DESERT MONITOR LIZARDS (SQUAMATA: VARANIDAE: Varanus: Psammosaurus) FROM THE MIDDLE EAST: FURTHER RECORDS OF NESTEROV’S DESERT MONITOR, Varanus (P.) nesterovi BÖHME, EHRLICH, MILTO, ORLOV ET SCHOLZ, 2015, FROM IRAQ, AND ADJACENT LOCALITIES OF Varanus (P.) g. griseus (DAUDIN, 1803) AND Varanus (P.) g. caspius (EICHWALD, 1831), WITH COMMENTS ON BIOGEOGRAPHY AND TAXONOMY

Abstract

We report on numerous additional records and observations of Varanus nesterovi Böhme et al., 2015, and its chorological relationships (parapatry vs. sympatry) to its close relatives, viz. V. griseus griseus and V. g. caspius. The new distributional data and locality records do not only hint on various threats for V. nesterovi, but document also its parapatry with griseus along a line from east of Mosul to east of Kirkuk, and with caspius near Shiraz. This chorological pattern of V. nesterovi's distribution range wedged between the ranges of the taxa griseus and caspius argues, next to morphological and molecular genetic grounds, strongly for full species status of the three taxa involved. We discuss a theoretically possible but questionable occurrence of V. caspius in Europe (south of Baku, Azerbajdzhan) in the framework of fossil varanids from European sites and we comment on the type locality of Psammosaurus caspius Eichwald, 1831.

Full text

DOI: 10.30906/1026-2296-2023-30-6-518-528
DESERT MONITOR LIZARDS (SQUAMATA: VARANIDAE: Varanus:Psammosaurus)
FROM THE MIDDLE EAST: FURTHER RECORDS OF NESTEROV’S DESERT
MONITOR, Varanus (P. )nesterovi BÖHME, EHRLICH, MILTO, ORLOV ET SCHOLZ,
2015, FROM IRAQ, AND ADJACENT LOCALITIES OF Varanus (P. )g. griseus
(DAUDIN, 1803) AND Varanus (P. )g. caspius (EICHWALD, 1831),
WITH COMMENTS ON BIOGEOGRAPHY AND TAXONOMY
Wolfgang Böhme,1Soran H. Ahmed,2Omar F. Al-Sheikhly,3Korsh Ararat,2
Markus Auer,4Farhad A. Khudur,2,5 and Christian Langner6
Submitted July 3, 2023.
We report on numerous additional records and observations of Varanus nesterovi Böhme et al., 2015, and its
chorological relationships (parapatry vs. sympatry) to its close relatives, viz. V. griseus griseus and V. g. caspius.
The new distributional data and locality records do not only hint on various threats for V. nesterovi, but document
also its parapatry with griseus along a line from east of Mosul to east of Kirkuk, and with caspius near Shiraz. This
chorological pattern of V. nesterovi‘s distribution range wedged between the ranges of the taxa griseus and caspius
argues, next to morphological and molecular genetic grounds, strongly for full species status of the three taxa in-
volved. We discuss a theoretically possible but questionable occurrence of V. caspius in Europe (south of Baku,
Azerbajdzhan) in the framework of fossil varanids from European sites and we comment on the type locality of
Psammosaurus caspius Eichwald, 1831.
Keywords: Varanus griseus group; Middle East and Central Asian distribution; biogeography; elevation of taxo-
nomic rank; conservation.
INTRODUCTION
Varanus nesterovi belongs to those remarkable recent
reptile discoveries originally made in natural history ar-
chives rather than in the field. The Zoological Museum of
the Russian Academy of Sciences in St. Petersburg held
three desert monitor lizards collected by P. V. Nesterov
on 8 June 1914 and catalogued as “Varanus griseus
Daud.” There was, however, a second label on the jar
reading “Varanus schimkewitschi Nesterov,” indicating
that their collector had already suspected the find of a
new species which he, however, due to political and
tragic personal instances, could never describe and pub-
lish, thus making the selected “label name” an unavail-
able nomen nudum. Only 100 years later, in 2015, after
reinvestigation of these remarkable specimens, we for-
mally described them and named them in honour of their
collector (Böhme et al., 2015). The locality of the
holotype, Biare = Byara, Sulaymaniyah Governorate
(35°13¢50¢¢ N 46°07¢15¢¢ E, 1086 m), currently Halabja
Governorate, and the locality of the two paratypes, Abu
Naft River near Nawtkhana village, in Mandali
(34°06¢26.9¢¢ N 45°32¢58.4¢¢ E), Diyala Governorate, are
very close to the border with Iran. The few further re-
cords of this rare monitor lizard, subsequently be traced
so far, were made on Iranian territory on somewhat lower
elevations (e.g., Dezful at 680 m a.s.l.) (Böhme et al.,
2015).
The second Iraqi record of V. nesterovi, this time by
photographic documentation, became known two years
1026-2296/2023/3006-0518 © 2023 Folium Publishing Company
Russian Journal of Herpetology Vol. 30, No. 6, 2023, pp. 518 – 528
1Leibniz Institute for the Analysis of Biodiversity (LIB), Museum
Koenig, Bonn, Germany.
2Department of Biology/College of Science, University of Sulaimani,
Iraq.
3Department of Biology, University of Baghdad, Iraq.
4Senckenberg Natural History Collections, Dresden, Germany.
5Mendel University, Brno, Czech Republic.
6Zoological Garden, Münster, Germany.

