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pp.31–38

Behavioral Observations and Descriptions of the Endangered Knobby
Newt Tylototriton wenxianensis and Their Application in Conservation
DA-JIE GONG* AND MAI MU
Department of Biology, College of Life Science, Northwest Normal University
Lanzhou, Gansu, China.
*Corresponding author E-mail: gongdj@nwnu.edu.cn
Abstract.- Tylototriton wenxianensis is an endangered species at the status of VU. This paper introduces observations
and studies including breeding, territorial, communication and antipredatory behaviors. The rules and mechanisms of
the behaviors are recorded separately and described in the paper. Relationships between the behaviors and environment are also explained for the purpose of describing how the environment acted on the behaviors and how the behaviors adapted to the environment. The paper ends with conservation plans in connection with behavioral ecology: A)
protect the particular habitat and avoid anthropogenic threats, B) artificial construction for natural migration and genecommunication and C) artificial breeding and re-introduction into nature.
Keywords.- Observations, Tylototriton wenxianensis, behaviors, habitats, conservation.

Introduction
Tylototriton wenxianensis (Caudata: Salamandridae) is
peculiar in China and poorly known. It is only foundalong the boundary between Gansu and Sichuan
Province, narrowly distributed in Wenxian, Qingchuan
and Pingwu (details see Table 1 and Fig. 1). It was
defined as a threatened species with the status of VU(IUCN, 2006).
A representative of the Family Salamandridae, T.
wenxianensis has rough and toxic skin with seasonalcolors. The bilateral warts stay longitudinal and clustered in the same size and the boundaries between them
are not clear enough. It is dark all over apart from the
red-orange fingers, toes and the venter of the tail (Fei,
1993).
The newt lives in the heavily forested mountains atapproximately 940 m. The adult is terricolous and usually wanders about the pool. It stays hidden under wet
gravel or small muddy caves covered by fallen leaves inthe daytime and appears at night for preying (Gong andMou, 2006).
Behavioral studies on tailed amphibians haverecently been reported. The studies were mainly focusedin the following fields: breeding behaviors, including
sexual recognition and sexual selection (Dawley, 1984),
modes of courtship and mating (Arnold et al., 1972,
1977; Salthe, 1967, 1974), sperm competition (Arnold,
1977; Halliday, 1998; Massey, 1988; Sparreboom, 1995;
Verrel et al., 1998), reproductive behavior (Fang, 1984;
Harris et al., 2002; Park et al., 2000), parental care
(Cramp, 1995; Nussbaum, 1985; Peterson, 2000), evolution of reproduction (Arnold, 1977; Veith, 1998; Verrell
and Krenz, 1998,); migratory behavior (Arntzen, 2002;© 2008 by Asiatic Herpetological Research

Douglas, 1979; Griffiths, 1996; Serbiolova, 1995;
Twitty, 1966); territorial behavior (Mathis, 1991; Mathis
et al.,1998; Mathis et al.,2000; Simons et al., 1997);
communications (Holliday, 1997; Houck, 1988; Houckand Sever, 1994; Rollmann, 1999; Verrell, 1989,1989a;);
antipredatory behavior (Brodie, 1990; Graves andQuinn, 2000; Maerz et al.,2001; Sih et al., 2000, 2003;
Storfer and Sih, 1998; Sullivan, 2002; Woody andMathis, 1997). All the above provide a source of reference for the behavioral study on T. wenxianensis.

Methods
Migratory behavior.- We chose the habitat in Qingchuan
as the site to observe, where a large population was
found (Gong and Mou, 2006). The newts are not active
year round except for the breeding season, so theirmigratory behaviours seem quite obvious and clear. We

Figure 1. Location of the habitats and observation sites.

