northern Navarre, western and central Pyrenees.
-S. s. gallaica
Variant coloration:Coimbra Portugal.
- S. s. gigliolii
(Eiselt and Lanza, 1956)
Range : South central and southern Italy.
S. s. hispanica
(Mertens and Muller, 1940)
The validity of this subspecies is questionable. May be synonymous with S. s. terrestris.
Range : Montseny Province of Barcelona.
- S. s. longirostris
(Joger and Steinfartz, 1994)
Range : Spain: Malaga and Cadiz provinces (Sierra de Ronda).
- S. s. morenica
(Joger and Steinfartz, 1994)
Range : Spain: Sierra Morenica.
- S. s. terrestris
Other forms:melanistic, erythristic,albino, high-yellow,orange.
Range : Most of France, northern and central Europe.
- S. s. werneri
(Sochurek and Gayda, 1941)
Range : Greece: Mount Pelion.
2- Salamandra algira algira
- Salamandra algira algira
Range : Morocco and Algeria.
- S. algira tingitana
(Donaire Barroso and Bogaerts, 2003)
Morocco.Specimens from Ceuta produce juviparous young, and melanistic specimens occur frequently. Examples from the Middle Atlas mountains produce larvae.
3- Salamandra corsica
Range : Corsica.
4- Salamandra infraimmaculata infraimmaculata
-Salamandra infraimmaculata infraimmaculata
(Joger and Steinfartz, 1995)
Southern Israel to southern Turkey.
- S. i. orientalis
(Joger and Steinfartz, 1995)
Range : Southern,central and eastern Turkey.
5- Salamandra lanzai
(Nascetti, Andreone, Capula, and Bullini, 1988)
Range : Italian western Alps.
- S. i. semenovi
(Joger and Steinfartz, 1995)
Range : Kurdistan and Zagros Mountains.
6- Salamandra atra atra
range : Alps and Dinaric Alps.
- S. atra aurorae
range : Asiago plateau, Italian Pre-Dolomites.
- S. atra pasubiensis
range : Mt. Pasubio and surrounding areas in the Venitian Alps of Italy
Special thanks to Mike East for providing photographs and for adding and correcting the information in this guide.
References and Additional Information
- AG Urodela, Registry for Genus Salamandra.
- American Museum & Natural History: Amphibian species of the world 3.0, An online reference.
- AmphibiaWeb Digital Library Project: University of California, Berkeley. Species accounts for Salamandridae.
- Bonato, L and Steinfartz, S. Evolution of the melanistic colour in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian Prealps. Italian Journal of Zoology, 72: 253 - 260 (2005)
-Clare. 2002. Caudata Culture Species Database Entry for Salamandra.http://www.caudata.org/cc/species/Salamandra/Salamandra_sp.shtml
- García-París, Alcombendas, Buckley and Wake. 2003. Dispersal of viviparity across contact zones in Iberian populations of fire salamanders (salamandra) inferred from discordance of genetic and morphological traits. Evolution 57:129.
- Mazzei. Amphibians and Reptiles of Europe.
- Schantz. Fire Salamander Website - Gallery.
- Schultschik. Current Systematics of the genus Salamandra.http://www.salamanderland.at/eigene.Schriften/Systematik.Salamandra.htm
Observations on Torrent Salamanders (Rhyacotriton) in Oregon and California
courtesy to : www.caudata.org By Henk Wallays
The salamanders of the genus Rhyacotriton have had quite a bumpy taxonomical journey. At their initial discovery, they were first included in the genus Hynobius (Gaige, 1917). Later, Dunn (1920) placed them into the ‘advanced’ salamanders, and they were thought to be a member of the Ambystomatidae (as was the case with Dicamptodon). In 1958, Tihen (1958) placed them into a separate subfamily, Rhyacotritoninae, and Regal (1966) placed them in a separate subfamily ofDicamptodontinae. They were later recognized as a true family by Edwards (1967). At that time only one species was known, Rhyacotriton olympicus. In 1992 Good and Wake reviewed this genus and discovered that it consisted of 4 distinct species.
Evolution, Taxonomy, and Range:
According to Good & Wake (1992), the original distribution of the genus started in the Northern Cascades in Central Oregon and later dispersed into the Coastal ranges, some 15 to 22 million years ago. This continuous range was disrupted in 3 main segments due to volcanic activity during the Miocene age. Later, the most Northern segment got further divided into 2 isolated areas by a massive river, created by the glacial expansion into the Puget Sound region. This resulted in 4 genetically isolated areas, each having a Rhyacotriton species.
Rhyacotriton variegatus.This species is the torrent salamander with the most Southern distribution and the longest area of distribution. It is found from mid-California up along Oregon to Lincoln City where its range finally meets with that of Rhyacotriton kezeri. Recently, there are some opinions expressed that this species needs revision and might turn out to be more than one species.
Rhyacotriton kezeri. The distribution of R. kezeri extends from around the Lincoln City area into Washington.
