International Society for the Study

and Conservation of Amphibians

(International Society of Batrachology)

SEAT

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Source : MNHN, Paris

AIDVTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

Morphological comparisons

of brown frogs (genus Rana)

from Sakhalin, Hokkaïdo and Primorsk

Masafumi MATsUI*, Anatolii M. BASSARUKIN**, Kiyoshi KASUGAI*,

Shingo TANABE*** & Sen TAKENAKA****

* Graduate School of Human and Environmental Studies, Kyoto University, Sakyo-ku, Kyoto 606, Japan

** Sakhalin Branch of Institute of Biology and Pedology, The Far Eastern Scientific Center,

Russian Academy of Sciences, Yuzhno-Sakhalinsk, Russia

*** Toji-in Minamicho 5-40, Kita-ku, Kyoto, 603, Japan

*#** Department of Natural Sciences, Hokkaido Tokai University, Minamisawa,

Minami-ku, Sapporo, 005, Japan

One species of brown frog from Sakhalin, previously assigned to Rana

temporaria, R. chensinensis or R. semiplicata, is very similar to R. pirica from

Hokkaido in body coloration and skin texture. However, results of morpho-

metric analyses indicate its intermediate nature between R. pirica and

R. dybowskii from Primorsk. Clarification of the taxonomic status of this frog

needs further study from different approaches.

INTRODUCTION

Two species of brown frogs (Rana temporaria group of BOULENGER, 1920) occur on

Sakhalin island (BANNIKOV et al., 1977). One is unambiguously identified as Rana

amurensis (BANNIKOV et al., 1977; KUZMIN et al., 1988), but the taxonomic status of the

other is at present uncertain. It is usually regarded as Rana chensinensis (ORLOVA et al.,

1977; KUZMIN et al., 1988) or sometimes as Rana semiplicata (BANNIKOV et al., 1977).

However, recent taxonomic studies revealed that the Asian brown frogs that have long

been collectively treated as Rana chensinensis actually include several distinct taxa (for

review, see MATSUI, 1991 and MaATsuI et al., 1993). As a result, the population of “Rana

Source : MNHN, Paris

2 ALYTES 12 (1)

chensinensis” from Hokkaido (KAWAMURA, 1962) was described as Rana pirica (MATSUI,

1991).

In describing Rana pirica, the senior author (MaTsUI, 1991) restricted the range of this

species to Hokkaido, but the close taxonomic relationship of the frogs on Sakhalin and

Hokkaido has long been accepted (OKADA, 1930, 1931, 1966; NAKAMURA & UÉNO, 1963,

1965), and their taxonomic relationships await clarification. As a part of our studies to

determine the taxonomic status of the brown frog from Sakhalin, we compared this

population morphometrically with Rana pirica from Hokkaido and R. dybowskii from

Primorsk.

MATERIALS AND METHODS

We examined a total of 71 preserved specimens of Rana pirica from Hokkaido (40

males and 31 females), 37 specimens of Rana sp. from Sakhalin (31 males and 6 females;

fig. 1) and 20 specimens of Rana dybowskii from Primorsk (15 males and 5 females)

(Appendix I). Seventeen body measurements were taken (Table I), mostly following

MaTsUI (1984). Briefly, they are: (1) snout-vent length (SVL); (2) head length (HL);

(3) head width (HW); (4) snout length (SL); (5) snout height (SH = height of snout,

measured at the level of the anterior corner of the eye, including the lower jaw);

(6) intercanthal distance (ICD); (7) eye length (EL); (8) tympanum diameter (TD =

greatest diameter); (9) internarial distance (IND); (10) interorbital distance (10D);

(11) upper eyelid width (UEW); (12) lower arm length (LAL); (13) hindlimb length (HLL);

(4) thigh length (THIGH), (15) tibia length (TL); (16) foot length (FL); (17) inner

metatarsal tubercle length (IMTL). All measurements were made to the nearest 0.1 mm

with dial calipers. For morphometric comparisons, only sexually mature adults were used,

and males and females were analyzed separately.

In order to examine interspecific and intersexual variation in each character

dimension, analysis-of-covariance (ANCOVA) procedure was performed using SVL as the

covariate. AÏl measurements were log-transformed in this procedure. Each character

dimension was then converted to a percentage ratio to SVL for further comparisons. For

these ratio variables and frequency data for degree of toe webbing and position of

tibio-tarsal articulation, Kruskal-Wallis tests with nonparametric multiple comparisons,

Wilcoxon rank sum tests, or Mann-Whitney’s U-tests were performed to detect the

presence or absence of differences in the frequency distributions. The significance level was

set at 0.05.

Programs designed and implemented by the S.A.S. (1989) package were used in the

multivariate analyses. The analyses were performed through the facilities of the Data

Processing Center, Kyoto University. À canonical discriminant analysis, performed by the

CANDISC procedure was conducted and canonical variate scores of individuals were

plotted on their respective axes. Further, a discriminant analysis (DISCRIM) was used to

assess the degree with which each individual specimen was correctly classified into its home

group.

Source : MNHN, Paris

Table I. - SVL (# + 1 SE, followed b

followed by ranges in parentheses

} ranges in parentheses, in mm) and percentage ratios of each character dimension to SVL (medians.

n brown frogs. See text for abbreviations.

Males Females

R. pirica Rana sp. from R. dybowskit R. pirica Rana sp. from R. dybowskit

( = 40) Sakhalin (n = 31) = 15) @ = 31) Sakhalin (n = 6) @=5)

SVL 53.6 + 0.7 54.6 + 0.8 56.1 +1.2 62.6 + 0.9 62.8 + 2.2 68.5 + 1.9

(44.1:62.9) (39.7-60.8) (50.9-67.4) (50.7-74.2) (56.3-71.4) (64.6-74.2)

HL/SVL 33.4 33.4 333 32.6 31.5 31.7

GL1-35.7) GL1-35.8) G1:5-35.7) (30:0-37.1) (5.6-33.2) (30.0-34.2)

HW/SVL 32.1 337 34.1 32.3 333 32.6

(29.9-35.7) (31.7-35.6) (29.8-36.8) (29.4-35.8) (28.2-33.8) (30.6-33.4)

SLISVL 133 13.5 13.7 12.4 12.8 133

(11.5-14.4) (11.9-14.9) (12.9-15.3) (10.7-13.3) (12.4-13.4) (12.6-14.7)

SH/SVL 9.8 9.6 9.5 103 10.1 9.8

(8.5-11.2) (8.3-11.0) (8.7-11.5) (8.7-11.8) (8.7-10.8) (9.3-10.2) Kad

ICD/SVL 12.8 12.2 11.9 11.8 12.0 11.4 >

(1.1-14.0) (0.7-13.8) (10.4-13.1) (8.9-13.6) (11.0-12.4) (10.9-12.9) 3

EL/SVL 9.9 10.0 9.9 8.5 9.4 é

(8.2-11.7) (8.6-10.8) (9.4-11.3) (7.7-9.7) (8.3-9.8) &

TD/SVL 6.5 7.0 6.4 +

Dé (5.58.7) (6.1:8.3) (636.7 es

IND/SVL 7.5 7.0 72 d

RS (6.3:8.6) (6.3:7.5) (677.8)

SV 7.0 68 6.2

Dr (6.4:8.1) (6.27.5) (6.2:6.7)

UEW/SV 7.6 7.5 8.0

b (6.6:9.4) (6.7:8.3) (7.6:8.8)

ISVL 48.6 48.9 47.0 463 45.1 42.1

LES (44.1-52.7) (46.7-53.1) (44.0-50.8) (41.9-49.8) (43.8-46.1) (40.6-44.7)

HLL/SVL 1723 172.6 179.1 157.9 161.0 169.6

(61.9-181.9) (160.0-186.2) (165.2-191.3) (141.0-170.2) (155.0-163.1) (159.9-172.4)

ISVL 48.5 48.9 50.2 46.7 46.2 47.1

ques (45.5-51.9) (45.9-52.1) (45:3-54.9) (42.0-50.0) (45.4-46.8) (45.7-49.9)

TL/SVL 50.9 51.5 53.7 47.9 47.6 518

(47.8-54.6) (48.7-54.4) (49.8-59.1) (42.0-52.2) (46.9-48.8) (48.9-54.3)

FL/SVL 57.0 583 60.5 51.6 53.1 54.1

(51:5-61.2) (53.7-63.2) (56.3-67.0) (45.9-55.9) (51.7-55.6) (52.6-57.0)

IMTL 5.6 6.0 5.8 5.1 5.8 5.8

ÊYE (4.7:7.0) (5.0:7.1) G3:6.7) (3.9:5.9) (5.662) (6.3:6.0

Source : MNHN, Paris

4 ALYTES 12 (1)

Table IL. - Variation in the point reached by the tibio-tarsal joint when the hindlimb is

bent forwards along the body in brown frogs. Figures indicate the number of

specimens (percentage frequency in parentheses).

Position of joint

Anterior edge Posterior Posterior

Species Tip of snout of eye to anterior edge| edge of eye

and sex or forward to snout of eye or behind

Males

R. pirica 0 8 (200%) | 32 (80.0 %) 0

Sakhalin 1 (32%) 6 (194%) | 24 (77.4 %) 0

R dybowskii 0 11 (733%) | 4 (267%) 0

Females

R pirica Q 0 22 (710%) | 9 (29.0%)

Sakhalin 0 0 4 (100 %) Q

R dybowskii 0 4 (80.0%) 1 (20.0 %) 0

RESULTS

GENERAL CHARACTERISTICS OF RANA SP. FROM SAKHALIN

Morphometric data for Rana sp. from Sakhalin are summarized in Table 1 together

with those for the other two species. Females of Rana sp. from Sakhalin are significantly

larger in SVL (% + 1 SE = 62.8 + 2.2 mm) than males (X + 1 SE = 54.6 + 0.8 mm;

t-test, p < 0.001). Lengths relative to SVL of EL, HL, SL, TD, IND, LAL, HLL,

THIGH, TL, and FL were significantly greater in males (Mann-Whitney’s U-test,

two-tailed, p < 0.05). Nearly half of these characters are those of hindlimbs. Thus, when

the hindlimb was bent forward along the body, the tibio-tarsal joint reached the point

beyond the anterior edge of eye in about one-fourth of the males, but the joint at most

reached the anterior edge of eye in females (Table IT), though a significant statistical

difference was not detected between the sexes (Wilcoxon rank sum test, two-tailed, p >

0.05).

Also, males of Rana sp. from Sakhalin tended to have slightly more developed toe

webbing than females (Table III), but, probably due to the small sample size of females,

significant sexual difference was detected only in the inner web of the second toe (Wilcoxon

rank sum test, two-tailed, p < 0.05).

GENERAL COMPARISONS OF RANA SP. FROM SAKHALIN WITH THE OTHER SPECIES

In general appearance, the Rana sp. from Sakhalin (fig. 1) more resembled Rana pirica

than Rana dybowski. Rana sp. from Sakhalin shared with R. pirica the following

characteristics that were different from R. dybowskii: thick dorsolateral fold; occasionally

Source : MNHN, Paris

MaATsUI et al.

Table II. - Variation in the degree of toe webbing in brown frogs, expressed by the

phalanges free of broad web. Figures indicate the number of specimens. in:

inner web; out: outer web. Bold characters indicate the position of median.

Rana sp. from

: R. pirica R.dybowskii

Position Sakhalin

Phalanx

of toe

l'out 0.5 4 0 Il 0 2 0

[l 27 6 25 28 13 4

1.5 0 0 4 3 0 0

2in il 0 0 1 0 0 0

15 20 1 10 1 S 2

2 11 5 26 29 10 2

2.5 0 0 3 1 0 0

2 out 0.5 16 1 16 6 12 3

1 15 5 22 25 3 l

15 0 0 2 0 0

3in 1.5 2 0 0 0 0 0

2 27 5 19 15 13 2

2.5 2 1 21 16 2 2

3 out 0.5 13 il 4 0 2 0

[l 18 5 35 29 13 3

1.5 0 0 l 2 0 1

4in 15 1 0 0 0 0 0

2 21 3 { 4 11 1

25 8 3 31 25 4 3

3; l 0 2 1 0 0

4 out Il 3, 0 2 0 ll 0

) 18 3 5 2 9 1

2 10 s} 32 22 5 3

2.5 0 0 l 6 0 0

Sin 0 2 0 0 0 0 0

0.5 23 4 22 12 12 4

1 5 2 18 17 3 0

1.5 0 0 0 1 0 0

Source : MNHN, Paris

6 ALYTES 12 (1)

Fig. 1. — Dorsal view of a male (left: KUHE 11623, SVL = 58.1 mm) and a female (right: KUHE

11649, SVL = 71.4 mm) of Rana sp. from Sakhalin, Russia.

presence of partial, light stripe on middorsal line; usually presence of heavy dark markings

covering the abdomen; dorsal skin with a few tubercles; ventral skin rugose; usually

presence of a few dark spots on flanks.

In R. dybowskii, in contrast, the dorsolateral fold is usually thin; the light stripe on

middorsal line is rarely present; dark markings on abdomen are absent or limited, even if

present; the dorsal skin is nearly smooth; the ventral skin is smooth; dark spots on flanks

are usually absent.

The degree of development of the outer metatarsal tubercle varied within each species,

but the tubercle was less frequently present in Rana sp. from Sakhalin than in R. pirica or

R. dybowskii. In Rana sp. from Sakhalin, the outer metatarsal tubercle was present only

in 25 % of the specimens examined, whereas in both R. pirica and R. dybowskii, about

70 % of the specimens possessed the tubercle.

