2004

Asiatic Herpetological ResearchVol. 10, pp. 168-175

First Description of Egg Sacs and Early Larval Development in
Hynobiid Salamanders (Urodela, Hynobiidae, Batrachuperus)
from North-Eastern Iran
MEHREGAN EBRAHIMI1, HAGI GHOLI KAMI2, MATTHIAS STÖCK3
1Department of Biology, Faculty of Sciences, University of Shiraz, P.O.Box:71454-1460,

Shiraz, Iran; E-mail: emehrgan@yahoo.com
2Department of Biology, Faculty of Sciences, University of Agricultural Sciences and Natural Resources,

P.O.Box:49165-386, Gorgan, Iran; E-mail: hgkami2000@yahoo.com
3Museum of Vertebrate Zoology, University of California, Berkeley, Department of Integrative Biology,

3101 Valley Life Sciences Building #3160,Berkeley, CA 94720-3160, USA; E-mail: matthias@berkeley.edu
Abstract. - Here we present the first life history data from a natural habitat on the clutch and from the laboratory on
the early embryonic development of an Iranian hynobiid salamander (the Eastern nominal taxon, Batrachuperus gorganensis). We compare these observations with those of other hynobiid species. The egg sac of B. gorganensis varied from 80 to 182 mm (mean 132.5), and in width from 16 to 22.5 mm (mean 18.9). Single sacs, the largest known
for the genus, contained 31-52 eggs (mean 37.4) in four rows per cross section. Egg sac data are more similar to thoseof Ranodon sibiricus (and B. mustersi) than to Eastern Batrachuperus (longdongensis, tibetanus, pinchonii,
karlschmidti, yenyuanensis), which have strings of single eggs. As is typical of all Batrachuperus, eggs of B. gorganensis are non-pigmented. Embryonic B. gorganensis larvae may exhibit rudimentary balancers, possibly like thoseof Ranodon sibiricus, but further investigation is necessary. Detailed morphometric measurements of the larvae of B.
gorganensis and color photographs of some developmental stages are presented.
Key words. - Hynobiidae, Batrachuperus, Batrachuperus gorganensis, Batrachuperus persicus, Batrachuperus
mustersi, Ranodon sibiricus, Iran, egg sacs, embryonic development, larvae, balancer, taxonomy, morphometrics.

Introduction
The southwestern-most representatives of the salamander family Hynobiidae occur within the narrow, EastWest oriented Hyrcanian Corridor (= Hyrcania) of
Northern Iran. The corridor is a “unique relict biogeographic area,” which is “well defined as the south-western and southern shores of the Caspian Sea ...” and is“one of the most clearly defined and delineatedprovinces in the Irano-Turanian region“ (Fet, 1994). The
Persian mountain salamander, Batrachuperus persicus
Eiselt and Steiner, 1970, was based on larvae collectedfrom the western part of Hyrcania (NW-Iran, mountains
near Asalem, Gilan province; Eiselt and Steiner, 1970).
Subsequently, specimens that metamorphosed in captivity were described and several additional localities werereported (see Schmidtler and Schmidtler, 1971; Steiner,1973).
The second nominal taxon, Batrachuperus gorganensis (Clergue-Gazeau and Thorn, 1979), wasdescribed from a single adult male type discovered in acave from the eastern edge of the Hyrcanian Corridor(Clergue-Gazeau and Farcy, 1978). Baloutch and Kami
(1995), Kami and Vakilpoure (1996) as well as Kami
(1999, 2004) published additional data on the biology
and distribution of Iranian hynobiid salamanders, all© 2004 by Asiatic Herpetological Research

assigned to B. persicus. Stöck (1999) provided a flowcytometric DNA measurement (34.77 pg, but based on
GC-biased DAPI-staining, see also Litvinchuk et al.,2004) and a Giemsa-stained karyotype (2n = 62) of
topotypic B. gorganensis (i.e. the eastern taxon). Stöckalso described external larval changes during the development from 40 mm until metamorphosis (100 mm),and reviewed previous papers (see also map with geographic coordinates of all previously published localities– Stöck, 1999: Fig. 1). After examination of topotypicsubadults and larvae of both taxa as well as morphometric comparisons with B. mustersi, B. pinchonii, andRanodon sibiricus, Stöck considered a clinal variation ofcharacters between the western (B. persicus) and theeastern nominal taxon (B. gorganensis) possible (seealso Thorn and Raffaëlli, 2001: 122). However, molecular and cytogenetic data as well as a continuous examination of the morphological variation throughout the
Hyrcanian corridor are lacking.
Both nominal Iranian taxa (cf. Anderson, 1985;
Brame, 1985: 567; Duellman and Trueb, 1986: 497;
Thorn and Raffaëlli, 2001) are considered two out ofcurrently nine species of the genus Batrachuperus(Frost, 2002). Risch (1984) regarded gorganensis tobelong to a separate genus (Paradactylodon; see Reilly,
1987). While the molecular phylogeny of eastern

