Asiatic Herpetological Research[April 1990

Stelliosacra

Vol.3, pp. 104-115

a Distinct Species of Asiaticfrom Tibet*(Smith 1935)

Rock Agamid

NATALIA B. ANANJEVA GUNTHER PETERS 2 J. ROBERT MACEY 3
AND THEODORE J. PAPENFUSS 3
1,,^Zoological Institute, USSR Academy of Sciences, Leningrad, USSR^Museum fur Naturkunde der Humboldt-Universitdt, Invalidenstrasse 43, Berlin, DDR^Museum of Vertebrate Zoology, University of California, Berkeley, California, USA

Abstract. -An examination of the type series of Agama himalayana sacra Smith (1935) and newmaterial collected in Tibet in 1988 has shown that this form should be considered a distinct species. In

following the recent revision of the genus Agamawith all other Asiatic rock agamids.

Key Words:(Moody 1980), Stellio sacra is included in the genus

Reptilia, Sauria, Agamidae, Stellio sacra, Tibet, systematics, distribution.

Methods

Introduction
Four agamid lizards were collected at the
beginning of the 20th Century near Lhasa,
Tibet.
They were described in a shortaccount by Smith (1935) as a subspecies ofthe Himalayan rock agamid Agama

himalayana sacra.

To

best of our knowledge all
specimens housed outside the Peoples
Republic of China were examined. Of thefour specimens in the type series, one
thelectotype and two paralectotypes are housed
at the British Museum (Natural History)[BMNH], and one paralectotype is housed
The examination of these type specimensat the Indian Museum, Calcutta (Zoologicaland new material collected in Lhasa and thevicinity of Lhasa, Tibet in 1988 by the thirdand fourth authors, led us to the conclusionthat it is not a subspecies of Stellio
himalayanus. Furthermore it is not a

Survey of India [ZSI]). Thirteen additional
specimens were collected during the 1988
joint Chengdu Institute of Biology
University of California Expedition and arehoused at the California Academy ofmember of the Stellio himalayanus species
complex which includes Stellio

Sciences (CAS).

badakhshanus, Stellio chernovi, Stellio
himalayanus, and Stellio stoliczkanus. The

Stelliosacra(Smith 1935), new combination

results of the examination of the agamids

Known Material

from Lhasa, Tibet confirm the opinion of
Ananjeva et al. (1981) that Stellio sacrashould be elevated to full specific status.

Material Examined:

According to the present revision of thegenus Agama (Wermuth 1967) into Agama,

Lectotype:

Stellio(Moody 1980; Sokolovsky 1975,1977) Trapelus, Psuedotrapelus, andXenagama (Moody 1980) the lizards
examined from Tibet should havethegeneric name, Stellio.

BMNH

1946.8.28.57

1904.12.28.1)(formerly
Locality: Near Lhasa, Tibet.

[Fig.

1.]

BMNH

1946.8. 28.58,
Paralectotypes:
1946.8.28.59 [Fig. 2], and ZSI
15740 [Fig. 3]. Locality: near Lhasa,

BMNH
Tibet.

CAS 170545.

*

Sino-Soviet- American Arid Asian Desert Regions

Research Paper no. 5.

fr\

1

fifin u.,Locality:

Elev.

3740 m,Sera Monastery, Lhasa (29° 39' N

91°

Asiatic Herpetological Research

April 1990

FIG. 1. Lectotype of Stellio sacraVol. 3, p. 105

BMNH 1946.8.28.57 (formerly 1904.12.28.1).

Sera Monastery, Lhasa (29° 39' N

06' E), Lhasa Municipality, Xizang (Tibet)

Papenfuss, CAS 170556-57 T. J.
Papenfuss and R. Macey. Date: 27 Sept.,

Autonomous Region, China. Collected by:

1988.

T. J. Papenfuss and R. Macey.
Sept., 1988.

CAS

170546-53.

91°

Date:

Locality:(See Table 1).

24
Other Material.

Elev.

