Amphibia-Reptilia (2016) DOI:10.1163/15685381-00003026
Alai! Alai! – a new species of the Gloydius halys (Pallas, 1776)
complex (Viperidae, Crotalinae), including a brief
review of the complex
Philipp Wagner1,∗, Arthur Tiutenko2, Glib Mazepa3,LeoJ.Borkin
4, Evgeniy Simonov5
Abstract. During a scientiﬁc ﬁeld expedition to the Alai-Pamir range ﬁve specimens of the genus Gloydius have been
collected in the larger Alai. A morphological and genetical examination of the specimens has shown that they are part of
the G. halys complex, but represent a new taxon which is characterized by the following unique character combination: It is
a slender and moderately stout small snake, up to 479 mm total length. The head has nine symmetrical plates on the upper
head, 7 supralabial and 8-9 infralabial scales. Body scales in 20-22 rows around midbody, 143-156 ventral and 35-45 usually
paired subcaudal scales. The cloacal plate not divided. The general coloration consists of various different tones of olive, tan
and brown, having a distinct head, but an indistinct body pattern with, excluding the tail, 26-29 transverse crossbands, which
are not extending to the sides of the body. The haplotype network shows the new species within the G. halys complex and
close related to both, G. h. halys and G. h. caraganus. So far the new described species is only known from the Alai range.
However, various Gloydius specimens are found in Kyrgyzstan and because of the complicated taxonomy those specimens
have to re-identiﬁed to clarify their status and the status of the new species.
Keywords: Alai, Central Asia, Crotalinae, Gloydius sp. n., Kyrgyzstan, Viperidae.
Snakes of the genus Gloydius Hoge and Roma-
no-Hoge, 1981 are venomous pitvipers and part
of the Asian radiation of the subfamily Crotali-
nae within the Viperidae. Recently, 13 species
are known: Gloydius blomhofﬁi (Boie, 1826)
from China, Korea, Japan and Russian Far East
(see Orlov et al., 2014); Gloydius brevicaudus
(Stejneger, 1907) from China, North and South
1 - Department of Biology, Villanova University, 800 Lan-
caster Avenue, Villanova, PA 19085, USA and Zoolo-
gische Staatssammlung München, Münchhausenstrasse
21, D81247 Munich, Germany
2 - University of Erlangen-Nuremberg, Schlossplatz 6,
D91054 Erlangen, Germany
3 - Department of Ecology and Evolution, University of
Lausanne, CH1015 Lausanne, Switzerland and Depart-
ment of Ecology and Genetics, Uppsala University,
75236 Uppsala, Sweden
4 - Zoological Institute, Russian Academy of Sciences,
Universitetskaya nab. 1, 199034 St. Petersburg, Russia
5 - Institute of Systematics and Ecology of Animals,
Siberian Branch of Russian Academy of Sciences,
Frunze 11, 630091 Novosibirsk, Russia and Tomsk State
University, Lenina Avenue 36, 634050 Tomsk, Russia
Korea; Gloydius halys (Pallas, 1776) ranging
from Azerbaijan and Iran through several coun-
tries of Middle Asia to eastern Siberia, Mon-
golia, and China; Gloydius himalayanus (Gün-
ther, 1864) from northeastern Pakistan along the
southern slopes of the Himalaya to northern In-
dia and Nepal; Gloydius intermedius (Strauch,
1868) restricted to Russian Far East, north-
eastern China and the Korean Peninsula accord-
ing to Orlov and Barabanov (1999). However,
according to Gloyd and Conant (1990) as well
as to Orlov in Ananjeva et al. (1998) G. inter-
medius is distributed from south-eastern Azer-
baijan, through northern Iran, southern Turk-
menistan, and north-western Afghanistan, to
north-western China and Mongolia, while G.
saxatilis (recognized as synonym of G. inter-
medius by Orlov and Barabanov ) from
eastern Siberia through northeastern China to
North and South Korea); Gloydius lijianlii Jiang
and Zhao, 2009 endemic to Daheishan Island,
China; Gloydius liupanensis Liu, Song and Lua,
1989 only known from its type locality in the Li-
upan Mountains of China; Gloydius monticola
(Werner, 1922) only known from mountains in
©Koninklijke Brill NV, Leiden, 2016. DOI:10.1163/15685381-00003026
2P. Wagner et al.
northern Yunnan, China; Gloydius shedaoensis
(Zhao, 1979) endemic to Shedao Island, China;
Gloydius strauchi (Bedriaga, 1912) known from
the Tibetan Plateau of “Tsinghai” (=Quinghai)
and western “Szechwan” (=Sichuan), China;
Gloydius tsushimaensis (Isogawa, Moriya and
Mitsui, 1994) endemic to Tsushima Island,
Japan; and Gloydius ussuriensis (Emelianov,
1929) ranging from far east Russia through
north-eastern China to North and South Korea,
as well as on Quelpart Island.
Gloydius was the subject of considerable tax-
onomic instability. Gloyd and Conant (1990)
published an intensive review of the genus Agk-
istrodon Palisot de Beauvois, 1799 (including
Gloydius at this time) providing intensive mor-
phological data and a new systematic approach
for this group. A few years later, Bour (1993),
based on his interpretation of the type local-
ity of the nominate subspecies, synonymized
G. h. caraganus with G. h. halys and intro-
duced the name Agkistrodon (=Gloydius)halys
mogoi for the populations of the former nom-
inate taxon. However, Chernov (1934), Anan-
jeva et al. (1997) as well as Orlov and Bara-
banov (1999) did not accept this restriction.
They revalidated G. h. caraganus, synonymized
G. h. mogoi with G. h. halys and stabilized
the latter with the designation of a neotype.
Moreover, Orlov and Barabanov (1999) radi-
cally revised the classiﬁcation made by Gloyd
and Conant (1990) and did not accept G. halys
and G. intermedius as two widely sympatric
species across Central Asia as suggested by
Gloyd and Conant (1990). Instead, they re-
stricted G. intermedius to eastern Asia and sug-
gested a single, widespread G. halys including
several subspecies: (1) G. h. boehmei (Nilson,
1983); (2) G. h. caraganus (Eichwald, 1831);
(3) G. h. caucasicus (Nikolsky, 1916); (4) G. h.
cognatus (Gloyd, 1977); (5) G. h. halys (Pallas,
1776); and (6) G. h. stejnegeri (Rendahl, 1933).
A taxonomic concept followed by Gumprecht
et al. (2004). Additionally, the re-examination
of the lectotype of G. intermedius by Orlov and
Barabanov (1999) has shown that it is identi-
cal with Gloydius saxatilis. Consequently, they
synonymized G. saxatilis with G. intermedius.