Desert Monitor Lizards (Squamata: Varanidae: Varanus:Psammosaurus) from the Middle East 519
Fig. 1. Varanus nesterovi in its natural environment at Mortka, Darbandikhan, Iraq. Photo by Korsh Ararat, 2016.
Fig. 2. Head portrait of the same individual on an oak trunk in its habitat. Photo by Korsh Ararat, 2016.

after the first description, but more than one century after
the collection of the type series in this country! It was the
botanist Bahaa Al-Midhatee on 22 July 2017, who suc-
ceeded to take photographs of a living specimen rather
close to the localities of the type specimens: Tilako vil-
lage near Sangaw (35°07¢42.8¢¢ N 45°04¢35.95¢¢ E), SW
Sulaimaniya Governorate. His very good photographs
showed clearly all diagnostic characters of this peculiar
lizard, viz. a spiny neck scalation, the shape of the nostril
and the tail compressed over its total length with the last
third being uniformly yellowish (Böhme, 2017).
In order to increase our knowledge of this newly
discovered species, we provide here numerous new dis-
tribution records. We discuss these new locality records
in the context of adjacent localities of its two nearest rela-
tives, currently still equipped with subspecific status, i.e.,
V. g. griseus in the west and V. g. caspius in the east.
RESULTS AND DISCUSSION
New records from Iraq
Whether this species was — undetected — already
observed in Iraq before its description by Böhme et al.
(2015) is a bit equivocal because Khalaf (1959) in his
short description of Iraqi desert monitors mentioned the
fading and even complete disappearance of the juvenile
dress in old individuals, which is much more typical for
V. nesterovi than for V. griseus. However, he gave no
localitiy data so that the basis of Khalaf’s statement re-
mains unclear.
Recently, more specimens of V. nesterovi have been
found in Iraq: in June 2016, a living individual was seen
and photographed in its natural environment (Fig. 1) by
Korsh Ararat at Mortka, Darbandikhan, Sulaymaniyah
Governorate (35°0¢45¢¢ N 45°58¢26¢¢ E). It climbed on an
oak tree trunk giving him the opportunity for a nice head
portrait showing the typical features: spiny neck scala-
tion and shape of the nostril (Fig. 2). In 2020, Mr. Mar-
wan Hassan was able to observe and film a specimen at
the same locality showing the entire lizard with its tail
shape and tail coloration (Fig. 3). Subsequently, Soran
H. Ahmed photographed a prepared monitor lizard,
stuffed and mounted by Farhad A. Khudhur in the Uni-
versity of Sulaimani. It was brought there as originating
from the Qaradagh area, and the key character of V. neste-
rovi, the extremely spiny neck scalation, is well visible
(Figs. 4 and 5) while the shape of the nostril, due to the
lack of the skull in the skin, cannot be determined in re-
spect of its original shape.
Due to the efforts of Mr. Aram Ghafoor, we received
further photographic documents, including several im-
pressive video sequences. In one of them it is visible how
an adult V. nesterovi enters a shallow water pool to catch
and eat a lake frog (Pelophylax cf. bedriagae)! In one of
these videos, it was even possible to film a juvenile speci-
men, documenting the juvenile dress of V. nesterovi for
the first time. This specimen was collected (Fig. 6) and
shows a quite similar juvenile dress as its two relatives in
the west and east. It is currently in private possession
(Soran H. Ahmed, personal communication).
520 Wolfgang Böhme et al.
Fig. 3. Another adult V. nesterovi at Mortka, Darbadikhan, showing
the unicolored yellow hindpart of tail. Photo by Marwan Hassan, 2020.

