Paramesotriton (Chang, 1935) Warty
Newts
Paramesotriton caudopunctatus
(Liu & Hu, 1973) Spot Tailed Warty Newt
Paramesotriton chinensis (Gray, 1859)
Chinese Warty Newt
Paramesotriton deloustali (Bourret,
1934) Tam Dao Warty Newt
Paramesotriton fuzhongensis (Wen,
1989) Fuzhong Warty Newt
Paramesotriton guangxiensis
(Huang, Tang, and Tang, 1983) Guangxi Warty Newt
Paramesotriton hongkongensis
(Meyers & Leviton, 1962) Hong Kong Warty Newt
Paramesotriton laoensis (Stuart &
Papenfuss, 2002) Laos Warty Newt
 Newts of the genus Paramesotriton are collectively referred
to as Warty Newts, a reflection of their tuberculate skin. Warty
newts are a robust group, all possessing thick, powerful limbs and tail,
triangular shaped heads, bright colored bellies, and characteristic
warty skin. Paramesotriton are closely related to the genera Cynops
and Pachytriton (Zhao, 1989), for which many similar characteristics are
shared, including breeding behavior, secondary characteristics (sexual
dimorphism), and habitat. Newts of the genus Paramesotriton are found in China, Vietnam, and Laos.
The natural habitat is generally mountain streams and brooks, with
the exception of P. hongkongensis, which prefers nearly
still waters. Warty newts are naturally territorial, especially
males, although females may also show aggression during breeding
times. A study of wild P. deloustali showed that newts were
separated by at least one meter of territory. A similar situation
likely occurs among the other species in the wild, as well.
However aggressive, Paramesotriton newts congregate or at
least seek out mates during the breeding season. During breeding
times aggression is at its peak as males compete for females, and
females compete for optimal nesting sites.
There is some controversy as to the precise classification of
warty newts, and the defining line between the species is
sometimes rather hazy. In general, there are seven proposed species, P. caudopunctatus, P. chinensis, P.
deloustali, P. fuzhongensis, P. guangxiensis, P.
hongkongensis, and P. laoensis. This set is widely
accepted, however, some would argue the validity of this arrangement. P. hongkongensis,
P. guangxiensis, and P. fuzhongensis are sometimes treated as synonyms
or subspecies of P. chinensis, despite some notable physical
differences. Also, many do not adhere to the validity of P. fuzhongensis,
and instead consider it a synonym of P. guangxiensis.
To further complicate matters, P. guanxiensis is
often treated as a synonym of P. deloustali, a theory that sends P.
fuzhongensis further into obscurity. Some have argued that Paramesotriton populations in northern Vietnam and southern Guangxi
Province, China, consist of P. guangxiensis, while the rare P.
deloustali are confined only to the Tam Dao region of Vietnam. On the other
hand, many propose that P. deloustali are found outside the Tam Dao
region, and even extend as far as southern China. Genetic differentiation may be
the only method capable of truly isolating their exact
ranges. P. laoensis, found in Laos, has been described
most recently, and is accompanied by limited information. This species is
markedly different in appearance from the others, possessing bright coloration
about the dorsum, and as far as is known, P. laoensis is allopatric with
all other species. Currently, the status of P. laoensis at the species
level is mostly accepted, however, this species often goes unrecognized in
phylogenetic studies.
Recent molecular data (Lu, et al., 2004) have shown that P. fuzhongensis
is indeed a legitimate species, distinct from P. guangxiensis. Further
studies based on morphological and molecular studies are needed to properly
define or differentiate P. deloustali and P. guangxiensis. P. caudopunctatus is
thought to be the most basal Paramesotriton species, a status
eluded to by its notable external and morphological characteristics. Lu, et al.
(2004) show that the Vietnamese and Chinese species form a monophyletic group.
Unfortunately, P. laoensis was not included in this study, however, this
species' status and phylogenetic relationship are addressed in Bryant &
Papenfuss (2002).
Paramesotriton caudopunctatus (Liu & Hu,
In Hu, Zhao, and Liu, 1973) Spot Tailed Warty Newt
Taxonomic Synonyms: Trituroides caudopunctatus (Liu & Hu, In
Hu, Zhao, and Liu, 1973), Paramesotriton caudopunctatus (Bischoff &
Böhme, 1980), Paramesotriton caudomaculatus (Seidel, 1981), Allomesotriton
caudopunctatus (Freytag, 1983) Paramesotriton (Allomesotriton)
caudopunctatus (Pang, Jiang, and Hu, 1992)
Vernacular Names: Spot Tailed Warty Newt, Spotted Tail Warty Newt,
Spotted Tail Warty Newt, Guizhou Warty Newt
 Paramesotriton caudopunctatus is a rather large species, reaching lengths up to
7 inches from snout to tail end. The overall coloration varies from light to dark brown,
and some are known to take on a blue or green hue. A red or orange dorsal
stripe is often present into adulthood. Unlike the dorsal stripes of P.
chinensis, those of P. caudopunctatus typically have a distinct black
coloration on both sides, that appears almost like a shadow. Occasionally, the dorsal surface from the back of the
head to the tail is black or dark
colored, while the remainder of the body and tail are brown. The back is
rather straight, with two distinctly flat areas on either side of the dorsal
ridge, marked at the outer edges by warty ridges, which give
the body an elongated-cube shape. The warty ridges
extend from the back of the head, along the
sides of the body, to the beginning of the tail, but may be reduced in smoother
individuals. P. caudopunctatus are almost fully aquatic as adults and juveniles, and
often possess
rather smooth skin compared to P. fuzhongensis, P. guangxiensis,
and P. chinensis. P. caudopunctatus
have conspicuously long snouts that one might describe as "dog-like" in
appearance when compared to the other species. This appearance is not
apparent in the other species of this genus, and results from a long, narrow skull (Sparreboom,
1981); a primitive characteristic of Paramesotriton. The digits are rather short and stubby on all four limbs. The belly pattern typically consists
of orangish-yellow, or pale peach lines and dots, as opposed to the large,
reddish-orange or yellow colored blotches and specs of the other species. Some may have greenish-gray colored bellies
and throats, with no hint of bright coloration at all, while others may have nearly solid
colored bellies of bright or pale orange. Males possess distinct spots
along the lateral side of the tail, concentrated at the distal part, at all times of the
year. The spots vary in size and color, but
there is usually an elongated spot at the very end of the rounded tail tip. The
tail spots are whitish-cream, pale peach, or orange colored, and clearly outlined in
solid or disconnected black markings. Tail spots usually become more distinct
during the breeding season. Female P. caudopunctatus occasionally possess similar spotting about the tail, which is usually reduced in comparison to a
typical male. In addition to the longer snout, the
tail spotting is unique to P.
caudopunctatus, and positively distinguishes this species from any other
in the genus.