Vol. 10, p. 169

Asiatic Herpetological Research

2004

ganensis) from a natural habitat. We add some data on
its development in the laboratory and compare theseobservations with the features known from other hynobiid species.

Results and Discussion

Figure 1. Locality: a. Sketch map of northern Iran with its
forested regions and the study area, the educational forest of Shastkola; b. Shastkola forest with districts I and II
and streams. The left arrow marks the Shastkola river; c.
District II of Shastkola forest shown in (b) with its 31 landparcels, the spring of Manzoulak is situated in parcel 14.

Batrachuperus has been examined (Fu et al., 2001),
molecular comparisons of their relationships with
Western Batrachuperus and Ranodon as well as between
the latter taxa are not yet available. However, unpublished results (Macey et al. in prep., with phylogram presented in Larson et al., 2003: 44, Fig. 2.4) provide evidence that Iranian Batrachuperus may be more closely
related to Ranodon (Kazakhstan, China) and B. mustersi (Afghanistan) than to the Eastern Batrachuperus
species (Tibet, China). Ranodon sibiricus has five toeson the hind limbs while B. mustersi,and B. persicus/B.
gorganensis have only four. However, the intraspecificvariation found by Reilly (1983) in B. mustersi, whichsometimes has five instead of four toes, demonstrates
how ambiguous this character can be.
In general, there is still a lack of biological data on
the Iranian taxa, e.g., their egg sacs and early larvae
have never been examined. Only remains of egg cases ofB. persicus were found by Steiner (1973) in late July in
the Central Elburz range and the reports on larvae startfrom a minimum size of 22.7 mm (Central Iran; Steiner,1973) and “about 30 mm” (NW-Iran, Schmidtler andSchmidtler, 1970). Here we present the first data on the
clutch of the Eastern Hyrcanian hynobiid taxon (B. gor-

Spawning site. - All clutches (Table 1) were found in
Cheshme-ye (= spring of) Manzoulak (36°42´ N, 54°21´E), which flows into an artificial pond (3.20 x 2.15 x
1.25 m). Its bottom is covered with a thick layer (ca. 15cm) of tree leaves. Some crabs (Potamon sp.) and larvaeofOdonata,
Trichoptera,
Coleoptera,
Ephemeroptera were observed. Manzoulak spring is oneof the important water resources of the Shastkola educational forest (3716 ha), which is situated 13 km SouthWest of Gorgan city between 36° 41´- 36° 45´ N and 54°
20´ - 54° 24´ E, on the northern slopes of the Elburz
mountains (Fig. 1). These mountains reach elevations
ranging from 240 m, above the Caspian Sea in the North
to 2168 m (Gholle-ye-leila Kouh). Shastkola forest isdivided into two organizational units, districts I (1698.6
ha) and II (2017.4 ha). The spring is situated in II [Fig.
1c, land parcel 14 (75 ha), 650 m to 830 m]. Outcrops
consist mainly of a sequence of sandstones and schist in
the lower part and Triassic sand stones in the upperparts. Shastkola forest has a Caspian climate, and thenearest climate station is Gorgan (155 m a.s.l.).
However, local climate at the breeding site differs considerably from that at the foothills and we currently haveno detailed climatic data on the site. Water temperatureduring collection was below 10°C. The forest tree andshrub species include Fagus orientalis, Carpinus betulus, Parrotia persica, Ruscus hyrcana, and Ilex aquifolia (for further information: Azadfar, 1994; Azadfar andDarghahi, 2002), which adds to the accumulating evidence that the Iranian hynobiid taxa inhabit humid
regions mainly overgrown with the plant assemblage
Fagetum hyrcanum in lower elevations in the West of
their range: Parrotio-Carpinetum; in eastern parts of the
Elburz N-slope: Carpinetum orientalis, Quercus
macranthera; see also ref. in Stöck, 1999).
Description of clutches, eggs and larvae. - About 70
clutches and four adult salamanders were discovered
when leaves were removed from the pond by G. Yolmeh
on January, 28, 2002, but only two of the clutches andone adult specimen (total length: 211 mm) were sent to
the Museum of Zoology of Gorgan University (ZMGU).
In the following year on February, 8, 2003, one complete
and two incomplete clutches were collected in the same
pond by M. Ebrahimi and G. Soltanpour. We presume
that spawning, at least at this site, happened between thesecond half of January and the first days of February.

