Asiatic Herpetological Research

April 1990

Studies on Hynobiid Salamanders, With Description of aVol. 3, pp. 37-45

New Genus

KRAIG ADLER AND ERMI ZHAO2
1^Cornell University, Section of Neurobiology and Behavior, Mudd Hall, Ithaca, New York 14853, USA
2

Chengdu Institute of Biology, P.O. Box 416, Academia Sinica, Chengdu, Sichuan, China

Abstract. -The types of Hynobius chinensis

Giinther, 1889, were reexamined and are redescribed; theCai, 1985, is relegated to the synonymy ofknown range in China is mapped. Hynobius yiwuensis
chinensis.

Hynobius retardatus Dunn, 1923, of Japan, differs markedly from all other Hynobius and is
The new genus is characterized and compared to the other eight

here placed in a new genus (Satobius ).

genera of hynobiids.

Key words:

Amphibia, Caudata, salamanders, Hynobiidae, Hynobius, Satobius, China, Japan.

We have examined Gunther's two

Introduction

As part of our work on a handbook of
Chinese amphibians and reptiles, we have
had to re-study certain Chinese hynobiid
salamanders described in the literature andconsider their relationships to extralimital
species, especially those native to Japan.
This paper is a result of these current
investigations and concerns the identity anddistribution of Hynobius
chinensis
Giinther, 1889, the taxonomic status of H.

yiwuensis Cai, 1985, andthe proper

generic status of a Japanese species, H.
retardatus Dunn, 1923.

Results and Discussion

syntypes (British Museum [Nat. Hist.]
numbers 1946.9.6.54 and .55, formerly
catalogued as 1889.6.25-26). Both arefemales, as judged from the external
appearance of their cloacae (the specimens
are too hardened to examine internally).

Based on our examination of the types,
H. chinensis is a true Hynobius as definedby Zhao and Hu (1984, English translation
1988). Since Gunther's (1889) originaldescription is brief, we provide thefollowing redescription: Head large, its
length from snout to gular fold longer thanits width (15.2 x 10.6 mm; 12.8 x 9.5mm); tip of snout rounded. Eyes are
dorsolateral in position, slighdy protruded;

and Distribution ofchinensis
Giinther, 1889
Hynobius
1. Identity

1889, Albert Giinther described
Hynobius chinensis from two specimens
collected in Hubei Province. Previously,
In

Hynobius species had been known only
from Japan and Korea, thus Gunther'srecords from central China were quite
isolated from the ranges of the then-known
species. Since 1889, however, no otherspecimens of Hynobius have been reported
from Hubei, despite much collecting there,leading Zhao and Hu (1983, English
translation 1988) to suggest that either the
locality data for Gunther's specimens arewrong or that chinensis is not, in fact, a
species of Hynobius.

1990 by Asiatic Herpetological Researchdiameter of eye shorter than the distancefrom its anterior corner to the tip of snout;
pupil rounded. Nostril between eye and tipof snout, and slightly closer to the latter;distance between nostrils slightly larger or"V"equal to the distance between eyes.
on
of
head.
No
labialshaped bulge
top

A

fold.

An indistinct gular fold. Angle ofjaw just behind the posterior corner of eye.
Both maxilla and mandible with tiny teeth.
Tongue elliptical, large, almost covering theentire floor of mouth. Series of vomerine"
teeth
1/ "-shaped, the outer branchcomprising 6-9 and inner branch 11-15
teeth; the angle formed by outer and innerbranches just beyond the anterior margin ofchoanae; inner branches much longer thanthe outer ones and extending backwards tothe level of the middle of eye ball; theVol. 3, p. 38

FIG. 1.

