Content uploaded by Mark Pestov
Author content
All content in this area was uploaded by Mark Pestov on Nov 13, 2020
Content may be subject to copyright.
Herpetological
Review
Volume 51, Number 3 - September 2020
438 ARTICLES
Acknowledgments.—We thank Andrew Holycross for providing
ASU p hoto voucher numbers. Ron Thom pson provided Primero
Conservation.org images. Dennis Caldwell drafted the map in Fig. 1.
Literature Cited
Card, D. C., D. R. Schield, R. H. Adams, A. B. Corbin, B. W. Perry, A. L.
Andrew, G. I. M. Pasquesi, E. N. Smith, T. Jezkova, S. M. Boback, W.
Booth, and T. A. Castoe. 2016. Phylogeographic and population
genetic analyses reveal multiple species of Boa and independent
origins of insular dwarfism. Mol. Phylogenet. Evol. 102:104-116.
Comision Nacional para el Conocimiento y Uso de la Biodiversidad. 2016.
EncicloVida. CONABIO. Mexico, Recuperado en 18 de noviembre
de 2019 de http://www.enciclovida.mx
Enderson, E. F., A. Quiiada-M., D. S. Turner, P. C. Rosen, a n d R. L. Bezy.
2009. The herpetofauna of Sonora, Mexico, with comparisons to
adjoining states. Check List 5:632-672.
Jacobs, S. 2018. Geographic distribution: Boa sigma. Mexico: Sonora.
Herpetol. Rev. 49:287.
Rorabaugh, J. C., and J. A. Lemos-Espinal. 2016. A Field Guide to the
Amphibians and Reptiles of Sonora, Mexico. ECO Herpetological
Publishing and D istribution, Rodeo, New Mexico. 688 pp.
---------
, D. Turner, T. R. Van Devender, V H. Cabrera, R. J. Maynard, R.
W . Va n D e ve n de r, R . A . V illa, P. Ham il to n, S . F. Hal e, C. Ag u ila r -M .,
A . B l an c o- G ., E. Wallace, an d C. He d gc o ck . 2019. H e r p e to f a u n a o f
t h e M e s a T re s R i o s a r e a in th e n o r th e r n S ie r ra M a d r e O c c i d e n ta l o f
S o n o r a, M e x i co . H e r p e to l . R ev . 50:251-259.
Sm it h , H. M. 1943. Summary of the collections of snakes and croco-
dilians made in Mexico under the Walter Rathb one Bacon Travel
ing Scholarship. Proc. U.S. Nat. Mus. 93:3 93-5 04.
Sob arz o, H. 1991. Vocabulario Sonorense. Instituto Sonorense de Cul-
tura, Hermosillo, Sonora, 261 pp.
Van D eve nde r, T. R., С. H. Low e , an d H. E. Lawl er . 1994. Factors influ
encing the distribution of the neotrop ical vine snake Oxybelis ae-
neus in Arizona and Sonora, Mexico. Herpetol. Nat. Hist. 2:27 -44.
-------------, G . Ya n es - A. , G . , A. L. R ein a-G . , M . V alenzuela- Y., M . P. Mo nt a-
Nez-A ., an d H. Silva-K. 2013. Comparison of the tropical floras of
the Sierra la Madera and the Sierra Madre Occidental, Sonora,
Mexico. In G. J. Gottfried, P. F. Folliott, B. S. Gebow, L. G. Eskew,
and L. C. Collins (compilers), Merging Science and Management
in a Rapidly Changing World: Biodiversity and Management of
the Madrean Archipelago III and 7th Conference on Research and
Resource Managem ent in the Southwestern Deserts, pp. 2 40-242.
2012 May 1-5, Tucson, Arizona. Proceedings RMRS-P-67. U.S. De
partment of Agriculture, Forest Service, Rocky Mountain Research
Station, Fort Collins, Colorado.
Herpetolog ical Review, 2020,51(3), 438-446.
© 2020 by Society for the Study of Amphibians and Reptiles
Biodiversity of the Herpetofauna of the
Muyunkum Desert, Kazakhstan
Th e Muyun kum De sert is situ ated in southern Kazakhstan.
Th is sandy de sert, with an area of 37,500 km2, stretches from
southe ast to northw est for a bou t 500 km. It is lo cate d in the
middle of th e N orth Tura nian deserts. Fr om the south and
west, it is bounded by th e Karatau Ridge, from the east by the
Chu-Iliysky M oun tains, and from the north b y the Betp ak-
Dala Desert. B ecause o f its location , this de sert is the lim it of
northward d istribu tion for a nu mber of reptile spe cies and is
ha bitat for isola ted pop ulations o f the psa mm oph ilou s species.