Later, Xu et al. (2012), following the taxo-
nomic concept of Gloyd and Conant (1990),
published a phylogenetic study, focused on the
Chinese species of the genus and showing G.
intermedius basal to a lineage including G. sax-
atilis/G. shedaoensis and their sister species
G. halys. Most recently, Hoser (2013) nomen-
claturally revised the entire Crotalinae and in-
troduced the generic name “Conantvipera”for
some Gloydius species. However, this author is
heavily criticized by the entire herpetological
community (see e.g., Kaiser et al., 2013) for his
nomenclatural acts which are published in his
self-made journal, without any reliable research
background and do not fulﬁll the criteria of the
code of ethics. Therefore, the name introduced
by Hoser (2013) is ignored in this publication
(although LB consider them as formally avail-
able and valid according to the Code of Zoolog-
ical Nomenclature until the International Com-
mission on Zoological Nomenclature consider
them as invalid).
This summary shows that Gloydius is a tax-
onomic complex group and, despite the inten-
sive reviews by Gloyd and Conant (1990) and
Orlov and Barabanov (1999) and the phyloge-
netic study by Xu et al. (2012), an integrative
revision is in need to clarify the relationships
between the species, their distributions and es-
pecially to clarify the status of the huge num-
ber of recognized subspecies. Herein, we follow
the taxonomic concept of Orlov and Barabanov
(1999), with some modiﬁcation concerning the
composition of the Gloydius halys complex and
the usage of the genus name Gloydius.
Although many expeditions, started from
Fedchenko, Severtsov and others, provided var-
ious herpetological collections and data, no
pitvipers were recorded in the Alai region un-
til Karpenko (1957). Later, these snakes were
recorded in several localities of the Alai re-
gion (Yakovleva, 1964; Eremchenko, 2007). In
2013, an international research expedition was
conducted to the Alai and Pamir region and
New species of Gloydius 3
was based for several days in a small valley
leading into the larger of the Alai valley. De-
spite the fact, that this is a famous area and
well-known for mammal and bird wildlife, the
knowledge about the other vertebrate groups is
limited. It is based on the results of William
Frederik Reinig, the zoologist of the German-
Soviet Alai-Pamir expedition which was orga-
nized by Willi Rickmer Rickmers in 1928 (see
e.g., Reinig, 1928, 1932; Rickmer Rickmers,
1929, 1930). However, Reinig was mainly fo-
cused on arthropods and especially the herpeto-
fauna of the Alai and Pamir is still only rudi-
mentarily known. Therefore, collections of am-
phibians and reptiles from this area are impor-
tant and done by the expedition.
The Pamir and Alai range is ecologically
characterized by high altitude steppes with
very pronounced temperature variation between
summer and winter and low precipitation
throughout the year. The Pamir has great alti-
tudinal variation ranging from 1740 to 7495 m
a.s.l., while the Alai range only up to 5544 m.
However, both are characterized by high at-
mospheric aridity, intense insolation, great sea-
sonal and large diurnal temperature ﬂuctuations,
low temperatures, scanty of no snow cover and
cold storms (Agakhanyantz and Lopatin, 1978;
Mani, 2007). The Pamir Mountains and the Alai
Valley are among the priority areas for nature
conservation. They are known as hotspot for
mountain species of Central Asia and the di-
versity includes endemic plant, insect, amphib-
ian, reptile and mammal species. Furthermore,
a study of the Conservation International Foun-
dation (2014) at the Pamir-Alai Transboundary
Conservation Area estimated that up to 51% of
the invertebrate fauna is endemic to the region,
while Karamhudoeva (2008) recognized about
15% of the Lepidoptera species as endemic for
the Pamir and Alai range. In Gloydius halys,the
Alai is of special interest because according to
Orlov and Barabanov (1999) it is an area where
the ranges of the two subspecies G. h. halys and
G. h. caraganus are overlapping. However, ac-
cording to Gloyd and Conant (1990) only the
latter is present in the Alai, but in its extreme
southeastern most distribution.
Therefore, the specimens were examined in
details and compared with other subspecies of
G. halys and the results are presented in this
Material and methods
The specimens were compared with the morphological data
provided by Gloyd and Conant (1990) and Orlov and Bara-
banov (1999) and furthermore with Gloydius specimens
housed in the ZFMK collection (see Appendix). The follow-
ing institute acronyms are used in this publication: ANSP,
Academy of Sciences in Philadelphia, PA, USA; BMNH,
The Natural History Museum, London, England; ISEA, In-
stitute of Systematics and Ecology of Animals, Siberian
Branch of Russian Academy of Sciences, Novosibirsk, Rus-
sia; MNHN, Muséum National d’Histoire Naturelle, Paris,
France; MNKNU, Museum of Nature at V. N. Karazin
Kharkiv National University, Kharkiv, Ukraine; NHRM,
Naturhistoriska Riksmuseet, Stockholm, Sweden; NMW,
Naturhistorisches Museum, Wien, Austria; USNM, United
States National Museum, Washington, DC, USA; UTA,
University of Texas in Arlington, Merriam Museum, Arling-
ton, TX, USA; ZMB, Museum für Naturkunde, Berlin, Ger-
many; ZFMK, Zoologisches Forschungsmuseum Alexander
Koenig, Bonn, Germany; ZISP, Zoological Institute, Rus-
sian Academy of Sciences, St. Petersburg, Russia.
For the comparison the following characters were used
and compared with the above mentioned sources: head
length, head width, head height, snout-vent length, tail
length (all taken with a digital caliper to the nearest of
0.1 mm); number of supralabial, infralabial, temporal, ven-
tral and subcaudal scales; shape of head, body and tail
scales; number of scale rows at fore body (one head length
posterior to the head), midbody and hind body (equivalent
of one head length anterior to the cloacal plate), all counted
straight across the body; number of crossbands on the body
excluding the tail, width of crossbands counted as scale
rows, extension of crossbands referring to the midbody scale
row counted from the ventral scales.
Digital X-ray images for skull illustration of the speci-
men ZMB 80360 were done using a Faxitron LX-60 Closed
Cabinet Digital Specimen Radiography System (Faxitron
Corp.) at the ZFMK. Figures used for X-ray comparisons
are modiﬁed from Gloyd and Conant (1990): G. blomhof-
ﬁi blomhofﬁi (KVK 832 without locality), G. intermedius
(ANSP 30146 from Adsh Mountain Range, Mongolia [ac-
cording to Orlov and Barabanov (1999) this specimen cor-
respondents with G. h. halys]), G. halys caraganus (ZIK
292 from near Kulsari, Kazakhstan), and G. himalayanus
(BMNH 19220.127.116.111 from Bakloh, India).