2004

Asiatic Herpetological ResearchVol. 10, p. 170

Figure 2 (A-J). A. Natural habitat of Batrachuperus gorganensis. Stream near Manzoulak spring, photograph taken:
March, 10, 2003. B. Clutch, preserved, collection date: March, 10, 2003, photograph taken: April, 6, 2003. C. Clutch,
collection date: March, 10, 2003, photograph taken: April, 4, 2003. D. The same clutch as in C, collection date: March,
10, 2003, photograph taken: April, 4, 2003. E. Fixed embryo, 7 days before hatching, photograph taken: October, 9,
2003. F. Fixed larva, 4 days after hatching, photograph taken: October, 9, 2003. G. Living larva, 18 days after hatching, photograph taken: April, 6, 2003. H. Living larva, 18 days after hatching, photograph taken: April, 6, 2003. I. Living
larva, 24 days after hatching, photograph taken: April, 6, 2003. J. Fixed larva, 34 days after hatching, photographtaken: October, 9, 2003.

However, in general, the breeding season in both Iranian
nominal taxa might last as long as in some other
Hynobiids (e.g. in Ranodon sibiricus from the end of
April until the beginning of August - Kuzmin, 1995:
111; from May to beginning of August in B. karlschmidti- Liu, 1950: 91).
During collection, some egg sacs broke. Originally,
the clutches were attached by their stalks to a stone at thecorner of the pond; no egg sacs were observed on its
walls. As is usual in Batrachuperus, each clutch consisted of two asymmetrical, colorless, cylindrical gelatinous
sacs that were connected by gelatinous filaments forming an attaching stalk and had free filaments at theirnonanchored conical ends. As in Ranodon sibiricus (seeKuzmin, 2001: 40), egg sac data varied between andwithin clutches (see Fig. 2 B-D; Table 1). Each cylindrical gelatinous sac contained four rows of eggs per crosssection and each egg was surrounded by a sphericalgelatinous envelope. As is known from Ranodon sibiricus (see Kuzmin and Thiesmeier, 2001: 42 and ref.
therein), these four egg rows produce a tetrahedral shape
immediately after clutch deposition. However, as theresult of an ingress of water, sacs later strongly increasein size. We believe that all discovered clutches had
already finished this first swelling, which usually gradually continues throughout the embryonic development
leading to a considerable enlargement of the sacs until
hatching. Tables 2 and 3 show comparisons of egg saclength and width of eight hynobiid species. Data on
length and width should be evaluated with some reservations, because the number of specimens examined ortime after clutch deposition is imperfectly known.
However, egg sacs of B. gorganensis probably are larger than those of any of the close relatives of the taxon(with the possible exception of B. longdongensis) andappear to contain among the highest number of eggs per

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2004

Table 1. Overview of described clutches of Batrachuperus gorganensis.
Date of Number ofcollection egg sacs

Length of gelati- Maximum
nous egg sac
width of(mm)
gelatinous
Left
Right egg sacs