Asiatic Herpetological Research

April 1990

Map of central China illustrating all known localities for Hynobius chinensis. The inset shows

the location of the more detailed map. Locality records in Zhejiang and Fujian provinces are noted by solid
circles; the star-shaped symbol in Hubei Province is the type locality (Yichang). Other localities mentioned

hollow circles; some major cities are added for reference. The river mapped in
Sichuan Province (upstream from Chongqing) that is continuous with the Chang Jiang (Yangtze River) is
known variously as the Dajin, Dadu, and Min.
in the text are noted by

Gunther (1889) statedposterior ends of two inner branches close
but do not meet at midline. Body short andType Locality:

limbs well developed and tips ofwhen limbs adpressed. Costal
meetdigitsgrooves 11, very prominent and meeting on

Pratt at Ichang [=Yichang]."

ventral midline. Fingers four, 2-3-4-1 inorder of length, the first finger almost equal

localities are mapped in Fig. 1.)

stout;

in length to the fourth.

Toes five, 3-4-2-5-

1 in order oflength. Digits flattened, free;
without palmar and tarsal tubercles; nocornified covering on palms, tarsa, fingers,
and toes. Tail length shorter than snout-

vent length; tail compressed, but cylindricalat the base and pointed at the end, without
crest on ventral edge and only slightly soon its dorsal side. Skin smooth. The

dimensions cannot be re-measured due tothe specimens' hardened condition; only
head length and width can be given(above). Further details about the types are

given in Zhao and Adler (1989). Cai et al.

(1985) have described the embryonicdevelopment and larval features ofchinensis.

BMNH
H

Lectotype: We hereby designate
1946.9.6.54 as the lectotype ofchinensis.

.

that "two specimens were collected by Mr.

(Here andbelow, modern spellings of place names,

where different, are given in brackets. All
A. E. Prattexplored China during 1887-1890 andsummarized his experiences in a book(1892). As Gunther noted in the appendix
to Pratt's book, herpetological specimens
were collected at various localities in Hubei

and Sichuan provinces, and he repeated the"Ichang" locality for the two Hynobius
Pope and Boring (1940),
however, stated that "Mell ('29) claims thatwerePratt's records from 'Ichang'collected in the mountains south of thespecimens.

River near Changyang."nosuch paper by Mell is listed
Curiously,in the bibliography of Pope and Boring's
monograph, nor can we find such astatement in any of Mell's publications, so
we are unable to verify the basis of Popeand Boring's information attributed to Mell.

Yangtze

In his book, Pratt (1892)

mentioned

finding salamanders only twice (pages 179
and 224) but both localities are in Sichuan:a lake above Ta-tsien-lu [=Kangding], atAsiatic Herpetological Research

April 1990

14,070 feet elevation, and at Kia-ting-fu[=Leshan], both of which are localities far
to the west of Yichang (see Fig. 1). Fourother new species of reptiles andamphibians were described by Gunther in
his 1889 paper, all having "Ichang" as thentype locality, and for these taxa the locality
has never been questioned, to ourknowledge. Thus, we are inclined toaccept the type locality as stated by
Gunther, until new evidence comes tohand, although this locality is about 700900 km west of the other known localities
for this species (see Fig. 1 and below).

Distribution:

Subsequent to Giinther'sreport, several other specimens of Hchinensis have been reported. Here wesummarize all known records (see Fig. 1):.

Fujian: Kuatun [=Guadun, 27° 42' N
117° 50' E, a town at Mt. Wuyi],Ch'ungan Hsien [=Chong'an County, 27°
46' N 118° 01'E] (Pope, 1931).
Hubei: Ichang [=Yichang, 30° 42' N 111°
17' E](Gunther, 1889).

Zhejiang: Dachen, 29° 28' N 120° 06' E,
140 m, and Chalin, (both in Yiwu County,
120° 04' E), Zhenhai, 29°
29° 18' N
121° 42' E, and Xiaoshan, 30°
57' N
10' N
120° 15' E, (Cai, 1985; types of
H. yiwuensis); Wenling, 28° 22' N 121°
22' E,

Vol. 3, p. 39

Unfortunately, he was unable to compare
his specimens directly with the types of H.
chinensis. Based on our comparison withthose types, however, we believe that thebase ourtwo taxa are synonymous.
conclusion primarily on the six features
used by Cai to diagnose his new form.