3700 m, at base of mountains approx. 3

km WNW (airline) of the Potala Palace,
Lhasa (29° 39' N
91° 06' E), Lhasa

Notethat thefollowing material isinon
Hu
et al. (1987) but noreported
numbers or reference to a museum
collection are mentioned.

It is probable that

Municipality, Xizang (Tibet) Autonomous

all or most of these specimens are housed atCollected by:
CAS
Region, China.
170546-49 R. Macey and T. J. Papenfuss,and CAS 170550-53 T. J. Papenfuss and
R. Macey. Date: 25 Sept., 1988.

the Chengdu Institute of Biology.

CAS 170554-57. Locality: Elev. 3990
m, 52.4 km south of Yangbajan (30° 13' N
90° 25' E), also at km 1900.8 from
Xining, on the Xining-Golmud-Lhasa Rd.,
Lhasa Municipality, Xizang (Tibet)
Autonomous Region, China. Collected by:
CAS 170554-55 R. Macey and T. J.

Three males, 3 females, and 3 juveniles.
Locality: Bomi (29° 50' N 95° 45' E),
Qamdo Prefecture, Xizang (Tibet)

Autonomous Region, China.
Six males, and 12 females.

Lhasa (29°39'

N

Locality:

91° 06' E), Lhasa

Municipality, Xizang (Tibet) Autonomous

Region, China.

Asiatic Herpetological ResearchVol. 3, p. 106

FIG. 2. Paralectotypes oiStellio sacra

Two males, 2 females, andLocality:1

April 1990

BMNH 1946.8. 28.58 and BMNH 1946.8.28.59.
juvenile.

Nyingchi (29° 32' N

94°

25' E), Lhasa Municipality, Xizang (Tibet)

lizards with a snout-vent length of 120and a tail length of 180-240
150

mm

mm

(Table 1).

Autonomous Region, China.
Gular Sac seems to be developed to aDistribution
Stellio sacra as presently understood, isrestricted to the river drainage of the
Yarlung Zangbo in the Lhasa Valley,

Xizang (Tibet) Autonomous Region, China(Fig. 4). Only four localities are known,

between3000

and4000 m.

greater degree than in other Stellio. Body
scales are small and granular. The scales
are not well differentiated. There is a veryon thefrom
middleof thethebeginsand continues as a poorly

slight but noticeable nuchal crest

head.

It

occiput

differentiatedvertebral

stripe.

The

previously assigned to Stellio himalayanus

longitudinal rows of enlarged and feebly
keeled scales on the vertebral region arearranged parallel to each other. There areneither groups of enlarged scales nor
separate enlarged scales on the dorsal lateralhimalayanus, however

regions.

all

Populations occurring in the Kunlun
Mountains of southern Xinjiang Uygur

Autonomous

Region,

China
their

were
present

taxonomic position is uncertain.
Diagnosis: Rock agamid with flattenedhead and body which is typical for thislizard group.

They are comparatively large

The males have a large patch of callous
on the belly.
The annuli andthescaleson the basalofsegmentation
scalesquarter of the tail are not prominent.

On theApril 1990

Asiatic Herpetological ResearchVol. 3, p. 107/\

FIG. 3. Paralectotype of Stellio sacra ZSI 15740.
lateral surface of the tail there are three tofour annuli in each segment.

back. The dark golden brown forms bands
across the back which are offset at thespine.

There is a small granular dark pattern on
the back. The center of the back tends tohave more black and toward the sides adark golden brown dominates.
Theelements
thisofseparate
pattern areconnected to heavily marked diffuse
transverse stripes. The narrow stripes form
two rows of the dark colored scales that
continue from the neck to the tail. Overall
the lizard is darkly colored but there are a

few randomly scattered yellow blotches on
the back (Fig. 5). Juveniles are lighter incolor tending more toward a dark goldenbrown with darker speckling all over theComparative Description

sacra
differs from Stellio
andrelatedforms in all thehimalayanus
Stelliodiagnosis characters. Stellio sacra is only
similar in body size to Stellio stoliczkanus

among the species examined. Other species
are notably smaller (Table 1; Anderson andLeviton 1969; Ananjeva et al. 1981).