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30

30

20

20

10

10

0

3-31*

4-4
4-2*

4-8
4-6

4-12
4-10

4-16*
4-14

4-20
4-18

4-24
4-22

4-28
4-26

5-2
4-30

5-6
5-4

Temperature/°C

Number of migratory newts

Asiatic Herpetological Research, Vol. 11

5-10
5-8

Date

Figure 2. Reproductive migration of Tylototriton wenxianensis. The polyline represents temperature, columns represent number of the migratory newts and “ * ” rainy days.

made continuous observations when they migrated forbreeding towards the pools nearby. We marked eachmigratory group with varisized loops and observed byradio tracking and GPS. Finally we made records of thetime, route length, size of groups and sex ratio in the
migration.
Reproductive behavior.- Reproductive behaviors
included mating and spawning. We observed the course
and took down the data of sex ratio, time, envioronmentand quantity of the spawns etc.
Conmmunication.- Communication happens along withcourtship. We tried to understand how the newts communicate with each other and why. The communications
included chemical signals and body-contacts (Jiang,
2004). Mechanisms of chemical signals from glands are
expected to be understood through dissection. Types ofthe body-contacts and their effects on courtship werestudied.
Territorial behaviors.- We measured the size of the territory occupied by 6 female newts and size of the tails
and bodies in contrast. Aggressive behaviors related tothe available prey in the environment were also
observed. Relationships between the data and phenomenon were discussed and concluded.
Antipredatory behavior.- The newts were less aggressive without structures for aggressivity (Jiang, 2004).

They moved too slowly to escape attacks from enemies
and to defend themselves. We made some model enemies and demonstrated the predation and antipredatory
behaviors artificially.

Results and Analysis
Behaviors.- We made continuous observations on the
migratory behaviors and found 186 adults including 170
females and only 16 males, the ratio was 170:16 (♀:♂)= 10.625:1. Breeding lasted from early April to late May
with a peak from April 8th to May 6th. The average temperature was 15.5±4°C (Fig. 2).
Like many other newts, T .wenxianensis does notseem to have significant secondary sexual characters. It
distinguishes and selects the opposite sex primarily bysight and sense of smell. The male seemed more active.
It courted the female by wagging its tail and dancing
round. Meanwhile, the female was secreting chemicals
and emitting a strong smell to attract the male. Soon themale bit the female on the tail (or hind limb), then they
circled around both with their tails wagging (Fig. 3).
Experiments show that the male prefers to choose a bigger one with more ovums and a stronger smell.
Mating did not accompany spawning in the meantime. It was the female that selected where to spawn.
The male moved to the pool and settled down earlier
than the female. However, the female left soon after
spawning, while the male stayed untill the end of the

Figure 3. Model of mating behavior. Shows the male (black) wagging its tail and dancing round.

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Table 1. Distribution and flora of T. wenxianensis along the boundary between Gansu and Sichuan Province.
Province

City

Country (town),
countyVillage

Latitude andlongitude

Altitude (m)

Flora

Gansu

Longnan

Bikou, Wenxian

Bifenggou

927

semitropics

Gansu

LongnanLiziba, Wenxian

Moziping

940

Gansu

Longnan
Hanjialiang
Wuxingsemitropicsintersemitropics
semitropicsintersemitropics
semitropicsintersemitropics
intersemitropics

Miaoshanli103.7°E
33.95°N
104.6°E
33.2°N
104.4°E
33.1°N
105.1°E
32.5°N
105.1°E
32.46°N
105°E
32.68°N
105.14°E
32.32°N
104.23°E
32.58°N
104.3°E
32.44°N
104.34°E
32.45°N

Sichuan

Guangyuan

Sichuan

Guangyuan

Sichuan

Guangyuan

Sichuan

Guangyuan

Sichuan

Mianyang

Sichuan

Mianyang

Liziba, Wenxian
Daba,
Qingchuan
Daba,
Qingchuan
Daba,
Qingchuan
Daba,
Qingchuan
Shuitian,PingwuBazi, Pingwu

Sichuan

MianyangBazi, Pingwu

Dawuji
Laowuji
Baishuling
Dafenling
Yinshanli

1100
1160
998
1092
1210
1134

semitropicsintersemitropics
intersemitropics

962

semitropicsintersemitropics
intersemitropics

970

intersemitropics

Figure 4. The sperm transmission. Swells show the colloidal secretion.

Figure 5. The courtship behavior. Shows the transmission of pheromones in the fantail.

Figure 6. Rigid body: the reaction to anti-predator. Shows
ribs jutting through the side warts.