In males, Rana sp. from Sakhalin did not differ from R. pirica, but both of them are

significantly different (nonparametric multiple comparison test, Qi > Q5os, 3 — 2:39)

from R. dybowskii in the position of the tibio-tarsal joint (Table II). Females showed a

similar tendency, but the difference was not significant. On the contrary, Rana sp. from

Sakhalin more closely resembled R. dybowskii than R. pirica in the degree of toe webbing

(Table II). In males, Rana sp. from Sakhalin had more developed webbing than R. pirica

in the inner webs of the second, third, and fourth toes and outer webs of the third and

Source : MNHN, Paris

MaTsUI et al. 7

fourth toes, but from R. dybowskii it differed only in the inner web of the second toe. In

females, significant difference was found only in the outer web of the fourth toe between

Rana sp. from Sakhalin and R. pirica.

UNIVARIATE COMPARISONS

Results of ANCOVA showed that all characters, except for IOD in Rana sp. from

Sakhalin and IMTL in R. dybowskii, had significant correlations with SVL in males. On

the contrary, many characters (HL, SH, EL, IND, IOD, UEW, LAL, HLL and FL in

Rana sp. from Sakhalin, HL, HW, SL, ICD, EL, TD, IND, IOD, LAL, HLL, THIGH,

TL, FL and IMTL in R. dybowskü, and UEW in R. pirica) showed insignificant

correlations with SVL in females, probably due to the small sample size and small

variation range in each sample. Significant deviation of the slope (x) from 1 (isomorphic

relationship: MATsUI, 1984) in the regression line was observed in some cases. In males,

SL, IND, LAL, THIGH and TL (fig. 2) in Rana sp. from Sakhalin, HL, HW, EL, IOD

and LAL in R. pirica, and HW and LAL in R. dybowskii, were all bradymorphic (x < 1)

to SVL. On the other hand, HL, SL, SH, EL, UEW, HLL, THIGH and TL in R. pirica

and SH in R. dybowskii were bradymorphic to SVL, but UEW in R. dybowskii was

tachymorphic (x > 1) to SVL in females. All the other regression lines significantly

correlated to SVL were insignificantly different from 1 in coefficient of the slope.

Among the three species, differences in the slope of regression lines were found only

in EL and THIGH in males. In EL, the slope in R. dybowskii was steeper than that of

Rana sp. from Sakhalin, which in turn was steeper than the slope of R. pirica. On the other

hand, the slope of THIGH in R. pirica was steeper than that of R. dybowskii, which in turn

was steeper than that of Rana sp. from Sakhalin. By contrast, much more numerous

differences were found among the three species in the position of the regression lines. In

males, HW, SL, ICD, IND, IOD, UEW, HLL, THIGH, TL, FL and IMTL were found

to differ in some combinations among the three species. Similarly, the position of the

regression lines for HL, SL, EL, UEW, HLL, TL and IMTL differed in females, but, as

noted above, these characters insignificantly correlated to SVL in one or two of the three

species.

In this way, the results of ANCOVA were rather complex, and particularly in females,

neither significant correlation of regression lines nor deviation from 1 in regression lines

were observed in many cases. Therefore, the relative sizes of morphometric characters were

simply expressed by percentage ratios to SVL. As a result, many characters showed

significant differences (Qu > Qoos, 3 — 2-39) among the three species.

Rana sp. from Sakhalin differed from R. pirica in seven of 16 characters compared:

two characters in both sexes, four in males, and one in females. In both sexes, Rana sp.

from Sakhalin had larger relative values in UEW (males: Q = 4.04; females: Q = 2.95)

and IMTL (males: Q = 2.93; females: Q = 3.35) than R. pirica. Males of Rana sp. from

Sakhalin had larger relative HW (Q = 4.08) and FL (Q = 2.99), and smaller ICD (Q =

2.49) and IOD (Q = 3.89) than R. pirica. Females did not differ in these characters, but

Rana sp. from Sakhalin had larger EL (Q = 2.45) than R. pirica.

Source : MNHN, Paris

8 ALYTES 12 (1)

TL (mm)

37.5 40 50 60 70

SVL (mm)

Fig. 2. — Allomorphic relationships between SVL and tibia length (TL) in males of Rana sp. from

Sakhalin (closed circles: regression equation, log TL — 0.850 log SVL —0.027, r = 0.931), R.

pirica (open circles: log TL = 1.010 log SVL — 0.310, r = 0.927) and R. dybowskit (open

triangles: log TL = 0.768 log SVL + 0.135, r = 0.894).

In five (one character in both sexes and four in males) out of 16 characters compared,

Rana sp. from Sakhalin differed from R. dybowskii. Both sexes of Rana sp. from Sakhalin

had smaller value in TL (males: Q = 3.08; females: Q = 2.51). Also, males of Rana sp.

from Sakhalin had larger IND (Q = 4.23) and UEW (Q = 3.36), and smaller HLL (Q

= 2.43) and THIGH (Q = 2.66) than male R. dybowskii (fig. 3), but females did not differ

in these characters.

The difference between R. pirica and R. dybowskii was the greatest and they differed

in 11 out of 16 characters compared: four characters in both sexes, five in males, and two

in females. In both sexes, R. pirica was smaller in TL (males: Q = 4.25; females: Q =

2.94), HLL (males: Q = 3.27; females: Q = 3.01), FL (males: Q = 4.28; females: Q =

2.51) and SL (males: Q = 2.45; females: Q = 2.74) than R. dybowskii. In males, R. pirica

was larger in IND (males: Q = 3.92) and ICD (Q = 3.31), but was smaller in THIGH

(Q = 2.44), HW (Q = 4.11), and TD (Q = 2.93) than R. dybowskii, while in females, R.

pirica was larger in LAL (Q = 3.11) and smaller in IMTL (Q = 2.90) than R. dybowskii.

Source : MNHN, Paris

MATSUI et al. 9

%TL/SVL

6 6.5 7 7.5 8 8.5 9

%IND/SVL

Fig. 3. — Relationships between relative tibia length to relative internarial distance in male samples

of Rana sp. from Sakhalin (closed circles), R. pirica (open circles) and_R. dybowskii (open

triangles).

Thus, Rana sp. from Sakhalin is characterized by having a relatively wide upper eyelid

and large inner metatarsal tubercle compared with R. pirica, and relatively short tibia

compared with R. dybowskii. In general, these results of univariate analyses, as in those

obtained from multivariate analyses, indicated an almost equally great morphological

differentiation of Rana sp. from Sakhalin from both R. pirica and R. dybowskii, but the

degree of differentiation is small compared with the greater difference between the latter

two species.

MULTIVARIATE COMPARISONS

Results from the CANDISC are presented in fig. 4. In both sexes, all F values from

the four criteria, e. g., Wilks’ Lambda, Pillaï’s Trace, Hotelling-Lawley Trace, and Roy’s

Greatest Root Statistics, were significant at p < 0.0001, indicating significant geographical

effect. In males, the eigenvalue of the first and second axes accounted for 3.453 and 1.379,

respectively, and the first canonical variable accounted for 71.5 % of the total amount of

Source : MNHN, Paris

10 ALYTES 12 (1)

-4 -3 -2 -1 0 1 2 3 4 5 6 7 8 9

CAN1

Fig. 4. — Plot of first against second canonical variates for samples of Rana sp. from Sakhalin (closed

circles), R. pirica (open circles) and R. dybowskit (open triangles). Top, males; bottom, females.

Source : MNHN, Paris

MATSUI et al. 11

Table IV. - Classification matrix based on linear discriminant function analysis in

brown frogs. Entries along the main diagonal are "correct" matches of

individuals to their source population.

Predicted group membership

Rana sp. from

Actual group Sakhalin R. pirica R.dybowskii

Males

Sakhalin 26 (83.9 %) 2 (6.5 %)

R. pirica 3 (7.5 %) 36 (90.0 %)

R. dybowskii 2 (15.4 %) 1(7.7 %)

Females

Sakhalin 4 (80.0 %) 0 (0 %)

R. pirica 2 (6.5 %) 28 (90.3 %)

R. dybowskii 1 (20.0 %) 1 (20.0 %)

eigenvalues, whereas in females, the eigenvalue was 7.629 and 1.461, respectively, and the

first variable accounted for 83.9 % of the total amount.

The absolute magnitudes of the standardized canonical discriminant coefficients are

proportional to the relative importance of each character in separating the three species

along each of the two canonical discriminant axes. In males, FL was the greatest

contributor to separation on the first axis (2.766), followed by TL (2.764) and HLL

(-2.651). On the second axis, LAL (1.220), HLL (-1.060) and IND (0.888) were the high

contributors. Similarly, variables with high contribution on the first and second axes in

females were LAL (-2.068), HLL (1.228) and TL (0.996), and TL (1.969), HL (-1.566) and

HW (-0.930), respectively.

In males, Rana sp. from Sakhalin was nearly completely distinguishable from R.

pirica, but completely overlapped R. dybowskii on the first canonical axis, while this

situation reversed on the second canonical axis. Thus, on the first two axes, Rana sp. from

Sakhalin was nearly completely distinguishable from R. pirica, and only slightly

overlapped R. dybowskii. In this way, the intermediate position of Rana sp. from Sakhalin

between R. pirica and R. dybowskii was clear in males. Separation was better in females,

and the three species did not overlap on these axes.

Reflecting the similarities of the three species in gross appearance, the DISCRIM

succeeded in assigning only 85.7 % (72 of 84) in males and 85.4 % (35 of 41) in females

of the individual specimens to correct home population (Table IV). In males, the

DISCRIM correctly classified 83.9 % (26 of 31) of Rana sp. from Sakhalin, with 9.7 % (3

of 31) of other specimens classified as members of R. dybowskii, and 6.5 % (2 of 31) as R.

pirica. On the other hand, in females, four of five (80.0 %) individuals of Rana sp. from

Sakhalin were correctly classified and all the remaining specimens were classified as

members of R. dybowskii with only a low posterior probability (48.1%).

Source : MNHN, Paris

12 ALYTES 12 (1)

DISCUSSION

Rana sp. from Sakhalin was first recorded from the southern part of the island in 1870

by DoBROTVORSKY and was identified as R. temporaria. Since then, the brown frog from

Sakhalin, together with that from Hokkaido (= R. pirica), have long been considered to

be conspecific with this European species (BOULENGER, 1886; STEINEGER, 1907; NIKOLSKY,

1905, 1918; OKADA & KAWANO, 1923; OKkADA, 1930, 1931; TERENT'EV & CHERNOV, 1949).

However, it is now generally accepted that R. temporaria has 2 n = 26 chromosomes

(Wirscui et al., 1958) and its range does not extend to the Asian regions (FROST, 1985).

By contrast, Rana sp. from Sakhalin is inferred to have 2 n = 24 chromosomes, like

Chinese R. chensinensis and its allies including R. pirica (SETO, 1965; GREEN, 1983; WEI et

al., 1990).

More recently, Rana sp. from Sakhalin and R. pirica have often been regarded as

populations of R. chensinensis (KAWAMURA, 1962; OKADA, 1966, as a subspecies of R.

temporaria; ORLOVA et al, 1977; FROST, 1985; KUZMIN et al., 1988), R. dybowskii

(NAKAMURA & UÉNO, 1963, as a subspecies of R. temporaria; NAKAMURA & UËNO, 1965,

as a subspecies of R. chensinensis) or R. semiplicata (BANNIKOV et al., 1977). Of these, R.

chensinensis was described from the Qinling Mountains of China (Davi, 1875), which are

geographically quite distant from Sakhalin or Hokkaido. On the other hand, R. dybowskii

was described from near Vladivostok, Russia (GÜNTHER, 1876). From this pattern of

distribution, the relationships of brown frogs from Sakhalin and Hokkaido were expected

to be closer to R. dybowskii than to R. chensinensis, and recent studies of the Hokkaido

population led to the description of R. pirica (MATSUI, 1991), leaving the taxonomic status

of Rana sp. from Sakhalin undetermined.

Taxonomic status of R. semiplicata is little understood. The species was described by

NikOLSKY (1918) from two specimens possibly from Poltavka, Maritime Territory, and it

was said to resemble R. tsushimensis and R. amurensis, both of which have 2 n = 26

chromosomes (ORLOVA et al., 1977). TERENTEV & CHERNOV (1949) claimed that

inaccuracies were contained in the original description, but from the description and

figures of a type specimen, this species is not related either to R. tsushimensis or to R.

amurensis, but seems to be synonymous with R. dybowskii (MATSUI, 1991). BANNIKOV et

al. (1977) included the coastal region of Russia and Sakhalin within the distribution range

of R. semiplicata, while ORLOVA et al. (1977) synonymized this species with R. chensinensis.

Mars (1991) suggested that “R. semiplicata” from Sakhalin (BANNIKOV et al., 1977) was

possibly conspecific with R. pirica.

Previous morphometric comparisons of R. pirica with R. dybowskii, R. ornativentris,

and topotypic R. chensinensis (MATsUI, 1991; MaTsuI et al., 1993) indicated the presence

of great differentiation between R. pirica and R. dybowski. However, the results of

morphometric analyses shown above indicate that Rana sp. from Sakhalin is not

completely identical to either R. pirica or R. dybowskü, but is morphometrically

intermediate between them. This result contradicts the taxonomic treatments made by

BANNIKOV et al. (1977) with regard the coastal and Sakhalin populations as conspecific, or

the prediction made by MATSUI (1991) to associate the Sakhalin population with R. pirica.