Total number of eggs
per sac
Left
Right

Date ofhatching

9+9+10+11=39 9+9+8+9=35
34
35

Fixed
Fixed

-

-

March, 19 to
24, 2003
All died
March, 19 to
24, 2003
Fixed
March, 21 to
24, 2003

51

74

64

71.1

28

93.3

January,
28, 2002

1
2

95
80

102
81

16.5
15.8

February,
2, 2003

1

150.5

172.6

-

31

38

2
3

182.5 destroyed
181
181.8

20.8
22.4

32
38

40
52

4
5

100.9

-

-

Small part of an egg sac, 18
Small part of an egg sac, 30

sac (Table 4). In addition, R. sibiricus and B. gorganensis share the common feature of four rows of eggs per
cross section in an egg sac, while all Eastern taxa (B.
longdongensis, B. tibetanus, B. pinchonii, B.
karlschmidti, B. yenyuanensis) seem to have only a single row of eggs in an egg case (judging from photographs and Zhao and Hu, 1988: 32). This featureremains obscure in B. mustersi, since neither Reilly’s(1983) description nor Nawabi’s (1965) photographresolve the question satisfactorily, but more than a single
row appears to be present. Egg number may be a taxonomic character but might not affect fecundity sincesome species seem to deposit more than one clutch per
breeding period (Liu, 1950; Reilly, 1983).
The diameter of the initially yellowish (as in all
Batrachuperus and R. sibiricus) eggs varied from 4.10
to 5.00 mm (mean 4.56; n = 10) when the clutch was
found. The early eggs had a white animal pole whereas
the vegetal pole was pigmented gray. Unfortunately,
early eggs were not photographed. The separation of
eggs in a row was less than the diameter of an egg. Egg
number varied from 7 to 11 per row (2 egg sacs examined) and from 31 to 52 in each sac (see Table 1). The
eggs have substantial amounts of yolk, as also is typicalof stream-breeding R. sibiricus (Kuzmin andThiesmeier, 2001).

Live hatchedlarvaeN%

Three complete clutches and one incomplete clutch
were kept in a refrigerator at 5-7°C (for conditions see
Stöck, 1999). The first larvae to hatch were found on
March, 19, 2003, and the latest specimens to hatch wereobserved on March, 25, 2003 (i.e. 40 to 46 days after
discovery, respectively). Some larvae at different developmental stages were fixed in 4% formalin and 70%
alcohol and measurements were taken from all specimens (summarized in Table 5). We report some larval
developmental traits and link them by inserting theapproximate stage (see inserted number in […]) of normal development for Ranodon sibiricus drawn by
Kuzmin and Thiesmeier (2001: 46, after the descriptionof Lebedkina, 1964).
Hatching larvae [stage 1] exhibited limb buds (Fig.
2F). Although no balancers were observed in hatchedlarvae (Fig. 2F), we believe that a rudimentary structure
observed in an embryo fixed 7 days before hatching(Fig. 2E) could be a balancer. These enigmatic,
ephemeral larval organs, found in three of the ten families of salamanders (Crawford and Wake, 1998), arepresent in many species of Hynobiidae (Crawford and
Wake, 1998: 115). In the presumed close relative of the
Iranian hynobiids, B. mustersi (see introduction), distinct balancers are well documented (Nawabi, 1965, seealso Reilly, 1983). Although such a well developed

Table 2. Egg sac length of eight* hynobiid species, comparison of available data on Batrachuperus mustersi from
Nawabi (1965), Reilly (1983), and Sparreboom (1979); on R. sibiricus (from Brushko and Narbaeva 1988, cited from
Kuzmin and Thiesmeier 2001); B. gorganensis (our data); B. longdongensis, B. tibetanus, B. yenyuanensis from Fei
and Ye (2001), B. pinchonii and B. karlschmidti from Liu (1950) and Fei and Ye (2001); in R. sibiricus numbers in parentheses show rare values in late stages of embryonic development (* some authors synonymized B. karlschmidti with
B. tibetanus, but see Fu et al. 2001 and Frost 2002).
B. gorganensis R. sibiricus B. mustersi
Mean
Max
Min
N

132.4818280
10?
100 (390)
65 (100)
10

96.62
150
60
13

B. longdongensis
200-

B. pinchonii
96
65-

B. karl- B. tibetanus
schmidti
96.6
96
140
75
102
5
-

B. yenyuanensis
125
70
-

2004

Asiatic Herpetological ResearchVol. 10, p. 172

Table 3. Egg sac width of eight hynobiid species, comparison of available data; same sources as in table 2; the only
available value for R. sibiricus is from an egg sac of 72-75 mm length.
B. gorganensis R. sibiricus B. mustersi
Mean
Max
Min
N