Males can be distinguished from females by their pronounced tail
spots, and wider head. Females also possess longer tails that usually
lack spotting.
Juveniles look very similar to adults, with defined orange-red dorsal
stripes, and light brown coloration. The long snout is also
distinguishable in juveniles, even in recently morphed individuals. In
captivity, juveniles can be raised in a semi-aquatic environment
identical to the adults, however, it is unknown if a terrestrial phase
is experienced in the wild.
P. caudopunctatus are found in the Guizhou (Kweichow) Province,
and the Guangxi Zhuang Autonomous Region, China. Territory overlaps with P.
fuzhongensis in the northern Xilin Hill area, Fuchuan Xian,
northeastern Guangxi. Males are territorial, and display aggression toward other males in their
vicinity, especially during breeding times. Females have a
tendency to guard eggs, and so may become more aggressive at this time, as
well.
Paramesotriton chinensis (Gray, 1859) Chinese Warty Newt
Taxonomic Synonyms: Cynops chinensis (Gray, 1859), Triton
chinensis (Strauch, 1870), Triton sinesis (Sauvage, 1876), Molge sinensis (Boulenger,
1882), Diemictylus sinensis (Stejneger, 1907), Triturus sinensis (Dunn,
1918), Triton chinensis (Wolterstorff, 1925), Cynops chinensis (Wolterstroff
& Herre, 1935), Trituroides chinensis (Chang, 1935), Triturus sinensis
boringi (Herre, 1939), Triturus chinensis (Pope & Boring, 1940),
Paramesotriton chinensis (Freytag, 1962), Trituroides hongkongensis (Myers
& Leviton, 1962), Trituroides chinensis (Anonymous, 1977),
Paramesotriton chinensis chinensis (Fei, Ye, and Huang, 1991),
Paramesotriton chinensis hongkongensis (Fei, Ye, and Huang, 1991)
Vernacular Names: Chinese Warty Newt, Chinese Newt
 Paramesotriton chinensis is perhaps the most varied in terms of coloration
and pattern. This species can be light gray to dark brown, and occasionally olive colored, with or without
orange or yellow dorsal stripes. The dorsal surface is often mottled with black, olive, or brown, and may have small yellow
or orange dots irregularly scattered about. Some may possess solid
colored dorsa, without dorsal stripes or bright colored markings. The
ventral pattern varies from irregular blotches, to small dots of orange or
yellow surrounded by contrasting black or dark gray coloration, which
extends to the throat and lower jaw area. Breeding males are easy to
differentiate due to the development of a bluish-white lateral sheen along
the tail. Males also possess shorter tails than females, and are often
slimmer, with less robust extremities.
Larvae are uniform black, including
the gills and feet. New morphs and juveniles are black or dark brown initially,
often with light spots on the bases of the limbs. Most individuals will
lighten up with age, becoming greenish-olive, gray, or brown dorsally.
P. chinensis are found in eastern China, including the provinces of Anhui
(Anhwei), Zhejiang (Chekiang), Hunan, Fujian, Guangdong, and Guangxi.
P. chinensis can be rather territorial and aggressive at times, and require a
spacious captive environment. In captivity, aggressive behavior may or may not be
present, depending on the individual, the season, and the sex ratio of the
inhabitants. Males and females
appear to develop increasingly aggressive behavior just before, and during the breeding
season. Fights are vicious, and can result in permanent wounds and
disfigurements. P. hongkongensis are often
treated as a subspecies or synonym of P. chinensis, although both
species display distinct physical characteristics and habitat preferences.
The tail fins of P. hongkongensis are higher than those of P. chinensis, often with
small waves on the upper side. P. hongkongensis also possess prominent
cranial ridges, and warty dorsolateral ridges, both of which are absent in
P. chinensis. The belly pattern of P. hongkongensis usually consists of smaller,
fleck-like orange or yellow markings, compared to the larger blotches of P.
chinensis. P. chinensis larvae are also markedly different in
appearance than P. hongkongensis larvae. P. chinensis
larvae are uniform black in coloration, including
the gills, whereas P. hongkongenis larvae possess cream colored
gills, snouts, and toes.
P. chinensis. Lightly mottled, tan
individual. Photo © Jessica J. Miller, jess@livingunderworld.org
|
P. chinensis. Moderately mottled, tan
individual. Photo © Jessica J. Miller, jess@livingunderworld.org
|
P. chinensis. Lightly mottled, tan
individual. Photo © Jessica J. Miller, jess@livingunderworld.org
|
P. chinensis. Two
variations of dorsal and ventral coloration. Photo © Jessica J.
Miller, jess@livingunderworld.org
|
P. chinensis. Light orange ventral
flecks, darker orange tail stripe. Photo © Jessica J. Miller, jess@livingunderworld.org
|
P. chinensis. Light orange ventral
flecks, darker orange tail stripe. Photo © Jessica J.
Miller, jess@livingunderworld.org
|
Paramesotriton deloustali (Bourret,
1934) Tam Dao Warty Newt
Taxonomic Synonyms: Mesotriton deloustali (Bourret, 1934),
Pachytriton deloustali (Chang, 1935), Paramesotriton deloustali (Chang,
1935)
Vernacular Names: Tam Dao Warty Newt, Tam Dao Newt, Vietnam Warty
Newt, Vietnamese Salamander
 Paramesotriton deloustali,
characterized by an
unusually large head, is the largest species of the genus. P. deloustali tend to have stunning undersides
comprised of a mosaic of carmine red or burnt orange blotches. In most
cases, the belly is highly red or orange with only a small portion of brown
or black weaved between the bright blotches. The tail is
also rather impressive, being long and thick, with a high tail fin. At a
glance, P. deloustali resembles P.
hongkongensis in general appearance, but again, the large head
and body size, and brilliantly patterned belly of P. deloustali are
unique.
Male P. deloustali can be differentiated from females by their
smaller size, longer limbs and head, and shorter tail. Also apparent in
breeding males is a bluish sheen about the lateral side of the tail, a
characteristic also observed in other Asian species.