Relationships of the Alai specimens and other taxa of
the G. halys-intermedius complex were inferred using par-
tial nucleotide sequences of the mitochondrial gene NADH
dehydrogenase subunit 4 (ND4). Total genomic DNA was
isolated from the liver or muscle tissues of the ethanol stored
4P. Wagner et al.
museum specimens using standard proteinase K and phenol-
chloroform protocols (Sambrook, et al., 1989; see table 1
for specimens and Genbank numbers). A target fragment
was ampliﬁed using primers Nd4 and tRNA-leu (Arevalo,
et al., 1994). PCR was performed in a 25 μl volume each
containing 15-60 ng DNA, 0.2 mM of each dNTP, 2.5 μlof
10×ampliﬁcation buffer (10 mM Tris-HCl pH 8.5, 50 mM
KCl and 2.5 mM MgCl2), 1 U of Taq DNA polymerase,
and 5 pmol of primers. Ampliﬁcation was performed in a
T100™ Thermal Cycler (Bio-Rad) with an initial denatura-
94°C, annealing at 54°C for 30 s, 60 s extension at 72°C, and
a ﬁnal extension of 5 min at 72°C. Sequencing was carried
out on an ABI 3730 automated capillary sequencer (Applied
Biosystems) with the ABI Prism Big Dye Terminator Cycle
Sequencing Ready Reaction Kit 3.1 using the same primers.
DNA sequences were aligned manually and checked for
unexpected stop codons using BioEdit 7.0 (Hall, 1999).
Aligned sequences were 641 bp long. MEGA 6 (Tamura,
et al., 2013) was used to calculate percentage differences
(p-distances) between obtained sequences. A haplotype
network was generated using the median-joining algorithm
(Bandelt et al., 1999) implemented in Network 18.104.22.168
The morphological comparison revealed differ-
ences in pholidosis to other Gloydius taxa. Fol-
lowing the key presented by Gloyd and Conant
(1990) the Alai specimens would refer to G.
monticola or G. h. cognatus. However, both taxa
range far away from the Alai valley and are su-
perﬁcially similar but very distinct in detail. The
specimens are distinct to most Gloydius taxa by
their low number of scale rows around midbody,
which is only similar in the combination of the
missing apical pits to G. monticola. But, the
Alai specimens have a higher number of scale
rows around midbody and a very distinct col-
oration. Recognizing the low number as unusual
but within the variation, in combination of the
missing apical pits, this would refer, because
of the low numbers of ventral scales and body
crossbands to G. h. cognatus. However, from
this taxon the Alai specimens are distinct in col-
oration having body crossbands not extending
to the lateral sides of the body and in some as-
pects of scalation (see below).
Nucleotide sequences of the ND4 gene were
obtained for two Alai specimens, G. h. halys,G.
h. caraganus and G. intermedius (table 1). Both
Alai specimens shared the same haplotype and
were fairly distinct from G. halys and other taxa
sequenced in this study (ﬁg. 1). Consequently
the specimens are described as a new species.
Tab le 1. Tissue samples used in this study with locality, voucher and GenBank accession numbers.
Taxon Locality Museum voucher GenBank No.
G. rickmersi sp. n. Kyrgyzstan, northern slope of the Alai ridge, MHNG 2752.69 KM078592
Tengizbai River, Kichelay
G. rickmersi sp. n. Kyrgyzstan, Kul-Otek near to one from Kyzyl-Eshme MHNG 2752.70 KM096379
G. h. halys Russia, Novosibirsk Region, Maslyanino District ISEA R397 JQ356857.1
G. h. halys Russia, Zabaykalsky Krai, Krasnochikoysky District ZISP TS2309 KM078593
G. h. caraganus Kazakhstan, Almaty Province ISEA R290 KM078594
G. intermedius Russia, Primorsky Krai, Ussuriysky Natural Reserve ZISP TS2314 KM078595
G. shedaoensis JQ687481.1
Figure 1. Haplotype network of selected members of Gloydius halys/intermedius complex based on ND4 sequences. Numbers
below “l” corresponds to the number of mutated positions (“l” without number corresponds to single mutation).
New species of Gloydius 5
Account of related taxa
The species of the genus Gloydius are recog-
nized here in the sense of Orlov and Bara-
banov (1999, see introduction for comments)
and all of the following species are recognized
herein as members of the G. halys complex, a
subgroup mainly including Central and Eastern
Gloydius halys halys (Pallas, 1776)
1776 Coluber halys Pallas, Reise durch ver-
schiedene Provinzen des russischen
Reichs. Kais. Akad. Wiss., St. Peters-
burg, Vol. 3: 703.
1820 Echidna aspis var. pallasii Merrem,
Versuch eines Systems der Amphi-
bien I (Tentamen Systematis Amphi-
biorum). J. C. Kriegeri, Marburg: 151.
1993 Gloydius halys mogoi Bour, Les voy-
ages de Peter Simon Pallas et l’origine
de Coluber halys (Serpentes Viperi-
dae). Bull. mens. Soc. linn. Lyon 62:
Neotype. ZISP 14784, an adult male from the
Borgaiskaya steep, 84 km West from Kyakhta
town, Burin-Khan mountain (designated by
Orlov and Barabanov, 2000).
Diagnosis. A moderately stout viper up to
530 mm total length in males, to 590 mm in fe-
males according to Gloyd and Conant (1990),
but up to 750 mm ﬁde Orlov and Barabanov
(1999). Snout seen in proﬁle recurved; supral-
abial scales usually 7-9. Apical pits absent. Dor-
sal scales in 23 (very rarely in 21 or 25) rows
around midbody; ventral scales between 164-
178 (147-187 ﬁde Orlov and Barabanov, 1999);
subcaudal scales paired, between 42-49 (29-56
ﬁde Orlov and Barabanov, 1999). Body with 33-
47 dark transverse bands, each 3-5 scales wide
and extending down to scale row 3 or 2; light
areas between blotches relatively narrow. Light
line above dark cheek stripe on 1-1.5 adjacent
rows of scales.
Distribution. Gloyd and Conant (1990) and
Orlov and Barabanov (1999) mentioned a distri-
bution from eastern Kazakhstan through south-
ern Siberia to Mongolia and a vertical distribu-
tion between 150 to at least 2300 m.
Gloydius halys boehmei (Nilson, 1983)
1983 Agkistrodon halys boehmei Nilson,
A new subspecies of the Asiatic pit
viper Agkistrodon halys Pallas, 1776
(Serpentes, Viperidae). Bonner zoolo-
gische Beiträge 34: 470.
