1999

Asiatic Herpetological ResearchVol. 8, pp. 29-37

Diel Activity of Ranodon sibiricus (Amphibia: Hynobiidae) in Relationship toEnvironment and Threats

Dag Dolmen Rudolf A. Kubykin 2 and Jo V. Arnekleiv
1

1,

Norwegian University of Science and Technology, The Museum; N-7004 Trondheim, Norway.

National

Academy of Science, Inst, of Zoology; Almaty 480032, Kazakhstan.
Abstract. -¥vt\d studies on the diel activity of Ranodon sibiricus were carried out in Dzhungaria, southeastern

Kazakhstan, in August 1994 and June 1995. Counts showed that adults and juveniles were strongly nocturnal,
which was also confirmed with the help of IR light/photocell equipment, whereas larvae were active both nightand day. The daily activity period of adults and juveniles started near the 5-lux limit and peaked around 23-01
hrs. Its length followed that of the night and was accordingly shorter in June than in August.

When the activity ofaquatic adults and juveniles started in the early night, about 5% came out of the water and walked on landalongside the brook. Most ranodons, however, were relatively stationary sit-and-wait predators in the water. The

UV

nocturnal activity of/?, sibiricus protects it against dehydration (terrestrial animals), strong
radiation anddiurnal predators. The most important task of the nocturnally and diurnally active larvae is to grow as fast as
possible to survive the first, hazardous hibernation. The difference in diel activity between adults/juveniles andlarvae reduces intraspecific competition and cannibalism within the population. Because of their secretive life,

metamorphosed stages of R. sibiricus are not easily accessible to predators or Man. Hence, the most important
threat is probably the large herds of cattle grazing in the area. Since adults and juveniles are well hidden by day,when the cattle wade the brooks, they are fairly safe, but cattle may pose a great threat to ranodon eggs andlarvae.

Key words. -Ranodon sibiricus. Kazakhstan, diel activity, rare species, environmental adaptation, threats.

Introduction

sible adaptations to the environment. Several otherquestions were also of interest. For instance, do theA threatened, secretive species

larvaeThe salamander Ranodon sibiricus Kessler, 1866 isanimals to reduce intraspecific competition and canni-

behave differently from the metamorphosed

Dzhungarian mountains of south-

balism (as in Triturus; see Dolmen 1988)? Does theeastern Kazakhstan and the extreme west of Xinjiang

nocturnality of the species really make it almost safe

province
range
very small and has
man-made
undergone manychanges during the past.

can R. sibiricus avoid the heavy impact of some 150

distributed in thein

China.

Itsis

The species is therefore threatened and is on the redlist of the

et al.

former Soviet Union/Kazakhstan (Bannikov

1978; Zhao and Adler 1993). Its habitat is cold,from terrestrial predators and Man? To what extent

cows which graze along and trample in the brooks
when crossing the study area twice a day in summer(Kubykin et al. 1995; Dolmen et al. 1997)?

tive way of life, individuals are not always easy to findA preliminary report on these studies was presented by Dolmen et al. (1995). Hereafter. Ranodon

even where they may be locally abundant. Althoughsibiricus is mostly referred to as ranodon.

clear mountain brooks. Because of their quite secre-

Shnitnikov (1913) found it easily, Brushko and Nar-

baeva

(

1988) emphasized the difficulties of finding R.

Principles of measuring activity

mountain rivers. Its secretive life isIn principle, there are at least three ways of mea-

also reflected in its nocturnal activity, an aspect briefly

suring the activity of an animal: 1 ) with an actographor other technical device such as light beam/photocell

sibiricus in thetouched upon by several authors (Shnitnikov 1913:
Paraskiv 1953; Kubykin 1986: Narbaeva and Brushko
1986; Wang 1990).

As part of a wider ecological study of Ranodon
sibiricus in Dzhungaria, we investigated its activity,
partly in 1993, but more thoroughly in 1994 and 1995,
to learn the details of its diel activity rhythm and pos-equipment, 2) by interval trapping of active animals,
and 3) by counting active animals. All three methods

have beenused for Triturus species
Europe (e.g. Himstedt 1971; Dol-successfully(Salamandridae) inmen 1983a,b; Griffiths 1985).