18.88
22.4
15.8
10

(23)
-

21.02
27.0
13.2
13

B. longdongensis
15-

organ is absent in Ranodon sibiricus (cf. Crawford and
Wake, 1998: 115), the presence of rudimentary balancers in the species appears possible. Regel (1968: 17,
see “pb” in her Fig. 7, transl. from Russian) wrote on R.
sibiricus larvae in developmental stage 3 (15-17 mm):
“On the lateral surface of the distal end of the palatinoquadratum cartilage an excrescence develops, which is arudimentary proc. balancer (Rusconi’s hook)”. Althoughbalancers are not cartilaginous structures, Kuzmin andThiesmeier (2001: 43) quote this as an indication of
rudimentary balancers in Ranodon. Based on theseobservations, we believe that at least rudimentary balancers will be found in B. gorganensis (and B. persicus?), and we recommend further comparative investigation with histological methods. Secondary loss of welldeveloped balancers may be an adaptation of the“limnophilous mountain brook type” of larvae in B. gorganensis / B. persicus (discussed in Stöck, 1999: 237).
According to available data, balancers may be absent inall species of Batrachuperus from China and Tibet(Table 4).
In larvae eight days after hatching (d.a.h.) [stage 23], the head had differentiated and forelimbs had ashovel-like form, but only hind limb buds were visible.
The yolk sac regressed and was very small in larvae 14
d.a.h. (April, 2nd, 2003). In larvae 16-18 d.a.h. [stage 67] (Fig. 2G-H), the yolk sac had disappeared, dark spots
became visible at the back, and the tail fin developed,
although development of the digits was incomplete(April, 4, 2003). In larvae 24 d.a.h. [stage 8-9] (Fig. 2I),

B. pinchonii
1219-

B. karl- B. tibetanus
schmidti
16.6
19
10
14
12
5
-

B. yenyuanensis
815-

digits were well developed in forelimbs, but the toes ofhind limbs were still incomplete (April, 12, 2003). In
larvae 34 d.a.h. [stage 12] (Fig. 2J) the forelimbs werecompletely developed, digits were clearly distinguishable, and tips of all digits and toes exhibited brown
horny claws. The ventral surface had a milky color, theupper part of the tail and the tail fin showed dark spots,
and the eyes appeared completely developed. Throughtime, the dark spots became more abundant at the tip oftail, and the mouth. Darkly pigmented dots appeared onpart of the head and trunk, the flanks, the upper parts ofthe tail, and the upper tail fin. At this stage, larvae werefed with nauplius larvae of Artemia. In larvae 46 d.a.h.,the belly remained spotless, but the flanks, dorsal bodyparts and upper parts of the tail showed dark spots.
Larvae 62 d.a.h. exhibited brownish spots on the bodyand dark spots became rarer (larger larvae had golden
dots on the gills and on parts of the body). The anterior
part of the upper tail fin extended to the occiput.
In addition to the clutches described above, morethan 20 large larvae with a total length of almost 10 cm
were collected in the leaves of the breeding pond, but
only one adult specimen was found. The pond wassearched completely, but no larvae smaller than about 10
cm were found. Therefore, we conclude that all theselarge larvae apparently stemmed from the preceding
year. The growth rate data of the larvae hatched from theclutches in our laboratory fit well with the later developmental stages reported by various previous authors fromnatural habitats (summarized in Stöck, 1999: Tab. 3).

Table 4. Number of eggs per sac, number of egg rows per cross section and presence of a balancer of eight hynobiid species, comparison of available data; same sources as in table 2, except for B. longdongensis in which the character was inferred from the ovary of voucher MVZ 208610.
B. gorganensis R. sibiricus B. mustersi
Mean
37.40
23.00
22.23
Max
52
44
31
Min
31
7
13
N
10
150
13
Rows of4
4>2
eggs percross
sectionBalancer rudimentary rudimentary? well developed

B. longdongensis
1(-2)?

B. pinchonii

No?

No?

23
5
1

B. karl- B. tibetanus
schmidti
12
25
7
16
5
1
1?

No

No?

B. yenyuanensis
13
6
1?

No?

Vol. 10, p. 173

d.a.h.