Source : MNHN, Paris

MaTsuI et al. 13

As noted by MaTsut et al. (1993), morphological differentiation seems generally

preceded by genetical differentiation among brown frogs of the R. temporaria group, and

the presence of even a subtle morphological difference between any two local populations

seems to suggest the possibility of their independent taxonomic status. Additional studies

from various approaches, especially biochemical ones, are necessary to elucidate the

taxonomic position of Rana sp. from Sakhalin, and such studies are now underway. On

the other hand, Rana sp. from Sakhalin is distinct from R. chensinensis in the characteristic

swelling of water in the oviduct during preservation, as are R. dybowsküi, R. pirica and a

population of R. chensinensis from northeastern China (the Hashima-frog) (MATSUI, 1991),

and these forms seem to constitute a monophyletic group.

ACKNOWLEDGMENTS

The Russian Academy of Sciences kindly permitted us to work in Sakhalin. We thank T. Saro

and S. NakaBayasu for help on the field trip, and T. HikiDA for help in computer analyses. T.

HikiDA also critically read the manuscript. Y. SHiBaTA provided literature and K. ADLER kindly

corrected verbal errors. We also thank E. BALLETTO and two anonymous referees for their valuable

suggestions. The research was partly supported by a grant from the U. S. National Geographic

Society to M. M. (No. 4505-91).

LITERATURE CITED

BANNIKOV, A. G., DAREVSKY, I. S., ISHCHENKO, V. G., RUSTAMOV, A. K. & SHCHERBAK, N. N., 1977.

— A guide-book of the amphibian and reptile fauna of the U.S.S.R. Moscow, Prosveshchenye

Publishing House: 1-414. [In Russian].

BOULENGER, G. A., 1886. — Note sur les grenouilles rousses d'Asie. Bull. Soc. zool. France, 11:

595-600.

= 1920. — A monograph of the South Asian, Papuan, Melanesian, and Australian frogs of the

genus Rana. Rec. indian Mus., 20: 1-126.

Davin, A., 1875. — Journal de mon troisième voyage d'exploration dans l'empire chinois. Paris,

Librairie Hachette: i-iv + 1-383.

Frosr, D. R. (ed.), 1985. — Amphibian species of the world: a taxonomic and geographical reference.

Lawrence, Allen Press: i-v + 1-732.

GREEN, D. M., 1983. — Evidence for chromosome number reduction and chromosomal homose-

quenciality in the 24 chromosome Korean frog Rana dybowskit and related species.

Chromosoma, 88: 222-226.

Günrer, À., 1876. — Description of a new frog from N. E. Asia. Ann. Mag. nat. Hist., (4), 17: 387.

KAWAMURA, T., 1962. — On the names of some Japanese frogs. J. Sci. Hiroshima Univ., (B 1), 20:

181-193, pl. 1-1.

KUZMIN, S. L., BORKIN, L. J., MUNHBAYR, H., VOROBYEBA, E. H., DAI KY, LS. & SEMENOV, D.

B., 1988. — Amphibians and reptiles of Mongolian People's Republic: general problems,

amphibians. Moscow, Nauka: 1-248. [In Russian].

Marsui, M., 1984. — Morphometric variation analyses and revision of the Japanese toads (genus

Bufo, Bufonidae). Contrib. biol. Lab. Kyoto Univ., 26: 209-428.

es 1991. — Original description of the brown frog from Hokkaido, Japan. Jpn. J. Herpetol., 14:

63-78.

Maïsut, M., Wu, G.-F. & SONG, M.-T., 1993. — Morphometric comparisons of Rana chensinensis

from Shaanxi with three Japanese brown frogs (genus Rana). Jpn. J. Herpetol., 15: 29-36

NAkAMURA, K. & UËNO, S.-I., 1963. — Japanese reptiles and amphibians in colour. Osaka, Hoikusha:

iix + 1-214, pl. I-XLIL. [In Japanese].

Source : MNHN, Paris

14 ALYTES 12 (1)

—— 1965. — Japanese reptiles and amphibians in colour. Second edition. Osaka, Hoikusha: i-ix +

1-214, pl. I-XLII. [In Japanese].

NikoLsky, A. M., 1905. — Herpetologica rossica. Mem. Acad. Sci. Imp. St.-Petersbourg, (8, CI.

Phys.-Math.), 17: 1-518.

1918. — Fauna of Russia and adjacent countries. Amphibians. Petrograd. [Translated from

Russian by Israel Program for Scientific Translations, Jerusalem, 1962].

OkaDA, Y., 1930. — Monograph of Japanese tailles batrachians. Tokyo, Iwanami-shoten: i-ii + i-vi

+ 1-234, pl. I-XXIV. [In Japanese].

ie 1931. — The tailless batrachians of the Japanese empire. Tokyo, Imp. Agr. Exper. Stat

1-215, pl. I-XXIV.

ne 1966. — Fauna Japonica. Anura (Amphibia). Tokyo, Tokyo Electrical Engineering College Press:

ixii + 1-234, pl. I-XXIV.

OkaDa, Y. & Kawano, U., 1923. — Notes on the distribution of Rana japonica and its allied species

in Japan. Zool. Mag., 35: 361-380, pl. VIIL. [In Japanese].

ORLovA, V. F., BAKHAREV, V. A. & BORKIN, L. J., 1977. — Karyotypes of some brown frogs of

Eurasia and a taxonomic analysis of karyotypes of the group. Proc. zool. Inst. Leningrad, 14:

81-103. [In Russian].

S.A.S., 1989. — User's guide. Version 6. Fourth edition. Vol. 1. Cary, North Carolina, S.A.S. Institute

Inc.

SETO, T., 1965. — Cytogenetic studies in lower vertebrates. II. Karyological studies of several species

of frogs (Ranidae). Cytologia, 30: 437-446.

Srenecer, L., 1907. — Herpetology of Japan and adjacent territory. Bull. U. S. nat. Mus., 58: 1-577,

pl. I-XXV.

TERENT'EV, P. V. & CHERNOV, S. A., 1949. — Key 10 amphibians and reptiles. Moscow, Nauka: 1-315.

[Translated from Russian by Israel Program for Scientific Translations, Jerusalem, 1965].

Wa, G., CHEN, F.-G. & XU, N., 1990. — An investigation for the karyotypic, C-banding and

Ag-NORS pattern on Rana chensinensis from type locality. Hereditas, (Beijing), 12: 24-26, pl.

I. [In Chinese].

WirscHi, E., KODANI, M. & MikaMo, K., 1958. — Comparative study of the chromosomes of

European, American, and Japanese frogs. Anat. Rec., 131: 610.

APPENDIX I

SPECIMENS EXAMINED

The samples are stored at the Graduate School of Human and Environmental Studies, Kyoto

University (KUHE) and in Sen TAKENAKA private collection (ST).

Rana sp. (n = 37). — Sakhalin, Russia: Uglegorsk (48°%40'N, 133°54'E): KUHE 11613-11627,

11644-11649, 11657-11670, 11680, 14138.

Rana pirica (n = 71). — Hokkaido, Japan: (1) Nükappu-cho (42°22’N, 142°19'E): KUHE 13871,

13876-13883, 13889-13891, 13893-13901; (2) Rankoshi-cho (42°52°N, 140°36'E): KUHE

13825-13828, 13833-13834, 13836, 13840-13842, 13844; (3) Kamishihoro-cho (43°2/'N,

143°12'E): KUHE 13141-13149, 13152-13154, 13156, 13158-13160; (4) Obihiro-shi (42°53'N,

143°11'E): KUHE 13745, 13747, 13750-13751, 13753-13755, 13758, 13760; (5) Tomakomai-shi

(42°40'N, 141936'E) KUHE 13845-13847, 13850-13852, 13854, 13857, 13859-13861, 13863-

13866.

Rana dybowskii (n = 20). — Primorsk, Russia: Lazo (43°21'N, 144°04'E): KUHE 11721-11732; ST

01-02, 06-07, 09, 14, 16, 18.

Corresponding editor: Emilio BALLETTO.

Source : MNHN, Paris

Alytes, 1994, 12 (1}: 15-29. 15

Site tenacity, within and between summers,

of Rana arvalis and Rana temporaria

Jon LOMAN

Department of Ecology, University of Lund,

Ecology Building, 223 62 Lund, Sweden

Adult moorfrogs Rana arvalis and common frogs Rana temporaria were

marked, released and recaptured during five summers in a 50 X 50 m large

moist meadow site. AIl capture sites were noted. The distribution of number

of captures per frog and the distance between successive capture sites were

analysed. It is concluded that most frogs that were captured at least twice in

the area occupied a permanent home range there. However, there was probably

some change of centre of activity over time. The size of these home ranges was

about 150 m° for R. arvalis and 330 m° for R. temporaria. There was no

difference between sexes, nor between large and small frogs. Some frogs

captured only once, especially R. arvalis, were probably temporary visitors.

Many frogs returned to the study area in successive years, usually to the same

part of the study area where they had spent the previous year.

INTRODUCTION

In the old days, an animal ecologist marked animals. Hopefully the animal could be

recaptured or observed and identified. A massive effort of this type yielded all sorts of

basic autecological information: estimates of individual growth rate, population densities

and individual movements. The prime example is the marking program of FIrcH (1958)

at the University of Kansas. No mammal, reptile or amphibian was allowed to remain

unmarked. I carried out a similar program for Swedish brown frogs (Rana arvalis and

Rana temporaria) on a small scale (50 X 50 m). I have previously published compilations

of growth rate (LOMAN, 1978) and population dynamics (LOMAN, 1984) emerging from this

work. The compilation of movements was not published at that time but follows here.

This study is an analysis of R. temporaria and R. arvalis marked and recaptured in

a restricted area during part of five summers. The results are analysed to provide evidence

for the nature of the frogs’ summer movements. In contrast to what had been possible if

radio transmitters were used, details in the movement patterns cannot be tracked. The

study aims at determining whether a restricted home range is used, at least for part of the

summer, and at yielding an estimate of the size of any home range used. I will also give

information of the between-year home range use of the frogs.

Source : MNHN, Paris

16 ALYTES 12 (1)

STUDY AREA AND METHODS

The study was conducted in a moist meadow habitat in southern Sweden (55°40'N,

13°30'E). The study site was a 50 X 50 m part of a meadow with a uniform vegetation

consisting of a thick layer of grasses and herbs, about 40 to 80 cm high. Scattered Salix

bushes also occurred. The site was about 200 m from the closest possible breeding sites for

the study species.

The study site was thoroughly searched during each summer from 1972 to 1976. The

study periods were July 11 to October 7 1972 (26 searches), July 2 to October 7 1973 (30),

July 8 to September 9 1974 (17), August 4 to October 6 1975 (25), and August 6 to August

30 1976 (13). Each search lasted for about one hour. I walked back and forth on fixed

paths in order not to disturb the habitat more than necessarily. These paths were about

2 m apart. All frogs seen were captured if they were considered possible adults (R. arvalis

at least 36 mm long and R. temporaria at least 46 mm long). Frogs of this size may breed

in the following spring (LOMAN, 1978). Only frogs actually measured to these sizes were

considered further. Each frog was individually marked by toeclipping and the capture site

noted to the closest 1 m coordinates. Wooden sticks were placed as a 10 m grid to facilitate

positioning.

Distances and time intervals between captures were significantly non-normal for both

species (Lilliefors test, WiLKkINsON, 1990: all cases P < 0.001). After log transformation,

the distributions for the R. temporaria data were not significantly different from normal

(Lilliefors test: times, P = 0.141; distances, P = 0.401). By this criterion, the moor frog

data differed somewhat from normality (Lilliefors test: times, P = 0.047; distances, P =

0.030). Nonetheless, the transformed data were used for both species in the significance

tests.

RESULTS

DISTRIBUTION OF CAPTURES PER FROG — ESTIMATING THE POPULATION’S SIZE

Most frogs were only captured once (fig. 1) and were thus potential transients. To

evaluate how common these were, I need an estimate of the total number of frogs at risk

of capture in the study area, pooled over all study years. I estimated the size of this

“population” (the sum of the five study years’ population sizes) by fitting a negative

binomial distribution to the data (CAUGHLEY, 1975: 154) and this yielded a surprisingly

good fit for the moor frog (Table 1). Extrapolating the fitted distribution to the zero class

(frogs available for capture but never captured) yielded values for the total population at

risk of capture. These were 1270 (3.5 times the total number captured) for R. arvalis and

598 (1.9 times the total number captured) for R. temporaria.

Source : MNHN, Paris

Rana arvalis Rana temporaria

individuals Individuals

i SS| ds — î

Number of captures Number of captures

M 1972 À 1973 1974 N 1975 À 1976

Fig. 1. — Distribution of number of captures per frog and year.

Source : MNHN, Paris

18 ALYTES 12 (1)

Table I. - Frequency distribution found and expected from a fitted negative binomial

distribution. The classes of animals captured 4 times or more are pooled for the

x-tests.

Rana arvalis Rana temporaria

Captures Observed Expected Observed Expected

1 282 286.5 215

2 60 63.0 71

3 16 13.8 20

4 2 3.0 8

5. 1 0.7 2

6 0 0.1 1

7 0 0.0 0

x 0.75

d.f. 1

P >0.10

DISTANCE BETWEEN SUCCESSIVE CAPTURES

The average distance between two captures of an individual within a year was 6.10 m

for R. arvalis and 9.45 m for R. temporaria (Table IN). These distances are significantly

different (t = 2.89, df. = 171 [pooled variances], P < 0.001). They are also both

significantly different from the average distance of 100 random distances, 26.50 m (R.

arvalis vs. random: t = 14.2, d.f. = 176 [pooled variances], P < 0.001; R. temporaria vs.

random: t = 10.7, d.f. = 193 [pooled variances], P < 0.001). These random distances were

formed as the distance between 100 pairs of rectangular distributed (in the interval 0 to

50) random coordinates.