Holotype. ZFMK 8648, from the Andarab
Valley in the province Baghlan, Afghanistan.
Diagnosis. A small viper up to 487 mm to-
tal length. Apical pits absent. Two pre- and two
postocular scales on each side. Seven supral-
abial scales and 11 sublabial scales. Dorsal
scales in 23 rows around midbody; ventral
scales 155; 35 subcaudal scales paired, cloacal
plate not divided. Body with 41 dark transverse
bands, each 3-4 scales wide and extending down
to scale row 7 or 8.
Distribution. Only known from its type local-
ity at 2500 m a.s.l.
Gloydius halys caraganus (Eichwald, 1831)
1831 Trigonocephalus caraganus Eichwald,
Zoologia specialis, quam expositis ani-
malibus tum vivis, tum fossilibus
potissimuni rossiae in universum, et
poloniae in specie, in usum lectionum
publicarum in Universitate Caesarea
Vilnensi. Zawadski, Vilnae: 170.
1931 Ancistrodon halys paramonovi Nikol-
sky, Eine neue Schlangenunterart aus
Turkestan [Ukrainian with German ti-
tle], Travaux Mus. Zool. (Kiev) 10:
115. Type locality: “in montibus
Tschimganensis altitudine 1500-
2500 m probe urbem Taschkent in
Turkestano” (=mountains near
Neotype. ZISP 2200, adult male from the
Mangyshlak Peninsula, eastern edge of the
Caspian Sea, Kazakhstan (designated by Orlov
and Barabanov, 1999).
Diagnosis. A relatively slender and moder-
ately stout viper up to 735 mm total length in
6P. Wagner et al.
males (740 mm and more according to Orlov
and Barabanov, 1999), to 530 mm in females.
Snout seen in proﬁle slightly recurved, supral-
abial scales usually 8 (71%) sometimes 7. Api-
cal pits absent. Dorsal scales in 23 (rarely 21)
rows around midbody; ventral scales between
149-167 (141-183 ﬁde Orlov and Barabanov,
1999); subcaudal scales paired, between 33-47
(16-51 ﬁde Orlov and Barabanov, 1999). Gen-
eral coloration is pale. Body with 36-50 dark
transverse bands, not extending low on the sides
and with relatively broad light areas between
Distribution. From northeastern shores of
the Caspian Sea through Kazakhstan, Uzbek-
istan, northwestern Tadzhikistan and Kirgizia to
Chinese border and probably extreme western
Remarks. Orlov and Barabanov (1999) as
well as Gumprecht et al. (2004) mentioned G. h.
halys and G. h. caraganus as sympatric in parts
of their distribution range. Therefore this sub-
species should be recognized on species rank,
probably including other G. halys subspecies
which needs to be clariﬁed by further research.
Gloydius halys caucasicus (Nikolsky, 1916)
1916 Ancistrodon halys caucasicus Nikol-
sky, Ophidia. Fauna of Russia and adja-
cent countries. Vol 2. Reptiles: 267 [in
1933 Ancistrodon halys persicus Rendahl,
Die Unterarten des Ancistrodon halys
Pall. Nebst einigen Bemerkungen zur
Herpetologie Zentralasiens. Arch.
Zool. (Stockholm) 25A (8): 11.
Neotype. ZISP 19017.1, adult male from
the vicinity of Kirovsk town, Lenkoran dis-
trict, Azerbaijan (designated by Orlov and Bara-
Diagnosis. A moderately stout viper up to
660 mm total length. Supralabial scales 7-8,
rarely 9. Dorsal scales in 23 (rarely 25) rows
around midbody; ventral scales between 142-
169; subcaudal scales paired, between 31-46.
Body with 33-42 dark transverse bands, each
4-6 scales wide and extending down to scale
Distribution. The subspecies is ranging from
southeastern Azerbaijan through northern Iran
to southern Turkmenistan (Kopet Dag Moun-
tains) and northwestern Afghanistan (see Orlov
and Barabanov, 1999; Wagner et al., in rev.).
Gloydius halys cognatus (Gloyd, 1977)
1977 Agkistrodon halys cognatus Gloyd, De-
scriptions of new taxa of crotalid sna-
kes from China and Ceylon (Sri
Lanka). Proc. Biol. Soc. Washington
Holotype. USNM 68586, from Choni on the
Tao River, Gansu province, China.
Diagnosis. A small viper of the genus, up
to 590 mm total length. Supralabial scales 7-8.
Dorsal scales in 23 (rarely 21) rows around mid-
body; ventral scales between 153-165; subcau-
dal scales paired, between 36-54. Body with 29-
43 dark transverse bands, each 4-5 scales wide
and extending down to scale row 4 to 2.
Distribution. Recently endemic to northern
Central China, but probably northwards into
Gloydius halys stejnegeri (Rendahl, 1933)
1933 Ancistrodon halys stejnegeri Rendahl,
Die Unterarten des Ancistrodon halys
Pall. Nebst einigen Bemerkungen zur
Herpetologie Zentralasiens. Arch.
Zool. (Stockholm) 25A (8): 18.
Lectotype. NHRM 1923 809.2780, from
Diagnosis. A small viper of the genus, up
to 625 mm total length. Supralabial scales 7-8.
Dorsal scales in 23 rows around midbody; ven-
tral scales between 147-165; subcaudal scales
paired, between 39-46. Body with 28-38 dark
transverse bands, each 3-6 scales wide and ex-
tending down to scale row 3 or 2.
Distribution. The taxon ranges from the edge
of the Gobi in southeastern Inner Mongolia
New species of Gloydius 7
southwards to the Shanxi and Hebei provinces
in China (Gloyd and Conant, 1990).
Gloydius intermedius (Strauch, 1868)
1868 Trigonocephalus intermedius Strauch,
Concerning poisonous snakes distribu-
ted in Russia. Trudy Perv. Siezda Russ.
Yestestv. Zool. 1: 294 [in Russian].
Lectotype. ZISP 2221, from “Prom. Tyr
(Amur) [=Cape Tyr, Amur River, Amurskaya
Region, Russian Far East]” (designated by
Orlov and Barabanov, 2000).
Diagnosis. A large viper of the genus, up to
at least 800 mm total length. Supralabial scales
7-8. Dorsal scales in 23, very rarely 21, rows
around midbody; ventral scales between 148-
175; subcaudal scales paired, between 34-52.
Body with 28-45 dark transverse bands, each 3-
6 scales wide and extending down to scale row
Distribution. According to Orlov and Bara-
banov (1999) the species is restricted to an
area from Russian Far East through northeast-
ern China to the Korean Peninsula.