Additional new localities from Iraq are: Sharbazher
area (35°39¢40¢¢ N 45°17¢45¢¢ E) and Sarzal village near
Qaradagh district (35°18¢28.71¢¢ N 45°23¢24.92¢¢ E), both
Sulayamaniyah Governorate; Halabja/Jaleela (35°11¢11¢¢ N
45°58¢26¢¢ E), Halabja Governorate; Koya/Bawaji
(36°04¢59¢¢ N 44°37¢47¢¢ E) and Taq Taq/Elenjagh
(35°53¢15.34¢¢ N 44°35¢21.39¢¢ E), both Erbil Governo-
rate. Very recently, Markus Auer and Christian Langner
saw two specimens (one of them with photographic do-
cumentation, SW of Kani Kirmanj village, Maidan District,
Diyala Governorate, 34°51¢25.5¢¢ N 45°38¢32.07¢¢ E). The
latter two places are at an altitude of ca. 500 m a.s.l., i.e.,
considerably lower than the type locality at 1086 m a.s.l.
(Böhme et al., 2015). Finally, next to further localities in
the Sulaymaniyah Governorate, an important northwest-
Desert Monitor Lizards (Squamata: Varanidae: Varanus:Psammosaurus) from the Middle East 521
Fig. 4. Dry, mounted specimen of V. nesterovi in the University of Sulaimani, prepared by Farhad A. Khudhur. Note the light-colored hindpart of
tail. Photo by Soran H. Ahmed.
Fig. 5. Head and neck of the specimen from Fig. 4 to show the diagnostic neck scalation. Photo by Soran H. Ahmed.

ern record was made at Akre (36°44¢N 43°59¢E), Duhok
Governorate.
Some of the voucher photographs for the new finds
and locality records refer to killed specimens and dramat-
ically show the various severe threats which the interna-
tionally protected lizards are facing. These are for in-
stance car traffic (Fig. 7) and predation by dogs (Fig. 8),
but also deliberate killing by local people (Fig. 9). There-
fore, our former appeal for conservation measures for
this relatively newly discovered monitor species (Böhme
et al., 2015) seems to be more than justified and urgent.
An unlabeled, likewise dry and stuffed desert moni-
tor which was found at the University of Baghdad, be-
longs, however, to Varanus griseus as can be seen by its
much smaller neck scalation and more slit-like nostril
shape (Fig. 10). Although locality data are missing, its
presumed geographical proximity to the range of V. nes-
terovi leads to the question after a possible contact zone
between these two taxa in Iraq.
Other Psammosaurus Voucher Specimens
from the Middle East
and the Question of Sympatry
In his great monograph on varanid lizards, Mertens
(1942) listed all voucher material accessible to him dur-
ing the wartime of WW II. This was also the case for
Varanus griseus (Daudin, 1803). From the large synony-
my/chresonymy list, he considered three taxa as valid
subspecies: these are, apart from the nominotypical form,
V. griseus caspius (Eichwald, 1831) and V. griseus ko-
niecznyi Mertens, 1954. The nominotypic form and
V. g. caspius are framing the distribution range of V. nes-
terovi at its western and eastern margins. Parapatry has
already been demonstrated for V. nesterovi and V. g. cas-
pius near Shiraz in Iran (Böhme et al., 2015) and is cor-
roborated here by a photographic voucher of V. nesterovi
on a market in Shiraz (ZFMK photo archive).
522 Wolfgang Böhme et al.
Fig. 6. First photograph of a juvenile of V. nesterovi, resembling the juvenile dresses of V. griseus ad V. caspius. Photo by Aram Ghafoor.
Fig. 7. Threats for a protected species: A Nesterov’s monitor killed by
car traffic.