Sex and size effects

There was no difference in the average distance between two captures, between the

sexes, in any of the species (Tables II and III). There was no effect of size, which

presumably would reflect age (LOMAN, 1978) (Table IT).

Successive captures of individual frogs

First to second versus first to third capture

If a frog was recaptured more than once, the second recapture tended to be about as

far from the original site as the first recapture (Table IV).

Time between captures

There was a slight tendency for capture sites based on captures far apart in time to

be further apart than those separated by only a short time (fig. 2). Based on all captures

Source : MNHN, Paris

LOMAN

Table IT. - Distance between successive captures of one frog. For frogs captured more

than twice, an individual mean value was used; a frog captured N times yielded

N-1 values; distance between capture 1 and 2, between 2 and 3, etc. The table

reports the average of these individual mean values.

Test for a sex difference

Rana arvalis

Females

Males

AIl

Rana temporaria

Females

Males

AIl

Random

0.55

Table III. - Four way ANOVA, testing simultaneously for effects of year, sex, frog

size, and time between capture and recapture on the distance between two

captures.

Species Source

Rana arvalis

Rana temporaria

Year

Sex

Size

Time int.

Year

Sex

Size

Time int.

pomp mme

Source : MNHN, Paris

20 ALYTES 12 (1)

Table IV. - Distance between first and second versus distance between first and third

capture of a frog. Values for frogs captured more than three times is based on a

mean value for each frog. A frog captured N times yielded N-2 values. Only

frogs captured at least three times are considered in this table. The test is a pair-

wise t-test and the probabilities given are one-tailed.

1st to 2nd site | 1st to 3rd site

N Mean | S.D. | Mean | S.D. t P

Rana arvalis 17

Rana temporaria 29

(including several values for some frogs, recaptured more than once), this effect was

significant for both species. When correcting for other factors, this increase in distance was

only found to be significant for R. arvalis (Table ID).

TIME SINCE MARKING FOR RECAPTURED FROGS

If marked frogs tended to leave the study area after some time, those frogs that were

recaptured would be a biased sample of all frogs marked. They would mainly be those that

had been marked (for the first time in that year) during the time shortly preceding the time

of attempted recapture. The time they had been marked should be less than the

“population mark age”, i.e. the average time all frogs marked that year had carried their

marks. Because successively more of those marked early would leave the study area, this

tendency should increase with time passed since marking started in that season. However,

there were no such tendencies (fig. 3).

BETWEEN-YEAR RECAPTURES

Return rate

Quite a large number of marked frogs were recaptured in the following year.

Information of growth rate obtained during the present field work and previously

published (LOMAN, 1978) was used to classify frogs as “large” or “small” adults. As large

adults were, in each year, those classified that already in the previous year were adults (i.e.

at least 36 and 46 mm respectively) and thus subject to capture and marking (if found)

already in that year. Overall, 33 % (R. arvalis) and 47 % (R. temporaria) of all “large”

adults found in one year had actually been captured and marked also in the preceding year

(Table V). This is only moderately less than the proportion expected (42 % and 67 %

respectively) if all frogs that were marked in one year and survived to the next also

returned to the study area.

Source : MNHN, Paris

MOVEMENT (M)

NVHOT

Fig. 2. — Relation of time between two captures and distance between the two sites. The correlation

is significant for both species (R. arvalis: r = 0.23, d.f. = 101, P = 0.010; R. emporaria: r =

0.16, d.f. = 137, P = 0.024). If the degree of freedom-values are reduced to the number of

different frogs involved, the values are still significant (R. arvalis: d.f. = 78, P < 0.025; R.

temporaria: d.f. = 95, P < 0.05). AII P-values are based on one-tailed tests.

A. arvalis #. temporaria

T T LS Tr 50 DT T T T

° 40 E +

‘ 2 sf . . -

Û È

= a !

. u :

| L E x 3

; à ; 2

rot ju jrs 0 sn 1 1

[e) 10 20 80 40 50 60 [e) 20 40 60 80 100

DAYS DAYS

Source : MNHN, Paris

Rel. mark age

(e]

0.0

A. arvalis A. temporaria 8

T T T T 8 T T = T T

: d u sé 2 : … =|

x : a ;

DE à ” s HET” .

‘ : CR È : Es ë

L | . 3. TE Gien. ë

FR z Ë 4 3

de - Ë È 2 + Él

CRC ' ; GE PT

+ Le £ ES . : Le . j na

te 1 —L 1 s [e) ” ar "1 1 Le

10 20 80 40 50 0 10 20 80 40 50 60

Population mark age (days) Population mark age (days)

Fig. 3. — Relation between the “relative mark age” and the average time all frogs marked in that year

had carried their marks (“population mark age”). “Relative mark age” is, for each capture, the

time since the recaptured frog was first marked, divided by the “population mark age” on the

day of that capture. There is a positive correlation for R. temporaria but it is far from significant

( = 0.045, d.f. = 105, P = 0.65). The slope for R. arvalis is negative, a direction that is excluded

from being significant a priori (ie. a one-tailed test).

Source : MNHN, Paris

LOMAN 23

Table V. - An analysis of the probability that a frog returns to the study area in

successive years. "Captures T-1" is the number of frogs captured and marked in

the study area in the year preceding T. "Population size T-1" is the estimated

number of frogs in that year. Population size was estimated with the method of

SCHUMACHER & ESCHMEYER (in SEBER, 1973: 139) (LOMAN, 1984).

"Proportion marked in T-1" is the quotient between these. "Large adults T" is

the number of frogs captured year T that were large enough to have been

catchable in year T-1 (LOMAN, 1978). "Recaptures T" is the number of frogs

captured in year T that were marked in the previous year. "Proportion

recaptures T" is the quotient between the last two figures. There were too few

captures of R. temporaria in 1973 and 1974 to warrant the computation of the

last quotient.

Population Proportion Large Proportion

Year Captures size marked adults Recaptures recaptured

T T-1 T-1 T-1 ju Ts EF:

Rana arvalis

1973 91 269 0.34 0.32

1974 48 54 0.89 0.46

1975 48 161 0.30 0.35

1976 56 96 0.58 0.26

Total 243 0.42 0.33

Rana temporaria

1973 0.83

1974 0.65

1975 0.62

1976 0.52

Total 0.67

Between-year site tenacity

The frogs that returned to the study site and were recaptured in a second year were

usually found close to the site where they had been found in the preceding year. The mean

distance between the two sites was 6.5 m (S.D. = 6.37, N = 28) for R. arvalis and 12.8

m (S.D. = 9.64, N = 12) for R. temporaria. If there were several captures in one year of

an individual, the arithmetic mean was used. These distances were not significantly

different from the average distance between two capture sites in one year (Table 11) (R.

arvalis: t = 0.30, d.f. = 104, P > 0.10; R. temporaria: t = 0.51, d.f. = 105, P > 0.10).

Source : MNHN, Paris

24 ALYTES 12 (1)

DISCUSSION

NATURE OF HOME RANGES

What was the nature of the summer movements of the frogs? The site tenacity was

striking. Distance between any two captures of a frog was significantly shorter than that

between two random points in the study area. This distance was found by simulation.

Therefore it is obvious that at least most frogs that were recaptured did have a home

range, in the sense of a “restricted area, more or less regularly used”. The most extreme

model for a summer home range would be a definite, small area, traversed throughout

almost daily. Such a model is supported by the fact that the distance between capture sites

1 and 3 was not larger than that between 1 and 2. Thus, sequences of distances between

three captures of a frog did not indicate any sort of directional movement.

However, there was a significant (but numerically rather slight, see the linear

regressions in fig. 2) tendency for captures far apart in time to also be far apart in space.

This can be interpreted as representing a gradual shift in home range location, or at least

in centre of activity within home range, over time.

If frogs tended to stay in a home range for some time, but then move much further

and at least leave the study area all together, this would “dilute”’ the marked population.

At the beginning of the study period each year, all marked (but see below) frogs would

be available for capture. The average time a recaptured frog had been marked would be

similar to the average time since marking for all marked frogs in the population. However,

later in the season, some of the frogs marked early would have left the study area and a

disproportionate fraction of those recaptured would have been frogs that settled (and were

marked for the first time) recently. The average time a recaptured frog had been marked

would at this time tend to be less than the average time since marking for all frogs in the

population. However, such a pattern was not discernible (fig. 3). This suggests that frogs

with a home range in the study area at the beginning of each study season (July-August)

tended to stay there at least until the end of this (up to September). However, if some frogs

moved very quickly through the study area, only being available for capture once, they

would not affect this pattern. They would be equally unavailable for capture at the

beginning as at the end of the capture season.

ON TRANSIENTS

Although the evidence is that most frogs captured at least twice were permanent

residents in the study area, this cannot be shown for all those captured only once. These

may have been unavailable for recapture because they were only temporarily present in the

study area. The question can be somewhat elucidated by analysing the distribution of

captures. It is appropriate to attempt fitting the distribution found to a negative binomial

(see first section of Results) as this distribution results from situations with different

capture probabilities for different individuals (CAUGHLEY, 1975). This was obviously the

Source : MNHN, Paris

LOMAN 25

case because the effort differed between different years. The fit was very good for R. arvalis,

so some confidence can be put in the expected number of individuals not captured at all

in this species, more than three times the total number captured. This capture efficiency

was surprisingly low. The individuals never captured are both those that remained resident

in the study area without being captured and those that only briefly entered it (some of

these latter could also contribute to the high number of frogs captured only once). It seems

however improbable that all these should have remained for long in the study area without

being captured. I judge a substantial number of those R. arvalis only captured once to be

transients. The evidence is less conclusive for R. temporaria because the fit was poorer and

because the estimated class of zero captures was smaller. Still, I would suggest also that

some R. temporaria were transients to the study area. Apart from possible true nomads,

these are frogs changing home range (though I have shown above that this is not a very

common behaviour), or frogs that make temporary excursions from home ranges outside

the study area. I have previously (LoMAN, 1981) shown that the presence of transient frogs

is necessary to explain the pattern of recolonisation of an area where R. temporaria were

removed. DoLE (1965a) has shown that leopard frogs (Rana pipiens) make long excursions,

outside their regular home ranges, during rainy nights. Such a phenomenon may explain

the pattern found here.

An alternative explanation to the presence of transients would be high mortality.

However, yearly mortality in the study population was previously calculated to 61 % (R.

arvalis) and 36% (R. temporaria) which appears to be normal for terrestrial ranids

(LomaN, 1984). Also, I would expect most mortality to take place during the spring and

autumn migration.

SIZE OF HOME RANGES

For both species, the data suggest that males and females as well as small and large

individuals, had home ranges of similar size. What was the actual size of the home ranges?

The small number of recaptures makes it impossible both to infer the shape of the home

ranges and the individual variation. However, a rough impression can be gained from the

following calculation. By means of simulations, the home range size that would give the

same average distance between two random points (in the simulated home range) as the

observed distance (Table II) between two capture sites can be calculated. These

calculations were based on two alternative models for the nature of the home range. With

a rectangular distribution model I calculated the diameter of the home range periphery.

This was 13.3 m for R. arvalis and 20.5 m for R. temporaria respectively. With a normal

distribution model, I calculated the diameter of a circle encompassing 95 % of the activity.

This was 15.5 m for R. arvalis and 24.0 m for R. temporaria respectively. Because we do

not know the actual distribution of activity in the home range and there are several forms

of sampling errors, the values calculated can only be considered a rough estimate of the

magnitude of the home ranges. One conclusion is that, at least for the R. temporaria which

had the largest home ranges, most frogs had home ranges that overlapped the border of

the study site. This should further tend to underestimate home range sizes. Also, it is

Source : MNHN, Paris

26 ALYTES 12 (1)

obvious from the information of density in the study area (see first section of Results and

LOMAN, 1984) that the home ranges of the studied frogs overlapped widely, both intra- and

interspecifically.

BETWEEN-YEAR MOVEMENTS

Surprisingly many frogs returned to the study area in successive years. The proportion

of all “large” adults that were already marked when captured for the first time in a year

was almost as large as the proportion of all frogs estimated to have been marked in the

study area in the previous year (see second section of Results and Table V). This

calculation suggests that 79 % (0.33/0.42) of all R. arvalis and 70 % (0.47/0.67) of all R.

temporaria returned to the study site. These figures rely heavily on the estimations of

population sizes. Because of biases and sampling errors in these estimates as well as

sampling errors in the other figures, it is only possible to state that a substantial proportion,

possibly almost all frogs alive, returned to the study site. Because of the arbitrary nature

of “return” (here defined as return to a 50 X 50 m square), a more precise answer, even

if it were possible to give, would only be of limited use. However, the tendency for many

frogs to return precisely to the same summer home range is clear; those that returned did

actually do so to the part of the study area where they spent the previous summer.

COMPARISON WITH OTHER STUDIES

It is striking that few studies of this subject seem to have been published recently. The

original list of references, compiled in 1981, contained quite a few comparable studies

(Table VI). One of the few recent studies of amphibian summer home ranges is that by

SINsCH (1988a). He employed a more sophisticated technique, trailing toads (Bufo bufo)

with special devices, leaving a thin thread after the animal. That technique has previously

also been used by DoLE (1965a), working with leopard frogs (Rana pipiens). However, it

seems that everyone now expects studies of animal movements to employ radio

transmitters. Though this certainly is justified, it emerges that little work, based on radio

transmitters, has been published on frog movements. Studies that do are work by VAN

GELDER and BUGTER (1987) on (one) R. arvalis, a report by FALLER-DOEPNER et al. (1991)

on the post-breeding migration of R. temporaria, a study of summer home ranges of Bufo

americanus (WERNER, 1991) and another study by SINSCH (1988b) on breeding behaviour

of Bufo calamita. K seems that the promise of radio transmitters has discouraged work

employing conventional capture-recapture studies. Actually, it is probably only recently

that transmitters small and long lived enough for useful work with frogs (but for the

largest species) have been available. Future work with radio transmitters will yield more

realistic descriptions of frog movements and home range use than is possible with

capture-recapture methods, the results of which must be analysed to provide indirect

evidence for the nature of the movements.