We

Vomerine Teeth. Cai described the lengthof the inner branch of vomerine teeth inyiwuensis as longer than that of chinensis,
yet our examination reveals no significant
difference.
Moreover, the shape of thevomerine teeth series is very similar and thenumbers of teeth in both the inner and outerbranches are within the same ranges in thetwo forms.

Head and Body Proportions. Cai statedthat in yiwuensis the head is much longer
than

broad,

incontrast

to

Giinther'smeasurements for one specimen in which
the length was only slightly greater (11 x
10 mm). Our re-measurement of Giinther's
specimens, taking snout to gular fold ashead length, shows that in both types ofchinensis the head is much longer thanbroad (see measurements in section 1,
above). Cai also claimed that chinensis andyiwuensis

differsomewhatinproportions, the latter being less stout.

body
Our

to notice significant
differences, when relative sizes are taken
into account.

comparisons failed

1500 ft., (Boring and Chang, 1933;

Adpressed

Chang, 1933).

Limbs.

In both types ofchinensis, the tips of the digits touch whenbut

Boring and Chang's specimens from
Zhejiang Province have been examined anddiagnosis states that in yiwuensis the tips

their identifications verified. Cai's species,

usually do not touch.

a synonym of
yiwuensis,
chinensis (see section 2, below). Thus,

specimens there is variation in this character

H

.

is

H

.

chinensis is the only Hynobius found in

mainland China, except for two species inthe extreme northeast; three other species
are endemic to Taiwan.

the

limbs

are

adpressed,

However, in Cai'sand in about one third of them the tips ofthe digits do meet
Costal Grooves.

In his diagnosis, Cai
stated that yiwuensis has 10 costal grooves,in contrast to chinensis which has
2. Taxonomic Status of
Hynobius yiwuensis Cai, 1985

Cai's

1 1 .

We

have confirmed the presence of 1 1 in thetypes of chinensis. However, in Cai's type
series there are a few individuals having 1 1

Cai (1985) described Hynobius
yiwuensis as new, based upon a series of

costal grooves, as Cai himself even noted

adults (16 males, 11 females), juveniles,
larvae, and eggs from Zhejiang Province.

specimens of chinensis from Wenling,
about 160 km southeast of Cai's type(1985,

p.

110).

In

a

seriesof tenAsiatic Herpetological ResearchVol. 3, p. 40

TABLE 1.

Comparison of the genera of hynobiid salamanders. See text for further details.

Bairachuperus*

Distribution

April 1990

Hynobius

Ltua

Onychodactylus

Pachy hynobius*

Pachypalaminus* Ranodon*

Salamandrella

Saiobtus

Asiatic Herpetological Research

April 1990

Vol. 3, p. 41

is a synonym ofDunn (1923a) noted that H.

chromosome number of 40; larval durationof one year or more (neoteny sometimes

retardatus was a "well-marked species"
and, as knowledge of this animal increased

occurs); and both terrestrial and aquatic
habits in adults during non-breeding
season.

inJapan(fuscusnigrescens).

insucceeding years,

its

distinctivenessfrom other Hynobius and hynobiids
generally became more apparent.
In 1932, Makino reported that H.
retardatus has a diploid somatic
chromosome number of 40, but since thecomparable numbers for most otherhynobiids were not known at that time, thesignificance of this very low number
was not recognized. In fact, retardatus has
by far the lowest number of chromosomesin the family Hynobiidae (Table 1). Only
full

in 1943, with Sato's magnificent review ofJapanese salamanders, in which he

These characteristics are discussed
below and, for easy access, are tabulated
for each of the nine genera of hynobiid
salamanders (Table 1).
Costal Grooves. These vertical grooves on
the side of the body correspond to theposition of ribs and, thus, to trunk
vertebrae; generally, the number of costal
grooves that can be counted is one less than
the number of trunk vertebrae.