Gloydius rickmersi sp. n. Wagner, Tiutenko,
Borkin and Simonov
Holotype. ZMB 80360 [ﬁeld no. PW P069],
adult female from Kul-Otek at the Sary-Buka
Valley in Kyrgyzstan, about 25 km (by air)
NE of Daroot-Korgon (Chong-Alai District)
near the town Kyzyl-Eshme in direction to the
Shuman-Bol pass and the Kichi-Alai River at
an elevation of 3000 m a.s.l. [N39.622132,
E72.284010], collected on 4.VIII.2013, at about
19.00 h and purchased by local boys to Philipp
Wagner. The new name was registered at
Zoobank under the LSID: 84644B3D-85BC-
Paratypes. MHNG 2745.46 (ﬁeld no. PW
P070), two pieces of a subadult female from
the same area as the holotype but about 200 m
higher in elevation, collected as dead body
on 7.VIII.2013 by Philipp Wagner. MHNG
2752.70, roadkilled at Kul-Otek near to one
from Kyzyl-Eshme, collected as dead body
by Glib Mazepa. MHNG 2752.69, roadkilled
at the northern slope of the Alai ridge, Ten-
gizbai River, Kichelay [N39.755, E72.202],
2798 m, collected by Spartak Litvinchuk. ZSM
211/2014, a juvenile male from the same area
as the holotype but lower in elevation towards
the entrance of the valley to the main Alai
valley, collected as dead body on the road on
16.VIII.2013 by Arthur Tiutenko.
Diagnosis. A slender, moderately stout small
snake of the genus Gloydius, with up to 479 mm
total length. Head slender, slightly triangular,
with nine symmetrical plates on the upper head,
7 supralabial and 8-9 infralabial scales. Body
scales in 20-22 rows around midbody, 143-
156 ventral and 35-45 usually paired subcaudal
scales. Cloacal plate not divided. Body, exclud-
ing the tail, with 26-29 transverse crossbands,
not extending to the sides of the body.
Description. A small snake of its genus, but
only one adult specimen is recently known.
Head slender, slightly triangular and with nine
symmetrical plates on top. Snout slightly up-
turned. Parietal scales large, longer than wide.
Two nasal scales present, with the nostril
pierced in the center between them. Loreal scale
about quadrangular. Six scales surrounding the
orbit, excluding the postfoveal. Two preocu-
lar, two postocular and three temporal scales.
Pit bordered anteriorly by a large prefoveal,
dorsally and ventrally by a narrow prefoveal,
and posteriorly by the angle of both of them.
Seven sublabial scales on each side, the sec-
ond is the smallest, the third is entering the or-
bit. Eight to nine infralabial scales. Body scales
elongated and mainly keeled, the ﬁrst two scale
rows bordering the abdominal plates mooth at
midbody. Apical pits absent. Body scales ar-
ranged in 22 [anterior]-20-22 [midbody]-15-17
[posterior] rows around the body. There are
143-156 ventral scales, excluding the terminal
scale. Cloacal plate not divided. 35-45 subcau-
dal scales usually paired, but sometimes the ﬁrst
two thirds of the tail bear unpaired, followed by
8P. Wagner et al.
Differential diagnosis. The new species is
distinct to all other species of the genus in
the combination of the following characters:
a slightly lower number of scale rows around
midbody (20-22 versus mainly 23, sometimes
21); a low number of transverse body cross-
bands (26-29 versus above 30 in all taxa be-
side G. himalayanus and the G. blomhofﬁi
complex), the mean number of ventral scales
(150 which is higher than in most of the G.
blomhofﬁ taxa, but distinctly lower than in the
G. halys-intermedius complex); and its small
size (479 mm total length versus more than
600 mm is most species, apart from G. b. du-
bitatus,G. b. ussuriensis,G. h. halys and G. h.
caucasicus which are about the same maximum
total length, and G. monticola and G. strauchi
which are smaller). In details, the new species is
distinct to taxa of the Gloydius blomhofﬁ com-
plex generally by the lack of apical scale pits,
but as well by having a higher number of ventral
scales (mean 150 versus 140.7 in G. b. blomhof-
ﬁi, 140.1 in G. b. dubitatus, 141.1 in G. b. brevi-
caudus, and 140.1 in G. b. siniticus; but apart
from G. ussuriensis with 151.2); in having a
lower number of infralabial scales (mean 8.5
versus 10.4 in G. b. blomhofﬁi, 10.0 in G. b. du-
bitatus, 10.0 in G. b. brevicaudus, 10.1 in G. b.
siniticus, and 9.8 in G. ussuriensis); in having
a higher or somewhat equal number of cross-
bands on the body (mean 27.5 versus 19.9 in
G. b. blomhofﬁi, 29.9 in G. b. dubitatus, 29.9 in
G. b. brevicaudus, 30.1 in G. b. siniticus, and
28.7 in G. ussuriensis); and by having a very
indistinct coloration of crossbands versus very
distinct crossbands in this complex. The new
species is distinct to taxa of the Gloydius in-
termedius complex (according to the concept
of Gloyd and Conant, 1990) by the absence
of apical scale pits; in having a lower number
of scale rows around midbody (mean 21 ver-
sus 22.8 in G. intermedius and G. saxatilis); in
having a lower number of ventral scales (mean
150 versus 158.5 in G. intermedius and 156.8
in G. saxatilis); and a lower number of body
crossbands (mean 27.5 versus 36.9 in G. inter-
medius and 38.7 in G. saxatilis). Furthermore,
the new species is distinct to all taxa of the Gloy-
dius halys complex in having a lower number of
scale rows around midbody (20-22 [mean =21]
versus usually 23 rarely 21 or 25), and very nar-
row indistinct crossbars along the body not ex-
tending to the lateral sides (versus distinct cross-
bars and extending to the ﬂanks in most taxa)
(see table 2). The new species is distinct to G. h.
halys by a lower number of ventral scales (143-
156, mean 150 versus 164-178, mean 172); by
a lower number of infralabial scales (mean 8.5
versus 10.7); a lower number of body cross-
bands (mean 27.5 versus 38.4); and a distinct
coloration (body crossbands not extending to
the ﬂanks versus extending to the ﬂanks). The
new species is distinct to G. h. boehmei by its
higher number of subcaudal scales (35-45 ver-
sus 35); its lower number of infralabial scales
(mean 8.5 versus 11); and a lower number of
body crossbands (mean 27.5 versus 41). Apart
from the above mentioned characters, the new
species is somewhat similar in coloration to G.
h. caraganus, but distinct in its lower number of
ventral scales (143-156, mean 150 versus 149-
167, mean 158); by its lower number of infral-
abial scales (mean 8.5 versus 10.8); and by its
lower number of body crossbands (mean 27.5
versus 44.4). The new species is distinct to G.