Nonetheless, most previous studies, like this one, have reported that frogs during the

nonbreeding season tend to occupy restricted areas, home ranges. Examples include: Rana

Source : MNHN, Paris

LOMAN 27

Table VI. - Published information on summer home range areas in frogs. Age

categories are adult and subadult. If only mean distance between capture is

given, this is found in the table, together with an estimate of home range area

(made with the "rectangular model" above). DOLE (1965b) and KRAMER (1974)

give direct estimates of home range area, computed with the minimum polygon

method. Sizes of adult frogs are from CONANT (1958), STEBBINS (1966) and

own data. ad: adult; sad: subadult. MD: Mean distance between captures (m).

MD | Area (m2)| N Reference

Rana pretiosa ad, sad | 50-100 12 (534) 23 | CARPENTER, 1954

Bufo boreas ad, sad | 60-125 13 (627) 19 | CARPENTER, 1954

Gastrophryne olivacea ad 20-35 33 | (4040) | 52 | FircH, 1958

Acris crepitans - 15-35 36 4810) 34 | PYBURN, 1958

Bufo terrestris - 40-75 13 (627) 27 |BELLIS, 1959

Rana sylvatica ad, sad 35-65 12 (534) |298|BELLIS, 1965

Rana arvalis sad 12 (534) 16 | HAAPANEN, 1970

Rana arvalis ad 35-55 5 (93) 35 | HAAPANEN, 1970

Rana temporaria sad 8 (237) 96 | HAAPANEN, 1970

Kana temporaria ad 45-70 8 (237) 23 | HAAPANEN, 1970

Bufo americanus ad 28-50 21 5 |WERNER, 1991

Bufo bufo ad 50-70 4 (59) 29 | HAAPANEN, 1974

Bufo bufo sad 5 (93) 22 | HAAPANEN, 1974

Pseudacris triseriata ad 20-35 490 9 |KRAMER, 1974

Rana pipiens, site 1 ad 50-85 370 28 | DOLE, 1965b

Rana pipiens, site 1 sad 280 17 | DOLE, 1965b

Rana pipiens, site II ad 50-85 90 18 | DoLE, 1965b

Rana pipiens, site I sad 80 4 |DoLE, 1965b

Rana arvalis ad 36-55 6 (133) 78 |This study

Rana temporaria ad 46-70 9.5! (330) | 95 |This study

Size (mm)

clamitans (MARTOF, 1953), R. pretiosa (CARPENTER, 1954; TURNER, 1960), R. sylvatica

(BELLIS, 1965), R. pipiens (DOLE, 1965a-b), R. arvalis and R. temporaria (HAAPANEN, 1970);

Bufo boreas (CARPENTER, 1954), B. terrestris (BELLIS, 1959), B. bufo (HEUSSER, 1968;

HAAPANEN, 1974; SINsCH, 1988), and B. americanus (WERNER, 1991); Acris crepitans

(PYBURN, 1958); Pseudacris triseriata (KRAMER, 1974); and several tropical species (INGER,

1969). The home range areas reported are similar in size to those found by me (Table VI).

According to some authors, the home range may change during the course of one

season: see BRECKENRIDGE & TESTER (1961) for Bufo hemiophrys, TURNER (1960) for Rana

pretiosa, and WERNER (1991) for Bufo americanus. 1 could not detect evidence of this. This

may mean that it was really of unusual occurrence in my populations. It could also be

because my study only lasted for part of the summer seasons. Also, it may be a matter of

how to interpret a pattern. Based on telemetry, VAN GELDER & BUGTER (1987) published

Source : MNHN, Paris

28 ALYTES 12 (1)

a detailed, and very instructive, map of the movements of a Rana arvalis during two

summer months. The frog spent the full period within an area of about 20 X 30 m. During

shorter periods it usually stayed within a more restricted area. However, it did also return

to previous centres of activity, qualifying the whole area as a home range. This would

probably correspond to the pattern I interpret from my data — essentially a restricted

home range but with some change of activity centre over time. However, depending of the

time scale used, it could also be characterized as changes of home range.

In their studies, MARTOF (1953), FircH (1958), TURNER (1960), BELLIS (1965) and

DoLe (1965b) found, as I did, a tendency for frogs to return to the same home range in

successive years.

I think my results provide a rough but useful picture of these two frog species’

summer movements. However, we miss information on how and when the summer home

range is established and also more detailed information on the use of space within the

home range, including possible exploratory movements outside the “regular” home range.

Also, it is not clear to what extent there are between-year shifts in home range and if such

shifts are restricted to any particular age classes. Much of this work will undoubtedly

benefit from the use of telemetry.

ACKNOWLEDGMENT

The final compilation of this study has been supported by the Swedish Council for Forestry and

Agricultural Research.

REFERENCES

BeLis, E. D., 1959. — A study of movement of American toads in a Minnesota bog. Copeia, 1959:

173-174

= 1965. — Home range and movements of the wood frog in a northern bog. Ecology, 46: 90-98.

BRECKENRIDGE, W. J. & TESTER, J. R., 1961. — Growth, local movements and hibernation of the

Manitoba toad, Bufo hemiophrys. Ecology, 42: 631-646.

CAUGHLEY, G., 1975. — Analysis of vertebrate populations. London, John Wiley & sons: 1-234.

CONANT, R., 1958. — A field guide to reptiles and amphibians of eastern North America. Boston,

Houghton Miflin Co.: 1-366.

DoLe, J. W., 1965a. — Summer movements of adult leopard frogs, Rana pipiens Schreber, in northern

i ology, 46: 236-255.

ee 1965b. — Spatial relations in natural populations of the leopard frog, Rana pipiens Schreber, in

northern Michigan. 4m. Midi. Natur., T4: 464-478.

FALLER-DOEPNER, U., REH, W. & SErrz, A., 1991. — Radio tracking of the R. temporaria. In: Proc.

Ath European int. Conf. Wildl. Telemetry, Aberdeen: 41.

Fircu, H.S., 1958. — Home ranges, territories, and seasonal movements of vertebrates of the natural

history reservation. Univ. Kansas Publ. M at. Hist., 11: 63-326.

GeLer, J. J vaN., & BUGTER, R., 1987. — The utility of thermo-telemetric equipment in ecological

studies on the moor frog (R. arvalis Nilsson): a pilot study. Beih. Schriftenr. Naturschutz

Landschaftspfl. Niedersachs., 19: 147-153.

Source : MNHN, Paris

LOMAN 29

HAAPANEN, A., 1970. — Site tenacity of the common frog (Rana temporaria L.) and the moor frog

(Rana arvalis Nilss.). Ann. Zool. Fenn., T: 61-66.

= 1974, — Site tenacity of the common toad, Bufo bufo (L). Ann. Zool. Fenn., 11: 251-252.

HEUSSER, H., 1968. — Die Lebenweise der Erdkrôte Bufo bufo (L.). Rev. suisse Zool., 75: 927-982.

INGER, R., 1969. — Organization of communities of frogs along small rain forest streams in Sarawak.

J. anim. Ecol., 38: 123-148.

KRAMER, D. C., 1974. — Home range of the western chorus frog Pseudacris triseriata triseriata. J.

Herp., 8: 245-246.

LoMaN, J., 1978. — Growth of brown frogs Rana arvalis Nilsson and R. temporaria L. in south

Sweden. Ekol. Polska, 26: 287-296.

== 1981. — Spacing mechanism in a population of the common frog Rana temporaria during the

non-breeding period. Oikos, 37: 225-227.

= 1984. — Density and survival of Rana arvalis and Rana temporaria. Alytes, 3: 125-134.

MarTor, B., 1953. — Home range and movements of the green frog, Rana clamitans. Ecology, 34:

529-543.

PyeurN, W. F., 1958. — Size and movements of a local population of cricket frogs (Acris crepitans).

Texas J. Sci., 10: 325-342.

SCHUMACHER, F. & ESCHMEYER, R., 1943. — The estimation of fish populations in lakes and ponds.

J. Tenn. Acad. Sci., 18: 228-249.

SEBER, G. A. F., 1973. — The estimation of animal abundance and related parameters. London, Griffin:

1-406.

SiNscH, U., 1988a. — Seasonal changes in the migratory behaviour of the toad Bufo bufo: direction

and magnitude of movements. Oecologia, 76: 390-398.

ee 1988b. — Temporal spacing of breeding activity in the natterjack toad, Bufo calamita. Oecologia,

76: 399-407.

STEBBINS, R. C., 1966. — À field guide to western reptiles and amphibians. Boston, Houghton Mifflin

Co.: 1-279.

WERNER, J. K., 1991. — A radiotelemetry implant technique for use with Bufo americanus. Herp.

Review, 22: 94-95.

WILKINSON, L., 1990. — SYSTAT: the sytem for statistics. Statistics. Evanston, USA, SYSTAT Inc.:

1-677.

Corresponding editor: Günter GOLLMANN.

Source : MNHN, Paris

Alytes, 1994, 12 (1): 30. Announcement

Dumerilia

We are glad to announce the creation of Dumerilia, a new herpetological journal,

published by the Association des Amis du Laboratoire des Reptiles et Amphibiens du Muséum

national d'Histoire naturelle de Paris (AALRAM). The name of this journal is a tribute to the

memory of André-Marie-Constant Duméxit. (1774-1860), one of the founders of herpetology, author

with Gabriel Biron (1808-1846) of the masterly book series entitled Erpétologie générale (1834

1854).

Dumerilia will be an occasional publication, open to papers in French or in English from

the staff of the Laboratoire des Reptiles et Amphibiens of the Paris Museum (including associated

and corresponding. members) and from other colleagues but dealing with the collections of this

laboratory or where are described new taxons, the Aolotypes of which at least are deposited in these

collections.

Papers sent 10 Dumerilia will be submitted to peer review by colleagues from the

international scientific community. The editorial team of Dumerilia for 1994 is as follows:

Chief Editor: Roger Bour (Paris, France).

Editorial Board: Aaron M. Barr (Villanova, U.S.A.); Robert Barwauzr (Paris, France): Edouard-

Raoul BrvGoo (Paris, france); James R: Buskrk (Oakland, U.S.A.): José M. Ci (Cas

Portugal); Marc Curvian (Montpellier, France): Alain Dunois (Paris, France): Heinz Gru.urrsen

(Wien, Austria); Ivan Inticu (Paris, France); Raymond F. Laurenr (Tucumän, Argentine):

Michel Lemir (Paris, France); Colin J. McCarriy (London, United Kingdom): Annemaric

Ouisr (Paris, France): Hidetoshi Ora (Okinawa, Japan); Georges Pasteur (Montpellier,

France): Peter Perrciarn (Orlando, U.S.A.); Jean-Claude Race (Paris, France): Beat Scnarn

(Genève, Switzerland); Jordi Sekra-Couo (Barcelona, Spain); Hussam Zaurr (Paris, France):

George R. Zuc (Washington, U.S.A.).

The first volume of Dumerilia,

Liste bibliographique des Reptiles actuels. IL. Chéloniens

by Patrick DAVID,

will be published during the first quarter of 1994.

It is a book (roughly 100 pages), the first of a series which is intended to cover all living

reptiles. Each volume of this series contains: (1) an introduction; (2) a list of all taxa (from class to

subspecies) currently recognized as valid, with for each taxon its scientific Latin name (with author

and date), its French and English common names, and a list of selected references to major works

through which much more references can be traced: (3) notes dealing with litigious points: (4) a

detailed bibliographic list of all references quoted (full citations).

The volume 1 of Dumerilia (Chelonians) can be ordered as follows:

- Payment in French Francs (130 FF for individuals: 260 FF for institutions):

(1) by direct postal transfer to our postal account: “AALRAM”, Nr. 36-962-64-U, La Source:

€) by cheques payable 10 “AALRAM” sent to: AALRAM, cho Alain Dumas, Laboratoire des

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(3) by cheques payable to *AALRAM”, sent to: AALRAM, c/o Patricia B. Zuc, Division of

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Source : MNHN, Paris

Alytes, 1994, 12 (1): 31-40. 31

Dispersal of Rana temporaria tadpoles

in large fishponds

Dominique AUGERT & Pierre JOLY

URA CNRS 1451 Ecologie des Eaux douces et des grands Fleuves,

Université Claude Bernard Lyon I,

43 boulevard du 11 novembre 1918, 69622 Villeurbanne Cedex, France

Dispersal of Rana temporaria tadpoles was monitored from hatching to

metamorphosis by systematic sampling at regularly distributed points of two

large fishponds in the Jurassian Bresse (Eastem France). Dispersal showed

two successive phases. During the first phase, shoals of tadpoles dispersed

from the breeding site towards different directions. Then, the shoals spread out

and the tadpoles colonized the shallower parts of the pond. They did not

colonize the entire pond banks but only the shallower ends, i. e. the largest

shallow areas occupied by a high density of vegetation. They emerged on the

side of a pond that was the shadiest. Despite great differences in vegetation,

mode and dynamics of tadpole dispersal did not differ between the two ponds.