1999

Asiatic Herpetological ResearchVol. 8, p. 30

The weather in both periods was mostly clear. The

Preliminary studies

1994 and June 1995 varied

In June 1993. the trapping technique was tried out for

air temperatures in August

ranodons in a small brook in the Borokhudzir valley

between 5.7 °C and 17.1 °C, and 3.5 °C and 21.0 °C,
respectively. The air humidity in August 1994 variedin

Dzhungaria. On 15 June, 10 simple funnel traps (1L "Cola traps") and two more complicated traps witheight entrances each (constructed by Arne Haug) werelong stretch of a small brooklet
placed in a 15m

(Brook 3)

in

which

six

ranodons and 50 double

clutches of eggs had been registered two days earlier.

Alternate traps had their entrances pointing

down-

stream and upstream. The traps were checked/emptied
every third hour throughout a day and night, but noranodons were caught. The water temperature varied

to 100% RH, with maximumThewater temperature at the site
humidityin
Brookvaried in August 1994
investigatedbetween 5.5 °C in the early morning and 19.3 °C in
the afternoon, and in June 1995 between 5.0 °C and20.0 °C. In Brook 5, the water temperature varied
between 6.2 °C and 14.8 °C in August 1994 and 4.8°C and 15.6 °C in June 1995. A more detailed description of the area is given by Dolmen et al. (1997).

from

than

less

50

at night.

1

between 3.8 °C and 7.6 °C. On 17 June, a similar setup was tried on a 15 m stretch of another, somewhat
larger brook (Brook 1) where, in addition to the ranodons living there naturally, 38 additional ones werereleased.

A few were caught during the next day andMaterial and Methods
Field investigations

The study took place in August 1994 and June 1995.

night, but never more than two each time.

Two persons equipped with good torches examined a

However, a study of a third brook (Brook 5) atnight revealed that many ranodons were going onto

total

land and wandering on the dew-wet banks beside thebrook. On a 500

m stretch at 23.00-24.00 hrs. (stan-

dard time), four

men with torches counted

odons that were active on wet land and1

14 ran1hiddenunder stones alongside the brook. Although aquatic
individuals were also active, the main movements up

and down the brook seemed to be taking place onland, not in the water. The trapping technique in water

was accordingly unsuitable for measuring the activityof ranodons.

For 1994 and 1995. it was therefore decided to use
the counting method for measuring the activity of ran-odons, and for comparison, data from the use of lightbeam/photocell equipment could serve as a control.

Description of the area

The study area, on the southern slopes of the River
Borokhudzir catchment in Dzhungaria at an altitudeof approximately 2200 m, consists of grazing for catbog, and juniper-covered hillsides. Two small tributary brooks to the River Borokhudzir (Brook 1 andtle,

Brook 5), approximately 200-300 m apart, were chosen for the study. Both were spring-fed, about 2-50
cm deep and 30-80 cm wide, and had gravelly or
stony bottoms. In a few places, turf almost overgrew
and covered the water. Boulders, where terrestrial ran-odons hid, could also be found on land alongside thebrooks.

The velocity of the water at different sites

along the two brooks varied from 0.0-0.2 and 0.0- 1 .0
m/s, respectively. Measurements were made in August

1994 when the flow was low.

of 90 m and 190 m, respectively, along the twobrooks (Brook 1 and Brook 5) every second hour for
three days and nights. The walking speed was approx-

Deep pools and shallow slowlywater
were
especially well examined, andflowingon land alongside the brooks.
also the nearest 2-3
imately 8 m/min.

m

No stones were turned, however, and no ranodons
were disturbed, except possibly by the torchlight. The

two brooks had been divided into sections of 5 or 10m, respectively, for statistical use. and the number ofranodons in each section were counted separately. The

numbers in different main developmental stages werealso kept apart: 0+ (young larvae, less than one year

1+ (old larvae and metamorphosed individuals
from the previous year), larger juveniles, and adults.