The body proportion data concerning
Stelliosacra and the Stellio himalayanus

species complex indicates small differences

Vol. 3, p. 108

TABLE 1.

Asiatic Herpetological Research

Characters of Stellio sacra specimens used for analysis.

April 1990

Asiatic Herpetological Research

April 1990

FIG. 4. Distribution of Stellio himalayanus and Stellio sacra.
Autonomous Region population of questionable taxonomic status.

The ventral head scales are also small
and not regular. The size of these scales
decreases from the nasal end toward thegular fold. On the sides of the head and on
the neck are situated small groups of verystrongly keeled spinose scales.

Stelliosacra

has nostrils shaped

longitudinally oval, similar to Stellio
chernovi and Stellio himalayanus. Whereas Stellio agrorensis, Stellio melanurus,
and Stellio tuberculatus

Vol. 3, p. 109

Question mark indicates Xinjiang

surrounding small scales. The scales onbase of the tail are keeled andthemucronate, but to a smaller degree than for

example in Stellio himalayanus.

The results of the comparative study ofAsiatic rock agamids enables the typicaldistinguishing characters of Stellio sacra tobe identified:1 .

The small size of the scales.

Stellio nuristanicus

have round nostrils.
round tympanum.

Stelliosacra has a

In Stellio sacra, the scales of the uppersurface of the fore and hind limbs arecomparatively small, but their size is similar
to the largest dorsal scales.

These scales

strongly keeled.
Single slightly
enlarged scales are distinguishable from theare

2. The presence of a short nuchal crest
made up of from two to three scale rows ofslightly enlarged, narrow, keeled, darkcolored scales. Such a nuchal crest isabsent in Stellio chernovi, Stellio
himalayanus, Stellio stoliczkanus and otherspecies. A nuchal crest is noted however in
Stellio melanurus, Stellio nuptus, Stellio
Stellio, and Stellio tuberculatus.

Asiatic Herpetological ResearchVol. 3, p. 110,-

April 1990«

sacra from elevation 3990 m, 52.4 km south of Yangbajan (30° 13' N
Autonomous
Region, China.
Xizang (Tibet)

FIG. 5.

Stellio

4. The90° 25' E),

rows of the enlarged dorsal

3. The ends of the occipital (nuchal) andneck scales are directed backwards but notscales are parallel but do not meet as inforeword; the parietal scales are oriented in

species of the "Stellio

a laterocaudal direction.

complex.

"himalayanus

TABLE 2. Comparative data on the relative proportions of the tail, limbs, and head of rock agamids of thegenus Stellio(5. sacraSpecies

and agamids of the 5. himalayanus complex).

Asiatic Herpetological Research

April 1990

Vol.3, p. Ill

FIG. 6. Habitat of Stellio sacra, elevation 3990 m, 52.4 km south of Yangbajan (30° 13' N 90° 25' E),
Xizang (Tibet) Autonomous Region, China.

5. In the region over the shoulders
there are two arched diffuse rows ofslightlyenlarged mucronate (spinose)

scales.
6. Theabsence of enlarged scales orofenlarged scales on the back ofregions
the body.

five scale rows.
9. The scalesof the back, shoulders,andsomethighsparietal scales have an
unusual microstructure. The margins of thescales are jagged resembling that of fringes.
This character was also observed in Stellio

annectens, Stellio erythrogaster, Stelliomelanurus, and Stellio nuptus.

7. The upper head scales distributed
between the nasal scales have a stretched

The length of these scales is twice

shape.
the width as also in Stellio

agrorensis,

10. There

is

a

patch

of

callous(granular) scales in the middle of the belly,

which is large in males.