INTRODUCTION

The question of tadpole dispersal is not very well documented. The literature has been

mainly concerned with the diel pattern of distribution in small ponds (BEISWENGER, 1977,

in Bufo americanus; GRIFFITHS, 1985, in Rana temporaria). The diel patterns observed in

these two species are similar and may be summarized as follows: during the morning, the

tadpoles migrate from the deepest zone of the pond towards the banks and aggregate in

the shallow areas while temperature is increasing; at dusk, they return to the deepest parts

of the pond. ANGELIER & ANGELIER (1968) noted aggregations of tadpoles of Rana

temporaria on the banks during sunny days. As suggested by these observations, this diel

cycle is probably thermo-dependent. Whereas water temperatures reach highest values

along the bank during the day, during the night they are highest at the bottom. However,

R. temporaria tadpoles may move from warmer to cooler water during dispersal

(GRIFFITHS & MYLOTTE, 1986).

Tadpole social behaviour during the entire larval period has been described

qualitatively in R. temporaria by SAVAGE (1961) and GUYÉTANT (1975). According to the

latter, tadpoles are gregarious after hatching. They stay for one week at the spawning site

where they feed on the gelatinous envelopes of the clutches. They remain near the banks

at less than 10 cm depth and disperse in the whole pond at the age of twelve to fifteen days;

three months after hatching, they come back towards the banks and form aggregations

Source : MNHN, Paris

32 ALYTES 12 (1)

before metamorphosis. According to SAVAGE (1961), R. temporaria tadpoles may remain

gregarious for three to six weeks; they then disperse into several separate shoals,

apparently based on food resources; eight to nine weeks after hatching they have a solitary

life characterized by few feeding movements. These studies were carried out in small

fishfree ponds.

The surface of the pond and the diversity of its microhabitat probably influence

tadpole dispersal. The previous observations concerned only small ponds where tadpoles

may wander quickly throughout the pond. Dispersal has not been described in large ponds

such as fishponds with a surface of several hectares, which provide a higher diversity of

microhabitats. Our purpose is to describe precisely the dispersal of tadpoles of Rana

temporaria during the whole larval period and during froglet emergence in two large

fishponds that attract a very large number of breeders (JoLy, 1991; AUGERT & JoLY, 1993).

This monitoring was based on systematic sampling of tadpoles at points regularly

distributed in the two fishponds.

MATERIAL AND METHODS

STUDY SITES

The two fishponds are located 9 km apart in Jurassian Bresse (Jura Department,

between Lons-le-Saulnier and Dôle, altitude 200-215 m). Jurassian Bresse is a mosaic of

deciduous forests (Quercus, Fraxinus and Carpinus), crops, meadows and fishponds

devoted to carp farming. These ponds are dried every winter and filled again with rain or

brook water. A 1.5 to 3 m deep central ditch leads to a draining point at the deepest end

(3 m deep maximum) of each pond. The opposite bank has one or several shallow “tails”.

Average pond depth is about 0.7 m. Adult Rana temporaria usually breed in a tail. Daguin

fishpond (3 ha) is surrounded by cultivated lands and meadows. Its tail is adjacent to a

little wood. Thévenon fishpond (3.5 ha) is within a forest. Conductivity measured during

the spawning period was 161 pS.cm'! at Daguin and 34.9 pS.cm'! at Thévenon. The higher

value at Daguin is due to the proximity of field and pasture. The aquatic vegetation of

each pond was mapped during June. Temperature values were regularly recorded at the

same hour at several points of the two ponds at a depth of 0.5 m.

SAMPLING MODE

In 1989, Rana temporaria tadpoles were collected in the two ponds on five dates

regularly distributed over time, from egg hatching to froglet metamorphosis (6 April,

19 April, 4 May, 22 May, 5 June). Samples along transverse transects were taken from a

boat. The distance between two successive transects was about 30 m. Samples were

uniformly distributed along each transect (about every 40 + 5 m). Hence the number of

samples per transect varied from five in the wider parts (one in the middle, two near the

Source : MNHN, Paris

AUGERT & JOLY 33

banks and two at intermediate points between the middle and the banks) to four and three

(middle and banks) in the narrower parts. Sixty-six and seventy-seven points were visited

at each date at Thévenon and at Daguin respectively. Three standardized sweeps of a

40 x 40 cm net (0.5 mm mesh) were made at each site from the bottom to the surface.

Sampling was always carried out by the same operator. Depth was measured at each

sampling point. From 5 to 9 June, we searched the edges of the two fishponds to determine

where the froglets left the ponds. Maps of tadpole dispersion were computed using

GraphMu on a Macintosh (THIOULOUSE, 1989).

RESULTS AND DISCUSSION

POND DESCRIPTION

The shallowest parts of the ponds (less than 1 m) showed two different shapes.

Whereas they were narrow at the lateral banks, they widened out in the tails (fig. 1).

Thirtheen and sixteen macrophyte species were recorded at Thévenon and Daguin

respectively (fig. 1). Six species were common to both ponds, Carex riparia, Glyceria sp.,

Iris pseudacorus, Utricularia vulgaris, Scirpus lacustris and Sparganium erectum. Floating

species (Trapa natans, Potamogeton lucens) occupied large areas at Daguin but were scarce

at Thévenon. The deepest part of Thévenon was covered by Characeae. This plant family

was absent at Daguin. The banks were colonized by Carex sp. at Thévenon and by Scirpus

lacustris at Daguin. The communities which occupied the pond tails were different:

Glyceria sp. and Carex sp. were the most frequent at Thévenon, Sparganium erectum,

Phragmites australis and Typha angustifolia were the most frequent at Daguin. Daguin is

a more eutrophic pond than Thévenon. Vegetation richness at Daguin probably is a

consequence of enrichment in mineral salts due to field drainage. Light levels were also

higher at Daguin.

Mean temperature values were not significantly different between tails, lateral banks

or central parts of the two ponds (two-ways ANOVA, see temperature values in Table I).

TADPOLE DISPERSION

Though beginning earlier at Daguin, tadpole dispersal presented some similarity

between the two ponds (fig. 2). The dispersal process went through several phases. About

one week after hatching, the tadpoles left the remains of the gelatinous mass. The first

tadpoles which were found far from the spawning site were aggregated in some particular

points (6 April at Daguin, 19 April at Thévenon). Their first movement was not a

diffusion-like dispersal from the spawning site but rather appeared as a group movement

of several thousand tadpoles, during about one week. In such shoals, physical contact

between tadpoles was frequently observed. After this shoaling movement, tadpoles

straggled more widely in the entire pond tails (19 April and 4 May at Daguin, 4 May at

Source : MNHN, Paris

(a) DEPTH (cm) (b) VEGETATION

DAGUIN

forest

—— tail

à £

— 50 m +— 50 m

THEVENON

| >

| E

| <

L2}

— 50 m +— 50 ml

LEGEND

sampling point

Carex (acuta and riparia) M \ymphea alba Scirpus palustris

Characeae sp. E Phragmites australis Ranunculus aquatilis Sparganium erectum

Giyceria (fluitans and maxime) HA Potamogeton lucens Roripa amphibia Trapa natans

ZA ris pseudacorus Æ Potamogeton natans El] Sagittaria sagittifolia Typha angustifolia

Lemna minor Œ Potamogeton pectinatus ÆA Scirous lacustris Utricularia vulgaris

Fig. 1. — (a). Depth isolines in the two ponds: the pond tails show a wide area of shallow water. (b) Maps of aquatic vegetation in

the two ponds, Daguin and Thévenon. Twenty aquatic plant species have been noted in the two ponds, 16 in Daguin and 13 in

Thévenon.

Source : MNHN, Paris

AUGERT & JOLY 35

Table I. — Temperature mean values (in °C) recorded at each date in the central

parts, the lateral banks and the tails of the two ponds.

——

Daguin Thévenon

Dates

Central parts | Lateral banks Tails Central parts | Lateral banks Tails

6 April 8.6 9.2 9.6 9.2 9.6 10.1

19 April 8.9 9.5 10.1 9.5 9.8 11.0

4 May 12.4 13.6 14.2 14.2 14.8 16.0

22 May 14.5 15.3 16.0 15.5 16.3 17.2

5 June 15.3 16.7 18.2 17.7 18.4 19.5

Thévenon). Physical contact between tadpoles became less frequent. Finally, two months

after hatching, they returned to the banks of the tails (22 May) where they emerged

(5 June).

In our study, the habitat parameters considered as influencing tadpole distribution

were depth, temperature and vegetation. It is clear that tadpoles did not colonize the entire

pond but only its shallower parts (< 75 cm) (fig. 3) and did not colonize all the banks but

only the ones situated in pond tails (fig. 2). They did not spread out further than 150 m

from the spawning site. Previous studies have shown that tadpoles prefer both shallow

water and high temperatures (NOLAND & ULTSCH, 1981; FLOYD, 1984; DuPRÉ &

PETRANKA, 1985). Tadpole distribution observed in our study may be due to depth rather

than to temperature which was on the average not significantly higher in the tails than in

the central parts or in the lateral banks of each pond.

Shifts in vegetation occupation occurred during tadpole development in both ponds

(fig. 4). At Daguin, at the beginning of April, the tadpoles reached highest densities in

Sparganium erectum, Scirpus lacustris and Phragmites australis (6 April). The tadpoles then

spread towards patches of Potamogeton lucens, Glyceria sp. and Utricularia vulgaris

(19 April; 4 May; 22 May). Before metamorphosis (5 June), they occupied only

Sparganium erectum and Glyceria sp. patches. They did not colonize Trapa natans,

Potamogeton praelongus or Sagittaria sagittifolia. At Thévenon, at the beginning of the

development (6 April), they were found in zones of Glyceria sp. Then, whereas some

tadpoles remained in G/yceria patches, a lot of them shifted (19 April) to patches of Carex

riparia, Utricularia vulgaris, and, to a lesser extent, Potamogeton pectinatus and Scirpus

lacustris. Before metamorphosis, they mainly occupied patches of Glyceria sp. and Carex

riparia (4 June). They avoided Characeae (fig. 4). As a whole, tadpoles mainly inhabited

species of shallow water but only those that occupy the tails. Contrarily to the lateral

banks, tails constitute large areas of shallow water and of high density of vegetation.

Source : MNHN, Paris

36 ALYTES 12 (1)

DAGUIN THEVENON

1 April 6 April

LEGEND :

x spawning site

sampling point

emergence zones

10 tadpoles

> 100 tadpoles

— 50m

Fig. 2. — Dispersion of tadpoles at each sampling date. The size of each circle is proportional to the

number of tadpoles collected according to a logarithmic scale. The size of the circles shown in

the legend are given as examples of tadpole numbers.

Source : MNHN, Paris

AUGERT & JOLY 37

100

DAGUIN POND

a) 20

£ Z

5 À >

nm |

ä Z

2 THEVENON POND

25-50 50-75 >75

DEPTH (cm)

Fig. 3. — Relationship between tadpole density (expressed in percentages) and water depth for the

five sampling dates. On the first date (6 April), tadpole dispersal has only begun in Daguin pond.

Source : MNHN, Paris

38 ALYTES 12 (1)

DATES

Utricularia vulgaris 6 April

DAGUIN POND A 19 April

E 4 May

Phragmites australis 22 May

O 5 June

Giyceria sp.

Potamogeton lucens

Scirpus lacustris

Sparganium erectum

Scirpus lacustris La

THEVENON POND

Potamogeton pectinatus

Utricularia vulgaris

Glyceria sp

Carex riparia

0 20 40 60 80 100

% SAMPLED TADPOLI

Fig. 4. — Relationship between tadpole density (expressed in percentages) and plant species for the

five sampling dates.

Source : MNHN, Paris

AUGERT & JOLY 39

Another factor, not measured in our study, which might influence tadpole dispersion,

was the presence of fishes which reached rather high densities in the two ponds (personal

observations). Indeed, as fishes are known to prey heavily on tadpoles (BEEBEE,

1983; KaTs, PETRANKA & SIH, 1988), they may restrict tadpole dispersion to the parts

of the ponds where vegetation provides a refuge against predation (CLAUSNITZER,

1983).

Metamorphosis also occurred in pond tails. The presence of vegetation and a gentle

slope may help to prevent tadpoles from drowning, what has been observed during

metamorphosis in R. temporaria (SAVAGE, 1961; AsHBy, 1969). At Thévenon, froglets

emerged on both sides of each of the two pond tails toward the forest. At Daguin, the

froglets left the pond on the side of the little wood which is adjacent to the fishpond tail.

Emigration in that direction may be due to an attraction of froglets by forest. In contrast

to KoskELA’s (1973) observations, metamorphosis occurred in sunny weather; hence

movement toward the forest may ensure favourable conditions for froglet life (AsHBY,

1969). According to SMIRINA (1980), froglet mortality is the highest during the first weeks

after metamorphosis. The hypothesis of an attraction of froglets by forest has to be tested

experimentally.

RÉSUMÉ

La dispersion des têtards de grenouilles rousses (Rana temporaria) a été suivie depuis

l'éclosion jusqu’à l'émergence, par un échantillonnage systématique en des points

régulièrement distribués, dans deux grands étangs de Bresse jurassienne (Est de la France).

Cette dispersion comporte deux phases successives. Au cours de la première phase, des

bancs de têtards se dispersent depuis le site de frai dans différentes directions. Ensuite, ces

bancs se diffusent et les têtards colonisent les parties les moins profondes de l'étang. Il est

remarquable qu’ils ne colonisent pas entièrement les rives, mais seulement les zones de

queue d’étangs, zones de faible profondeur les plus vastes comportant une forte densité de

végétation. Ils émergent du côté le plus ombragé de l’étang. Malgré de fortes différences

de végétation, le mode et la dynamique de dispersion des têtards ne diffèrent pas entre les

deux étangs.