Th e first references to th e herpeto fau na of the Muyu nkum
Desert date from the second h alf of the last century. Paraskiv
(1956) listed a total of 16 species for the Muyunku m D ese rt and
surrounding areas. However, mos t of the fin ding s were mentioned
without precise g eograp hical attributio n. In su bse quent years,
resea rch studies have been fragmentary. In 1967 , ce nsu ses of
Eremias gra mm ica (Lichten stein, 1 823), E rem ias velox (Pallas,
1771), and Ph ryn oce pha lus m ystaceus (Pallas, 1776) were
cond ucted on the southern periphery of the M uyunkum D esert
(Vtorov and Pereshk olnik 1970). In 1978, data on new findings
and abund anc e of five reptile s pecies were obtaine d (Brushko
1995; Kubykin and Brushko 1998; B rushko and Kubykin 2000 ). In
the late 1980s, Erem ias lineola ta (Nikolsky, 1896) was recorded
for the first time on the eastern edge o f the desert; and there w ere
also findings of H em orrho is ravergieri (Menetries, 1832) on the
no rthern edge (Go lubev 1990). In 1994, studies of Teratoscincus
scincus (Schlegel, 1858) in the southeastern p erip hery of the
desert were repo rted (Borkin et al. 2007). By the sa me authors,
10 reptile sp ecies were also re corded there. Two invento ries of
Testudo horsfield ii Gray, 1844 on the sou thern edge o f the d esert
were con ducte d in 2003 and 200 5 (Bond aren ko et al. 2 008 ). The
localities of som e species of lizards and sn akes are given in the
survey monograp hs by Sin daco and colle agues (Sin daco and
Erem che nko 2008; Sind aco et al. 2013); however, due to the small
scale of the ma ps used, it is difficu lt to estim ate the distribution
of sp ecie s in this region. Thus, m ost o f the available info rma tion
on reptiles is frag mentary and refers to areas peripheral to the
Muyu nkum Deser t. There is no c omplete list of reptiles of this
area and the data on th eir distribu tion and p opulation density
are in sufficien t. In this regard, the aim o f this work is to com pile
a list of the herp etofau na of the Muyunku m De sert and also to
cla rify the distributio n and relative abunda nce of the reptiles.
MARINA A. CHIRIKOVA*
YULIA A.ZIMA
Institute of Zoology of the Republic of Kazakhstan, al-Farabi 93,
050060 Almaty, Kazakhstan
MARKV. PESTOV
The Dront Ecological Center, 603001 Nizhny Novgorod, Russia
VLADIMIR A. TERENTYEV
Association for the Conservation of Biodiversity of Kazakhstan,
Beibitshilik 18,010000 Nur-Sultan, Kazakhstan
Corresponding author; e-mail: m.chirikova@mail.ru
Herp etological Rev iew 51 (3), 2 02 0
ARTICLES 439
Mate ri als an d M e t h o d s
Study Area.—Th e Muyunkum Des ert con sists of hilly-ridged
sands of m ora ine and river sediments fro m the Chu River. The
higher sou the rn p art with sand dunes, up to 640 m in height,
de creases grad ually in the no rth and northw est to 110 -130 m.
Th e northern p art of the desert adjoining the Chu Valley co n
sists ma inly of a hilly-ridged plain 15 -30 km wide. N umero us salt
mar shes and pans are located c loser to the Chu River. T he river
dries up in the sum mer. In the river valley, there are thick ets of
willow (Salix sp.), ole aster (E laeagnussp.), and othe r ve geta tion
typ ical for riparian zones. In the w estern part of the desert, fixed
du nes prevail. On the dry aligne d valleys, White E arth Wo rm
wood (Artemisia terrae albae), Erkek (Agropyron sp.), and Black
Saxaul (Haloxylon ap hyllum) d ominate on dry fla t valleys and
the areas betw een dunes, and are covered with dry liche n. The
central part of the dese rt is ch aracterized b y high sand ridges
(5 0-7 0 m high). Vegetation on slo pes is rep resented by different
sp ecies of C alligonum (Calligonum spp.), Sand Aca cia (Ammo-
dendron sp.), W hite Saxaul (Haloxylon persicu m), and M am
moth Wild Rye (Leymu s racem osu s). In the sou the astern pa rt of
the desert there are nu merous places of groundwa ter disc harg e
and lak e-m arshy low lands with bordering reed beds and sedge
mead ows. The northeastern part of the Mu yunkum is covered
with the saxaul trees (Bizhanova 199 8; Dzhanaliyeva 1998; Be -
dareva2009).
Within the Muyu nkum Desert area, there is the Umbet state
hu nting farm, the A ndysay sta te wildlife area, as well as two gas-
co nden sate fields and a ur anium mine.
Timing and Distance o f Researc h Trips.— Expeditions took
place d uring 20 17 -201 9 (31 August 3 1-6 Septem be r 2017, 1-6
September 2018, 4-1 6 May 2019, and 1 1-22 lune 2019) (Fig. 1).
Field work was u ndertak en during th e period s o f maximum
rep tile activity (spring, su mm er and a utumn mo nth s). Th e total
length of the expedition routes was > 20 00 km.
Methods.—To identify reptiles and assess their nu mbers,
walkin g routes (line-transect) were u sed. Fo r obligate
ps ammop hile s of P. mystaceus, T scincus, E. g ram m ica occupying
on ly limited s ections of bare san d, cou nts were con duc ted by
the tria l-ar eas m eth od. Th e width o f walking rou tes w as 3-5
m, dependin g on the vegetation density. The faun istic records
were usually sup plemented with photographs of spe cim ens and
ha bitats. Coordina tes of all localities were recorded usin g GPS.
The results are given in terms of desc riptive sta tistics: minimum
(min) and m aximum (max) va lues, average (M), and average
error (m). The c alculation s were p erform ed in Statistic a 10.0.