Stellio nuristanicus, and Stellio tuberculatus

where as the scales of Stellio chernovi andhimalayanus have a roundish,Stelliopolygonal shape.
8. Onlya single row of small scales isnoted between the suboculars and thesupralabials. Other species have two to

1 1
The males have a large patch of analpores and from six to seven rows of callous
scales before the cloaca.
.

Coloration

The dorsal coloration of

the

type

Asiatic Herpetological ResearchVol. 3, p. 112

April 1990

specimens consists of diffuse components,
each scale is either totally dark or light incoloration. The parietal region and along
the middle of the back (two scale rows ofabsolutely dark scales) are more dark incoloration. From the middle of the back tomelanurus, Stellio nuptus, and Stellio
tuberculatus. The characters which enable
us to bring together Stellio sacra and these
species should be considered as

the sides there are light brown transverse

characters are long limbs, the presence of a

stripes.

This

colorationispoorlydeveloped in some specimens.

The subadult specimen has a similarsmall speckled pattern and in addition has
alternating light and dark transverse stripes
coming from the vertebral ridge toward thesides of the body. This pattern is similar tothatof juvenile specimens of Stellio

tuberculatus.

species as Stellio

plesiomorphic

agrorensis,

similarities.

StellioThese

small gular sac and nuchal crest, thepolyannular structure of the caudal
segments, the juvenile color pattern, and soon. This assumption about a plesiomorphic
condition is based on the idea that theoligomerization (decreasing of number of
elements) of homologic organs (herepholidosis elements) is one of the main
directions of agamid evolution (Dogel
1954).

As far as we can tell from the available
specimens of Stellio sacra, no contrasting
pattern and coloration on the surface of thegular region, neck and nuchal region are
noted.
Such a contrasting pattern andcoloration are typical for specimens of
Stellio himalayanus and related species.

Characters, such as the orientation of thescale axes in nuchal and neck regions andin parietal scales is also of great interest.

Itis suggested (Peters unpublished data) thatthe more common condition of the scale,i.e. the orientation of the point backwards,
is typical not only for most of the species in

Natural History
Stellio sacra
hills

the genus Stellio (including Stellio sacra ),but also for the majority of agamids andare common in the rocky

surrounding the Lhasa Valley.

Theywere seen only on slopes covered withlarge boulders (Fig. 6). Often a single
adult male occupied a pile of boulders inassociation with several females andStellio sacra is an agile, alert
thatis difficult to approach closely.
speciesjuveniles.

The only other reptile that occurs on the
rocky slopes is the gecko, Cyrtodactylus

tibetanus.

Another

agamid,Phrynocephalus theobaldi is abundant on
the sandy soil at the base of the rocky hills.

Discussion
The large review of the rich Indian faunathat Smith (1935) dealt with explains why

Agamid, Stellio sacra, wasdescribed
as a subspecies of theoriginally

Hence it is acondition.
The contrary
plesiomorphic
direction, where the point of the scale isforwards, is observed in Stellio melanurus,
Stellio nuptus and a number of African
species. In accordance with Peter's pointof view, this should be considered as anapomorphic condition. However there is aproblem in interpreting such a character as
the presence of the scales with the jagged
lizards

asamargins. This scale structure is found inspecies with both types of scale orientation,
including Stellio sacra as well as Stellio
erythrogaster, Stellio melanurus, Stellio
nuptus and the African species, Stellio
annectens. The interpretation of suchcharacters makes all possible explanationsof the relationships of these species

the Sacredcontroversial.

Himalayan agamid, Stellio himalayanus.
The study presented here concludes that

to

sacra

not related to the Stellio
himalayanus complex. On the other handthe above descriptions of the morphologicalcharacters do not allow an interpretation of
the direct relation of Stellio sacra to such

Stellioiswhole.

These facts and the inability at this time
determine even a single obvious

synapomorphy for all Asiatic rock agamidsof the genus Stellio (with or without Stelliomelanurus and Stellio nuptus ) does not
allow

an

acceptablephylogenetic

relationship to be developed. This problem

Asiatic Herpetological Research

April 1990

Vol. 3, p. 113

TABLE 3. The distribution of the species in the genus Stellio.