h. caucasicus by its slightly lower number of
ventral scales (mean 150 versus 157); the lower
number of infralabial scales (mean 8.5 versus
11); a lower number of body crossbands (mean
27.5 versus 36.4); a distinct coloration (body
crossbands not extending to the ﬂanks versus
extending to the ﬂanks); and by the absence
of apical scale pits. The new species is distinct
to G. h. cognatus by its slightly lower number
of ventral scales (mean 150 versus 159); the
lower number of infralabial scales (mean 8.5
versus 10.2); a lower number of body cross-
bands (mean 27.5 versus 35.9); and a distinct
coloration (body crossbands not extending to
the ﬂanks versus extending to the ﬂanks). The
new species is distinct to G. h. stejnegeri by
slightly lower number of ventral scales (mean
New species of Gloydius 9
Tab le 2. Variation in selected Gloydius species. Values in the new species refer to this study and include the range followed by the mean value in brackets. Values of other taxa refer above
to Gloyd and Conant (1990), followed by their mean value in brackets, while values below refer to Orlov and Barabanov (1999). Own data in square brackets. AP: apical pits; CP: cloacal
plate; IL: infralabial scales; o: or; r: rarely; SC: subcaudal scales; SM: scales around midbody; SL: sublabial scales; ToL: total length (in mm); TVB: transversal bands; V: ventral scales;
∗the holotype has paired/unpaired subcaudal scales.
G.rickmersi sp. n. G.caraganus G. h. halys G. h. boehmei G. h. caucasicus G. h. cognatus G. h. stejnegeri G.intermedius
ToL  735 590 487 575 745 625 710
(–) (–) (–) (–) (–) (–) (–)
740 714 487 660 590 625 800
SM [20-22] 22-23 23 (r21) – 21-23 23 (r21) 21-25 23 (r21 o 25)
 (22.9) (22.9) (–) (22.9) (22.5) (23) (22.8)
23 (r21) 23 (r21 o 25) 23 23 (r25) 23 (r21) 23 23 (r21)
V [143-156] 149-167 164-178 – 142-169 153-165 147-165 149-165
 (158) (172) (–) (157) (159) (156) (158.5)
141-183 141-187 155 142-169 153-165 147-165 148-175
SC [35-45] 33-47 42-49 – 31-46 36-54 39-46 38-48
 (38) (45) (–) (38) (43) (42) (39.9)
46-51 29-56 35 31-46 36-54 39-46 34-52
SL  7-8 7-9 – 7-9 7-9 7-9 6-8
(7.7) (7.8) (–) (7.5) (7.3) (7.4) (7.2)
7-8 7-9 7 7-9 7-8 7-8 7-8
IL [8-9] 9-12 10-12 – 9-13 9-11 10-12 9-11
[8.5] (10.8) (10.7) (–) (11.0) (10.2) (10.7) (10.3)
––11 – – – –
TVB [26-29] 36-50 33-47  33-42 29-43 28-38 28-45
[27.5] (44.4) (38.4) (–) (36.4) (35.9) (31.6) (36.9)
––– – – – –
SC paired∗paired paired [paired] paired paired paired paired
CP not divided not divided not divided [not divided] not divided not divided not divided not divided
AP absent absent absent [absent] present absent present present
10 P. Wagner et al.
Figure 2. Holotype (ZMB 80360) of Gloydius rickmersi sp. n. from Sary-Buka Valley, Kyrgyz Republic. This ﬁgure is
published in colour in the online version.
150 versus 156); the lower number of infralabial
scales (mean 8.5 versus 10.7); a lower number
of body crossbands (mean 27.5 versus 31.6);
a distinct coloration (body crossbands not ex-
tending to the ﬂanks versus extending to the
ﬂanks); and by the absence of apical scale pits.
Description of the holotype. A small and
slender adult male (ﬁgs 2, 3) with a total
length of 479 mm (body length 414.5 mm, tail
length 64.5 mm). Head slender, slightly trian-
gular, 18 mm in length, 13.35 mm wide and
8.2 mm heigh. Crown with nine symmetrical
New species of Gloydius 11
Figure 3. Detailed pholidosis of the holotype (ZMB 80360) of Gloydius rickmersi sp. n. from Sary-Buka Valley, Kyrgyz
plates. Internasal scales more than two times
wider than long, their posterior margins curving
slightly backwards. Internasal and prefrontal
scales somewhat thickened, and with a slight de-
pression of the suture between the frontal and
prefrontal as well the prefrontal and internasal
scales, giving the appearance of a slightly up-
turned snout. Parietal scales large, about 1.5
times longer than wide. Rostral slightly higher
than wide, with an obtuse angle above. Two
nasal scales present, with the nostril pierced in
the center between them. Anterior scale larger
than the posterior one. Loreal scale subquad-
rangular, slightly wider than high. Six scales
surrounding the orbit, excluding the postfoveal.
Two preocular scales, the lower one narrow and
forming the posterior dorsal border of the pit.
Two postocular scales, the lower one crescent-
shapes and extending forward beneath the eye.
Pit bordered anteriorly by a large prefoveal,
dorsally and ventrally by a narrow prefoveal,
and posteriorly by the angle of both of them.
Seven sublabial scales on each side, the second
is distinctly smaller than the others, the third
is entering the orbit. Eight infralabial scales on
the right, and nine on the left side. Temporal
scales form a horizontal row and consist of three
scales on each side, decreasing in size poste-
riorly. Medium-sized mental scale, triangular,
sides about equal. First pair of infralabial scales
in short contact behind the mental. One pair of
chin shields, 1.5 times longer than wide. Gu-
lar scales along the median from chin shields to
ﬁrst ventral scale in ﬁve rows, from infralabial
to infralabial in ten rows. Body scales elongated
and mainly keeled. The keel is in the centre and
12 P. Wagner et al.
nearly as long as the scale. Apical pits absent.
The ﬁrst two scale rows bordering the abdom-
inal shields smooth, the second row becoming
keeled at the last third of the body towards the
tail. Body scales arranged in 22 rows interiorly
(at ventral scale no. 15), in 21 rows at midbody
(no. 79 =half of the SVL) and in 17 rows pos-
teriorly (no. 143). There are 149 ventral scales,
and additional two preventral scales. Cloacal
plate not divided. First 18 subcaudal scales un-
paired (totally two third of the tail length), fol-
lowed by 24 paired scales, resulting in a total
number of 42 subcaudal scales.