old),Aquarium investigations
In

December 1994, additional investigations of theactivity of

two ranodon individuals, an adult and alarge juvenile, were carried out in Almaty. An experi-mental aquarium was made of transparent plexiglass

shaped like a circular channel. The outer diameter was
36.5 cm, the inner diameter 15 cm and the height 10
cm. The aquarium was filled half full of water broughtfrom Dzhungaria and some large stones acted as shelter for the animals and to enable them to climb up into
the air. The aquarium was equipped with an infra-red

beam/photocell system (Visolux LS 4-GaAs).
and
The beam was set to a diameter of about 2
light

mm

focused near the bottom of the aquarium, where theranodons usually walk around. Interruptions of thebeam were recorded by an electric counter (Visolux
LU GaAs), which was read every second hour
throughout eight days and nights. Food was not pro-Asiatic Herpetological Research

1999

placed inside a building, close to a north-facing win-

The chi-square test was used to find out whether
or not the activity patterns had a significant diurnal

dow. The(06-18 hrs.) or nocturnal (18-06 hrs.) predominance,

vided

duringthelightexperiment.

The aquarium wasVol. 8, p. 31

regime was therefore natural,i.e.

approximately like outside. The temperature was constant and slightly lower than the usual room temperature.

The same equipment, connected to a 12V car battery, was used for an adult and a large juvenile in
D/hungaria in June 1995. The aquarium was protected from disturbance and strong sunshine by being

placed halfway under a car and partly covered with ablanket. The water was replenished (in daytime) once

and the chi-square

testwith two variables withoul

expected values was used to test any differences in
activity pattern between, for example, adults and lar-

Results
The field activity studies

and nights. On the second evening, the onset of activ-

The activity of Ranodon sibiricus was definitely nocturnal, but with some modifications with respect to thedevelopmental stage and the month of the year. The

ity was studied in more detail at 5-minute intervals foranimals in all stages were amazingly clever at hiding

two hours.

during parts of the day and night. Several were seen atnight on a gravelly bottom along stretches of theduring the experiment, which ran for barely three days

Environmental factors

brook which we had searched very thoroughly in day-Light intensities between

and 10 lux normally act astime without finding any.

Zeitgeber for activity in plants and animals (cf. Dol-men 1983b). The 5-lux value is therefore often used in
activity studies as a limit

between day and night. In

our study it was determined subjectively, evening andmorning.

The 5-lux

value

in

Dzhungaria occurs

roughly 10-15 minutes before (or after) the 0.1-0.5
lux values, which corresponds well with the lightintensity

required

instance. Thiswasreading a newspaper, forconfirmed with a Hartmann andforBraun ECLX 4 luxmeter.

approximately symmetrically around the darkest timeof night (ca. 24.00-00.30). All hours mentioned hereafter refer to standard time, notsummer time. An

important difference in the understanding of diagrams
based on the two methods used for measuring activity
isthat a)same patterns in both brooks, although the peakapparently varied slightly from day to day between 21
and 01 hrs. (Fig. 1).

Adults and juveniles had similar patterns, and seen
together the rise in activity level from 19 to 21 hrs.

was always

the counting

method measures

activity at the time of counting,thewhereas b) the IRsignificant(P<0.002: WilLikewise, the1 ).

decrease in activity from 03 to 05 hrs. was always sig-

Brook 5, where most animals were regisThe variation in the time of peak
activity from day to day was usually not significant.
tered (P<0.002).

Brook 1, the larvae showed a similar rise inactivity from 19 to 21 hrs. When 0+ and 1+ larvae areIn

counted together, the increase is significant (P<0.002
or P<0.02) on two of three dates. The activity of 0+
larvae in Brook 5 was not as well defined as in Brook

being much more diurnal. In contrast to adults andjuveniles, and also to the larvae of the other brook,1,

beam/photocell method measures the activity during
the two hours preceding the time when the counter iswhich wereread.

squaretest),

allmainly nocturnal (P«0.001; chi-the larvae of Brook 5

showed a muchofdiurnality (nocturnality tested:higher degree
0.20<P<0.30). Indeed, traces of diurnal activity can

Statistics

Wilcoxon's

statisticallycoxon's signed-ranks test, see Fig.
nificant in

The times for reading the Visolux counter were set

here

In August 1994, the diel activity showed basicallythematched-pairs

signed-ranks

test

(two-

was used to test any significance levels in theactivity measured in the field, the number of ranodons

also be seen in the 0+ larvae of Brook 1.

tailed)within the different brook sections being counted foran hour and comparison being made with the situation

two hours later.

The same test was used for the aquarium study,
but it was based on the ranodon counts for all the days
within a certain time interval combined, and thesewere compared with the corresponding counts in thenext time interval.