The lowest distance between V. g. griseus and V. nes-
terovi was so far between Haditha and Byara, the type lo-
cality of the latter species. It is now further reduced by
the distance beween Mosul (NMW voucher specimens)
and Akre (video record by Korsh Ararat, ZFMK photo
archive), which is just ca. 100 km air distance (Nos. 1
and 4 on Fig. 11). Approximately on a prolonged line
from Haditha to Mosul northwards, there are two
northeasternmost localities in the Kurdish part of Turkey,
viz. Silopi (Ilgaz et al., 2008), and Cizre (photographic
record, fide Soran H. Ahmed), in the triangle between
Iraq, Turkey and Syria.
The two nearest V. nesterovi localities to the next
V. g. caspius distribution point are Gachsaran and Shiraz
as opposed to Ziroud at the Persian Gulf coast, both ca.
250 km apart (Fig. 11 and Table 1). As can be seen from
the map (Fig. 11), all localities of V. nesterovi are situated
along the southwestern margin of the Zagros Mountains
(from ca. 500 m a.s.l.: Maidan/Diyala, to Byara, nearly
1100 m a.s.l.). Those of V. g. caspius (only some mar-
ginal points are included here; for detailed records see
Malakhov and Chririkova, 2018) are reaching from the
coast (Ziroud) towards far inland, e.g., Ghom (930 m
a.s.l.). The V. g. griseus localities, in contrast, are con-
fined to desertic habitats in the Mesopotamian plain.
A Surviving Desert Monitor Lizard in Europe?
Currently, no extant monitor lizard species is known
from European territory as geographically defined by an
eastern border following the Ural Mountains and the Ural
River, through the Caspian Sea to the eastern end of the
main chain of the Great Causasus from which it follows
westwards to the Black Sea. From there it runs through
the Bosporus, the Marmara Sea and the Dardanelles to-
wards the Aegean Sea where the islands on the Anatolian
shelf are excluded (Mertens and Wermuth, 1960; Böhme,
1981). However, fossil representatives of this family
were present in Europe until the middle Pleistocene (near
Athens, Greece: Georgalis et al., 2017).
Several varanid paleospecies have been described
from European and neighbouring fossil sites. For an
Desert Monitor Lizards (Squamata: Varanidae: Varanus:Psammosaurus) from the Middle East 523
Fig. 9. Threats for a protected species: V. nesterovi shot by a local, his
AKM Kalashnikov machine gun serving as a scale is 88 cm long.
Fig. 8. Threats for a protected species: two dogs having killed a Neste-
rov’s monitor.

overview see Molnar, 2004, e.g., V. marathonensis Weit-
hofer, 1888; V. hofmani Roger, 1898; V. atticus Nopsca,
1908; V. deserticolus Bolkay, 1913; V. darevskii Levsha-
kova, 1986; V. lungui Zerova et Chkhikvadze, 1986;
V. amnhophilis Conrad et al., 2012; V. mokrensis Ivanov
et al., 2018; Varanus sp., Vasilyan and Bukhianidze,
2020; Èernanskýet al., 2020). This number was greatly
reduced by subsequent synonymizations. Those de-
scribed before Estes (1983) were already synonymized
by this author with V. marathonensis, but also the pre-
sumed giant species V. amnhophilis from Samos Island
shared this fate (Villa et al., 2018). The latter authors rec-
ognized only two European paleospecies, viz. V. mara-
thonensis and V. mokrensis. This seems problematic as
long as only osteological characters alone are available.
Levshakova (1986) and Stanner (2004) considered
Psammosaurus as composed of three species, viz. the ex-
tant V. griseus and the two fossil taxa V. darevskii and
V. marathonensis. Ivanov et al. (2018) as well as Villa et
al. (2018), in contrast, assigned the last-named species to
the Indian clade where also V. bengalensis is accommo-
dated (Empagusia group of Varanus). Varanus darevskii
from the Pliocene of Tadzhikistan, however, is still re-
garded as a close relative or even predecessor of the ex-
tant desert monitors, and, particularly, on biogeographi-
cal grounds, of V. g. caspius.
When searching for museum specimens from the
Middle East, my attention was directed to a specimen in
the Vienna Natural History Museum (NMW 12459) from
“südlich von (= south of) Baku” (40°24¢N 49°53¢E) do-
nated to the museum by Erich Sochurek on 11 October
1952. This locality is situated northeast of the Great Cau-
casus chain, i.e., geographically indeed a European site
and would document the first record of an extant monitor
lizard on our continent. However, because this conspi-
cious, big lizard was never recorded before from this re-
gion (Bannikov et al., 1977; Alekperov, 1978; Ananjeva
524 Wolfgang Böhme et al.
Fig. 10. Head of stuffed, dry desert momnitor (V. griseus) in the University of Baghdad. Photo by Omar M. Sheikhly.
Fig. 11. Map of the Middle East showing all records of V. nesterovi
(open circles), and the respectively nearest localities of its related geo-
graphical neighbours V. griseus (open squares) and V. caspius (open
lozenges). Map prepared by Morris Flecks.