Coloration. The general coloration consists
of various different tones of olive, tan and
brown. The head pattern is distinct, while the
body pattern is very vague. Pattern of the head
with a dark brown, somewhat triangular spot sit-
uated medianly on the internasal and prefrontal
scales. Followed by a pair of somewhat quad-
rangular spots, nearly in contact to each other,
and one on each side of the head, occupying
most of the supraocular scales posteriorly and
parts of the frontal and parietal scales. A second
pair of rhomboidal spots, not nearly in contact,
is situated on the posterior part of the parietal
scales. This pair is followed by a pair of elon-
gated blotches on the back of the head, becom-
ing conﬂuent posteriorly and enclosing a sand-
hour like light area in the median area of the
hind head. A dark brown, somewhat white bor-
dered, cheek stripe extends from the orbit to be-
yond the angle of the jaw. The upper white line
occupies the lower edges of an adjacent row of
horizontal temporal scales, while the lower ex-
tends from the lower postocular across the lower
edge of the ﬁrst temporal scale and the last three
supralabial scales. Supralabial scales speckled
with gray, infralabial scales dark brown with
whitish spots. The body coloration consists of
a series of very indistinct, irregular, often in-
complete, and narrow pale olive dorsal cross-
bands which are incompletely and indistinctly
dark brown dark bordered and do not extend to
the lateral sides of the body. Crossbands longi-
tudinal along the median line two, rarely three,
scale rows long, and across the body seven scale
rows wide. Totally in 29 crossbands along the
body, and in 6 along the tail. The belly is dirty
white with a gray speckling. A series of dark
brown regular ventrolateral blotches is present
on each side along the entire body.
Skull. The skull of Gloydius rickmersi sp. n.
is overall similar to the skulls of G. h. halys
and G. h. caraganus (ﬁg. 4), and distinct to
G. blomhofﬁi and G. himalayanus which have
more elongated and ﬂat skulls. It is similar to
G. h. caraganus in the shape of the premaxilla
bones and the elongated quadrate, but distinct
in the dentition of the lower maxilla and the
wider angle of the quadrate bones. In the latter
characters it resembles G. h. halys from which
it is distinct in the shape of the premaxilla. The
dentition of the upper jaw is similar to taxa of
the G. halys complex but distinct to G. blomhofﬁ
which has much smaller teeth. The new species
has more teeth in the lower jaw than G. h.
caraganus but a similar number as compared
with G. h. halys.
Variation. Both rather complete paratypes
generally agree with the holotype. MHNG
2745.46 has a body length of 371 mm and a tail
length of 56 mm. The head scalation is identical
to the holotype, but the infralabial scales are not
countable. There are two preventral, 154 ven-
tral and 35 subcaudal scales which are entirely
paired. Body scales around the body in 21 rows
at the anterior, in 20 rows at the midbody and 15
rows at the posterior portion of the body. The
26 body and 4 tail crossbands are much more
indistinct and narrow than in the holotype. ZSM
211/2014 has a body length of 312 mm and a tail
length of 71 mm. The scalation is identical with
the holotype but differs in the following aspects:
There are 9 infralabial scales on both sides, two
preventral, 141 ventral and 45 entirely paired
subcaudal scales. Body scales around the body
in 22 rows at the anterior, in 22 rows at the
midbody and 17 rows at the posterior portion
of the body. Body and tail coloration similar to
the holotype, but the crossbands are too indis-
tinct and not countable. The belly coloration is
New species of Gloydius 13
Figure 4. Digital X-ray image (left: dorsal; right: lateral view) of the holotype of Gloydius rickmersi sp. n. (ZMB 80360),
compared with ﬁgures modiﬁed from Gloyd and Conant (1990). For further details see Material and methods.
uniform cream, lacking the gray speckling and
dark brown blotches as in the holotype.
Distribution. So far the new species is only
known from its type locality and one locality
at the northern slope of Alai range (Kichelay
at the Tengizbay River, beyond the Tengizbay
Pass, see ﬁg. 5). However, within the small val-
ley of the type locality it has a broader distribu-
14 P. Wagner et al.
Figure 5. Distribution of Gloydius rickmersi sp. n. in the Kyrgyz Republic. 1: Kul-Otek at the Sary-Buka Valley about 25 km
(by air) NE of Daroot-Korgon (Chong-Alai District) near the town Kyzyl-Eshme in direction to the Shuman-Bol pass and the
Kichi-Alai River. 2: Northern slope of the Alai range, Tengizbai River, Kichelay, 2798 m.
tion and occurs from the entrance of the valley
near the Alai lowlands, up to the end of the val-
ley in higher elevation. A specimen recognized
as G. halys from “Daraut River Canyon” by
the PATCA Project most probably refers to the
herein described new species. Other specimens
recognized from eastern parts of the Alai and
from the entire country have to be re-examined
to clarify their identity. Orlov and Barabanov
(1999) mentioned the altitudinal distribution of
G. h. halys and G. h. caraganus as distinct.
While the previous one is referred to elevations
up to 3000 m, the latter is mentioned as in-
habiting deserts and hilly habitats not higher
than 1000 m a.s.l. However, Gloyd and Conant
(1990) are providing different ranges with G. h.
caraganus ranging up to at least 4000 m and
G. h. halys between 150 and 2300 m. Gloydius
rickmersi sp. n. was collected between 2800-
3000 m a.s.l. and therefore within the range of
G. h. caraganus but higher than of G. h. halys
according to Gloyd and Conant (1990). How-
ever, the upper end of the range of G. h. halys
is much higher than G. h. caraganus following
Orlov and Barabanov (1999).
Relationships. The haplotype network (ﬁg. 1)
shows G. rickmersi sp. n. within the G. halys
complex. It is close related to both, G. h. halys
supported by the p-distances of 4.8-5.0%, and
to G. h. caraganus showing a not considerably
higher p-distance of 5.1%. In comparison, the
p-distance between G. h. halys and G. inter-
medius is only 3.6%.
Habitat and ecology. The holotype was col-
lected at dusk on the road slightly above the
base camp (ﬁg. 6), while the topotypical para-
type (MHNG 2745.46) was collected as dead
body, probably killed by a predator next to
a small stream about 200 m above the base
camp. According to Yakovleva (1964) Gloydius
h. caraganus is very variable in Kirgizia regard-
ing the habitat preference and specimens were
found on plains, in steppe foothills, meadows,
open shrubby areas, the rocky banks of rivers,
on mountain slopes and open forests. However,
according to both Gloyd and Conant (1990) and
Orlov and Barabanov (1999) not all Gloydius
specimens in Kirgizia can be recognized as G.
h. caraganus and therefore the mentioned habi-
tats could refer to a variety of taxa. It can be
assumed that the herein described species does
not has a similar variability, and is restricted
to higher altitudinal habitats but should inhabit
both, the rocky and drier areas as well as the
meadows in the valley. Two dead bodies were
found directly next to small streams, which sup-
ports the wet meadows as habitat as well. Sev-
eral old dead bodies were found on the road
along the valley and it can be suggested that
the road is frequently used for body-temperature
regulation before the night. Especially as the
holotype was collected at dusk and according
New species of Gloydius 15
Figure 6. Habitat of Gloydius rickmersi sp. n. at the type locality in the Sary-Buka Valley. This ﬁgure is published in colour
in the online version.
to reports by local people, G. rickmersi sp. n.
seems to be nocturnal.