The typical number of costal grooves inincluded a special comparative study of
their chromosomes, could the matter beproperly evaluated, and Sato himself(1943, p. 489) suggested that retardatus
might be worthy of generic rank.
Unfortunately, Sato's premature death inAugust 1945, during the atomic bombingof Hiroshima (for a biography of Sato, see

5. retardatus is 11.

Adler, 1989), prevented him from pursuing

Adpressed Limbs. As a relative measure oflimb length, the minimum distance between
the tips of the digits is determined with thelimbs adpressed along the sides of the
body. Distances are then measured inintercostal spaces, the fleshy folds between

this matter.

In subsequent years, additional

data on anatomy, karyology, and biology

have accumulated which further support the
separation of retardatus into a new genus,
which we name:In the Japanese speciesof Hynobius the modal numbers of grooves

range from 11 to 13 (Misawa, 1989) andsome mainland species have as many as 14
For species in other(Dunn, 1923b).
hynobiid genera, the modal numbers of
costal grooves range from 10 to 14.

adjacent costal grooves.

Satobius, new genus
In metamorphosed adults of 5
retardatus the intercostal distance between
the digits of adpressed limbs is +1 to +4;that is, because the limbs and toes are very.

Type

Species: Hynobius

retardatus

Dunn, 1923a.
Content: A single species.

A

long, the digits actually overlap by one tofour intercostal spaces. Proportionately,

genus of hynobiid(family
Hynobiidae)
characterized by very long limbs and toes

these are the longest limbs found in any
member of the entire family. In species ofHynobius, this measurement ranges from 5of digits of adpressed limbs overlap
to +4 intercostal spaces in adults); avery long tail (in adults, 100 to 118% of

to +3; among hynobiids other thanHynobius and Satobius, the longest limbs
are found in Pachypalaminus (-2 to 0), buthead and body length combined); a longneck and small head; no premaxillary
fontanelle or basi branchial radii; two shortseries of vomerine teeth arranged intransverse arcs between the internal nares;
vomer sutured to anterior end of

the toes are shorter than those of Satobius.

Diagnosis:
salamanders
(tips

+1

parasphenoid; lungs present; diploid (2«)

The tail of 5. retardatus, as measured
from posterior angle of the vent, is longer
than the combined measurements of headTail.

plus body length. In adults, tail length
varies from 100 to 118% of head-bodyAsiatic Herpetological ResearchVol. 3, p. 42

length, whereas in all other hynobiids it isshorter except in Ranodon (about 100-120% of head-bodylength)andthe total width of theinChromosomes.

significantly shorter than the head-bodylength (60-80%); only in the adults of H.
nigrescens does the tail occasionally equal

diploid

Apparently in all

hynobiids, tail length relative to head-bodyincreases with overall size, so the numbers
given here are all taken, for comparative

purposes, from large adults.

Vomer and Vomerine Teeth. The paired
vomer bones (or prevomers for those who
deny homology to the vomer of mammals)of the palate bear teeth near their posterior

The relationship between thevomers and the parasphenoids lying
posterior to them varies among hynobiid
edges.

1).

first

Makino (1932) was theto report that S.

retardatus has achromosome number of 40,

asconfirmed by others (Azumi and Sasaki,
No other
1971; Morescalchi, 1975).
is
known
to
have
fewer
than 56.
hynobiid
In Japanese species of Hynobius, the socalled pond-type species are 2n=5 6(retardatus is a pond breeder) and themountain brook types are 2/i=58 (except H.
okiensis where 2n=56); H. kimurai isordinarily 2n=56, except in one population(2n=60) which may represent a separate
species (Morescalchi, 1975; Morescalchi etal., 1979; Ikebe et al., 1989). The Korean
H. leechii is 2n=56 (Makino, 1934), but
the chromosome number is not known for

any of the Chinese species of Hynobius.

In S.

retardatus, theposterior edge of the vomer is sutured togenera (Table

two series (versus

30% in 5. retardatus).

Onychodactylus (about 100-115%). The
tail lengths of Hynobius species arehead-body in length.