Figure 7. The raised tail and warning color on the ventral
surface of the tail.

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Figure 8. The protuberant glands and warning color on
the back. Shows harmful and toxic skin secretion, glandiform warts, spine glands, caudal glands, glandiform
warts, spine glands, caudal glands, tergal glands, etc.

Figure 9. Playing dead. Shows the newts played deadwith upward bottom and rigid body.

Figure 10. Mimicry: newt playing dead, with venter
upward and body rigid.

Figure 11. The breeding pool (after spawning). Vicinity ofWuxing Village, Sichuan Province, southwest China.

Table 2. Relation between spawning territory, food and size of six female newts.
Areas of spawning territory (m2) Percent of total (%)

No.

TL (mm)SVL (mm)Food environment

1♀

0.64

10.39

58

71

poor

2♀

0.81

13.15

59.2

72.2

poor

3♀

0.55

8.93

55.7

70.1

poor

4♀

1.26

20.45

65.9

73.2

medium

5♀

2.3

37.34

67.4

74.6

high

6♀

0.6

9.74

56.1

71.4

poor

Table 3. The antipredatory behaviors of T. wenxianensis, grouped by different types. The observations were made in
the field and in artificial conditions. a. harmful skin secretion; b. toxic skin secretion; c. parotids; d. glandiform warts; e.
spine glands; f. caudal glands; g. tergal glands; h. tergal colors; i. ventral colors; j. rigid body; k. roll of body; l. exposed
abdomen; m. raised tail; n. raised jaw; o. arched body; p. larger head; q. swagging tail; r. curved forehead; s. extended
nerve.
Imitative
toxicant body

Types
No.

a

b

c

d

WarningcolorsefghParticularskeleton

Warning postures
i

j

k

l

m

n

o

p

q

r

s

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plantsfood

breeding behavior. Tactile signals were transmitted
through touch on the mouth, collisions and friction
between bodies.
In the long course of evolution, series of antipredatory behaviors had formed against captures, such as
escapes, cryptic coloration, rigid body (Fig. 6), aposematic coloration (Fig. 7, Fig. 8), camouflage (Fig. 9),
warning postures (Fig. 7), imitative toxicants, particularskeletal adaptations (Fig. 6), chemical defenses andplaying dead (Fig. 10) etc. (Table 3).

stonespool
soil and mud

Figure 12. The artificial breeding box made of glass, 110
× 40 × 60 cm (lateral view). Shows the constitution andfeatures of the breeding ecotope under artificial control.

Relationship between the behaviors and environment.The female finished spawning in a short time withoutmale involved. Spawning lasted 2–3 days smoothly inrainy seasons without interference. They got to thespawning pools no sooner than they selected the propersites. Spawning began one day later. The female
spawned one a time lasting half a hour, and the wholecourse 5–6 hours. The course maintained a even paceslowly without a peak.
In Qingchuan, three pools were found with female
newts and a mass of spawn. The three conditions forspawning sites were: A) ground covered with plants (theplants may be divided into three layers: the upper consisting of tall sparse laurisilvae, bushes in the middle andwet weeds at the bottom), B) spawning sites with semipermanent pools, and the water soaked out after rain, C)
it chose a mesa with loose soil as the spawning site on
the hillside, covered by fallen leaves (Fig. 11).
The newt preyed mainly on insects, earthworms andsnails. Like other amphibians, it stayed hidden in the day
and preyed in the night. The behaviors changed sensitively with the temperature and sunlight.
In summers, plants flourished and numerous insects
appeared, when the newt acts frequently. After thunderstorms, earthworms come out from the soil and supply
food to the newts. In winter, the newts have to hibernate
in response to the lack of food and low temperature.