Du ring preparation of ma ps, we used our ow n data, literatu re
data, and data from the c ollectio ns of the Institute o f Zoology
of the Republic of Kazakh stan (IZK), the Zoological Museum of
Mosc ow Sta te University (ZMMU), the Zo ological Institute of the
Ru ssia n A cadem y o f Scien ces (ZISP) (Appendix 1), the National
M useum of Natural History, N atio nal Acad emy of Scie nce s
of Ukraine (NMNH) (Sh cherbak et al. 1997 ), and the Kyrgyz
Biological-So il Institute (E rem chenko et al. 199 2). T he maps are
ba sed on Bin g Maps satellite im agery in the SAS Plan et program.
A well-known and informa tive method of the rank/abundance
plot (or W hittaker plot) was used to disp lay the d istribution of
sp ecies abu ndan ce (Magu rran 2004 ).
Ta xon omy and nomen clature of rep tiles was ado pted from
the l atest sy stema tic lists o f Kazak hstan (D ujseb ayev a20 13) and
Re ptile Datab ase (Uetz et al. 2019). The populatio ns of toads in
Muyunkum require taxono mic c larific atio n. A ccordin g to the
la test revision, Bufotes p errini M azepa et al., 201 9 inha bit areas
no rth of Karatau (Dufre snes et al. 201 9). U ntil spe cial studies
are implem ented , we will c ons ider tha t this species inhabits
Muyunkum De sert. Zoo geographical affiliation s w ere identified
according to the c lassifica tion o f Rustamov (1981 ) and Vigna
Taglianti et al. (1999).
Resu lt s
Species co mp osition.— We found one spe cies of am phibian
and 14 specie s of reptiles. Two species (Natrixtessellata [Laurenti,
1768] and Pseud opus apodus [Pallas, 1775]) w ere record ed in
Herp etological Rev iew 51 (3), 2 02 0
PHOTOS BY M. PESTOV (A, С D, F, G, H), V. TERENTYEV ( B), AND V. KOLBINTSEV (E)
440 ARTICLES
Fig. 2. Some reptile species of the Muyunkum Desert: A) Testudo horsfleldiv, B) Teratoscincus semens', C) Trapelus sanguinolentus; D) Phryno-
cephalus mystaceus; E) Eremias scriptcr, F) Eremias gramm ica; G) Gloydius halys; H) Eryx tataricus.
Herp etological Rev iew 51 (3), 2 02 0
ARTICLES 441
Fig . 3 . Distribution of reptiles in the Muyunkum Desert (yellow dots: literature data; red dots: data collected during this study).
* PI a ty ceps karelini
Д Hemorrhois ravergieri
о Psam mophis lineolatus
О Pseudopus apodus
0 Natrix tessellata
® Eryx ta trie us
□ Gloydius halys
о Mediodactylus russowii
aAlsoph ilax pipiens
0 Teratoscincus scincus
® Crossobam on eversmani
О Testudo horsfieldii
О Eremias velox
A Eremias intermedia
Eremias lineolata
Erem ias scripta
Eremias grammica
Eremias arguta
areas adjacent to the de sert (Table 1, Fig. 2 ). Pseud opus apo dus
is inclu ded in the Red B ook of Kazakhstan; Testudo horsfie ldii
and Eryx tataricus (Lic htenstein, 1 823), in Appendix II CITES
(Convention on In terna tiona l Trade in En dan gere d Specie s of
Wild Fau na and Flo ra). T he C entral Asian turtle is listed in the
IUCN Red L ist (Vulnerable, VU).
Distribution.—T he m os t com mo n specie s are T horsfield ii
and T sang uinolentus (Fig. ЗА, С). In contra st, Mediodactylus
russowii (Strauch, 1887), E rem ias scripta (Strauch, 1867), Gloydius
halys (Pallas, 1776), and H. ravergieri (Fig. 3B, E, G, H) are rare.
Phryn oceph alus mystaceus, E. interm edia, E. lineolata, E. velox,
Eryx tataricus, and P. lineolatus (Bran dt, 1838) were observed in
the central, southern, and eastern parts of the Muyunkum Desert,
and are practically not se en at all in the w estern part. Eremias
grammica was registered in the southw est and sou theast p arts of
the d esert b ut not in the centra l an d northern parts (Fig. 3E). The
tadpoles o f Bufotesperrini were observed in artesian well areas in
the north western part o f Muyun kum and in a small lake situated
in the western part of the d esert; two adult sp ecimens were found
in the central and western parts of the d esert.
Habitats.—W ith the exc eption of op en d unes and lake-marshy
lowlands, T hor sfieldii has be en found in all hab itats, including
in the vicinity of s alt pans. Trapelus sang uinolentus was found in
the semi-fixed and fixed sands with calligo num bushe s and saxaul
trees, saxaul fo rest, and at the edge o f salt pans. Teratoscincus
scincus, P. m ystaceus, E. lineolata, E. scripta, and E. grammica
inhabited o pen dunes and/or the a rea of exposed sand on the
semi-fixed sand massifs (Fig. 4A, B). Teratoscincus scincus and P.
mystaceus were also found on roads at the place s with disp ersed
sand, at a distan ce of up to 3 km from an open dune (Fig. 4F).
Eremias velox, E. intermedia, and P. lineolatus were record ed on
semi-fixed sands with small patc hes of bare sand, calligonum ,
Herp etoiogical Rev iew 51 (3), 2 02 0
PHOTOS BY M. CHIRIKOVA (А, С E, G, H) AND YU. ZIMA (B, D. F)
442 ARTICLES
Fig. 4. Habitats of the Muyunkum Desert: A) open dunes; B) semi-flxed sands with exposed patches; C, D) fixed sands; E) saxaul forest; F)
roads with dispersed sand; G) salt pans; H) unpaved lake-marshy lowlands with reed beds and sedge meadows.