R. variegatus, Cape Ferello, Oregon.
Rhyacotriton cascadae. This species lives around the Oregon Columbia River Gorge into the Cascade Mountains. Due to its dispersed distribution, this is by far the most variable and least studied species, and it may turn out to be a conglomerate of several species.
Rhyacotriton olympicus.The distribution of this species is restricted to the Olympic Peninsula of Washington. Its distribution area has no contact zone with other Rhyacotriton species. This is the sole species that I have not (yet) been able to see in nature. The demarcation line between the dorsal and ventral coloration is wavy. The back and sides do not carry spots. The belly usually has large dark spots or mottling.
Personal observations :
In 1998 and 2001, I traveled to Oregon and California with Russell Kurtz. During these trips, 3 Rhyacotriton species were observed in their natural habitat. We also visited the contact zone between R. variegatus and R. kezeri at Lincoln City, Oregon (not mentioned in this article). These field trips were performed during springtime, at cooler temperatures, while some habitats were still snow-covered.
Humboldt County, California: Rhyacotriton variegatus :
With the aid of Brad Norman, we were able to enter a study area of the University of California in Humboldt County, where ecological studies have been performed on both Plethodon elongates and Ascaphus truei. The general idea was to find tailed frogs in order to photograph these on slides (I had failed to find adults before). The whole scenery lay beside a large river. We focused our search around a small stream coming down from the slopes in the woods, finally emptying into that river. The first 15 meters of this stream resulted into some larval Dicamptodon tenebrosus that were hiding among stones and fallen leaves. They measured up to 9 cm in length and are known to be serious predators. We went up a little more following the stream along the emerald green moss carpets. After 20 more meters of stone-turning, we stumbled upon our first Rhyacotriton variegatus. The air temperature was 9°C, the water was a bit colder, 8-8.5°C. Unlike the Dicamptodonlarvae, the animals were found in the calmer side-pockets of the stream hiding between some of the fallen leaves or underneath a stone with overhanging moss. All of them were uniformly dark colored, with a belly that carried much green & yellow and with some black spots scattered around. The cloaca of the adult male was very swollen, proving that we were there at breeding season. Finally, after a chase of about 10 minutes, I was able to capture an Ascaphus truei tadpole, but this was again as close as I got to these frogs. Apparently they avoid European tourists.
R. variegatus, Humbolt Co., California
R. variegatus habitat, Humbolt Co., California.
R. variegatus, Humbolt Co., California
Cape Ferello and Humbug Mountain trail, Oregon: Rhyacotriton variegatus
As we went up to Northern Oregon, we intended to look for more Rhyacotriton populations. With this idea in mind, we stopped both at Cape Ferrello, Oregon and Humbug Mountain. On a northern exposed slope, under the canopy of a primary coniferous wood, we found a small stream searching its way down. The sides were filled with large moss-covered stones and fallen logs of redwood trees. We knew that we were again in Rhyacotriton land and started searching again along the sides of the stream. Hiding in these side pockets, we found some adult Rhyacotriton variegatus. Again, the larger larval Dicamptodon tenebrosus (>10 cm) were lying under stones hidden in the stream. They would certainly not refuse a healthy Rhyacotriton menu. The coloration of the adult Rhyacotriton differed considerably from the Californian animals: they had a less dark color.
R. variegatus, Cape Ferello, Oregon.
R. variegatus, Humbug Mt., Oregon.
Beside this small stream, we noticed a wet heap of small pebbles. A small seepage came through and underneath these pebbles, flowing downwards to feed the stream in which we found the adults. On top of the pebbles lay fallen fern leaves and pieces of wood. While removing some of this debris, the richness of the invertebrates living on these particles was surprising. Quite a large number of crustacean-like invertebrates (resembling Gammarus) jumped away upon removal of the debris. The pebbles were quite small (2-4 cm), and the holes were miniscule. Replacing and moving some of the pebbles frequently revealed larval Rhyacotriton variegatus wriggling their way out of the disturbed area and trying to dig underneath the pebbles further away. The black dots on their backs made them difficult to spot among the stones once they stopped moving. We could only find them while they were moving or by the reflection of their shiny wet backs. The larvae were practically the same size as the pebbles. From a logical point of view, it looks as if these larvae fled the stream in order to be protected from possible predators. They are able to hide well within the small holes against threatening intrusion and at the same time still benefit from the cold spring water. In about 15 minutes, we were able to capture 7 specimens, with many more larvae and juveniles escaping, wriggling through the pebbles. After taking some shots and measurements, we set them back in their habitat unharmed.
So it looks as if the larger Rhyacotriton more often live along the edges of the stream, whereas the larvae prefer to stay out of the flowing water, for example in a seepage.