breeding season. The analysis from Douglas (1979):
First, to choose the optimum condition for spawning andto reduce risks in breeding migration; Second, to control
the male genotype effectively through sexual selection.
Mating behavior happened on land, while spawningunderwater. We analyzed the reason: the adult newt
mainly lives a terricolous life. Sometimes it approaches
the pools or streams for food. The observations on the
behaviors in captivity indicate that the adults neededalmost no water unless they felt too dry. They had adapted to the climate overland. In addition, in reproductive
seasons, inorganic conditions overland near the pools
may help them to secrete pheromones and sex hormones
which could be significantly reduced underwater.
The newt took no attack on its kin, but became quite
different towards non-related individuals. The newt recognized its kin through chemical signals. In the hot summer, the adult admitted the posterity into its territory
sometimes and food became insufficient. Occupying territory usually accompanied attacks and fights. That
depended on food quantity, size of body and environment (Table 2). The newt with longer tail and bodyoccupied larger territory, closely related to the foodquantity.
Communications were drived mainly by chemicalsignals and body contact. Chemical signals are necessary for the accurate and regular courtship behavior. The
newts distinguished the sex and attracted each other bytransmission of pheromones (Fig. 4) and secretion of
hormones (Fig. 5), which later brought about the mating
behavior.
Body contact was also important, particularly inConservation plans.(1) Protect the particular habitat and avoid anthropogenic threats.
•Figure 10 shows the particular breeding habitat. The
newts enter the pools and ground nearby only in

Table 4. Relationship between the conditions and hatching rate under artificial control.
No.

Temperature Relative
Water pH*(°C)
humidity (%)

Sunlight
Incubation No. of larvae No. that
periodperiod (day)hatched out(hour/day)

Hatching
rate (%)

1
2
3

5~15
10~20
15~25

30~40
40~50
50~60

5.2
5.4
5.6

0.5
2
3

32
28
23

40
50
45

5
11
28

12.5
22
62.2

4

20~30

60~70

6

1.5

20

52

40

76.9

*Huang (2007) reported that pH 5.2~6.0 was most suitable for the development of larvae.

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reproductive seasons, so it is of great significance toprotect the similar pools. The pools were more or less
50 cm deep, 10 m2 in size and at an elevation of 1000
m. Pieces of rocks piled around. Soil and mud aroundthe pool was covered densely by rotted leaves. The
vegetation nearby were mainly made up of shrubs andarbors (see Table 1).
Human activities, especially farming and poaching
severely threatens breeding. Recommendations aboutplans to avoid these threats: A) build up fences aroundthe mating sites and breeding pools to prevent thenatural enemies and poachers from breaking in, B)
reduce farming and grazing by livestock, especiallyaround irrigation water from the pools, C) make (andenforce) laws to punish poachers.

(2) Artificial construction for natural migration andgene communication.
•Protect the whole habitat and ensure the natural
migratory behavior. The migratory routes are uaually
blocked by farmland etc., so it may help keep theproper migration to build up artificial passages acrossthe farmland.
Connect the adjacent pools by building canals asmuch as possible in order to ensure the communication of gene from different populations, especiallyfrom breeding groups. The measure may help avoid
inbreeding depression and loss of genetic diversity ina small population and also help with evolution of thespecies.
(3) Artificial breeding and re-introduction into nature•In view of the high sex ratio (♀:♂ = 10.625:1 on
average, n = 186) in the breeding season and the low
rate of hatchability (46.54%; n = 1272) in nature, it isquite feasible to increase the hatching rate and reduce
the sperm competition between the males in artificial
conditions. We directly gathered the females andeugenic males with longer bodies (longer tails andbodies seem dominant in the sex competition apt tosurvive under natural selection (Jiang, 2004) andhigher sperm density, and also keep the sex ratio at
1:1 in a artificial glass box (Fig. 12) to avoid competition and injury as well as to ensure high-quality
inheritance.
Studies showed that factors influencing the hatchingrate were natural enemies (such as snakes), climatechanges and pollution (Tian et al., 1997). It is aneffective approach to raising the survival rate andenlarging the population by hatching out the larvae in
artificial conditions and re-introducing them into nature.
The conditions include temperature, humidity, sunlightperiod and pH of water etc. (Table 4). Re-introduce the2008

juveniles into nature when they complete metamorphosis and get ready to land.

Acknowledgments
We wish to thank K. R. Zhang for assistance and valuable help with our investigation of distribution of T.
wenxianensis and J. R. Teng who provided us the specimens of T. wenxianensis.

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Submitted: 17 November 2006
Accepted: 22 September 2007

2008

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