Herp etological Rev iew 51 (3), 2 02 0
ARTICLES 443
Tab le 1. Summary list of herp etofauna from the Muyunkum Desert and surrounding areas.
Species Muyunkum Surrounding Literature This study
Desert areas data
Bufotes per riniMazepa et al., 2019 + + + +
Testudo horsfieldii Gray, 1844 + + +
Ablepharus deserti Strauch, 1868 + +
Crossobamon eversmanni (Wiegmann, 1834) + +
Alsophylax pipiens (Pallas, 1827) + +
Mediodactylus russowii (Strauch, 1887) + + +
Teratoscincus scincus (Schlegel, 1858) + + +
Trapelus sanguinolentus (Pallas, 1814) + + + +
Phrynocephalus helioscopus (Pallas, 1771) + +
Phrynocephalus mystaceus (Pallas, 1776) + + +
Eremias velox (Pallas, 1771) + + + +
Eremias intermedia (Strauch, 1876) + + + +
Eremias lineolata (Nikolsky, 1897) + + +
Eremias scripta (Strauch, 1867) + + +
Eremias arguta (Pallas, 1773) + +
Eremiasgrammica (Lichtenstein, 1823) + + +
Pseudopus apodus (Pallas, 1775) + +
Platyceps karelini (Brandt, 1838) + +
Hemorrhois ravergieri (Menetries, 1832) + + +
Psam mophis lineolatus (Brandt, 1838) + + +
Natrix tessellata (Laurenti, 1768) + +
Eryx tataricus (Lichtenstein, 1823) + + +
Gloydius halys (Pallas, 1776) + + + +
and sand acacia shrubs. Mediodactylus ru ssowii was detected on
a saxaul tree at a large ope n dune on the sou thwest edge of the
desert and in a saxaul forest in the eastern part of the M uyunkum
(Fig. 4 E). Eryx tataricusv/as observe d m ainly in the sands fixed by
cereals, shru bs of sand acacia , and saxaul trees. Pseudopus apod us
was reco rded on the road alon g a farm field near the M uyun kum
Desert.
Number o f individuals a nd pop ula tion density.—In terms
of num bers, E. velox, T. sang uinolentu s, and T. h orsfie ldii (in
decreasing order) were the most frequently enco untere d (Fig. 5).
Th ey were follow ed by the following reptile species: R mystaceus, T.
scincus, P. lineolatus,E. tataricus, and E. interm edia. Mediodactylus
russowii, E. scripta, G. halys, and H. ravergieri were recorded o n a
few occas ions .
Fig. 6 shows the density graphs for som e reptile species. T he
density o f T. horsfieldii in the southw este rn pa rt of the sands is
slightly higher (16/ha) than in the eastern one. On altered h abitats
ne ar abandon ed sheepfold she ds, a higher density (up to 26.64/
ha) was noted. The po pula tion density of T. sanguinolen tus
averaged 20.5/ha, b ut in some a reas reached 40-53.2/h a. Like the
Central Asian Tortoise, the density of the steppe a gam a turned out
to be lower in the eastern part o f the desert.
Th e po pula tion den sity of P. my staceus was 5.28-30.0/ha. In
some pa tches of exposed sand in the central part of the Muyunkum
De sert, there were up to 10 spe cimen s per 4 0 m2, which, when
rec alculated, is m ore than 1000/ ha (due to the scale in Fig. 6, we
do not give these data).
We calculated the density of T. scincus at on ly several localities
in the southw estern and southeastern parts of the san ds. On a
large dune in the southwe stern part o f the sand s, there were five
Fig . 5. The rank/abundance or Whittaker plot of reptiles in the Muyun
kum Desert and surrounding areas. The ordinates on the logarithmic
scale (log10) show the num ber of individuals, and the x-axis shows the
ranked sequence of species from the most to the least abundant.
adults and 16 sem i-ad ult individuals, which yielded a density
estim ate of 84/ha. By con trast, in the sou theastern area at three
different sites we obta ined density estim ates of 19.9/ha, 5.2/ha,
and 39.9/ha.
Eremias velox w as the dominant species in the eastern part
of the sands. Its density varied betw een 1.3-49.9/h a (Fig. 6). At
the sam e tim e, areas of very high den sity were observed on the
order of 240-40 0/ha (due to the sca le in Fig. 6, we did no t provide
these data).
Herp etological Rev iew 51 (3), 2 02 0
444 ARTICLES
Fig. 6. Population density of some species o f reptiles in the Muyunkum Desert.
□
□
□
Test и do horsfieldii
Tг up chi\ sanffuinolen/us
Phrynmephulus helioscupus
Eremias intermedia
Eremias velox
Eremias lineolata
Eremias sc rip!и
Eremias grummicu
Eremias g ramm ica w as re cord ed at two lo calities in the
southw est and so uth eas t parts of the desert, w ith the p opu latio n
de nsity o f 13.7 ± 3.7 (4.9-19.9) and 9.9 6/ha, respectively. Erem ias
scriptav/as rare and the population density in the southe ast pa rt
of sands was 2.46/ ha. E rem ias lineolata is a co mm on spe cies,
with density estimates of 7.5-24.0/h a. In the sou thern p art o f the
sands, there were patch es of sand on which we observed up to 9
sp ecimen s in an area o f 25 m2.