ACKNOWLEDGEMENTS

We thank M. MorEsTiN, owner of the ponds, for having made this study possible. We are

grateful to Monique JOLY and Fabrice MARTINET for the determination of the aquatic plant species

and to Odile GROLET, Marianne OUALI and Pa RiErA for their help in the field. We are also

indebted to Prof. Louis CAILLÈRE, Dr. GRIFFITHS and two anonymous referees for their criticisms on

first drafts of the manuscript and to Glyn THOIRON for reviewing the English language.

Source : MNHN, Paris

40 ALYTES 12 (1)

LITERATURE CITED

ANGELIER, E. & ANGELIER, M. L., 1968. — Observations sur le développement embryonnaire et

larvaire de Rana temporaria L. (Batracien, Anoure). Annl. Limnol., 4: 113-131.

Asumy, K. R, 1969. — The population ecology of a self-maintaining colony of the common frog

(Rana temporaria). J. Zool., 158: 453-474.

AUGERT, D. & JOLY, P, 1993. — Plasticity of age at maturity between two neighbouring populations

of the common frog (Rana temporaria L.). Can. J. Zool., T1: 26-33.

BEEBEE, T. J. C., 1983. — The natterjack toad. Oxford, Oxford University Press: 1-159.

BEISWENGER, R. E., 1977. — Diel pattern of aggregative behaviour in tadpoles of Bufo americanus,

in relation to light and temperature. Ecology, 58: 98-108.

CLAUSNITZER, H. J. , 1983. — Zum gemeinsamen Vorkommen von Amphibien und Fischen.

Salamandra, 19: 158-162.

Durré, R. K. & PETRANKA, J. W., 1985. — Ontogeny of temperature selection in larval amphibians.

Copeia, 1985: 462-467.

FLoyp, R. B., 1984. — Variation in temperature preference with stage of development of Bufo

marinus larvae. J. Herpet., 18: 153-158.

GRIFFITHS, R. A., 1985. — Diel pattern of movement and aggregation in tadpoles of the common

frog, Rana temporaria. Herpet. J., 1: 10-13.

Grirrrrus, R. A. & MYLOTTE, V. J., 1986. — Observations on the dispersal of common frog tadpoles

Rana temporaria from the spawn site. Brit. herpet. Soc. Bull., 18: 21-23.

GuyéranT, R. , 1975. — Etude des interactions intraspécifiques chez les tétards de quelques amphibiens

anoures. Conséquences physiologiques. Thèse d'Etat, University of Besançon: 1-190.

Jouy, P., 1991. — Variation in size and fecundity between neighbouring populations in the common

frog, Rana temporaria. Alytes, 9: 79-88.

KaTs, L. B. , PETRANKA, J. W. & Sin, A., 1988. — Antipredator defenses and the persistence of

amphibian larvae with fishes. Ecology, 69: 1865-1870.

KoskeLA, P., 1973. — Duration of the larval stage, growth and migration in Rana temporaria L. in

two ponds in northern Finland in relation to environmental factors. Ann. Zool. Fennici, 10:

414-418.

NOLAND, R. & ULTsCH, G. R. , 1981. — The role of temperature and dissolved oxygen in

microhabitat selection by the tadpoles of a frog (Rana pipiens) and a toad (Bufo terrestris).

Copeia, 1981: 645-652.

SAVAGE, R. M. , 1961. — The ecology and life-history of the common frog (Rana temporaria

temporaria). London, Pitman and Sons: 1-221.

SMiiNa, E. M. , 1980. — On growth rate and survival of common frogs (Rana temporaria) during

the first years of life. Zoo!. Zh., 59: 1331-1332.

THIOULOUSE, J:, 1989. — Statistical analysis and graphical display of multivariate data on the

Macintosh. Computer Applications in the Biosciences, 5: 287-292.

Corresponding editor: Günter GOLLMANN.

Source : MNHN, Paris

Alytes, 1994, 12 (1): 41-47. 41

Age structure and survival rate in Alpine newts

(Triturus alpestris) at high altitude

Robert SCHABETSBERGER & Alfred GOLDSCHMID

Zoological Institute, University of Salzburg, Hellbrunnerstrasse 34, 5020 Salzburg, Austria

Individual ages of 67 Alpine newts from a karst plateau in the North-

‘eastemn Calcareous Alps of Austria (Totes Gebirge, 1650 m altitude) have been

assessed using skeletochronology of cross sections of the humerus. Both sexes

start to reproduce at the age of 10 years, longevity exceeds 20 years and the

oldest animal survived to an age of 22 years.

Survival rates estimated from age structure and mark-recapture experi-

ments appeared lower in males, causing a female-biased sex ratio. Males may

be subject to increased mortality during the spring migration.

INTRODUCTION

Skeletochronological studies of Triturus have shown that age at first reproduction is

delayed and longevity increased with increasing altitude and latitude of the breeding site

(DoLMEN, 1983; CAETANO et al., 1985; CAETANO, 1990; CAETANO & CASTANET, 1993).

Decreased ambient temperatures lead to reduced metabolic rates, which in turn cause

slower growth and delayed sexual maturity (JORGENSEN, 1992). Age and survival in the

Alpine newt were studied on the southwest plateau of the Totes Gebirge (1650 m), the

largest karst massif (590 km?) in the Northeastern Calcareous Alps of Austria. This massif

generally receives more than 2200 mm precipitation annually and snow cover is present

more than 196 days each year (DINÇER et al., 1972); breeding sites were covered with ice

from the beginning of November until mid-June during the study period. Some of the

breeding sites are former dolines, where temperatures are reduced by the settling of cold

air.

MATERIAL AND METHODS

Alpine newts were collected from Lake Dreibrüdersee (1643 m) and from a pond

(1650 m) situated approximately 1 km east from the lake in a neighbouring valley. A

mountain range with peaks up to 1950 m separates the two breeding sites. Lake

Source : MNHN, Paris

4 ALYTES 12 (1)

Dreibrüdersee fills a former doline (137 m diameter) and is surrounded by steep rocky

slopes.

Individual ages of 36 females and 8 males from Lake Dreibrüdersee, and 23 males

from the nearby pond, were assessed by skeletochronology using cross sections of the

humerus. Mean snout-vent lengths of breeding females and males in Lake Dreibrüdersee

were 57.06 + 0.22 mm (95 % C.L.) and 49.53 + 0.32 mm respectively. Only males found

dead were used for age determination. Live females were anaesthetized with MS 222 and

the entire sample was fixed in 10 % formalin. Diaphysial regions of the humerus were

decalcified with 40% EDTA, dehydrated in Isopropylalcohol, stored over night in

Methylbenzoat, and embedded in Paraplast. Cross sections 10 mm in thickness were

stained with DELAFIELD-Hematoxylin (ROMEIS, 1968) for several hours and then

destained with hydrochloric acid.

The mean survival rate (D) for the age classes 14 to 19+ in females from Lake

Dreibrüdersee and 14 to 17 in males from the pond were calculated from the age structure

by using the ROBsoN & CHAPMAN formula (KREBS, 1989: 432-433):

2ù

PSN ET

whe:

® = average finite annual survival rate or probability of survival; finite survival rates can

range from 0 to 1; a survival rate of 0.65 means that in average 65 % of the population

survive from one year to the next;

N, = number of individuals in age class x;

ÈN = number of all animals from age classes x to x+i N, + N,4, + Nu + + Nous

for females from Lake Dreibrüdersee this is: N,4 + Nis + … + Ni,

T = sum of the coded ages times their frequencies when coded age is found by setting the

youngest included age class to 0, the next age to one and so forth: ON, + IN,,, + 2N,42

+ +iN,,;; for females from Lake Dreibrüdersee this is: N,; + 2N,6 + … + 5N,,.

Animals from age classes 10 to 13 were omitted from the calculations because they

were underrepresented in the sample (fig. 2).

This formula is only valid for a stationary population in which recruitment

and survival are relatively constant. As these assumptions are hardly fulfilled in

natural populations, the calculated values should be treated as rough estimates which

allow comparison of average survival rates between sexes and between different

populations.

Adults (855 females, 277 males) in Lake Dreibrüdersee were marked individually by

tattooing with Alcian blue between 1986 and 1992 (Jouy & Miaup, 1990). Population size

and finite annual survival rates were estimated with a multiple mark-recapture census

(JoLLy-SEBER method, KREBs, 1989: 37-43). In contrast to the RoBsON & CHAPMAN method

this model does not assume a constant survival rate.

Reliability of population size estimates was assessed using less complex models

(PETERSEN and SCHUHMACHER methods, KREBS, 1989: 16-36) which work only in a closed

Source : MNHN, Paris

SCHABETSBERGER & GOLDSCHMID 43

LS SN

Fig. 1. — Cross section of the humerus of a 15 years old Alpine newt female. a: LAG formed during

the first 10 years of life. b: LAG become narrow after the newt starts to reproduce. Fig. 1b is

taken from another cross section of the same bone shown in fig. la.

population. During the summer period the percentage of marked animals in consecutive

samples increased linearly with the total amount of newts marked in this year

(SCHUHMACHER-ESCHMEYER plot; KREBs, 1989: 35), showing that a constant population

size was reached for a short time interval after migration into the lake.

RESULTS

No marked animals from Lake Dreibrüdersee were found in other breeding sites. Due

to the isolated position of the lake and the high recapture rates of individually marked

newts, any interchange with other populations seemed unlikely. There was no other

breeding pond available in the vicinity of Lake Dreibrüdersee.

Both sexes started to breed late in life, first entering the breeding site at the age of 10

years. The long winter period resulted in clear lines of arrested bone growth (LAG), which

were very close together after the animals had started to reproduce (fig. la). After 10 years

age, bone growth was strongly reduced, resulting in a darkly staining outer zone of narrow

LAG (fig. 1b). No animals younger than 10 years age were caught in the breeding sites

indicating that the younger newts in this population lead an entirely terrestrial life.

Source : MNHN, Paris

44 ALYTES 12 (1)

Ç DBS

d'DBS

Number of animals

5 10 1S 20 25

Age (years)

Fig 2. — Age structure of the breeding populations of Alpine newts from Lake Dreibrüdersee (DBS)

and a nearby pond.

Animals 14 years old constituted the largest age class (fig. 2). Longevity exceeded 20

years in one female and one male newt from Lake Dreibrüdersee. The average annual

survival rate derived from the age structure was estimated to be 0.67 + 0.003 (b + S.E.)

for females from Lake Dreibrüdersee and 0.45 + 0.012 for males from the pond, meaning

that on average 67 % of the females and 45 % of the males survived from one year to the

next.

Estimates of annual female survival rates (JOLLY-SEBER method) were above 0.70

(Table I). Approximately 60 % of the females (D + S.E. = 0.609 + 0.043) and 40 % of

the males (D = 0.397 + 0.087) present in 1986 survived to 1988. Assuming a constant

probability of survival in this two-year interval yields annual survival rates of 0.78 and

0.63 for females and males, respectively. From 1988 to 1992, the sample size of males was

too small for application of the JorLy-SEBER model. The results indicate that males

suffered from higher mortality than femal

The newt population in Lake Dreibrüdersee consisted of approximately 500-800

females and 150 males, resulting in a biased sex ratio of 1/3.7 to 1/4.8 males to females

(Table 1). Estimates of population size with different mark-recapture models were within

the confidence limits of each other.

Source : MNHN, Paris

SCHABETSBERGER & GOLDSCHMID 45

Table I. - Population size, sex ratio and survival rate of Alpine newt adults in Lake

Dreibrüdersee estimated with mark-recapture experiments. Sex ratio: d/$. No

animals were marked in 1987.

Population size (N) Survival rate

D+S.E.

PETERSEN [P] / JoLLv-SEBER

ScHunMAcuER [S] N+S.E.

(95 % C.L.)

1986 $: 773 (541-1314) [P] 1986-1988:

: 0.609 + 0.043

g': 0.397 + 0.087

1988 $: 517 (466-580) [S] 546 + 40 0.824 + 0.053

d': 149 (125-184) [S]

Sex ratio: 1/3.5

: 708 (609-845) [S]

d': 156 (122-232) [P]

Sex ratio: 1/4.6

807 + 58 0.747 + 0.090

1990 - 664 + 86 0.965 + 0.186

1991 : 682 (321-1443) [P] 596 + 102

DISCUSSION

Alpine newts in temperate environments start to reproduce at three to four years of

age and live for a maximum of 10 years (SMIRINA & ROËEK, 1976; SMIRINA & SUFIANIDU,

1985; MiauD, 1990). The present study has shown that first reproduction can be

significantly delayed due to cold temperature conditions. Adults have to use a very short

summer period for reproduction. The average yearly ambient temperature at Lake

Dreibrüdersee is around 2° C, resulting in slow growth rates. Growth is further reduced

with the onset of first reproduction, as more energy has to be put into gonadal

development. Strongly reduced bone growth after 10 years age and the fact that no

younger animals were caught in the breeding sites indicate that Alpine newts cannot

reproduce earlier in life under these environmental conditions.

Animals from Lake Dreibrüdersee were bred under laboratory conditions simulating

lowland temperature regimes. They reached body size of breeding animals within two to

three years, showing that reduced growth was not genetically determined in this

population (pers. obs.). Food availability was high in Lake Dreibrüdersee and reduced

growth did not result from limited resources in the lake. During the second half of their

aquatic period, individually marked females regained biomass lost during oviposition

(pers. obs.).