At the type locality Asymblepharus alaicus
yakovlevae Eremchenko, 1983 as well as Bufo
pewzowi Bedriaga, 1898 were recognized. In
the habitat of the specimen from the north-
ern slope of the Alay Range Eremias nikolskii
Nikolsky, 1905 and again Bufo pewzowi were
Etymology. Nearly 50 years after his death,
this species is named in honor to Willi Rickmer
Rickmers for his contributions to our knowledge
about the Alai and Pamir regions and his out-
standing work as organizer of the ﬁrst German-
Russian expedition to this area.
Two studies are providing morphological data
for this comparison: Gloyd and Conant (1990)
and Orlov and Barabanov (1999). Both studies
are strongly differing in their taxonomic con-
cepts and therefore as well in the morphological
characters provided for the different taxa. As
the previous one provides more details of more
taxa, the specimens from Alai were mainly
compared with the data provided by Gloyd and
Conant (1990), but also with the few characters
given from Orlov and Barabanov (1999). The
examination and comparison with both studies
resulted in differences to all Gloydius taxa, but
especially to both G. h. caraganus and G. h.
halys. Consequently, the specimens from Alai
were hence described as a new species. It was
Orlov and Barabanov (1999) who already men-
tioned that G. h. halys and G. h. caraganus
are sympatric in some areas of their distribu-
tion range. We agree with this, and think that the
morphological differences are strong enough to
recognize G. h. caraganus as full species. How-
ever, a further integrative revision is in need to
clarify the status of the other involved taxa.
According to the unresolved systematics of
the G. halys/intermedius complex the actual dis-
tribution of Gloydius in the Kyrgyz Republic
is varying between the different authors and
needs to be reviewed. Although several au-
thors mention Gloydius halys as not rare in
Kyrgyzstan (e.g., Yakovleva, 1964; Terent’ev
and Chernov, 1965; Eremchenko et al., 1992).
Moreover, Yakovleva (1964) mentioned that
16 P. Wagner et al.
among the, at her time, four known subspecies,
only G. h. caraganus occurs in Kirgizia and
stated that the taxon has been found in almost
all mountain foothills and also penetrated into
mountain ranges. However, Gloyd and Conant
(1990), differentiating both groups by the pres-
ence or absence of apical pits, mentioned that
some of Yakovleva’s (1964) specimens could be
misidentiﬁed G. intermedius and consequently
recognized G. h. caraganus and G. intermedius
from Kyrgyzstan. Later, Orlov and Barabanov
(1999, map 1), not recognizing apical pits as
a character to distinguish both groups, showed
both G. h. halys and G. h. caraganus on a
rough distribution map including Kyrgyzstan,
but failed to present point localities or the re-
spective material. These authors restricted G.
intermedius to NE China, extreme SE Russia
and the Korean Peninsula. Very recently, Sin-
daco et al. (2013) provided a rough scale distri-
bution map with point localities, but only refer-
ring to G. halys as species and excluding G. in-
termedius. A summary of these localities refer-
ring to the different taxonomic concepts is given
in ﬁg. 5.
The different populations of the polytypic
G. halys are distributed over a large area from
western to eastern Central Asia. However, only
three taxa, G. h. halys,G. h. caraganus and G.
h. caucasicus are distributed over larger areas.
Gloydius h. halys and G. h. caraganus seem to
occur in Kirghizia, while other taxa are geo-
graphically restricted. Especially G. h. boehmei
and G. h. caucasicus can be found in reasonable
geographic distance to the Alai, and are sepa-
rated from this valley by high mountain ranges
like the Pamir and the Hindu Kush. In the Kyr-
gyz Republic Gloydius occurs within an a ver-
tical range between 550 m in the “Chuiskaya
Dolina [=valley]” (Yakovleva, 1964) and at
least 4000 m on the southeastern slopes of
the Ferganskiy Mountains (Chernov, 1959). Al-
though, Gloydius rickmersi sp. n. seems to be
a today Alai endemic and isolated high alpine
relict population from a formerly wider dis-
tributed species. Although Gloydius taxa are
very variable in their elevational distribution
(e.g., G. h. halys: 150 to 2300 m; G. h. cauca-
sicus: sea level to at least 3000 m; G. h. cara-
ganus: below sea level to 4000 m ﬁde Gloyd and
Conant ; to 1000 m ﬁde Orlov and Bara-
banov ) an endemic alpine species in the
Alai is not surprising as it is known as an alpine
biodiversity hotspot with even several endemic
vertebrate taxa (e.g., Ellobius alaicus, Roden-
Acknowledgements. This research was ﬁnancially sup-
ported by the National Geographic Society (grant no.
GEFNE81-13). We are grateful to the National Academy
of Sciences of Kyrgyz Republic and the Academy of Sci-
ences of the Republic of Tajikistan for the support of the
scientiﬁc expedition. We thank Spartak Litvinchuk for pro-
viding a paratype. We are grateful to Frank Tillack for his
useful comments on the manuscript and to Roberto Sindaco
for providing additional distribution data. PW, AT, GM and
LB are grateful to all members of the Alai-Pamir expedi-
tion for the great time in the ﬁeld and especially to Camilla
Hansen for her help and her support of the expedition.
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Appendix. Material examined
G. h. boehmei.AFGHANISTAN. ZFMK 8648, Andarab val-
G. h. caraganus.K
AZAKHSTAN. ZFMK 16385, Tsches-
kaskan. KIRGHIZIA. ZFMK 44983, Frunse.
G. h. caucasicus.I
RAN. ZFMK 89231-33, Laar valley.
G. h. halys.K
AZAKHSTAN. ZFMK 59366, without further
locality. MONGOLIA. ZFMK 59365, Chuzhird.
“G. intermedius”. MONGOLIA. ZFMK 44228 Gobi, Dalan.
RUSSIA. ZFMK 51068-69, Buchta Mekovodnaja;
ZFMK 44984-85, Tschingan; ZFMK 49054-56, Ussuri