Distribution

Species
Stellio adramitana

Arabia(Anderson 18%)

Stellio agrorensis (Stoliczka 1872)
Stellio annectens (Blanford 1870)

Afghanistan, Pakistan, India
Africa (Somalia, Ethiopia)

Stellio atricollis(Smith 1849)(Anderson and Leviton 1969)

Eastern and southern Africa

Stellio badakhshanus

Afghanistan

Stellio caucasius(Eichwald 1831)

Caucasus, Tadjikistan, Turkmania, Turkey, Iraq,
Iran, Afghanistan, Pakistan

Stellio chernovi

Tadjikistan, Turkmania, Uzbekistan

Stellio cyanogaster (Ruppell 1835)

Somalia, Ethiopia(Ananjeva, Peters,
and Rzepakovsky 1981)

Iran, Turkmania

Stellio erythrogaster(Nikolsky 1896)
Stellio himalayanus (Steindachner 1869)

Tadjikistan, Uzbekistan, Afghanistan, Pakistan,
India

Stellio lehmanni(Nikolsky 1896)(Blyth 1854)
Stellio microlepis (Blanford 1874)

Tadjikistan, Turkmania, Uzbekistan, Afghanistan

Stellio nuptus

Iraq, Iran, Afghanistan, Pakistan

Stellio melanurusIran, Pakistan
Iran(De Filippi 1843)(Anderson and Leviton 1969)

Stellio nuristanicus

Afghanistan

Stellio phillipsii(Boulenger 1895)
Stellio sacra (Smith 1935)
Stellio stellio (Linnaeus 1758)
Stellio stoliczkanus (Blanford 1875)
Stellio trachypleurus (Peters 1982)
Stellio tuberculatum (Hardwicke and Gray 1827)

Ethiopia
Tibet

Stellio yemenensis

Arabia(Klauzewitz 1954)
Stellio zonorus (Boulenger 1895)

and the fact thatmany other forms are poorly studied but
also by the obvious existence of parallel
trends in different developmental lines

within this lizard group.
In order to arrive at a more trust worthy

hypothesis of the relationships of rockagamids, it is necessary to carry out

biochemical investigations.

Mongolia, China
Ethiopia
India, Nepal, Afghanistan, Pakistan

Somalia

may be explained not only by the small
sample size of Stellio sacra

Greece, southwest Asia, northern Egypt

The

first

preliminary results of such investigations
are that of Joger and Arano (1987) andAnanjeva and Sokolova (in prep.). This
type of data will be highly interesting tocompare with the results of comparative

morphological studies.

monophyletic group. Rock agamids of thegenus Stellio are distributed from Greece
and the Nile River Delta in the west to theGobi Altai of southern Mongolia, and thewestern deserts of China in the east. In thewestern part of this region, Greece to the
Nile River Delta, Stellio stellio is found.
The southern portion of this region fromsouthwestern Iran to Pakistan Stelliomelanurus and Stellio nuptus occur.
These species are probably of African
origin (Peters unpublished). In southern
Iran and Afghanistan along with Stelliomelanurus and Stellio nuptus also occur
Stellio agrorensis, Stellio badakhshanus,
Stellio caucasius, Stellio erythrogaster,
Stellio himalayanus, Stellio microlepis,

and Stellio nuristanicus.

The distributional patterns of Asiaticrock agamids of the genus Stellio (Table 3)

and their chorological isolation seems to
support the idea that Stellio is a

The Asiatic rock agamids of the genus
Stellio sensu stricto are absent

from themountainous regions of western Indostan

Asiatic Herpetological ResearchVol. 3, p. 114

and the eastern portion of the Arabian
Peninsula. They are also not found acrossthe Red Sea in Ethiopia and Somalia.

These

areas

are

poorlystudiedherpetologically and it is possible that they
do in fact occur in these regions.