P. deloustali are
one of the rarest amphibian species, and have been seldom bred in captivity.
P. deloustali is listed in the Red Data Book of Vietnam
as Endangered (2000), as Vulnerable in the IUCN Red List (2000), and is
found in
Appendix II of the
UNEP
- WCMP World Conservation Monitoring Program. P. deloustali is
also protected
in Vietnam by Government Decree No. 40/2002/ND-CP (2002), which prohibits any
exploitation. However, despite P. deloustali's government and international
protection, individuals inevitably show up in the
pet trade, especially in local shops in Vietnam and surrounding countries.
P. deloustali are found in the Tam Dao mountain range
(Vinh Phuc, Thai Nguyen, and Tuyen Quang Provinces), Xin Man (Ha Giang
Province), Van Ban (Lao Cai Province), Ba Be District (Bac Kan Province),
and Cho Don District (Bac Kan Province). P. deloustali is also said
to occur in the Lang Son Province.
Populations in the Ba Be, Cho Don, and Bach Thong Districts, Bac Kan
Province, are concentrated in 5 communes; Dong Phuc, Quang Khe, Bang
Phuc, Tan Lap, and Quang Bach, in and around Ba Be National Park. Field
studies show that in these districts, P. deloustali is found in
shallow, slow-moving streams with sand and gravel bottoms. P. deloustali
males are territorial, and may claim areas of a meter of more in the wild. Although most do
not regard P. deloustali as economically important, some locals from
the fore mentioned areas occasionally use a dried, or powdered form of P.
deloustali for medicinal purposes.
Until recently, the mountainous regions of Phja Bjooc and Tam Dao,
Vinh Phuc, Thai Nguyen, and Tuyen Quang Provinces, and the Ba Be
National Park area, Bac Can (Bac Kan) Province, Vietnam, were considered wild
paradises of sorts, where escape from the thriving life of nearby Hanoi could be
reached. These mountain ranges are watershed forests of the streams flowing
into Cau River and Ba Be Lake. The climate of Tam Dao in particular is warm and humid, with ambient
temperatures averaging around 86°F during the summer, and around 50°F during the
winter months. Unfortunately, P. deloustali's serene Tam
Dao habitat has been radically changed since the late 80's and early 90's,
as a result of political and economic changes. During these times, more construction and industrialization
commenced in an effort to increase tourism, in addition to increased logging of the
surrounding forests. The increased industrialization of the mountain ranges introduced toxins and pollution into the
environment, and significantly reduced P. deloustali's
habitat. In 1997, after chemical runoff from a nearby hotel
construction site polluted the salamanders only known habitat, an
artificial water basin in the Tam Dao mountain Range, P. deloustali
were thought to be extinct in the wild. Fortunately, populations were later
discovered elsewhere. Recently, the Tam Dao
mountain range has been proclaimed a Natural Reserve, which has
halted logging of the surrounding forests, and decreased nearby
construction. However, P. deloustali's Tam Tao range still includes a nearby developing village, which could well
impact the livelihood of P. deloustali in this area.
Wild P. deloustali populations were discovered around the Na Nheo village situated in the Hoang Lien
mountains, Lao Cai Province, Vietnam, in 2002. Some have speculated that
these populations, and any Vietnamese populations found outside of Tam
Dao, are not P. deloustali, but are un-described salamanders, or
possibly unknown populations of the Chinese species P.
guangxiensis. These two species are very similar morphologically and
genetically, and more work is needed to define their actual ranges. In any event, these
populations are treated as P. deloustali for the most part, and no
studies have yet presented knowledge otherwise. Specimens collected in 2002 were all
found around rivers and streams near the afore mentioned village, at
elevations of 600 - 900 meters, and from one stream at 1000 meters. A
population of P. deloustali was also discovered near the Ban Xe
village, Na Chi commune, Xin Man District, in 2002. There are also a few
captive breeding projects, which have so far proven successful.
In
the wild, P. deloustali are thought to remain aquatic
year-round, without a hibernation or aestivation period. Adults are found
mostly in the sluggish pools of moving streams, although some may also venture into faster flowing waters.
Territorial behavior has been observed in males in the wild, as well as in
captivity. In the Tam Dao range, breeding activity has been documented during the winter season, when
temperatures quickly drop.
Paramesotriton fuzhongensis (Wen, 1989) Fuzhong Warty Newt
Taxonomic Synonyms: None
Vernacular Names: Fuzhong Warty Newt, Fuzhong Newt, Wanggao Newt
 Paramesotriton fuzhongensis often possess
variable dorsal patterns, and
considerably warty skin. The dorsal
coloration is usually a dark brownish-olive, sometimes mottled with a second, either lighter or darker, color. This
is not always the case, as some individuals may be a solid brown or olive
color. P. fuzhongensis sometimes have tiny dots of orange-red or
yellow color irregularly dispersed about the dorsum. A light orange or yellow dorsal
stripe is also apparent in some newts. The belly pattern usually consists of large, disconnected
orange or reddish blotches, which extends onto the throat and chin. During the
breeding season, male P. fuzhongensis develop the bluish-purple sheen along the tail that is common in
other Asian species. The skin also becomes smoother, and sometimes lighter colored. The eyes of P.
fuzhongensis are light colored, and the snout ends in a rather blunt
and squared manner. Males can be
differentiated from females by their shorter, and higher tails, smaller
reddish-orange spots on the throat region, and longer vent (Wen, 1989).
Larvae of P. fuzhongensis are black or dark brown, usually with
reddish colored gills. Some may also have cream colored dots about the
dorsum and abdomen. Juveniles are medium to dark brown dorsally, sometimes with
orangish-red
dots about the dorsum, and orangish-red dorsal stripes. The short, squared
snout is obvious at an early stage in development.
P. fuzhongensis is often treated as a subspecies or form
of P. guangxiensis, despite some notable physical differences. Lu,
et al. (2004) define P. fuzhongensis as a valid species based on
molecular studies, which supports Wen's (1989) original classification at
the species level. According to Wen (1989), P. fuzhongensis can
be differentiated from P. guangxiensis by their longer and narrower
heads, longer and lower tails, and the possession of black mottling or
blotching about the dorsum in many individuals, which is almost never
observed in P. guangxiensis. Wen (1989) notes that P.
fuzhongensis possess more granular skin, such that the warts are more
frequent and in higher concentrations when compared to P. guangxiensis, and that the forelimbs of P.
fuzhongensis reach as far as between the eye and nostril when drawn
forward, while those of P. guangxiensis do not reach as far. Based on X-Rays of skeletal
structure, it appears as though the forelimbs of P. guangxiensis,
when compared to P. fuzhongensis, are
positioned farther back relative to the eye position, which may be
observable in life.