The period of activity of this species fitted very
well with the light and dark periods, i.e. activity wasconnected with the part of the 24-hour period when
the light intensity was below 5 lux. This was approxi-

mately 20.00-05.30 hrs. at that time of year. Except
for young larvae, any activity outside that period was

negligible.

A pattern similar to that of August 1994 wasrevealed in June 1995. The period of darkness (<5Vol. 8, p. 32

Asiatic Herpetological Research

Hrs.

09

1999

Hrs
11 13 15 17 19 21

23 01 03 05 07 09

09 11 13 15 17 19 21 23 01 03 05 07 09

j9 11 13 15 17 19 21 23 01 03 05 07 09

09 11 13 15 17 19 21 23 01 03 05 07 09

4525

403S -

5
10-

15

20-

11 13 15 17 19 21

23 01 03 05 07 09

09 1 1 13 15 17 19 21 23 01 03 05 07 09

Figure 1 Diel activity of Ranodon sibiricus, based on
counts in Brooks 1 and 5 in August 1994. Levels of sta.

*tistical significance:

P<0.05 or P<0.02;**

P<0.01 orP<0.002.

was shorter than in August, i.e. about
21.00-04.30 hrs., and the activity peaks of adults andlux) in Junelarge juveniles

were narrower, at 23 (21) or 01

hrs.

I

11

13 15 17 19 21 23 01 03 05 07 09

09 11 13 15 17 19 21 23 01 03 05 07 09

Figure 2. Diel activity of Ranodon sibiricus in Brooks 1
and 5 in June 1995. For explanation, see Fig. 1.

(Fig. 2).

The youngest stage(1 + )

showed activity peaks
varying from 21 to 03 hrs. and also some diurnal
activity.

At least three times, larvae (one each time)

were even registered directly exposed to sunshine, at
1 1,

13 and 17 hrs. Nevertheless, on the whole, the lar-vae definitely had a nocturnal pattern.

hrs.) and a minor peak at 22 hrs. (i.e. 20-22 hrs.). withlittleor no activity during the light hours. The rise inactivity

from 16-18 to 18-20 hrs. and from 22-00 to

00-02

hrs.

was

significant(P<0.05),as

was thedecrease from 00-02 to 04-06 hrs. (two time intervals
seen together) (Fig. 3).

The activity period of the species again fitted very
well with the (shorter) dark period. The rise in activity
from 19 to 21 hrs. was always significant (P<0.002 or0.02) in Brook 5, both for adults/juveniles and 1+ larvae, as was the decline from 03 to 05 hrs (P<0.002,
0.01, 0.02 or 0.05),

had a main peak at 02 hrs. (i.e. peak activity at 00-02

and usually even 01-03 hrs. for

adults and juveniles in both brooks (P<0.002, 0.02 or
0.05). The day-to-day variation in peak activity seenat midnight was not significant.

The activity period of the animals at this time ofyear was clearly longer than the activity measured insummer, as was the period of darkness (<5 lux),which lasted from approximately 17.30 to 08.00 hrs.

The aquarium study in June 1995 confirmed theresults of the field investigations, although there wereminor deviations, probably for methodological reasons. Activity was negligible until 21 hrs. when a risein activity occurred resulting in a peak at 21-23, 23-01

or 01-03 hrs. (Fig. 4).

The aquarium studies
In the aquarium investigations in December 1994, theIn the more detailed aquarium investigations on 20

activity,

June, the activity started about 15 minutes before theA

5-lux limit was reached in the open. However, since

cumulative curve for the whole period of investigation

the aquarium was partially covered (see Methods), theranodons also showed a definite nocturnal

with peak activity at 20-22, 22-00 and/or 00-02 hrs.

Vol. 8, p. 33

Asiatic Herpetological Research

1999

Hrs.

Hrs.

14

12

16

20

1!

22

00

02

04

06

08

10

12

13

15

17

19

23

21

01

03

05

07

09

11

13

09

11

13

No.

19-20 June 1995

150-1

100

50

350
Figure 3.Diel activity of two Ranodon sibiricus speci-

300-

mens kept in an aquarium under natural lighting conditions for eight days and nights in December 1994,250-

based on the IR light/photocell method. Level of statis-

200-

*

tical significance:

P<0.05.