et al., 2006; Sindaco and Jerjomchenko, 2008), the reli-
ability of this locality information must be viewed with
scepticism. Erich Sochurek (1923 – 1987) was an Aus-
trian amateur herpetologist who was not only famous for
several herpetological collecting travels but also for the
import of reptiles from various countries around the
world. Neither his obituary nor his publication list give
any hint of a personal stay of Erich Sochurek in Azerbai-
jan (Tiedemann and Grillitsch, 1988) so that it can be ex-
cluded that he collected this specimen himself, the more
as the label says “don.” (= donavit, Latin for donated),
and not “leg.” (= legit, Latin for collected).
Desert Monitor Lizards (Squamata: Varanidae: Varanus:Psammosaurus) from the Middle East 525
TABLE 1. List of the localities indicated on Fig. 11
No. Country Region Locality Latitude Longitude
Varanus griseus
1 Iraq Gov. Duhok Mosul 36.3350° N 43.1189° E
2 Gov. Salah Al-Din between Haditha and Baiji 34.6333° N 43.0667° E
3 Kuwait Kuwait City 29.3667° N 47.9667° E
Varanus nesterovi
4 Iraq Gov. Duhok Akre, Gov. Ninawa 36.7177° N 43.9647° E
5 Gov. Erbil Koya, Bawayi 36.1391° N 44.5699° E
6 Taq Taq, Elenjagh 35.9561° N 44.5440° E
7 Gov. Sulaymaniyah Akhjalar 35.7426° N 44.8967° E
8 Tilako village, near Sangaw 35.1286° N 45.0767° E
9 Sarzal village, Qaradagh 35.4427° N 45.3086° E
10 Qaradagh 1 35.3148° N 45.3434° E
11 Qaradagh 2 35.3095° N 45.3658° E
12 Qaradagh 3 35.3349° N 45.4057° E
13 Qaradagh 4 35.3846° N 45.4140° E
14 Qaradagh 5 35.3364° N 45.4322° E
15 Qaradagh 6 35.3691° N 45.4552° E
16 Arbat 35.4514° N 45.5512° E
17 Barznja 35.4525° N 45.7169° E
18 2 km E Sewsenan 35.2085° N 45.5160° E
19 Shekh Tawil 35.0191° N 45.4623° E
20 Bawakhoshen 35.1657° N 45.6202° E
21 Darbandikhan 35.1282° N 45.6193° E
22 Darbandikhan, Gullan Mountain 35.0992° N 45.6265° E
23 between Darbandikhan and Mortka 35.0835° N 45.7491° E
24 Gov. Halabja Bamo town 34.9553° N 45.7877° E
25 Barwin 34.9151° N 45.8373° E
26 Halabja, Jaleela 35.1955° N 46.0286° E
27 Byara village (type locality) 35.2306° N 46.1208° E
28 Gov. Diyala SW of Kani Kirmanj village 34.8571° N 45.6422° E
29 Abu Naft River near Nawtkhana village 34.1072° N 45.5496° E
30 Iran Ilam Prov. Mehran City 33.1167° N 46.1667° E
31 Dehloran 32.6833° N 47.3000° E
32 Abdanan 32.9500° N 47.4500° E
33 Khuzestan Prov. Dezful 32.3833° N 48.4667° E
34 Kohgiluyeh and Boyer-Ahmad Prov. Gachsaran 30.2000° N 50.7833° E
35 Fars Prov. Shiraz (market) 29.6100° N 52.5425° E
Varanus caspius
36 Iran Ghom Prov. Ghom 34.6500° N 50.8833° E
37 Bushehr Prov. Ziroud 28.1189° N 51.3692° E
38 Fars Prov. between Neyriz and Darab 28.7519° N 54.5444° E
39 Kerman Prov. 50 km NW Dowlatabad village 29.0167° N 56.8833° E
40 Azerbaijan Baku 40.4000° N 49.8833° E
41 Turkmenistan Dardzha Peninsula (type locality) 39.7563° N 53.7454° E