Incre ased occurrence of P lineolatus was also observed in the
southern and southeastern parts o f the desert (7.9-12.0/ ha). D en
sity estim ates fori?, tataricusv/as slightly lower—6.0 and 9.0/ha.
Dis c u ss i o n
The results of the litera ture review and o ur obtain ed data
showed that one spe cies of am ph ibia n and 16 s pec ies of reptiles
were fou nd to inh abit the Muyunkum D esert; and another
six spe cies pres ent in surro unding areas. We did no t record
six species m entio ned in the litera ture relating to the region:
Crossobam on eversm anni (Wiegma nn, 1834) (Paraskiv 1956;
Go lubev 1990; Brushko 1995; Borkin et al. 2 007 ), Alsophylax
pipiens (Pallas, 1827) (Brushk o 1995; Shche rbak et al. 1997),
Phryno cep halus heliosc opus (Pallas, 1771) (Brushko 1995),
Ablep haru s deserti Strauch, 1868 (E rem chenk o e t al. 1999),
Eremias arguta (Pallas, 1773) (Shcherbak 1974), and Platyceps
karelini (Brandt, 1838) (Paraskiv 1956). Information on records
of Varanus griseu s Daudin, 1 803 h ave a ppeared p erio dica lly
(Brushko 1995; Chirikova et al, 2019), bu t o ur stud ies show that
the V. griseus does not in habit the Muyunkum.
Th e num be r o f reptile s pec ies in the M uyunkum Des ert
is 31.4 % o f the to tal numb er o f reptile specie s o f Kazakhstan
(Dujseb ayeva 2013). T he herp eto fau na is quite h om oge neo us
as to its origin: 87.5% of it is com posed of Tura n elem ents. Only
G. halys and H. raverg ieri belon g to othe r comple xes. Th at is,
the herpetofau na of this desert is mainly ch aracterized by true
de sert s pec ies (Lowe 1 989). Unlike the Kyzylkum (Brushko
1995; Zima and Chirikova 201 0), such Sah ara-Sind ian species
as V. g riseus and Spalerosophis diadem a (Schlegel, 1837) are not
found in Muyunkum. Due to the n orthe rn loc ation of the desert,
there are no species of th e Iranian-Afghan, Kazakh-Mongolian ,
or o ther complexes typical of more southern deserts of Central
Asia (Bo gdanov 196 5; S hch erb ak 1994). Phrynocephalusguttatus
(Gme lin, 1789) and P hryn ocepha lus intersca pularisL ichtenstein
and M arte ns, 1 856, the repres entatives of the Central Asian
fauna, are also not found in Muyunkum . However, a ccording
to D unayev (2009), P guttatu s lived in th e M uyunkum in the
Lower an d M iddle Quatern ary Period . Th e M uyunkum Desert is
po orer than the sand y deserts o f the Northern Caspian region
(Nakaren ok 2002) and the N orthern Aral region (Lobachev
et al. 1 973; Dinesman 1953), where s teppe elem ents o f the
herp eto fau na are includ ed.
In ge neral, th ree species—E. velox, T. sanguinolentus, and
T. h orsfieldii— dominate in the Muyunkum D esert (Figs. 3,
5). Trapelus sang uin olentus and T. horsfieldii do minate in the
western part of the sand s and E. velox in the central a nd eastern
parts. The d ensity of T. ho rsfieldii in some area s was higher tha t
ea rlier estim ate s (Brushko and Kubykin 1981; Bondarenko et al.
20 08). V torov and Pereshk olnik (1970) reported al ow d ens ity of
E. velox (2/ha) in the sou thern area o f Muyunkum com pared to
the cong ene ric E. gram mica (18/ha). B ecau se their counts were
co ndu cte d in Septemb er, such results can be explained as the
result of decrea sed activity. However, our survey s—cond ucted
at peak activity— in dicate th at in sou the astern pa rt of the desert,
the nu mb ers o f E. velox are de clining , while E. g rammica is
increas ing, and in the sou thw estern part E. velox is replaced by
E. gra mm ica (Fig. 6).
Th e po pulation dens ity estim ate we obtain ed for T. scincus
in th e southeas tern part of the de sert is com parable with data
ob tained earlier by B orkin e t al. (2007) for the s outhe rn edge
of the san ds. In the southe astern part, the density was sligh tly
higher. We a ttribu te this to fe wer suitable habitats in this p art of
the desert and as a result, a high con centration of lizards on the
areas t hat are s uita ble for them .
We fo und T. horsfieldii, T. scincus, T. sanguinolentus, E.
lineolata, E. velox, E. grammica, P lineolatus, and E. tataricus
in new lo cations, exte nding o ur un der stan din g of th eir
Herp etological Rev iew 51 (3), 2 02 0
ARTICLES 445
distr ibutio ns in this desert (Fig. 3) (Brushko and Kubykin 1981;
Brushko 1995; Borkin et al. 2 007 ; Kubykin and Brushko 1998).
Regar ding d istribution patterns, species can be divided into the
following groups: 1) ubiquitou s; 2) pa tchy wide spread; 3) known
from single records ; 4) living in areas ad jac ent to the Mu yunkum
Desert and able to pe netrate its edges.
Testudo horsfieldii and T. sangu inolen tus belong to the first
group. Th ese species are widespread in the M uyunku m Desert,
inhabit various habitats, an d mak e up a dense popu lation. T he
no rthernm ost popula tion o f T. horsfieldii inhabits this area.