150-

the activity
light intensity there was a bit lower and5 lux. It
probably started at a light value very close toincreased gradually over the next hour (Fig. 5).

Observations of aquatic and terrestrial ran-

odons

100

50-

20O
21-22 JuneBy far the majority of active ranodons were foundon land. The frequencies of finds
made on land relative to the total number of ranodon
counts were only 6.1% (N=425) and 3.6% (N=279)
for August 1994 and June 1995, respectively. The
in

20-21 June

the water, notnumber of ranodons counted on land alongside thebrooks was thus relatively low, but sometimes
increased when the number counted in the water
increased. Animals walking on the moist grassy

150

10050

—
13

i

1

l

15

17

19

—

i

1

21

23

1

01

—

1

1

1

03

05

07

Figure 4. Diel activity of two Ranodon sibiricus speci-

mens kept in an aquarium under natural lighting conditions for nearly three days and nights in June 1995,

ground beside the brooks had a wet skin and werebased on the IR light/photocell method.

considered not to be really terrestrial, but aquatic ani-

was again reached in the morning, the activity had
decreased to approximately zero. This means thatmals which had only

left

the water temporarily forsome reason.

even though the species is clearly nocturnal, it uses all

ranodons were active to the same degree at
Not
same
the
time, not even near midnight. In June 1993,
allwhen we counted ranodons on land at 23.00-24.00
hrs., i.e. when the activity was at its highest, 14 wereseen walking along the bank, but another 1 1 really ter-

specimens (with a dry skin) were still passive
and very sluggish, hiding under stones.
restrial

Discussion

Most earlier authors have noted the extreme nocand the more diurnal

turnal habits of the ranodons

habits of the larvae (Shnitnikov 1913; Kubykin 1986;

Narbaeva and Brushko 1986). Metamorphosed specimens, both in water and on land, hide under stones
during the day, but come out at night and often move
upstream or downstream for some distance. Wang

(1990) stated thatinwestern Xinjiang, China, ran-odons also hides among plants and under stones durTheing the day and is active between 24 and 06 hrs.

The activity pattern
Ranodon sibiricus proved to be highly nocturnal, withvery

the time it can get for activity, i.e. mainly hunting.

little, if any, activity during the day.

An exception

likeXinjiang province,
Beijing time,

which

is

the rest of China,

follows

three hours earlier than sun

was the young larvae (0+) in August, in part also the
one-year old larvae (1 + in June, which showed a cer-time in western Xinjiang. The true activity time ofranodons in western Xinjiang should therefore betain amount of diurnal as well as nocturnal activity.
The 5-min. reading in the aquarium study showed inaround 21-03)

detail that the ranodons gradually started their activity

near the 5-lux limit in the evening. When that limit

hrs.,

i.e.

similar to that found in ourinvestigations in eastern Kazakhstan.

20

19

1999

Asiatic Herpetological ResearchVol. 8, p. 34

21

24

22

01

for a day or more, and aquatic ranodons, which have a
wet skin and which only temporarily come out of thewater (Dolmen et al. 1997).

No.
5 mm.

Shnitnikov (1913) and Paraskiv (1953) also noted
and adult ranodons leave the water orthat juvenile
20 Junetheir terrestrial hiding places at twilight and in dark-

ness, and are then active and are found on the banks ofbrooks, where they often

move long distances (cf.

Kubykin 1986; Narbaeva and Brushko 1986).
However, as these authors pointed out. not all ran-10-

odons leave the water at night, at least not at the sametime. The majority, in fact, stay in the water and may

5-

only occasionally come up. We found that

19

RanodonFigure 5. Details of the onset of activity of two
sibiricus specimens kept in an aquarium under naturallighting conditions on 20 June 1995,

light/photocell method. Activity

based on the IR

values are shown ascounts per 5 minutes (dots) and per 2 hours (trianthe 5-lux limit of lightgles). The activity started nearwhen theweather had been dry for many days (August 1994
and June 1995), relatively fewer ranodons were foundon land, walking around at night or hiding understones, than when the weather was wet (June 1993).