1
1
1
1
4
4
4
4
8
8
8
8
14
14
14
14
16
16
16
16
19
19
19
19
24
24
24
24
34
34
34
34
46
46
46
46
54
54
54
54
62
62
62

Asiatic Herpetological Research

2004

distancemin.
max snout numbereyehind distance hight to ant. of recdist.
total head body
body
tail
head diame- eye to foreleg leg between dorsal point ognized mass
length length length1 length2 length width(g)tersnout length length feetfine dorsal costal
fin grooves

MEAN
19.28
MIN
19.05
MAX
19.61
SD
0.28
N = 4 (1)
MEAN
19.30
MIN
18.50
MAX
19.79
SD
0.70
N = 3 (1)
MEAN
21.76
MIN
21.27
MAX
22.20
SD
0.30
N = 8 (1)
MEAN
24.61
MIN
23.13
MAX
25.53
SD
0.92
N = 7(1)
MEAN
27.06
MIN
26.45
MAX
27.93
SD
0.43
N=9
MEAN
24.87
MIN
20.90
MAX
36.90
SD
4.28
N =15 (3)
26.67
MEAN
22.94
MIN
30.15
MAX
SD
2.58
N = 6 (2)
30.46
MEAN
29.04
MIN
32.74
MAX
1.71
SD
N = 6 (2)
31.69
MEAN
28.12
MIN
33.80
MAX
2.03
SD
N = 6 (3)
31.78
MEAN
25.90
MIN
36.20
MAX
3.69
SD
N = 8 (3)
34.65
MEAN
31.82
MIN
37.20
MAX
2.70
SD

2.40 11.05 11.63 7.49
2.25 10.80 11.40 7.06
2.50 11.40 11.80 7.80
0.11 0.25 0.17 0.37

2.54
2.00
2.95
0.40

0.73
0.67
0.80
0.06

0.89
0.77
0.97
0.09

0.65
0.50
0.80
0.15

1.04
1.00
1.05
0.03

4.01
3.50
5.00
0.67

0.05
0.05
0.05
0.00

2.67 11.00 11.73 7.37
2.40 10.50 11.20 7.00
2.90 11.40 12.10 7.60
0.25 0.46 0.47 0.32

2.45
2.30
2.65
0.18

0.77
0.70
0.85
0.08

0.90
0.90
0.90
0.00

0.65
0.55
0.80
0.13

1.12
1.10
1.15
0.03

3.50
3.00
4.00
0.50

0.05
0.04
0.06
0.01

2.79 12.11 12.73 8.86 3.19 0.96 1.06 1.34
2.25 11.80 12.20 8.40 2.75 0.90 0.90 1.00
3.60 12.50 13.00 9.40 3.50 1.02 1.22 1.70
0.53 0.27 0.29 0.37 0.26 0.04 0.11 0.23

1.33
1.10
1.50
0.12

5.09
4.75
5.50
0.23

0.07
0.06
0.08
0.01

3.28 13.68 14.44 10.15 3.61
2.65 12.76 13.50 9.50 2.90
3.70 14.00 14.90 10.70 4.10
0.39 0.41 0.53 0.44 0.47

1.03
0.95
1.10
0.06

1.22
1.00
1.45
0.14

2.14
1.50
2.60
0.35

0.70
0.55
0.90
0.14

7.97
7.45
8.25
0.26

1.45
1.35
1.60
0.09

5.90
4.00
8.00
1.29

11
14

0.10
0.09
0.12
0.01

4.03 14.65 15.54 11.47 3.86
3.25 14.10 15.10 10.36 3.50
4.40 15.54 16.65 12.00 4.00
0.35 0.41 0.48 0.53 0.17

1.19
0.95
1.30
0.10

1.36
1.00
1.55
0.16

2.72
2.00
3.10
0.32

1.39
1.00
1.55
0.19

8.37
8.15
8.60
0.16

1.49
1.40
1.55
0.05

6.56
5.05
8.15
0.86

12
14

0.12
0.11
0.15
0.01

3.28 13.83 14.56 10.09 2.83
2.35 11.80 12.50 8.00 2.00
5.40 19.79 20.53 16.28 5.30
0.80 2.04 2.10 2.32 0.88

1.08
0.75
1.80
0.29

1.08
0.75
1.70
0.28

1.86
0.75
5.20
1.21

1.41
0.25
4.00
1.22

8.56
7.25
9.70
0.88

1.22
0.95
2.15
0.30

5.29
3.00
8.00
1.69

13
14

0.06
0.04
0.21
0.04

3.70 14.75 15.63 10.77 3.11
3.00 13.30 14.10 8.90 2.65
4.80 16.40 17.20 13.00 3.75
0.76 1.26 1.22 1.49 0.43