P. fuzhongensis, and P. guangxiensis for that matter, are
rather understudied in captivity and in the wild, and their taxonomic
designations could possibly change in the future.
A recent trend in identifying warty newts has developed as a result of uncharacteristic species showing up
in the pet trade, where unidentifiable
species are ambiguously designated P. fuzhongensis or P. guangxiensis.
This has added to existing doubts about the taxonomic status, or even the legitimacy, of these
two species. In reality, there are probably some undescribed
species or subspecies that are simply placed in the most similar appearing
species, for lack of better knowledge.
P. fuzhongensis are found in northeastern Guangxi Province,
China. The natural habitat consists of mid-slope streams, surrounded by
broad-leaf forests (Wen, 1989). The Guangxi Zhuang Autonomous Region is
considered subtropical, and humid; typical monsoon climate. The monsoon
season occurs from April to September, during the summer months. Winter is
short, and mild. Guangxi's topography consists of mountains, hilly
landscape, and plains regions. From northwest to southeast, a gradient of
descending elevation is formed, and covers an average range of 3,000-6,000
at the highest peaks, down to 300-1,500. Average temperatures in Guangxi
range from 74°F to 85°F during the warmest month, July, and 42°F to
60°F during the coldest month, January.
P.
fuzhongensis has been described by some keepers as only slightly to
moderately aggressive, while other keepers have noted typical territorial/aggressive behavior,
especially in breeding males.
Paramesotriton guangxiensis (Huang,
Tang, & Tang, 1983) Guangxi Warty Newt
Taxonomic Synonyms: Trituroides guangxiensis (Huang, Tang,
and Tankg, 1983), Paramesotriton guangxiensis (Zhao & Hu, 1984),
Paramesotriton guangxiensis (Zhang & Wen, 2000)
Vernacular Names: Guangxi Warty Newt, Guangxi Newt
 Paramesotriton guangxiensis are robust, with large heads and thick
limbs. The dorsum is typically uniform dark brown, and the venter consists
of variations of bright blotches of reddish-orange, over a dark background. P. guangxiensis
is often lumped into the same species as P. fuzhongensis, although P.
fuzhongensis has been designated at the species level. Still some do
not adhere to this placement. Whether
or not species level taxonomic classification is accurate in the case of P.
guangxiensis, there are significant enough differences in terms
of physical appearance to differentiate this "type" from other Paramesotriton
newts,
and so remarks seem warranted. In general, P. guangxiensis can be
differentiated from P. fuzhongensis by their shorter and broader
head, shorter and taller tail, and lack of black mottling or spotting about
the dorsum (Wen, 1989). Wen (1989) notes that P. fuzhongensis possess more
granular skin, such that the warts are more frequent and in higher
concentrations, as well as longer, less laterally compressed tails. Wen
(1989) also notes that the forelimbs of P. fuzhongensis reach as far
as between the eye and nostril when drawn forward, while those of P.
guangxiensis do not reach this length. Based on X-Rays of skeletal
structure, it appears as though the forelimbs of P. guangxiensis,
when compared to P. fuzhongensis, are
positioned farther back relative to the eye position, which may be
observable in life. Males can be differentiated by females by their shorter
tails, slimmer physique, and less robust extremities.
A recent trend in identifying warty newts has developed as a result of uncharacteristic species showing up
in the pet trade, where unidentifiable
species are ambiguously designated P. fuzhongensis or P. guangxiensis.
This has added to existing doubts about the taxonomic status, or even the legitimacy, of these
two species. In reality, there are probably some undecided
species or subspecies that are simply placed in the most similar appearing
species, for lack of better knowledge.
Paramesotriton guanxiensis are found in southwestern Guangxi Province,
China. P. guangxiensis are only known from their type locality, at
Paiyang Mountain, Mingjiang, Ningming County, Nanning Prefecture, although
it is speculated that newts found in northeastern Cao Bang province,
Vietnam, are also P. guangxiensis. The
Guangxi Zhuang Autonomous Region is considered subtropical, and humid;
typical monsoon climate. The monsoon season occurs from April to September,
during the summer months. Winter is short, and mild. Guangxi's topography
consists of mountains, hilly landscape, and plains regions. From northwest
to southeast, a gradient of descending elevation is formed, and covers an
average range of 3,000-6,000 at the highest peaks, down to 300-1,500.
Average temperatures in Guangxi range from 74°F to 85°F during the
warmest month, July, and 42°F to 60°F during the coldest month, January. Some
have argued that P. guangxiensis extend into northeastern Vietnam,
in the provice of Cao Bang, where
they are mistaken for P. deloustali populations. In fact, it has
been suggested that any Vietnamese populations outside of the Tam Dao
region of Vietnam are actually P. guangxiensis, not the Vietnamese
species P. deloustali. Others have suggested that
populations of P. guangxiensis in southern China are actually P.
deloustali. To further complicate matters, some do not adhere to the
validity of the two species, and instead treat the two as synonyms.
Molecular studies by Lu, et al. (2004) showed that P. guangxiensis
and P. deloustali are indeed very close in relation, if not
synonymous, however, more
work is needed in terms of classification and distribution.
Paramesotriton hongkongensis (Meyers and Leviton, 1962) Hong Kong Warty Newt
Taxonomic Synonyms: Trituroides hongkongensis (Myers &
Leviton, 1962), Paramesotriton hongkongensis (Freytag, 1962),
Paramesotriton chinensis hongkongensis (Fei, Ye, and Huang, 1991)
Vernacular Names: Honk Kong Warty Newt, Hong Kong Newt
 Paramesotriton hongkongensis
is a robust species, with thick limbs and tail. This species
can be distinguished from the other Paramesotriton species by the
presence of prominent cranial ridges, and the least warty skin of the genus. P. hongkongensis is almost always a
solid, medium to dark brown color
without any significant mottling or bright coloration on the dorsum. The tail may fade
into a light tannish-orange, with dark specks. The belly pattern is
variable, but usually consists of small to medium sized, disconnected blotches, or
large flecks of
orange or reddish, on a dark gray or black background. The belly pattern continues up the
throat to the lower jaw. Most individuals also have the same orange or
reddish color along the bottom edge of the tail. The tails of aquatic adults are
notably compressed laterally, which is often more prominent in
males. During the breeding season,
males develop significantly smooth skin, swollen cloaca, and a distinct bluish-purple sheen along the
lateral sides of the tail. Males also possess shorter tails than females,
and a smaller, less-robust physique. P. hongkongensis usually have thick, warty ridges starting from
the back of the head, running along the sides of the body, and onto the
proximal portion of the tail.