Therefore, the first conclusion would be that this liz-
ard bearing still the obliquely striped juvenile dress in its
adulthood would just be a misplaced or wrongly labelled
specimen of V. g. caspius, for which this retention of the
juvenile dress is typical. This taxon has a vast distribution
from central Iran (Ghom) and the eastern coast of the
Caspian Sea in the west to Tadzhikistan in the east (Ehr-
lich, 2013; Chirikova et al., 2019). Baku and the Apshe-
ron Peninsula which are forming the opposite coast of the
region of the type locality of V. g. caspius, viz. Dardsha at
the Turkmenbashi Gulf (Eichwald, 1831: “Hab. in littore
orientali caspii maris, ad sinum balchanensem, in penin-
sula Dardsha”). It seems that Mertens (1942) believed the
Dardsha peninsula and the Balchan Bay to be two differ-
ent places because he restricted Eichwald’s locality to the
peninsula’s name by adding: “außerdem (= moreover) in
littore...” etc. and repeated this (as “Terra typica restri-
cta”) three times (Mertens, 1954, 1959, 1963). This re-
striction, however, is unjustified because the Balchan
Bay is (or respectively was: see Scobel, 1901) just a wa-
ter arm separating the Dardsha peninsula from the adja-
cent landmass (see Fig. 11).
The western margin of the V. g. caspius range is thus
separated from the questionable locality Baku by an air
distance of ca. 280 km across the Caspian Sea. However,
as can be seen from Fig. 11, the sea depth is rather shal-
low between these sites separating two much deeper bas-
ins from each other, so that a dry connection during the
Pleistocene is well imaginable. According to older and
current maps, the sea level is today markedly lower than
1870 because the former Balchan Bay dissappeared mak-
ing the Dardzha peninsula today an integrative part of the
mainland which currently includes even the former
off-coast Tscheleken Island. Moreover, the steppe region
south of Baku (Mugan Steppe) fits the ecological require-
ments of V. g. caspius according to the species distribu-
tion model by Malakhov and Chirikova (2018). How-
ever, it is also possible that the Baku specimen from the
Vienna Museum is a misplaced individual which may
have been inadvertendly transported from the Turkme-
nian bank of the Caspian Sea to the Baku area. There are
even some — undocumented — rumors that a Russian
zoologist has tried to establish a monitor population in
the Mugan steppe which was, however, prohibited by lo-
cal farmers (M. Auer, unpublished information) but the
truth content of these rumors cannot be verified here. So,
irrespective of the habitat suitability in the Mugan steppe
south of Baku, there is no reliable record of the existence
of the Caspian desert monitor in Azerbajdzhan.
Taxonomic Rank of the Psammosaurus Taxa
According to De Lisle (2009), Eichwald’s type (holo-
type by implication) was kept in the “Evermann Mu-
seum” (misprint for Eversmann) of the University of
Kazan (EMKSU), Tatarstan, Russia, but could not be lo-
cated there. This was corroborated by Doronin et al.
(2017) who found a skeleton of this taxon in the Kazan
Natural History Museum which was described by
Eichwald (1838), but did not fit the collection data of
Eichwald’s (1831) earlier taxonomic description (an
English translation of the original description is provided
at the end of this paper). The holotype of Psammosaurus
caspius Eichwald, 1831 must therefore be regarded lost
(Doronin et al., 2017). For the reader’s convenience, we
add an English translation of Eichwald’s description at
the end of this paper.
Apart from the distinct morphological differences be-
tween the nominal taxa griseus,nesterovi, and caspius
(shape of nostril, neck scalation, dorsal color pattern, tail
shape and tail coloration), also the chorological pattern of
the three parapatric forms speaks for their full specific
rather than subspecific status. This becomes even more
likely as the two taxa framing the range of V. nesterovi on
both sides turned out to be genetically considerably dis-
tant from each other in the analysis by Brennan et al.
(2021).
CONCLUSION
Our review of the Middle East desert monitors
(Psammosaurus clade of Varanus) revealed the center of
speciation of this Saharo-Sindian group to be in South-
west Asia. In addition to the three Middle East species
V. griseus,V. nesterovi, and V. caspius, there is also a
fourth recent species of this clade, viz. V. koniecznyi
Mertens, 1954, an inhabitant of southern Pakistan and
northwestern India (Mertens, 1954; Böhme, 2003). The
SW Asian origin of the Psammosaurus clade is further-
more most plausible by V. darevskii, the extinct predeces-
sor and Pliocene ancestor of V. caspius (Levchakova,
1986) from Tadzhikistan. This is congruent with the
phylogenetic tree published by Portik and Papenfuss
(2012) where the clade containing V. griseus is the sister
clade of the Polydaedalus group of Varanus, a strictly
African clade with only one outlier (V. yemenensis)inthe
southwestern Arabian Peninsula. The new distributional
data provided in this paper add to our picture of the
chorology of V. nesterovi as an endemic inhabitant of the
southwestern Zagros Mountain range. Its distribution
area in submontane to montaine steppe habitats is
wedged between those of its two much more widespread
526 Wolfgang Böhme et al.