Bo ndaren ko and Du jsebayeva (2012) assum ed th at T. horsfieldii
within Muyunkum included two subreg ional groups: “A shchikol-
Chu” and “Talas-Muyunkum.” Having evaluated all available
informa tion , we con clud e that the distribution o f the Central
Asian Turtle is u niform throughout the M uyunkum .
Bufotes perrini, P. mystaceu s, T. scincus, E. velox, E. intermedia,
E. lineolata, E. tataricu s, and P lineolatus are m em bers o f the
second group. The se s pecies are clo sely linked to ce rtain biotopes,
which in turn d etermine s the mo saic nature of their distribu tion.
The third group in cludes Crossob amon eversm anni
(W iegma nn, 1834), M. russowii, E. scripta, E. grammica, P.
karelini, H. ravergieri, and G. halys. All the se species are know n
on ly from single records m ainly from th e p eriphery o f the sands
(Fig. 3) (Paraskiv 1956 ; Brushk o 1995; Bo rkin et al. 2007 ; IZK and
own colle cted data). The localities o f C. eversm ann i, M. russowii
and E. scripta in th e M uyunkum Desert are the n orthe rnmost for
these s pecies (Sindako and E rem che nko 2008). Gloydius halys
was know n fro m the no rth east periphe ry of the Muyunkum (IZK,
Kubykin and Brush ko 1998); and we also found it in the eastern
pa rt o f the d esert (Fig. 3G). M ost likely G. halys enter ed the
Muyunkum De sert from the Karatau and Zhamb yl mo untains,
and from the B etp ak-Dala De ser t where it is a com mon species
(Kubykin and Brus hko 1 998). H em orrhois ravergieri was recorded
along the n orthe astern edge of the d esert and c an b e foun d
along the Chu River valley (G olub ev 1990) and in lake-ma rshy
low land s in th e eastern p art of the desert.
Th e fo urth group includes A. pipiens, P. helioscop us, A. deserti,
E. arguta, P. apodus, and N. tessellata. Most of thes e species are
sclerobionts, with single records known from surrounding
areas or the edge o f the sand m assif (Paraskiv 1 956; Brushko
1995; Sh che rba k et al. 1997; Erem che nko et al. 199 9; this study)
(Fig. 3). Du e to th e peculia rities o f their biology, these s pecies
do not pene trate deep into sandy area s and they should no t b e
considere d typ ical inhab itants of the Muyunkum Desert.
Th e un even d istr ibution o f reptiles in the M uyunkum is
primarily due to climatic features. In the w estern p art o f sands,
the average annual pr ecip itatio n is less and sun shine duration
is lo nger com pared to the ea ster n part (M edeu 2010). T his
also dete rmines th e c ompo sition of ve getation: Black Saxaul
co mm un itie s are lo cated in the w este rn pa rt, W hite Saxau l
co mm un itie s in co mb ina tion w ith p sam mo phytic-shrub
co mm un itie s in the cen tral and n orthern p arts o f the d esert
(Beda reva 2009 ). An other im portant fac tor is surfac e relief: the
pr esence o f sand dunes and se mi-fixed sands with patc hes of b are
sand in the southern part of the Muy unkum De sert positively
influ ences the distrib utio n o f psam moph ilous reptile species,
in p articular E. gr am mica and P. mystaceus. M ore wid espread
pastu re farming in the southe astern an d cen tral parts also has
a positive impa ct on the d istribu tion of s ome r eptile species.
Becau se of livesto ck grazing, san d is dislodged and fur ther wind
erosion occu rs, whic h leads to the form ation o f suitable habitats
for psammop hile s (Na karenok 2002).
Based on th e results of this study, it appears tha t herp etofaunal
diversity o f this d esert is relativ ely im poverished c omp ared to
sandy dese rts of th e N orthern Caspian, the N orthern Aral region,
and sou thern sand deserts of Central Asia. The distribution of
rep tiles in the M uyunkum Desert is uneven due to diffe rences in
clima te, relief, and degree of grazing in the we stern, eastern, and
southern p arts o f the desert. Additional surveys are needed to
cover a wider range o f ha bitats, such as lake-marshylo wla nds and
cla y areas, to be tter understan d the herpetofau na o f Muyunkum.
Acknowledgments.—We thank Vassiliy Fedorenko, Rustam Kara-
balayev, and An drey Gavrilov for their assistance during our field work;
Vassiliy Fedorenko for help in the creation of the maps; and Tatjana
Dujsebayeva, Valentina Orlova, and Konstantin Milto for the oppor
tunity to work with collection catalogues; and Vladimir Kolbintsev for
allowing the use of one of his photographs. This work was carried out
within the Central Asian Desert Initiative (CADI), which is a part of
the International Climate Initiative (IKI). The Federal Ministry for the
Environment, Nature Conservation and Nuclear Safety (BMU) sup
ports this initiative on the basis of the decision adopted by the Ger
man Bundestag.
Litera tu re C ited
Bedare va, О. M. 2 0 0 9 . Ecosystems of the middle deserts of Kazakhstan
and their inventory by remote sensing methods. Unpubl. Ph.D. th e
sis, University of Kaliningrad, Kaliningrad, Russia. 3 72 pp. (in Rus
sian)
Bizh anova, G. 1998. Anthropogenic transformation of sand desert veg
etation in Kazakhstan. Unpubl. Ph.D. thesis, Institute of Botany and
Phytointroduction, Almaty, Kazakhstan. 50 pp. (in Russian)
Bog da no v, O. P. 1965. Ecology of Reptiles of Central Asia. Nauka, Tash
kent. 257 pp.