Larvae are dark brown or black dorsally, with cream colored snouts,
gills, and toes, and sometimes with a cream colored dorsal stripe that
extends down the tail. Juveniles are initially uniform black or dark brown,
and develop lighter color with age.
P.
hongkongenis is often treated as a subspecies of P.
chinensis, however, many do not adhere to this classification based on the disjunctive
distribution of the two, and differences in physical appearance and habitat
preference. P.
hongkongensis are larger, and more robust than P. chinensis, and
possess two unique characteristics: Warty cranial ridges, and less warty skin
than any other Paramesotriton species. Furthermore, P.
hongkongensis are typically one solid color, dorsally, whereas P.
chinensis may be mottled with darker or lighter coloration. The belly
pattern is also different in that P. hongkongensis typically have
large flecks of bright coloration, and P. chinensis usually have
large, irregular blotches. The larvae of P. hongkongensis also
possess cream colored attributes, as described previously, whereas P.
chinensis larvae are uniform black. The habitat preference is also
different, with P. hongkongensis uniquely preferring a subtropical
pond type environment.
In captivity, P. hongkongensis may show extreme aggression at
times, however, some have described peaceful residence in spacious tanks.
Aggression is likely related to the breeding season, when males become more
aggressive in their competition for females. Males housed together without females may remain compatible, and
show no signs of aggression.
P. hongkongensis are found in sluggish pools within moving
streams in the mountains of Hong Kong. Although once found along the Fujian-Guangdong coast region of China, P. hongkongensis are now restricted
to Hong Kong. Some argue that P. hongkongensis was never found on
the mainland coast, and populations once described here were actually P.
chinensis. One documented habitat of P. hongkongensis is the
Tung Lung Stream, and surrounding streams, Lantau North County Park (Lantau
Island), Hong Kong. Tung Lung Stream is
situated atop the Tung Chung Valley, at altitudes of up to 800 meters above
sea level, and splits into approximately 10 waterfalls (Chan, 1999). The untamed surrounding mountains are
very steep, consisting of sheer cliffs and gorges. The streams are surrounded by dense
forests, of which grow a couple
of subtropical orchid species, Pholidota chinensis, and Coelogyne
fimbriata, indicating an environment of warm, humid climate (Chan, 1999). In general, Hong Kong is a subtropical environment, with a
monsoon season from March to May, followed by a humid summer with
temperatures as high as 95°F, and finally a cooler, drier winter. However,
the climate experiences considerable fluctuations, and provides milder
micro-climates. It can be assumed that
the temperatures are lower at higher elevations, however, the presences of
subtropical orchids indicates that at least the Tung Lung Stream habitat of
P. hongkongensis may be at least warmer, and more humid than temperate
zones. This temperature
model coincides with the warmer temperatures some have suggested for P.
hongkongensis in captivity.
Paramesotriton laoensis (Stuart & Papenfuss,
2002) Laos Warty Newt
Taxonomic Synonyms: None
Vernacular Names: Laos Warty Newt
 Paramesotriton laoensis, the most recent discovery, is found in northern Laos. P.
laoensis possess the typical warty skin, triangular shaped head, and
thick limbs and tail common to warty newts. The dorsum of P. laoensis,
however, is markedly different from any other species, consisting mostly of
tannish-yellow coloration with some intermittent black areas, and a tannish dorsal
stripe that extends from just beyond the snout to just after the cloacal
region
(Stuart & Papenfuss, 2002) . The tannish color make the newts cryptic in their habitat, which
consists of un-shaded streams. The limbs, feet, and tail are dark black,
making this newt especially striking. The belly consists of
yellowish-orange, or reddish blotches mottled with dark black coloration. The bright belly color continues up the throat, and sometimes down the
bottom side of the tail. This species possesses cranial ridges, in a similar
fashion as P.
hongkongensis, that are typically tannish-yellow colored. P. laoensis
also have markedly large
paratoid glands projecting from the back of the head. P. laoensis have a thick upper lip that overlaps
the lower, which aids salamanders in feeding in water, and indicates a
highly aquatic lifestyle. Unlike other members
of this genus, P. laoensis possess a reduced tongue pad, lacking a
free posterior region, a characteristic of Pachytrtion
(Stuart & Papenfuss, 2002). P. laoensis was placed in the genus Paramesotriton based on skull
shape, maxillary orientation, and number of trunk vertebrae (Stuart &
Papenfuss, 2002). Based on these
observations, this new species shares more morphological characteristics
with Paramesotriton than with Pachytriton, Cynops, Echinotriton,
and Tylototriton. Males and females are approximately the same size,
ranging between 6.0 and 7.4 inches.
P. laoensis were discovered in the Phoukhout District, Xiang Khouang
Province, Laos. This species is also found at the base of Phou Sang Kat
Mountain, and the Saysamboun Special Zone. P. laoensis inhabit
shallow streams with light currents. Because of their cryptic coloration,
they are seem moving about during the day, as well as at night
(Stuart & Papenfuss, 2002).
P. laoensis. Photo © Bryan L. Stuart
|
P. laoensis. Photo © Bryan L. Stuart
|
Warty Newts in Captivity
This section is meant as a general guide to the captive care of warty newts,
and is based on the experiences of keepers
and breeders of the more common species.
 In
general, Paramesotriton species, especially males, are
territorial and aggressive toward other members of the same
species or genus,
so ample territory is required when housing more than one individual in the
same tank.
Aggression is not always limited to same-sex individuals, as
attacks have been noted between male-female pairs, as well as male-male, and
female-female pairs. Territorial individuals can smell and see other newts in the tank, and will often actively
pursue others in their territory if they catch a whiff of an intruder. Territorial individuals will attack intruding newts, biting and
thrashing the limbs and tail, and sometimes dragging them under rocks or
into other hiding spots. Aggressive behavior in Paramesotriton species
is one sided, resulting in attacks, and not necessarily fights.