Environmental adaptations
The nocturnal/crepuscular activity pattern in amphibi-

intensity (see text for further explanation).

ans is probably ultimately an adaptation to the nightWe

had no opportunity

tonewlyhatched larvae. In 1995, in the most favourable brookinvestigate(Brook 1), the first larvae hatched during the secondand day temperature and air humidity cycles. The vulmakes it
nerability of the animals to dry conditions
hotnecessary for them to hide during the driest andhalf of June. According to Paraskiv (1953), newlyhatched larvae remain in calm, shallow water welltest hours, and even when, like the ranodons, they stayilluminated by the sun, where they hide among stones

thus primarily nocturnal, although their activity pattern may vary ontogenetically (e.g. Noble 1954;

on the bottom. However, they often come out and lie
on the stones, and are active by day, not avoiding thesunshine (cf. Kubykin 1986). We found that at leastof this diurnal activity lasted throughout the sumpartmer - although the main activity was nocturnal in- and even to some extent in the nextyear
Augustin water, this pattern is maintained. Most urodeles are

McDiarmid

1994).

The activity is proximately con-trolled by the light/dark cycle. Different temperaturemuch as 10-

regimes, even a rise in temperature of as
15 °C, seemed to have little, if any, influence on theactivity pattern of Ranodon sibiricus.

Relative humid-

A well-defined phase shift from diurnal to noc-

least the moisture on the ground, could beity, or atturnal activity, as described, for example, for Triturusimportant for whether or not amphibians
land, however. All records of ranodons walking on(June).

cristatus larvae (see Dolmen 1983a,b), probably does

not take place. Paraskiv (1953) mentions that older
larvae, at the time of their metamorphosis when theirgills arewill walk on

made when the air humidity was above
80% RH, always between 21.00 and 03.00 hrs.
land were

becoming resorbed and they are going onto

land, avoid the sunshine and keep to shady places.

Water phase and land life
The metamorphosed ranodons stay in water or onland, or alternate more or less regularly between these
two media. In daytime, but also sometimes at night,

we foundterrestrial

individuals

in

damp

hollows

under stones and the like within a few metres of thewater. We made no attempt to specifically measure theof ranodons in
activity of such animals. The activity
the brooks increased at night, however, and some also

went up on to land. We are therefore dealing with two
anicategories of ranodons on land: true terrestrial
land
on
which
and
skin
a
which
have
stay
mals,
dry

Another factor which could represent a threat to

Ranodon sibiricus, living at high elevations (ca. 17002700 m), is the strong UV radiation. Except for a very
few larvae of the youngest stages (0+ in August and1+ in June), no animals were seen voluntarily exposFor cold-stenothering themselves directly to the sun.
mic animals like R. sibiricus, it is probably even more
for most

important to be noctumally active thanother amphibians. High temperatures, i.e. more thanitis

22 °C, are reported to be lethal, at least to ranodon larvae (Brushko and Narbaeva 1988). Temperatures wereas low as 6.0-11.5 °C in the air and 4.8-1 1.0 °C in thewater during the peak aquatic and land-walking activityof ranodons at night. Maximum air temperaturesduring the day were 17.1-21 °C, and water tempera-

Asiatic Herpetological Research

1999

17

15

13

Hrs. 11

19

21

%

23

01

_i

Brook

40 H

i

03

05

_i

i

07 09

09

11

Brook

—
N=166
— 0+ larvae N=168
ad., |uv.

30
20 H-- 1+ larvae

17

15

19

23

21

01

03

05

07 09

i_

15-18 Aug. 1994

1

13

Vol. 8, p. 35

18-21 June 1995

1ad., juv.

N=75

1+ larvae

N=52

N=28

10

10

Brook 520

N=425ad., juv.

ad., juv.

N=279

1+ larvae N=409

0+ larvae N=151

30
40

50
Figure 6. Relative degree of activity (%) of Ranodon sibiricus adults/juveniles and larvae in Brooks 1 and 5 (cf. Figs.
1-2). Young larvae (0+, and in part 1+) exhibit a markedly greater degree of diurnality than older life stages.
tures

reached19.3-20.0

°C.

A

few

larvaewerecounted during the warmest periods, too.

night when light intensity is below 5 lux, which again
isrelatively short at midsummer. The need for protec-

We made no specific study of the possible influ-

tion against climatic factors, as described above, andence of moonlight on the activity of ranodons. However,possibly from (diurnal) predators thus seems to begreater than the need for extra growth in metamor-

notes on the weather, etc,

phosed ranodons.

when we compared our data on activity with
we found no trend of, forinstance, decreasing activity when the half moon (atthe most) was sometimes shining. The light regime
under such circumstances was also always well below

the 5-lux limit.