Bon da renko, D . A., and T. N. Du isebay eva . 2012. Central Asian turtle,
Agrionemys horsfieldii (Gray, 1844), in Kazakhstan (its distribution,
hab itat division, and population density). Curr. Stud. Herpetol.
12:3-26. (in Russian, summary in English)
---------
, E. A. Pere go n ts ev , and G. B. Mu kh ta r. 2008. Assessment of the
current state of populations of the Central Asian tortoise (Agrione
mys horsfieldii Gray, 1844) in the landscapes of Southern Kazakh
stan. Russ. J. Ecol+. 3:222-226. (in Russian)
Bor kin, L. J., V K. Er em che nk o, an d A. M. Pan fil ov . 2007. On the ecology
ofthe gecko Teratoscincus scincus. Curr. Stud. Herpetol. 7:16-56. (in
Russian, summary in English)
Bru sh ko, Z. K. 1 9 9 5 . Lizards of Desert Regions of Kazakhstan. Konzhyk,
Almaty. 232 pp. (in Russian, summary in English)
-------------, an d R. A. Kub ykin . 1981. The number of Central Asian turtles in
some areas of the Dzham bul region (Kazakhstan). In I. S. Darevsky
(ed.).Voprosy gerpetologii [Problems of Herpetology]. Nauka, Len
ingrad. 24 pp. (in Russian)
-------------, a n d
-------------
. 2000. Distribution and ecology of snake-arrows
(Psammophis lineolatum Brandt, 1838) in Kazakhstan. Selevinia
1-4:130-137. (in Russian, summary in English)
Chir iko va, M. A., Y. A. Zi ma , М. V Pestov, В. M. Gu b i n , a nd A. V Ko valen
ko . 2019. Investigations on distribution, occurrence and threats of
Varanus griseus caspius (Eichwald, 1831) at the north -eastern pe
riphery of its geographical range (Kazakhstan). Russ. J. Herpetol.
26:185-200.
Din esma n, L. G. 1953. Amphibians and reptiles of the south-east of the
Turgai canteen of the country and the northern Aral Sea region. Tr.
Inst, of Geography, Academy of Sciences of the USSR 54:384-422.
(in Russian)
Dufr es n es , C ., G. M az epa, D . J abl ons ki, R. C alia ri O l ive ira, T. W en sel ee rs ,
D . A . S ha banov, M . A uer , R . E rn s t, С. Koc h , H. E. R amIrez- C hav es, K.
Pat ric k M uld er , E . S im ono v, A. T iu te nko , D . K ry vokhy zha, P. L . W e n -
ne kes, О. I. Zinenko , O.V Ko rshu nov , A . M . A l-Jo ha ny , E . A . P er eg ontsev,
R . M asroo r, C . B e tto - C olliar d, D en oEl М . , L . J. B or ki n, D . V Sk or ino v,
Herp etoiogical Rev iew 51 (3), 2 02 0
446 ARTICLES
R . A. Pasynko va, L . F. Mazanaeva , J. M. Rosa no v, S. Du bey , a nd S. Lltv in-
ch u k . 2019. Fifteen shades of green: The evolution of Bufotes toads
revisited. Mol. Phylogenet. Evol. 141:106615.
Dujs eba yeva, T. N. 2013. Classes Amphibians and Reptiles. In A. F.
Kovshar (ed.), Vertebrate Animals of Kazakhstan, pp. 57-82. Atamu-
ra, Almaty, (in Russian)
Dun aev, E. A. 2 0 0 9 . Systematics and paleogeography: conceptual syn
thesis using the example of Phrynocephalus (superspecies guttatus)
(Reptilia: Agamidae). In A. V Sviridov and A. I. Shatalkin (eds.), Evo
lution and Systematics: Lamarck and Darwin in Modern Studies,
pp. 275-298. Archiver of the Zoological Museum of Moscow State
University. KMK Scientific Press, Moscow, (in Russian, summary in
English)
Dzhana liev a, G. M. (ed.). 1998. Physical Geography of the Republic of
Kazakhstan. Kazakh University, Almaty. 266 pp. (in Russian)
Ere mch en ko , V К, A. M. Pan filov, and E. I. Tza rinen ko. 1992. Abstract of
the research on cytogenetics and systematics of some Asian species
of Scincidae and Lacertidae. Ilim, Bishkek. 183 pp. (in Russian)
-------------,
-------------
, L . J. Bork in , an d H . H e lf en b er ge r. 1999. New data on
distribution of the desert lidless-skink (Ablepharus deserti Strauch,
1868) in Kazakhstan and Kirghizstan with notes on ecology and cy
togenetics. News of NAS KR 3-4:1 06-109 (in Russian with English
summary)
Gol ube v, M . L. 1 9 9 0. New finds of reptiles and amphibians in Kazakh
stan. Vestn. Zool. 5:76-78. (in Russian)
Kub yk in R. A., a n d Z. K. B rushk o . 1998. Contemporary spreading and
information on Agkistrodon halys caraganus Eichwald, 1831 (Rep
tilia, Crotalidae) numbers in Kazakhstan. B . KazNU. Series biologi
cal 6:9-13. (in Russian, summary in English)
Lob ac he v, V S., Yu. D . C hug un o v, an d I. N. Chukanin a. 1973. Features of
herpetofauna ofthe Northern Aral Sea. Vestn. Herp. pp. 116 -118. (in
Russian).