When one newt is caught by another, it is usually rendered incapable of
retaliation, and in some cases will consequently take to the land portion of
the tank for long periods of time. Continual attacks may lead to death, but most
often results in the victim of attack becoming stressed and seeking refuge on
land, rarely with serious wounds. Interestingly, some individuals can be kept peacefully in an adequate
sized tank, without incident, while other individuals of the same species
may choose to remain aggressive even in large tanks. Aggression in captivity
may be associated with the breeding season, when females defend breeding
sites, and males compete for females. In captivity, territorial behavior may
be influenced by sex ratio, where males are only aggressive
toward other males in the presence of females. However, same-sex aggression
has been observed in males and females while housed in isolation from the
opposite sex. Some Paramesotriton keepers may find it beneficial to
house individuals separately, only introducing isolated male-female pairs
during the breeding season. Others may keep one male and one or more females
together, but again, aggression is not always observed among members of the
same gender. P. caudopunctatus, for example, show mild parental care
toward eggs, and females have been known to attack intruding males and other
females during the breeding season. Contradicting reports among Paramesotriton keepers seem to be a
common occurrence, and as of yet there is no universal recipe to keeping and
breeding all species, consistently. For example, many would say that P.
caudopunctatus are non-aggressive, while an equal number of keepers may say they are one of the more aggressive species.
Again, such experiences may be directly correlated to sex ratio, and/or the
current season, both of which can result in variable observations in
captivity. In any event, a keeper should be able
to tell the preference of their captives by their behavior. One or more
aggressive individuals may require separate living quarters, whether they be
males, females, or both. It is also worth mentioning that at least P.
deloustali are known to occupy approximately one meter of territory in
the wild, a fact that emphasized the importance of large tanks with Paramesotriton.
I have compiled a table below of my own personal
experiences with four Paramesotriton species, which summarizes their
affinity for aggression with brief descriptions of occurrences.
| Table
1.1 - Personal experiences with four species of Paramesotriton;
affinity
for aggression with descriptions of occurrences. |
| |
Sex Ratio |
Territorial |
Observed in |
Occurrences |
| P.
caudopunctatus |
2 females
3 males |
YES |
Male - Male
Female-Female
Female-Male |
Males violently aggressive
toward other males during the breeding season in the presence of
females. Males tolerate other males during the off season. Males not aggressive toward female at
all. Females violently aggressive toward males and other females
when breeding. |
| P. chinensis |
3 females
2 males |
YES |
Female - Female
Male - Male |
Males aggressive
toward males in presences of females at all times of the year, but
especially during the breeding season. Females aggressive toward
females when isolated from males, but only when temperatures begin
to rise from winter cooling. During winter cooling period, when
temperatures average 55°F - 62°F, females isolated from males are passive. |
| P.
hongkongensis |
2 females
2 males |
YES |
Male - Male
Female - Male |
Females mildly aggressive toward males outside of breeding season, but not
aggressive toward other females. Males moderately aggressive toward
males at all times of the year, but especially during the breeding
season. |
| P. fuzhongensis |
2 females
1 males |
NO |
n/a |
Single male is
not aggressive toward females. Females are not aggressive toward
each other or the single male. Other keepers have noted extreme
aggression between breeding males. |
P. hongkongensis and P. fuzhongensis have
been bred in captivity by keeping a temperature schedule around 68°F - 74°F during the summer months, and 59°F -
65°F
during the winter months, however, the actual temperatures encountered in the wild are debatable. P. hongkongensis in particular are known to prefer warmer
temperatures, and should not be kept lower than 55°F for long periods of
time. P. chinensis and P. caudopunctatus can be
maintained at a gradient temperature schedule of 63°F - 73°F during the warmer months, and
60°F - 55°F
during the winter months without incident, however, some have suggested P. chinensis,
and P.
guangxiensis for that matter, should never be kept above 70°F
during the warmest times of the year. Temperatures lower than 45°F may result in
members of all species seeking refuge on land.
 Paramesotriton species
are often described as stream-dwellers, however, this is really only
accurate for P. caudopunctatus. P. chinensis, P. fuzhongensis, P.
guangxiensis, and P. deloustali require adequate aeration, but
not necessarily fast-flowing waters. P. hongkongensis actually prefer
an even less turbulent environment, comparable to Cynops species.
Captive preferences of P. laoensis are not fully known, but may possibly
consist of a
mild stream-type environment. Most filter outputs can be positioned to create
moderate aeration throughout the tank. Except with P. caudopunctatus,
the filter should not create a powerful current, but should be
manipulated to create the equivalent of a mild ripple. For P. caudopunctatus, the filter can be used to create a
stream environment by circulating a moderate flow of water throughout the
tank. It is recommended to create areas in the tank that are shielded from the water movement so that the inhabitants have a choice to
reside between strong and weak flow. This is more natural, as
shallow streams often have pockets of slow moving or still water, as well as
objects such as rocks, wood, or other debris, that serve as hiding spots,
and where water flow would also be reduced. Rocks, driftwood, or thick
vegetation in some areas work well to slow or block the water flow in
captivity. P.
hongkongensis may exit the water completely if the current is too powerful in
the tank. For this species, an adjustable flow filter should be used, with
the output set on low. Some have suggested keeping P. hongkongensis
similar to Cynops ensicuada, that is, in still or slow moving waters,
with warmer temperatures. Like all captives, warty newts will indicate their preferences by their behavior.
Large
tanks can be made more natural with a shallow water area that flows into
a deeper pool at one end, and that is surrounded by a natural embankment. The pool
area can include a trickling waterfall, as well, which makes a nice,
aesthetic addition, and also provides additional oxygenation.
The water level
can be 6-10 inches deep. Like all newts, Paramesotriton
sp. are sensitive to water
chemistry, and a sudden change from semi-aquatic to terrestrial behavior may
indicate a problem with the water quality. It is recommended to keep a
freshwater test kit on hand, and regularly check the pH, ammonia, nitrite,
and nitrate levels. P. hongkongensis and P. chinensis do not
seem picky about exact pH, but a safe range for all species would probably be between
6.8-7.4. As mentioned earlier, the water area can support a
filter with a carbon and
sponge insert, or a canister type biological filter for larger tanks. The
filter is used to aid in keeping the water clean of chemicals and pollutants, as
well as providing aeration. Gravel is the standard form of submersed substrate, and live aquatic plants can be
included as well. Thick vegetation should be used in moderation, and even less
so with the stream-type environment of P. caudopunctatus. As mentioned earlier, thick vegetation can be used
to create some areas in the tank that are shielded from strong water flow;
however, some open areas should also be allowed.