Ranodon sibiricus is clearly more nocturnal than,
Dolmenfor instance, the Triturus species studied by(

1983a,b).

It

can also be seen that the activity period

However, a marked difference could be seen in theof metamorphosed ranodons and larvae,

diel activity

especially0+ larvae aged

1-2

months (Fig. 6). The

youngest stages seem to be more or less active bothday and night. Still younger larvae than those we havestudied are possibly even more diurnally active (cf.
Paraskiv 1953).
probable explanation of the pro-A

which is in conformity with the length of the night. A

longed activity of the larvae, compared to older
stages, is their need to eat and grow as much as possi-

similar phenomenon was shown for the more crepus-

ble during the short frost-free season in thecular Triturus vulgaris andBeing large may perhaps increase their chancesof surviving their first, most hazardous, winter. Grow-

of R. sibiricusinJune was shorter thaninAugust,cristatus in

Norway(Dolmen 1983a,b). Moreover, although normally twopeaked, in northern latitudes at midsummer the crepT.

ing is therefore probably their most important task.

Alarger size also makes their migration up or down theuscular peaks of the Triturus species fused, and theactivity curve thus revealed only one, long midnight

brook to hibernation sites inpeak when real

protects them from predatory fish (see below).

nights (<5 lux) disappeared. Espe-moun-tains.

cially T. vulgaris, being the more crepuscular species,

By

stayingduringthelateautumn easier andsummerinthelower,

warmer part of small brooks, in shallow water, andtakes advantage of these long periods of twilight innorthern latitudes in that its hunting day thereby

alternating

becomes longer and its growth better.

1953: Dolmen et al. 1997), the metabolism of the lar-

The activity period of the strictly nocturnal Ranodon sibiricus, however, becomes shorter at midsum-

vae is raised and kept at a near optimal level. Theremer.

advantageous for an animal to have theopportunity to seek food for as many hours as possiIt

isble; theranodons in fact spends the whole night. Its

activityisare,between shade and sunshine (Paraskiv

however, certain limits for a cold-stenothermic

animal like Ranodon sibiricus. Small larvae are sensitive towarm weather, and 22 °C is said to be lethal(Brushko and Narbaeva 1988).

light

Such a difference in diel activity and microhabitat

regime through the season, i.e. the period per day andbetween adult urodeles and their larvae is not uncom-

limited,however, by the varyingVol. 8, p. 36

Asiatic Herpetological Researchmon (McDiarmid

1999

Dolmen (1983a,b; 1988)

Ardea cinerea are said to be natural predators of

as a means to reduce intraspecific com-

1994).

between different developmental stages andranodons (Brushko and Narbaeva 1988), likewise
black storks Ciconia nigra (Shnitnikov 1913). We

as protection against cannibalism, which may be com-think that some of the many species of birds of preymon in salamanders and which is also known to occur
in Ranodon sibiricus (Kubykin 1986: Wang 1990:

which occur in great numbers in the mountains (e.g.

Kuzmin 1991a). In species whose adults and larvae
live sympatrically, a behavioural mechanism whichodons if they had the opportunity. However, of theseinterpreted
petitionitbuzzards Buteo spp.) will also occasionally take ran-favours the avoidance of cannibalism could presum-

only the wild boar is nocturnal.
Adults and large juvenile ranodons therefore seem toably be of adaptive value.

have few, if any, really important predators. However,terrestrial predators,

two kinds ofHunting strategy

fish(Diptychus maculatus

and D.

dybowskii (Cypriniformes)) are present in the River

During our preliminary studies

in

June

1993, themethod we used for trapping ranodons in water failed
and we concluded that most translocations of Ran-

Borokhudzir, at least in the main river. Fish, which areusually nocturnal or crepuscular,

may therefore bepotential predators of ranodon larvae. However, sinceodon sibiricus, at least at that particular time of year,
probably took place on land. However, only a small

these fish are absent, or extremely rare, in the small,

proportion of our many records of active ranodons inare hardly a threat to the ranodons there.