The distribution of Asiatic rock agamids
seems to be a single unit. Most problematicthe origin of such species as Stelliomelanurus, Stellio nuptus, and Stellio
stellio, which may be of African origin as it
was already mentioned above.
is

April 1990

ANANJEVA, N. B., G. PETERS, AND V. T.
RZEPAKOWSKY.

1981.

[New species of rockagamid from Tadjikistan Agama chernovi

sp.

nov.]. Proceedings of the Zoological Institute ofthe

USSR Academy of Sciences, Leningrad

101:23-27. (In Russian).

ANDERSON, S. T. AND A. E. LEVITON.

1969.

Amphibians and reptiles collected by the Street
Expedition to Afghanistan, 1965. Proceedings of
the California Academy of Sciences, series 4.
37(2):25-56.

DOGEL, V. A. 1954.

[Oligomerization of theorgans as one of the maintendencies of animal evolution].
LeningradState University. 368 pp. (In Russian).

homological

It is possible that this assumption will becorroborated during further research inbiochemical phylogeny. The preliminary
results of Joger and Arano (1987) on
Stellio stellio is a first step. The origin of
the genus Stellio is interesting and the data
about the early divergence of Stellio in
Stellio sensu stricto (Asiatic species) andthe Stellio atricollis species group fromAfrica and southern Arabia is useful (Joger

and Arano 1987).

It is possible that Stellioa paraphyletic group of species. The
Arabian - African species group is Stellioisadramitana, Stellio annectens, Stellio
atricollis, Stellio cyanogaster, Stellio
phillipsi,

Stelliotrachypleurus, Stellio

yemenensis, and Stellio zonurus (Table3). These species are similar to the Asiatic
species Stellio melanurus and Stellio
nuptusin anumber of characters. This

allows an assumption that they are related.

HU, S., E. ZHAO, Y. JIANG, L. FEI, C. YE, Q.
HU, Q. HUANG, Y. HUANG, AND W. TIAN.
(incorrectly stated in English as Hu, S, L. Fei,

Q. Hu, Q. Huang, Y. Huang, Y. Jiang, W.
1987.
Tian, C. Ye, E. Zhao).
AmphibiaTheof the scientificseriesof
Xizang.
Reptilia
expedition to the Qinghai-Xizang Plateau. The
comprehensive scientific expedition to Qinghai-

Xizang Plateau, Academia Sinica. Chengdu
Institute of Biology, Academi (sic.) Sinica. 153
pp. (If Chinese).

JOGER, U. AND B. ARANO. 1987. Biochemical
phylogeny of the Agama genus group. Pp. 215Van Gelder, Strijbosch.and Bergers
218. In
Proceedings of the 4th Ordinary General
Meeting of the Society of European(eds.).

Herpetologists, Nijmegen.

MOODY, S. M.

1980.

Phylogenetic and historicalbiogeographical relationships of the genera in the

Acknowledgments

Agamidae (Reptilia: Lacertilia). Ph.D. Thesis.
University of Michigan. 373 pp.

We

are most greatful for the loan of typematerial of Stellio sacra from Dr. A. C. G.
Grandison, Dr. E. H. Arnold and Dr. A. F.
Stimson of the British Museum of Natural
History, London, Great Britain, and Dr.

Shara of the Indian Museum, Calcutta,
India.

PETERS, C. 1982. Eine neue Wirtelschwanzagame
aus Ostafrika (Agamidae:
Agama ).
Mitteilungen aus der Zoologishen Sammlung desMuseums fur Naturkunde in Berlin 58(2):265-

268.

SMITH, M. A.

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Leningrad 101:3-20. (In Russian).

SOKOLOVSKY, V. V.

1975.

[Comparative

karyological study of the lizards from the family

Agamidae. II. Karyotypes of five species of thegenus Agama ]. Cytologica 16(l):91-93. (InRussian).

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SOKOLOVSKY, V. V.

1977.

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