Lowe, С. H. 1989. The riparianness of a desert herpetofauna. In D. L.
Abell (tech. coord.), Proceedings of the California Riparian Sys
tems Conference: Protection, Management, and Restoration for the
1990s, pp. 143-148. USDA Forest Service General Technical Report
PSW-110.
Mag ur ra n, A. E. 2004. Measuring Biological Diversity. Blackwell Pub
lishing, Oxford. 256 pp.
Me d eu , A. R. (ed.). 2 0 1 0 . National Atlas ofRepub lic of Kazakhstan. Vol.
I: Natural Conditions and Resources. Institute of Geography RK, Al
maty. 1 5 0 pp.
Nak areno k, E. G. 2002. Ecological aspects of the formation of herpe
tofauna of the Northern Caspian and the trends of its modern de
velopment. Unpubl. Ph.D. thesis, State Pedagogical University of
Nizhny Novgorod, Nizhny Novgorod, Russia. 23 pp. (in Russian)
Par askiv, K. P. 1956. The Reptiles of Kazakhstan. Academy of Sciences
KazSSR, Alma-Ata. 228 pp. (in Russian)
Rus ta mo v, A. K. 1981. Zoogeographic relations of the herpetofauna of
Central Asia and the Caucasus. Bull. MOIP, Dep. Biol. 86:31-36.
Shch erbak, N. N. 1994. Zoogeographic analysis of the reptiles of
Turkmenistan. In V Fet and К. I. Atamuradov (eds.), Biogeogra
phy and Ecology of Turkmenistan, pp. 307-32 8. Kluwer Academic
Publishers, Dordrecht, Netherlands.
---------
. 1974. Racerunners of the Palearctic. Naukova dumka, Kyiv. 294
pp.
---------
, A. A. Tokar, a nd I. V K ir ilen ko . 1997. Gecko lizards (Reptilia:
Sauria, Gekkonidae). The catalog of the collections of the Zoological
Museum of the National Museum of Natural History at the National
Academy of Sciences of Ukraine. Zoological Museum NMNH NAS
of Ukraine, Kiev. 45 pp. (in Russian)
Sinda co , R., a n d V R. Ier em enk o. 2008. The Reptiles of Western Palearc
tic. 1. Annotated Checklist and Distributional Atlas of the Turtles,
Crocodiles, Amphisbaenians and Lizards of Europe, North Africa,
Middle East and Central Asia. Edizioni Belvedere, Latina (Italy). 579
pp.
---------
, A. Ven chi, an d C. Gr ie co . 2013. The Reptiles ofWestern Palearc
tic. 2. Annotated Checklist and Distributional Atlas of the Snakes of
Europe, North Africa, Middle East and Central Asia, with an Update
to the Vol. 1. Edizioni Belvedere, Latina (Italy). 543 pp.
Uet z, P., P. Fre e d , and I. HoSek (eds.). 2019. The Reptile D atabase, http://
www.reptile-database.org, accessed 22 August 2019.
Vigna T ag lia nti, A ., P. A u d is io , M. Bio n di, M. Bol og na , G. Carp an et o, A .
De B ias e, S . Fa tto rini, E. Pia tt ell a, R. Sin daco, A . V en chi , and M. Zapp a-
ro li . 1999. A proposal for a chorotype classification of the Near East
fauna, in the framework of the Western Palearctic region. Biogeo-
graphia 20:31-59.
Vto ro v, P. P., and S. L. Peresh kolnik . 1970. Counts of reptiles at several
points in Central Asia. Zool. Zh. 49:4 68-478. (in Russian)
Zim a, Y. A., and M. A. Chirik ova. 2010. To the fauna of reptiles of the
South-Eastern Kyzylkum. Selevinia. pp. 96-98. (in Russian, sum
mary in English)
App end ix 1
List of inventory numb ers of collections.
IZK: Testudo horsfieldii: 97/3127, 109/1252-1255; Trapelus sanguino
lentus: 55/744-755, 106/1231-1239, 106/1242-1243, 267/2867-2868,
351/3857-3858; Phrynocephalus helioscopus: 83/981-988, 371/4139;
Phrynocephalus mystaceus: 82/978-980, 107/1244-1246, 271/2912—
2915, 190/1941-1944, 803/1; Mediodactylus russowii: 18/4795; Eremi
as velox: 269/2879-2902, Eremias intermedia: 77/972-973, 716/1-2,
754/1-10, 777/1-7; Erem ias gram mica: 81/977, 755/1-4; Eremias line-
olata: 739/1-5, 802/1-3; Natrix tessellata: 207/2153-2154; Hemorrhois
ravergieri: 104/1227-1228; Platyceps karelini: 105/1229; Eryx tataricus.
9/1226; Psammophis lineolatus: 800/1.
ZMMU: Eremias velox: R-6863, R-6866, R-7947; Eremias intermedia:
R-6862; Eremias lineolata: R-6860; Eremias scripta: R-6861.
ZISP: Eremias intermedia: 13479; Gloydius halys: 14812; Eryx tataricus.
17299.
Herp etological Rev iew 51 (3), 2 02 0