spring-fed brooks in which the ranodons breeds, they

1994 and 1995 were made on land. We therefore conclude that most of their activity, like hunting (notlonger translocations), is nevertheless aquatic. Hence,

The influence of cattle
The negative influence of cattle on Ranodon sibiricus

maintained by Paraskiv ( 1953), too; they do not movementioned by several authors (Kubykin
Narbaeva
and Brushko 1986; Brushko and Nar1986;

about much in the water. This contrasts with the Euro-baeva 1988; Wang 1990; Dolmen et al. 1997). Dam-ranodons are probably largely sit-and-wait hunters, as

Dolmenhas beenmechanical when the

1983a,b), whichlive in the stagnant water of ponds and small lakes.

age

For a running-water species like R. sibiricus, it may

excrements may also pose a threat to eggs and small

be more economic to sit and wait for drifting preythan to spend energy looking for them.

larvae.

pean Triturus species (seeOur preliminary results on food items showed that
Ranodon sibiricus is an opportunist when it comes toiscattletrampleinthebrooks, crushing ranodons and destroying eggs. Their

In 1993, up to about 150 cows, watched by shepherds on horseback, twice daily crossed the brooks ofthe study area. There were also flocks of horses, sheep

choice of prey, and thus supports the results of stom-and goats. Corresponding numbers of domestic ani-

ach analyses made by Kuzmin (1991b). There wasalso a tendency for ranodon stomachs to be at theirmals were seen in 1994 and 1995. Most crossing offullest after midnight.

Based on the undigested stom-brooks took place in daylight at about 8 hrs. and 20
hrs., and at thattime the adult and juvenile ranodons

ach contents of a few specimens caught in daytime,

were still relatively safe in their hiding places. Never-

Shnitnikov (1913) suggested that the ranodons feeds

theless,by day. However, apparently no ranodons caught by
him at night were examined.

been crushed under the feet of cows. On other occa-

Diel activity and predators

During the day, metamorphosed ranodons cleverly
hide under boulders and in earth cavities in the brooksions,

we twice found dead ranodons which had

we came across mechanically damaged eggs,

presumably the victims of wading cattle. Larvae, too,
may suffer from cattle. However, we believe that thedamage to metamorphosed ranodons is not reallylarge (cf. Kubykin 1986).

bed and are often difficult to catch. Because of theirstrict nocturnality

and their ability to hide in daytime,

Conclusions

night searches with good torches are the best way ofRanodon sibiricus is a nocturnal species in the juve-

looking for ranodons in a brook. Searches in daytime

nile

may be strenuous, and often negative if the population

larvae is more evenly distributed through the day anddensity is not large. At night, however, the animals arenight.

easy to see in small and medium-sized brooks.

and metamorphosed stages probably reduces intraspe-

can be expected that the activity cycles of apredator and its prey will coincide. Wild boars Sus

cific

It

scrofa and grey herons

and adult stages, whereas the activity of young

The difference in diel activity between larvae

competition and cannibalism. In daytime, theranodons hide cleverly in the brook bed or on land.

This secretive life makes them difficult to find, both

Asiatic Herpetological Research

1999

for diurnal predatorsand people. At night, however,Vol. 8. p. 37

salamander). Pp.

187-191. In E.V. Gvozdev(ed.),

the ranodons are easy to see. Their nocturnality also

Redkie Zhivotnye Kazakhstana. Nauka Publ., Alma-makes adult and juvenile ranodons less vulnerable toAta (Almaty).

mechanical damage from grazing cattle, but eggs andKubykin, R.A.; D. Dolmen; J.V. Arnekleiv and D.
Mikutavicius 1995. Influence of cattle-breeding onlarvae may easily be harmed.

density of Ranodon sibiricus (Hynobiidae) in spawn-

Acknowledgments

ing places (Dzhungarski Alatau Mts.).

Marina V. Basargina was our excellent interpreter andgood helper during all three field trips. Richard Binns
has kindly improved the language of the manuscript.

P.

35. In N.

Ananeva (ed.). Second International Asian Herpetological Meeting, 6-10 Sept. 1995, Ashgabat. [Abstr.]

Kuzmin, S.L. 1991a. A review of studies on amphibian and reptilian feeding ecology in the USSR. Herpe-

tozoa 4:99- 115.

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