Source : MNHN, Paris

46 ALYTES 12 (1)

A long phase of hibernation followed by a comparatively short period of activity in

summer probably did not support the development of double LAG reported for other

Triturus populations (CAETANO et al., 1985; CAETANO & CASTANET, 1993).

The average female survival rate calculated with the ROBsON & CHAPMAN formula is

close to the estimate of 0.65 published by MiAUD (1991) for a lowland Alpine newt

population in south-east France. Nevertheless, mark-recapture experiments yielded higher

survival rates above 0.7 (Table 1). This difference resulted from not including age classes

10 to 13 in the calculation of survival rate with the ROBsON & CHAPMAN formula. Some

of the 10 to 13 years old specimens did not enter the breeding sites. Additionally, the age

of animals older than 15 years may have been underestimated, because LAG became

increasingly more difficult to distinguish. For this reason mark-recapture experiments gave

a more realistic description of survival.

Males seemed to have a higher mortality rate than females. These different survival

rates in males and females clearly need to be investigated further. In their detailed studies

of Alpine newts at similar altitudes in the Austrian Alps, FABER (1991) and GUTLEB (1990)

also found skewed sex ratios of 1/1.45 and 1/1.22 males to females respectively. Higher

mortality during the spring migration could be the reason for the lower survival rates in

adult males. Adult males started spring migration before the females. This was also found

for other newt species (BLAB, 1978; HARRISON et al., 1983; GRIFFITHS, 1984). During

migration at the beginning of June, temperature often drops below freezing. At this time

males enter numerous small water bodies on their way to the breeding site. These puddles

easily freeze and the newts suffocate. Of 50 dead newts collected in the spring 48 were

males.

RÉSUMÉ

L’âge individuel de 67 Tritons alpestres d’un plateau karstique des Alpes calcaires du

nord-est de l’Autriche (Totes Gebirge, altitude 1650 m) a été mis en évidence par

squelettochronologie s'appuyant sur des coupes transversales d’humérus. En raison de la

longue période d’hiver, les animaux des deux sexes commencent à se reproduire à l’âge de

10 ans. L’espérance de vie s’élève à 20 ans, l'animal le plus vieux observé ayant atteint l’âge

de 22 ans. Le taux de survie, estimé en fonction de la structure d'âge de la population et

d'expériences de marquage-recapture, semble plus bas chez les mâles, ce qui entraîne un

biais du sex-ratio en faveur des femelles. Ce phénomène pourrait être dû à une mortalité

élevée des mâles pendant la migration de printemps.

ACKNOWLEDGEMENTS

We wish to thank Pierre JOLY and Claude MiauD for showing us the tattooing technique and

Heidi LANGER for the help with sectioning and staining the bone samples. Günter GOLLMANN and two

anonymous referees made numerous constructive suggestions that improved the manuscript.

Source : MNHN, Paris

SCHABETSBERGER & GOLDSCHMID 47

LITERATURE CITED

BLAB, J., 1978. — Untersuchungen zur Ükologie, Raum-Zeit Einbindung und Funktion von

Amphibienpopulationen. Schr.-Reïhe Landschafispfl. Naturschutz, 18: 1-141.

CaëraNo, M. H., 1990. — Use and results of skeletochronology in some urodeles (7riturus

marmoratus, Latreille 1800 and Triturus boscai, Lataste 1879). Ann. Sci. Nat., 11: 197-199.

Cagrano, M. H. & CASTANET, J., 1993. — Variability and microevolutionary patterns in Triturus

marmoratus from Portugal: age, size, longevity and individual growth. Amphibia-Reptilia, 14:

117-125.

CAETANO, M. H., CASTANET, J. & FRANCILLON, H., 1985. — Détermination de l’âge de Triturus

marmoratus marmoratus (Latreille 1800) du Parc National de Peneda Geres (Portugal) par

squelettochronologie. Amphibia-Reptilia, 6: 117-132.

DINGER, T., PAYNE, B. R., YEN, C. K. & ZÔTL, J., 1972. — Das Ti

der Karstmassive der nordôstlichen Kalkhochalpen (Ergebnisse von Isotopenmessungen). Steir.

Beitr. z. Hydrogeologie, 24: 71-109.

DoLEn, D., 1983. — Growth and size of Triturus vulgaris and T. cristatus (Amphibia) in different

parts of Norway. Holarct. Ecol., 6: 356-371.

FaBer, H., 1991. — Laichplatzôkologie alpiner Tümpel. Dissertation, Universität Graz: 1-188.

GRIFFITHS, R. A., 1984. — Seasonal behaviour and intrahabitat movements in an urban population

of smooth newts, Triturus vulgaris (Amphibia: Salamandridae). J. Zool. Lond., 203: 241-251.

Guris8, B., 1990. — Populationsôkologische Untersuchungen am Bergmolch (Triturus alpestris) im

Kärniner Nockgebiet (Firstmoor 1920 m). Diplomarbeit, Universität Wien: 1-56.

HARRISON, J. D., GiTTiNs S. P. & SLATER, F. M., 1983. — The breeding migrations of smooth and

palmate newts (Triturus vulgaris and Triturus helveticus) at a pond in mid Wales. J. Zool. Lond.,

199: 249-258.

Jouy, P. & Mau, C., 1990. — Tattooing as an individual marking technique in urodeles. Alytes, 8:

11-16.

JORGENSEN, C. B., 1992. — Growth and reproduction. In: M. E. FebEr & W. W. BURGGREN (eds.),

Environmental physiology of the amphibians, Chicago & London, Univ. Chicago Press: 439-467.

Kkens, C. J., 1989. — Ecological methodology. New York, Harper & Row Publishers: 1-652.

MiauD, C., 1990. — La dynamique des populations subdivisées: étude comparative chez trois

Amphibiens Urodèles (Triturus alpestris, T. helveticus er T. cristatus). Thèse de Doctorat,

Université Claude Bernard Lyon I: 1-20.

ss 1991. — La squelettochronologie chez le Triturus (Amphibiens, Urodeles) à partir d'une étude

de T. alpestris, T. helveticus et T. cristatus du sud-est de la France. Zn: J. L. BAGLINIÈRE, J.

CASTANET, F. CONRAD & F. J. MEUNIER (eds.), Tissus durs et âge individuel des vertébrés,

Colloques et séminaires ORSTOM-INRA, Colloque national, Bondy, France, 4-6 mars 1991:

363-384.

ROMEIS, B., 1968. — Mikroskopische Technik. München & Wien, R. Oldenbourg Verlag: 1-757.

SMiRINA, E. & Roëek, Z., 1976. — On the possibility of using annual bone layers of Alpine newts,

Triturus alpestris (Amphibia: Urodela) for their age determination. Vest. Cs. Spol. Zool., 40:

232-237.

SMIRINA, E. & SUFIANIDU, T.., 1985. — On the life span of neotenic and metamorphosed Alpine newts

Triturus alpestris from high mountains of Greece. Zool. Zh., 64: 311-315.

te Gebirge als Entwässerungstypus

Corresponding editor: Günter GOLLMANN.

Source : MNHN, Paris

Alytes, 1994, 12 (1): 48. Announcements

Meetings

SixTH MEETING OF THE TrITURUS GROUP

SchloB Ulmerfeld, 120 km west of Vienna, Austria.

9-12 September 1994.

For information please contact:

Dr. Robert SCHABETSBERGER, Institut für Zoologie der Universität Salzburg,

HellbrunnerstraBe 34, 5020 Salzburg, Austria.

Tel.: +662 8044 5600.

Fax: +662 8044 5698.

WORKSHOP ON POPULATION BIOLOGY OF AMPHIBIANS

Vienna, Austria.

14-17 September 1994.

For information please contact:

Dr. Walter HôbL or Dr. Günter GOLLMANN, Institut für Zoologie der Universität

Wien, AlthanstraBe 14, 1090 Wien, Austria.

Fax: +431 31336 700.

SECOND INTERNATIONAL SYMPOSIUM ON ECOLOGY AND GENETICS OF EUROPEAN

WATER FROGS

Wroctaw, Poland.

19-25 September 1994.

For information please contact:

Dr. Maria OGtELSKA, Zoological Institute, University of Wrocltaw, ul. Sienkie-

wicza 21, 50-335 Wroclaw, Poland.

Tel.: +48 71 22 50 41.

Fax/tel.: +48 71 22 28 17.

E-mail: Ogielska at PLWRUWI1.Bitnet.

SECOND INTERNATIONAL ASIAN HERPETOLOGICAL MEETING

Ashgabat, Turkmenistan.

25-30 August 1995.

For information please contact:

- Dr. Sakhat M. SHAMMAKOV, Institute of Zoology, Academy of Sciences of

Turkmenistan, Azady Street 59, 744000 Ashgabat, Turkmenistan.

- Dr. Theodore J. PAPENFUSS, Museum of Vertebrate Zoology, University of

California, Berkeley, California 94720, U.S.A.

Tel.: 510 642 3567.

Fax: 510 643 8238.

BIBL. DU

MUSEUM

re 4 Source : MNHN, Paris

AILTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1994

Chief Editor: Alain Dupois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

naturelle, 25 rue Cuvier, 75005 Paris, France).

Deputy Editor: Günter GOLLMANN (Institut für Zoologie, Universität Wien, AlthanstraBe 14, 1090

Wien, Austria).

Editorial Board: Ronald G. ALTiG (Mississipppi State University, U.S.A.); Emilio BALLETTO (Torino,

Italy); Alain CoLLENOT (Paris, France); Tim HALLIDAY (Milton Keynes, United Kingdom):

W. Ronald HEYER (Washington, U.S.A.); Walter HôDz (Wien, Austria); Pierre JoLy (Lyon,

France); Masafumi Maïsui (Kyoto, Japan); Jaime E. PÉFAUR (Mérida, Venezuela); J. Dale

ROBERTS (Perth, Australia); Ulrich SINSCH (Koblenz, Germany); Marvalee H. WAKE (Berkeley,

USA).

Technical Editorial Team (Paris, France): Alain Dumois (texts); Roger BoUR (tables); Annemarie

OHLER (figures).

Index Editors: Annemarie OHLER (Paris, France); Stephen J. RicHarDs (Townsville, Australia).

GUIDE FOR AUTHORS

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with

amphibians. Beside articles and notes reporting results of original research, consideration is given for

publication to synthetic review articles, book reviews, comments and replies, and to papers based

upon original high quality illustrations (such as color or black and white photographs), showing

beautiful or rare species, interesting behaviors, etc.

The title should be followed by the name(s) and addresses) of the author(s). The text should

be typewritten or printed double-spaced on one side of the paper. The manuscript should be organized

as follows: English abstract, introduction, material and methods, results, discussion, conclusion,

French or Spanish abstract, acknowledgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or Table IV. Figures should

not exceed 16 X 24 cm. The size of the lettering should ensure its legibility after reduction. The

legends of figures and tables should be assembled on a separate sheet. Each figure should be

numbered using a pencil.

References in the text are to be written in capital letters (SOMEONE, 1948; So & So, 1987;

EveryBoDY et al., 1882). References in the literature cited section should be presented as follows:

BourRET, R., 1942. - Les batraciens de l'Indochine. Hanoi, Institut Océanographique de l’Indochine:

ix + 1-547, pl. I-IV.

GRAF, J.-D. & PoLLs PELAZ, M., 1989. - Evolutionary genetics of the Rana esculenta complex. In:

R. M. DAWLEY & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The

New York State Museum: 289-302.

INGER, R. F., Vois, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some

Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet., 12: 121-145.

Manuscrits should be submitted in triplicate either to Alain Dumois (address above) if dealing

with amphibian morphology, systematics, biogeography, evolution, genetics or developmental

biology, or to Günter GOLLMANN (address above) if dealing with amphibian population genetics,

ecology, ethology or life history.

Acceptance for publication will be decided by the editors following review by at least two

referces. If possible, after acceptance, a copy of the final manuscript on a floppy disk (3 % or 5 4)

should be sent to the Chief Editor. We welcome the following formats of text processing: (1)

preferably, MS Word (1.1 to 6.0, DOS or Windows), WordPerfect (4.1 to 5.1, DOS or Windows) or

WordStar (3.3 to 7.0); (2) less preferably, formated DOS (ASCII) or DOS-formated MS Word for the

Macintosh (on a 3 % high density 1.44 Mo floppy disk only). |

No page charges are requested from the author(s), but the publication of color photographs is

charged. For each published paper, 25 free reprints are offered by Alytes to the author(s). Additional

reprints may be purchased.

Published with the support of AALRAM L

(Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum National d'Histoire Naturelle, Paris, France).

Directeur de la Publication: Alain DuBois.

Numéro de Commission Paritaire: 64851.

Alytes, 1994, 12 (1): 1-48.

Contents

Masafumi MATSUI, Anatolii M. BASSARUKIN, Kiyoshi KASUGAI, Shingo TANABE & Sen

TAKENAKA

Morphological comparisons of brown frogs (genus Rana)

from Sakhalin, Hokkaido and Primorsk ................................ 1

Jon LOMAN

Site tenacity, within and between summers,

of Rana arvalis and Rana temporaria .…. 15

Dominique AUGERT & Pierre JOLY

Dispersal of Rana temporaria tadpoles in large fishponds ............... S1

Robert SCHABETSBERGER & Alfred GOLSCHMID

Age structure and survival rate in Alpine newts

(Triturus alpestris) at high altitude 41

Announcements

Dumerilia 30

Meetings 48

Alytes is printed on acid-free paper.

Alytes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 1° trimestre 1994.

Source : MNHN, Paris