The exception is P. hongkongensis, which can be
housed in rather densely vegetated aquariums because of their preference for
calmer waters. Paramesotriton species seem to appreciate stacked rock
hiding places, where they can hide among the crevices. Bare bottom tanks are not
recommended with newts as large as Paramesotriton, as they can prevent
the newts from gripping the bottom. If desired, small
guppies can be added to the water, but are not suggested if breeding is to
be attempted, as they will eat the eggs and larvae. Fish also carry
parasites and disease, so there are some risks with if they're included. See
article
0020 - Captive Habitats: Ideas, Instructions, and Examples and the
Captive
& Wild Habitats section of the
Photo
Gallery.
Paramesotriton species are mostly aquatic, however, a land area
for short term refuge should still be provided. Large, mossy rocks or driftwood
protruding from the waters surface are commonly used as the land area for Paramesotriton
adults. Stacked rocks, driftwood, or other non-toxic substance should also be used to create underwater
hiding places. Newts that spend a significant amount of time out of the
water should be provided with dark hiding places on land, as well. The land area also provides a refuge for newts
fleeing from an aggressor, as well provides spots of reduced water flow. As mentioned earlier, a more natural setup would
consist of shallow water area surrounded by a natural embankment, such that
water collects in a deeper pool at one end.
Juveniles are generally terrestrial, and can be housed in a woodland-type setup,
with damp, chemical-free soil, live plants, and plenty of small hiding
places. Moist, bleach-free paper toweling, or chemical-free pillow moss can
also be used for terrestrial substrates. P. caudopunctatus juveniles are known to do well in a
semi-aquatic setup, where they will spend almost 100% of their time submerged.
As with all amphibians, tap water should be treated with a water
conditioner or primer prior to the introduction of amphibians. Never used plain
tap water, de-ionized, distilled, or reverse osmosis waters alone with amphibians.
Also, aquatic and semi-aquatic tanks should be cycled (nitrogen cycle) prior to the introduction
of amphibians. See
article
0006 - Water Quality and Amphibians and
article
0009 - Introduction to the Nitrogen Cycle for more information about
water quality and the importance of the nitrogen cycle.
Breeding Warty Newts
Paramesotriton species have seldom been bred in captivity, however,
breeding successes seem to be increasing in number. The article,
Hongkong
Warzenmolch - Paramesotriton hongkongensis, notes that P. hongkongensis enters breeding mode as the
temperatures rise after a mild cooling period of no lower than 53°F.
A hibernation period may not be necessary , but a cooling perioed should be followed by warming
temperatures to simulate the changing seasons. Arnaud Jamin notes in his
article,
Paramesotriton
sp. for Batraciens,
that P. caudopunctatus, P. chinensis, and P.
fuzhongensis (P. guanxiensis) can be kept around 68°F during
the warmer months, and as low as 41°F during the winter months, and that P.
hongkongensis can be kept at temperatures as high as 77°F during the
warmer months, but not lower than 59°F during the colder
months. At least a few keepers have bred P. hongkongensis by
providing a temperature schedule similar to the fore mentioned. On the other hand, some have said
that a cooling period lower than 53°F is too low for all
Paramesotriton species, and will inhibit egg development and
courtship. All in all, the exact temperatures required for Paramesotriton spp. are
quite debated.
Courtship is thought to be similar to Cynops species, consisting
of tail-fanning, and increased aggressive behavior. Tail-folding and
whipping has also been observed, in a similar display to that of Triturus
males during the breeding season. Females
usually deposit their eggs among aquatic
vegetation, sometimes in rows enclosed in thin leaves. In captivity, java
moss may be used by some newts for egg deposition. P.
caudopunctatus females have been known to guard eggs that are
deposited under submerged rocks in a similar fashion as Pachytriton
females.
Larvae can be housed in moderate to well-oxygenated waters, depending
on the species, for which an air
stone, bubble curtain, or filter should suffice to simulate. Some have suggested
that P. hongkongensis
eggs and larvae should be kept in the 70°F - 75°F range, and 69°F - 73°F for P.
caudopunctatus and P. fuzhongensis.
Morphs of P. hongkongensis, P. chinensis, and P. fuzhongensis
usually seek out terrestrial refuges, and may not adhere to a semi-aquatic tank setup. P. caudopunctatus
juveniles can be successfully raised in a semi-aquatic environment, with
temperatures in the low to mid 70's. P. hongkongensis
morphs
should also be kept slightly warmer, around 70°F - 75°F. The young of most species are uniform black or
brown, with bright belly coloration developing over time, except P.
caudopunctatus, which develop the characteristic orange dorsal stripe early
on, and are usually tannish brown in coloration. Sexual maturity
is reached by the second year in most species.
Feeding
in Captivity
 Paramesotrion species are voracious hunters, and can become
accustomed to a regular feeding regimen. Paramesotriton spp. have a keen sense of smell, but also hunt
vision and the lateral line system to detect food. Most individuals will
take food items on land and in the water, allowing for the consumption of a
diverse array of food items. Established individuals typically have no
problems taking food from forceps, and eagerly await their dinners near the
surface of their water. Paramesotriton
species tend to lunge at anything moving, including fingers, nets,
siphons, other newts, etc., especially those accustomed to regular feedings.
Fortunately, bites do not cause pain, but may be startling, and frantically jerking your finger
away
could damage the newts teeth. Impact trauma can also occur from erratic
movement of fingers, or objects, while in the mouth of newts, as they tend
not to let go, and could be accidentally banged into other structures in the
tank. If one latches firmly onto a finger, or other object,
gently scooping the little assassin up out of the water is usually an
effective deterrent. If that doesn't work, holding onto the abdomen region,
just behind the front limbs, and very gently pulling the finger or object
from the mouth usually entices the newt to release its hold. When feeding
with forceps, newts may sometimes try to devour the feeding device, as well.
The photo at right shows a Paramesotriton
hongkongensis coming out of hiding to grab a wax worm.