April 1990

the parasphenoid and overlaps very little, if

some specimens, onto the palatal
surface of the parasphenoid (Inukai, 1932;at all in

Sato, 1943). The two series of vomerine
teeth extend broadly between the internalnares in two slight arcs which nearly meet

midline; the overall length of thisofteeth along the midline is about
patch
30% of the width of the series.
at theRecent studies by Japanese and Italian
workers show that the karyology of S.
retardatus is even more different from thatof Hynobius species than the low numberof chromosomes would suggest.
The
combined lengths of the chromosomes in
the genomes of Hynobius species and inretardatus are nearly equal at the same
degree of condensation, and the amount of
nuclear

DNA is also approximately equal(Morescalchi,

The general pattern of these teeth is likethose in the genera Onychodactylus andRanodon, and quite unlike that in Hynobius
where the vomers (and the vomerine teethon their posterior surface) extend onto theparasphenoids to a degree varying from1975).

Despitethesesimilarities, there are important differencesthat set retardatus apart from species ofHynobius. Kuro-o et al. (1987, as

modified in 1989) were able to compare

chromosome pairs in four Hynobius (3Japanese and 1 Korean species) andIn some Hynobius thisretardatus, using the R-banding techniqueleechii and(RBG method), allowing them to identify

lichenatus) but in most it extends onto thepalatal surface for from one-third to as
much as one-half the length of the

18 of 28 pairs in Hynobius and 16 of 20 inretardatus. Based on this analysis,
chromosome pairs 2 and 8 of retardatus areparasphenoids (e.g., H. formosanus andsonani) (Sato, 1943). Since the vomerine
teeth are located on this edge of the vomer,in these Hynobius the vomerine series has a

not at all represented in the genomes ofthese Hynobius species, whereas pairs 2,

species to species.

overlap

is

small(e.g.,

H.

pattern wholly unlike that in retardatus,
beginning at the nares and extending far
posteriorly on the palate in the shape of alyre (see Table 1). In H. formosanus, forexample, the length of the vomerine teeth

along the midline of the palate is fully twiceand 22, found in all four
Hynobius, were lacking altogether in thatof retardatus; other chromosome pairs were
12,

20,

completely (11 pairs) or partially (3 pairs)
homeologous. To summarize, among thesefour Hynobius
species homeologies
totalled about 90%, but this value fell to

65% when retardatus was included. DNA

Asiatic Herpetological Research

April 1990

was

also performed by theseauthors, using highly-repetitive DNAs asprobes on Southern blot hybridization,
which showed that retardatus was
distinctively different from the five

analysisHynobius species tested (4 Japanese and 1
Korean).

Breeding

Biology.

The reproductive

biology of 5. retardatus has been studied indetail (Sasaki, 1924; Makino, 1933; Sato,
1989 and references cited therein). The
larval period is normally less than a year,
but at high elevations can take more than
one year. Neotenous individuals are wellknown (Sasaki, 1924). Adults breed in
ponds and are terrestrial during nonreproductive periods. However, unlike
species of Hynobius which remain on land
while not breeding, retardatus often visits
the water during non-breeding periods(Sasaki, 1924).
Distribution. Satobius retardatus is foundonly in Hokkaido, the northernmost of themain Japanese islands. No Hynobius are
known from Hokkaido and, among
hynobiids, only Salamandrella keyserlingii
is found there.
Relationships. Historically, 5. retardatus
has been said to be most closely related to

H. nigrescens of neighboring Honshu
Island, Japan (Sato, 1943). Indeed, thetwo species are superficially similar in that,as adults, both are blackish in color withlittlepattern and both possess relatively

long tails.
examination,

However,theseonclosertwo

species arefundamentally different. H. nigrescens is alarge-headed and short-necked species, as
are all Hynobius, when compared with S.
retardatus. The skull of nigrescens, inparticular the vomer-parasphenoid
relationship and the pattern of vomerine
teeth, are typical of Hynobius and unlike