relatives: V. griseus in the west and V. caspius in the east.
The identified areas of their parapatric occurrence in both
contact zones argue for full species status of the four re-
cent taxa forming the Psammosaurus clade, next to the
distinct morphological differences by which they are dis-
tinguished from each other. Further support for this con-
cept can be expected by closer infra(sub)specific analy-
ses of both V. griseus and V. caspius which seem also to
be composed of several morphologically well distin-
guishable populations within their huge distribution
ranges while a molecular genetic revision of the entire
Psammosaurus group is highly warranted but still pend-
ing. The recent genetic analysis of the genus Varanus as a
whole by Brennan et al. (2021) has demonstrated already
a considerable distance between the two nominal taxa
griseus and caspius. The much smaller distribution range
of V. nesterovi wedged between the two former and its
various threats described here, highlights also the ur-
gency of immediate conservation measures.
Acknowledgments. Our first thanks go to Mr. Aram
Ghafoor for providing us with rich photographic material in-
cluding video films giving first insights into the natural history
of Nesterov’s monitor lizard. He also secured the first juvenile
of this species, thus enabling us to compare its color pattern
with that of the two geographically adjacent neighbouring taxa.
More photos were made by Mr. Marwan Hassan. Further, we
thank Frank Tillack and Georg Gassner of the Natural History
Museums in Berlin and Vienna respectively, for information on
Middle East desert monitors under their care, and Georg
Gassner together with Alice Schumacher also for photographs
of the corresponding voucher specimens. We received valuable
photographic records of V. caspius by Dr. Jiøi Moravec, Prague,
and by Julia Zima, Almaty. Important but hardly accessible lit-
erature references have been sent to us by Prof. Dr. Aaron M.
Bauer, Villanova, PA, and by Dr. Igor V. Doronin, St. Peters-
burg. Additional helpful advice came from Dr. Daniel Jablon-
ski, Bratislava, and from Willi Miesen, LIB (ZFMK) Bonn.
Thanks are due also to Abdul-Karim Abdul-Rahman Rashid,
major of Kani Kirmanj, for his support during fieldwork of
M. A. and C. L. For technical support for the completion of the
manuscript we are again indebted to Ursula Bott and Morris
Flecks, both LIB (ZFMK) Bonn.
APPENDIX. Translation of Eichwald’s (1831)
Original Latin Description of Desert Monitors
(p. 190) “164. Psammosaurus, Fitz., Varanus Merr., body
overall covered with small scales, arranged in transverse series;
very long tail round, later slightly compressed, as for swimmers
(natatoria), head pyramid-shaped, with all teeth pointed, elon-
gated and posteriorly curved.
+P. caspius, m. [= mihi = by me], with smaller scales on
head; larger on dorsum, suboval, medially elevated [keeled?],
with concave furrows at margins. Lives on the east coast of the
Caspian Sea, at the Balchasian Gulf, on Dardsha Peninsula,
close to water, — (p. 191) — which it is said to visit; resides in
rabbit holes of the sandy desert, also in self-digged burrows;
runs very fast; feet nonetheless short, with very long toes, and
strong claws.
2. P. griseus, Fitz., Monitor scincus, Cuv., Tupinambis
griseus auct., tail round, compressed towards the tip, with com-
pressed teeth, pointed. Lives in the deserts of Egypt, where ma-
gicians know various feats with them, however after having re-
moved the teeth. It is the Scincus of the Greek, and the
Crocodilus terrestris of Herodot, and the Uaran al hard of the
Arabs.
165. Varanus, Merr., body with small scales, tail keeled,
keel formed by a double series of erected, approximated scales;
teeth anteriorly conical, posteriorly blunt. Lives in Egypt, up to
6 feet long; devoures crocodile eggs; its images are visible on
the sarcophagi of the Egyptians.”
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