R. variegatus habitat.
North Tillamook, Southern Clatsop County, Oregon: Rhyacotriton kezeri
In order to find Rhyacotriton kezeri we went up to the Northern region of Oregon where we encountered difficulties in locating a suitable habitat. After quite some searching we discovered an interesting stream, but since the speed of the water was quite quick we were not too confident. Instead of searching along the edges, we first focused on the little standing side pockets of the stream. These calm water areas were still sparsely filled with fresh and cold water. Slow removal of overhanging stones finally revealed some Rhyacotriton larvae. After capturing about 5 larvae, we finally started to look around the borders of the stream. Two adult specimens were separately caught, a male and a female. Images of the animals and habitat were taken, after which the animals were released.
Mount Hood National Forest: Rhyacotriton cascadae :
This species proved to be the most difficult one to find. We had been searching at various spots both along the Columbia River Gorge and up on top of the valley in primary woods: none of them revealed specimens. Since most of the spots in the valley were situated in areas of deciduous woods, this didn’t surprise us. But when even the cooler streams on top of the valley in primary coniferous woods did not show these animals, we were loosing confidence. We continued to drive around the neighbourhood up and down the roads in and out of forested areas. Some parts had been intensively logged, and certain places that looked promising lay half-open in the sparse spring sunlight.
Finally we went into Mount Hood National Forest. The road kept going up, and soon we came across the ‘snow-frontier’: little white patches were still lying beside the road and in the woods. In the first 100 meters of this cold front, we noticed a stream coming down, going underneath the road. The current of the water was quite fast, so we started looking in the promising side-pockets of standing water and some slowly seeping water areas. At times we were lying down in the snow. Russell firstly discovered a larval Dicamptodon tenebrosus again. Since we had always found both species together, this strengthened our belief that we had discovered a proper spot. I had been checking up on what looked like an interesting side-pocket and noticed some movement when replacing an overhanging stone: a Rhyacotriton larva fled from the light. Finally we had found a habitat of Rhyacotriton cascadae. In that same side pocket I turned up 4 larval specimens of various lengths (3 to 5 cm). In the next half hour, we were able to capture about 10 of them. While I was shooting slides of these younger specimens in their habitat, Russell worked his way deeper into the stream, unrolling stones into the side-pockets and started finding adults. These were not found in the calmer water parts, but were more hiding alongside the edges of the streams under moss or stones. Their coloration was strikingly beautiful: the belly was clear orange, with a little bit of white, but no black spots. Most of all, though, their very reddish coloration and the large round black dots made out much of their beauty. Since I had broken my thermometer, we could not measure the water temperatures, but this was surely one of the coldest regions we had visited.
Torrent salamanders require such a specific habitat that they can hardly be called abundant, but when the right spots are present, they sometimes seem to be there in large numbers. The majority of the Rhyacotriton habitats we visited consisted of small streams or seepages in shadowed areas of primary coniferous woods with a bottom of small pebbles/gravel. The adult specimens were mostly found along the edges of the streams, whereas the larvae and juveniles prefer seepages with gravel/pebbles or remain in calmer side pockets of the streams that are still sparsely fed with fresh cold water. In all the places the temperatures were very low. We did not find animals in water temperatures higher then 11°C (mostly in the range of 5°-9°C), so they like it cold. For this reason, it is also best not to handle them too long. After a while they kind of remain passive (heating up?); setting them back in the water slowly revives them.
We didn’t find animals in streams with large rocks and stones, where the gaps between the stones surpassed 3 to 5 cms. The size of the stones ranged from less then 1 centimeter to a few centimeters. These tiny holes are at the same time a good protection against larger predators and allow them to dig in deeper when temperatures increase, for example during the warmer summer months. So far, very few nests have been discovered for any torrent salamander species, but they seem to deposit deeper in such stones (Nussbaum, 1969). Requiring such a specific habitat, these species are indeed quite fragile and together with Ascaphus truei (the tailed frog) they are considered ‘keystone’ species. They benefit from a protected status.
As for natural enemies, they are certainly preyed upon by the larger Dicamptodon tenebrosus larvae living in the same habitat. Their largest threat however consists of something completely different: logging. A primary coniferous wood functions as a large umbrella. The thick canopy prevents sunlight from heating the ground, and the ground moisture also buffers against higher temperatures. In fact, when entering such a wood with a short-sleeved T-shirt, it feels like you are entering a refrigerator; temperatures are lower and the air humidity is very high. For example, during our trips, the air temperatures outside the wood ranged from 12-17°C. Inside the woods we measured temperatures ranging from 7-9°C.
Taking away trees directly around a habitat or even on top of a habitat can drive a Rhyacotriton population to extinction. Erosion sets off in the stream, with silt filling the necessary holes between the pebbles/gravel. This means no more hiding places against predators, and exposure to warmer temperatures.
The larvae are most fragile, being by definition restricted to the water. If such a ‘logging disaster’ happens, their long larval life becomes an important adverse limiting factor. Since these animals are tied to the water, they can only go up higher or descend the stream. While the latter choice would probably be the easiest one, it is likely to end in a trout-inhabited stream. The adults, on the other hand, would need to find another typical Rhyacotriton habitat again, before they warm up or dry out. They can perhaps get away from the stream, but do require cold and very humid conditions while on the move.