1.09
0.90
1.30
0.17

1.18
0.85
1.65
0.27

1.78
1.25
2.50
0.44

0.60
0.50
0.75
0.13

8.93
8.00
9.70
0.86

1.30
1.10
1.50
0.14

4.94
3.50
7.75
1.53

4.68 16.33 17.12 13.10 4.15
4.20 15.20 16.10 12.10 3.80
4.90 16.90 17.60 14.20 4.50
0.25 0.60 0.53 0.79 0.26

1.43
1.25
1.65
0.16

1.67
1.50
1.80
0.13

3.15
2.30
3.80
0.62

1.91
0.80
2.85
1.00

8.95
8.40
9.50
0.42

1.58
1.50
1.70
0.08

6.92
6.00
8.50
0.97

4.42 16.58 17.56 13.95 4.18
4.00 16.00 16.80 12.30 3.30
4.75 17.57 18.68 15.30 5.00
0.25 0.56 0.68 1.12 0.72

1.40
1.15
1.60
0.17

1.59
1.20
1.90
0.27

3.53
2.10
4.55
1.03

2.19
1.00
3.70
1.24

8.92
8.50
9.50
0.48

1.45
1.25
1.65
0.15

7.83
7.00
9.00
0.67

4.71 17.24 18.13 14.08 4.23
3.35 14.90 15.70 10.50 3.00
5.50 19.24 20.35 16.66 5.50
0.73 1.65 1.78 2.26 0.94

1.35
1.05
1.65
0.22

1.53
1.25
1.90
0.26

3.32
1.95
4.75
1.03

1.72
0.50
4.10
1.14

8.66
7.60
9.40
0.51

1.61
1.40
2.05
0.22

7.23
5.00
9.50
1.57

12
13

0.13
0.08
0.22
0.05

5.50 18.52 19.43 15.60 4.90
4.60 16.70 17.60 13.50 4.30
6.00 20.35 21.27 16.65 5.40
0.78 1.83 1.84 1.82 0.56

1.38
1.15
1.50
0.20

1.53
1.25
1.75
0.26

4.12
3.35
4.60
0.67

2.40
1.60
3.20
0.80

9.40
8.70
10.00
0.66

1.92
1.55
2.25
0.35

7.90
7.50
8.60
0.61

12
12

0.24
0.16
0.32
0.08

0.08
0.07
0.11
0.02
12
12

0.13
0.11
0.16
0.02
0.10
0.07
0.13
0.02

2004

Asiatic Herpetological ResearchVol. 10, p. 174

Table 5 (previous page). Morphometric data on early larvae of Batrachuperus gorganensis.
d.a.h. = days after hatching; SD = standard deviation; Min = minimum; Max = maximum;
N = number of larvae measured (the value in parentheses shows from how many clutches the measured larvae originated)

According to these studies, larvae from the first season
may reach a total length of 30 to 62 mm (persicus,
between June and August) and 41 to 50 mm (gorganensis, mid of June), respectively, and must overwinterbefore they form “large larvae”, which can enter metamorphosis in the following year.

Conclusions and Future Research
Our initial studies support the relationship of the threetaxa Ranodon sibiricus, Batrachuperus mustersi and the
Iranian hynobiid(s), i.e. B. persicus and B. gorganensis.
These taxa have apparently more features in common
than with the Batrachuperus taxa East of Tibet. In addition to the essential need for molecular data, it would bevaluable and interesting to study egg deposition in thenatural habitats in more detail and to collect data
throughout the range of the Iranian hynobiid salamanders, especially from the central and western portions.
During proof reading, one of us (HGK) provided information that MMTT 452, 453, and 454 are eggs of B. persicus, found near the road from Asalem to Khalkhal, collected May, 20, 1975, by M. Thireau, R. Khazaie and R.
G. Tuck.

Acknowledgments
M. E. and H. G .K. wish to thank G. Yolmeh for providing material and initial information on the spawning
pond, as well as G. Soltanpour for help during the fieldwork. M. S. thanks David Wake for references andadvice, Tate Tunstall for translation from Chinese, S. L.
Kuzmin for data on R. sibiricus, G. Wogan and J.
Vindum for references, Robert Hijmans for cartographic
support, J. Robert Macey and Theodore Papenfuss for
permission to include unpublished information about
relationships of Hynobiidae, João Alexandrino for talking about natural history, James Parham for (im-)
patience with the final version of the manuscript, andRobert Bingham for checking the English.

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