Vol. 3, p. 43

Several of the diagnostic characteristics
of S. retardatus are similar to those of
members of the R a no don group ofhynobiids, which includes Batrachuperus,
Liua, Onychodactylus, and Ranodon (Zhaoand Hu, 1984). The suturing of the vomer
and parasphenoid, the pattern of vomerine
teeth, and the relative lengths of limbs andtail in retardatus are similar to the condition
in one or more members of the Ranodon
group, but in most other respects Satobius
is clearly a member of the Hynobius groupof genera: Hynobius, Pachypalaminus(synonymized with Hynobius by Nishio etal.,1987) andSalamandrella(fordiscussion of these two groups of genera,
Within thissee Zhao and Hu, 1984).
group, the closest living relative ofretardatus may be H. leechii of Korea andnortheastern China, according to theirchromosome structure; indeed, as Ikebe et(1989) point out, based on C-banding
patterns, retardatus is more similar to thenorthernmost populations of leechii rather
than those in the southern part of theal.

Korean peninsula.

We suggest that the ancestral stock
leading to S. retardatus was derived from a
Hynobius-likc ancestor and arrived inJapan from the mainland very early, before
the formation of the Tsugaru Strait which
later isolated Hokkaido from the southern
Japanese islands. Satobius differentiated inJapan but was later excluded from the more
southern Japanese islands by a later
invasion of hynobiids from the mainland.
Thus isolated by the Tsugaru barrier,Satobius further differentiated, yet retainedsome of its primitive characters, including a
few today found only in the Ranodon

group.

The long isolation from Hynobius has
led also to the chromosomal differentiation

described earlier (although retaining somesimilarities to H. leechii on the mainland).

the situation in Satobius. Furthermore, thenumber of chromosomes and details of

The great reduction in chromosome number

chromosome structure, based on both C-

in retardatus may be due in part to fusions,banding and R-banding methods, aresince the total composite lengths andcontent are approximately the same as for

distinctly different. In short, we believethat the similarities between nigrescens and

DNA

the

retardatus are due to convergence and do

Hynobius genome (Kuro-o et al.,However, in view of the many1989).

not reflect close phylogenetic relationship.

differences based on the C-banding and R-

Asiatic Herpetological ResearchVol. 3, p. 44

banding studies mentioned above, thekaryotype of retardatus cannot be explainedby a simple mechanism of fusions alone.

April 1990

Bulletin 8:75-80.

DUNN, E. R.

1923a.

New species of Hynobius

from Japan. Proceedings of the California

Etymology. We take great pleasure innaming this new genus for Ikio Sato (1902who first recognized its
1945),
distinctiveness from Hynobius. The nameSatobius is derived in part from the stem ofHynobius (hynis [or hynnis], Greek forplowshare, the cutting part of a plow, andbios, life).

Academy of Sciences, ser. 4, 12:27-29.

DUNN, E. R.
family1923b.

The salamanders of theHynobiidae.

Proceedings of theAmerican Academy of Arts and Sciences
58:445-523.

GONTHER, A.

1889.

Third contribution to ourknowledge of reptiles and fishes from the Upper
Annals and Magazine of
Yangtsze-Kiang.

Acknowledgments

Natural History, ser. 6, 4:218-229.

Our research has been supported by the

USA

National Academy of Sciences(Committee on Scholarly Exchange with
the People's Republic of China). We thankBarry Clarke (British Museum [Nat. Hist.])and Robert C. Drewes and Jens V. Vindum(California Academy of Sciences) for theloan of specimens.
We thank Satoshi

Amagai (Cornell University) for sometranslations of articles in Japanese.

IKEBE, C, M. KURO-O, T. OHHASHI, H.

YAMADA, K.-I. AOKI, AND S.-I. KOHNO.
1989.

Evolution of chromosome 10

INUKAI, T. 1932. Urodelenarten aus Nordjapan
mit besonderer Beriicksichtigung derMorphologie des Schadels.

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Transactions of the Sapporo Natural History
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