In the presence of food items, some individuals may become exceptionally
aggressive and cause injury to other newts in the tank. Because of
this, it may be necessary to feed individuals in separate areas of the tank,
or in separate containers to avoid accidental, or intentional biting and attacking. This is not the
case for all individuals, just something to watch out for. Paramesotriton species are one of the least
picky caudate species, as well, accepting frozen/live bloodworms, tubifex
worms, black worms, chopped earthworm, waxworms, maggots, etc. A list of
store-bought foods commonly available for captive individuals comprises
the following table, Table 1.1.
|
Table 1.1 - Commonly Available Food Items
acceptable for Paramesotrition spp. |
|
Aquatic Larvae: |
Terrestrial Juvenile |
Semi-Aquatic Adult: |
|
» Black worms
» Blood worms (live/frozen)
» Brine shrimp (newly hatched)
» Copepods
» Daphnia
» Earthworms (finely chopped)
» Fish pellets (finely crushed)
» Grindal worms
» Microworms
» Tubifex worms
» Paramecium
» Scuds (Amphipods)
» White worms
|
» Black worms
» Aphids
» Blood worms (live/frozen)
» Crickets (gut-loaded)
» Earthworms (chopped)
» Fly larvae (maggots)
» Flour beetles and larvae
» Fruit flies and larvae
» Grubs
» Rice beetles and larvae
» Tubifex worms
» Waxworms (small)
|
» Black worms
» Blood worms (live/frozen)
» Earthworms (chopped)
» Fly Larvae (maggots)
» Grubs
» Tubifex worms
» Waxworms (small)
» Crickets (gut loaded)
|
Amphibians should be given a variety of different live foods to ensure proper
nourishment and health. Chopped earthworm, waxworms, frozen/live
bloodworms, and maggots seem to work well as
semi-staple foods for adults. Some
individuals may have trouble digesting the tough outer "skin" of maggots, and
may vomit them back up, or excrete them undigested.
If undigested maggots are seen in the tank, they should not be fed to such
individuals. Crickets do not provide adequate nutrition, due to an imbalance
in the calcium/phosphorus ratio, and should not be offered as a staple item.
However, pinhead crickets possess a more balanced calcium/phosphorus ratio,
and are therefore an acceptable supplement to the main diet of juveniles.
Another problem with crickets is that they possess a hard exoskeleton, which
can cause serious digestive problems to newts who have gorged themselves.
This condition is called Gastric Overload, and results when partially
digested food remains in the stomach of the amphibian, causing strain on the
circulatory system and surrounding organs, and toxic buildup associated with
putrefying foods. Juveniles seem to develop well with a varied diet of
frozen/live bloodworms, live black worms, chopped earthworm, fruit fly
maggots, fruit flies, and occasional pinhead crickets. Feeding only one or
two types of food to developing juveniles could result in Metabolic Bone
Disease, or other nutritional disorders, due to lack of complete nutrition,
which is why it is important to feed juveniles a variety of foods. Adults and larvae should also receive a diverse, nutritional
diet, as they can develop nutritional disorders as well. Larvae can be fed
typical microfoods, such as daphnia, scuds (amphipods), copepods, black
worms, bloodworms, finely chopped earthworm, mosquito eggs, paramecium, etc.
For more
information about foods, and nutritional disorders, see
article
0002 - Amphibian Foods and Feeding Tips, and article
0001 - Common Amphibian Ailments, respectively.
Journal & Research References
Lu, Shunqing, et al. 2004. Molecular phylogeny of the genus Paramesotriton
(Caudata: Salamandridae). Biochemical Genetics 42(5/6): 139-148.
Nguyen, Van Sang, et al. 2003. Feasibility Study for a Conservation
Program for the Vietnamese Salamander Paramesotriton deloustali in Ba
Be and Cho Don Districts. PARC Ba Be/Na Hang.
Rehak, Ivan. 2002. Pačolek vietnamskỷ – vymirajici klenot z Tam Dao.
ZIVA Journal, 5/2002.
Stuart, Bryan L. and Theodore J. Papenfuss. 2002. A new salamander of the genus Paramesotriton
(Caudata: Salamandridae) from Laos. Journal of Herpetology 36(2): 145-148.
Sparreboom, M. 1981. Een onbeken de salamander uit China: Paramesotriton
caudopunctatus. Lacerta (jaargang nr. 8): 102-108.
Wen, Yetang. 1989. A new species of the genus Paramesotriton (Amphibia:
Caudata) from Guangxi and a comparison with P. guangxiensis. Chinese
Herpetological Research 2(2): 15-20.
Literature References
Fei, Liang. Atlas of Amphibians of China.
Technology Press, Zhengzhou, China. 1999.
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.
Zhao, E. China Red Data Book of Endangered Animals: Amphibia and Reptilia.
Beijing, China: Science Press; Endangered Species Scientific Commission, P.R.C.,
1998.
Zhao, E., et al. Studies on Chinese Salamanders.
Society for the Study of Amphibians and Reptiles, 1988.
Internet References
AmphibiaWeb. (2003). http://amphibiaweb.org/.
(Accessed: 2001-2003).
Chan, Danial Y.M. (1999). Tung Lung Waterfalls, Lantau North County Park.
Waterfalls of Hong Kong.
http://www.hkwaterfall.net/hk_falls/hkwf_tlg.htm
(Accessed: 2003).
Frank, Peter. Hongkong Warzenmolch - Paramesotriton hongkongensis.
http://www.agurodela.de/arten/paramesotriton_hongkongensis.php
(Accessed: 2002).
Hung, Le Manh, Tran Theiu Du, and Vu Huu Trac. (2002). A rapid field
survey of Xin Man and Yen Minh Districts, Ha Province, Vietnam. Birdlife
International Vietnam Program.
http://www.wing-wbsj.or.jp/~vietnam/pdf/reportHG.pdf
(Accessed: 2002).
Jamin, Arnaud. Paramesotriton sp. Batraciens.
http://www.batraciens.net/paramesotriton_sp.html
(Accessed: 2002).
Tordoff, Andrew W., et. al. (2002). A rapid field survey of Van Ban
District, Lao Cai Province, Vietnam. Birdlife International Vietnam
Programme.
http://www.wing-wbsj.or.jp/~vietnam/pdf/reportVBan.pdf
(Accessed: 2002).
Travel, Guangxi. China Internet Information Center.
http://www.china.org.cn/english/travel/41300.htm
(Accessed 2004).
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