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The distribution of the critically endangered salamander Paradactylodon (Afghanodon) mustersi (Smith, 1940) in Afghanistan


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The Afghanistan Mountain Salamander, Paradactylodon (Afghanodon) mustersi (Smith, 1940), is an evolutionary old species, listed as Critically Endangered by IUCN Red List. The species is endemic to the Hindu Kush mountain range with records from only a few localities from four central-eastern Afghan provinces (Kabul, Ghazni, Parwan, and Wardak). Due to the long-term complicated political situation in the country which makes zoological research almost impossible, the current distribution and the presence of P. mustersi at previously known localities has remained unassessed for 40 years. We carried out recent, sporadic surveys between 2017 and 2021 to detect P. mustersi in three tributaries of the Paghman stream on the southern slopes of the Hindu Kush Mountains and the two nearby provinces (Panjsheer and Parwan), where the presence of the species was expected. We confirmed the occurrence of P. mustersi at all survey sites, and present the first record of the species for Panjsheer Province. We also confirmed that the species is currently endangered by human-mediated factors like habitat disturbance (increased visitors attendance, water pollution, construction activities), especially in the Paghman area.
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The distribution of the critically endangered salamander
Paradactylodon (Afghanodon) mustersi (Smith, 1940)
in Afghanistan
Ahmad Samim Ayobi1, Rafaqat Masroor2, Abdul Basit3, Daniel Jablonski4
1 Department of Horticulture, Agriculture Faculty of Parwan University, Parwan, Afghanistan
2 Zoological Sciences Division, Pakistan Museum of Natural History, Garden Avenue, Shakarparian, Islamabad, Pakistan
3 Tawakh, Anuba District, Panjsheer Province, Afghanistan
4 DepartmentofZoology,ComeniusUniversityinBratislava,Ilkovičova6,Mlynskádolina,84215,Bratislava,Slovakia
Corresponding author: Daniel Jablonski (
Academic editor: Ben Wielstra
30 April 2022
3 June 2022
30 June 2022
The Afghanistan Mountain Salamander, Paradactylodon (Afghanodon) mustersi (Smith, 1940), is an evolutionary old species, listed
as Critically Endangered by IUCN Red List. The species is endemic to the Hindu Kush mountain range with records from only a
few localities from four central-eastern Afghan provinces (Kabul, Ghazni, Parwan, and Wardak). Due to the long-term complicated
political situation in the country which makes zoological research almost impossible, the current distribution and the presence of
P. mustersi at previously known localities has remained unassessed for 40 years. We carried out recent, sporadic surveys between
2017 and 2021 to detect P. mustersi in three tributaries of the Paghman stream on the southern slopes of the Hindu Kush Mountains
and the two nearby provinces (Panjsheer and Parwan), where the presence of the species was expected. We conrmed the occurrence
of P. mustersi at all survey sites, and present the rst record of the species for Panjsheer Province. We also conrmed that the species
is currently endangered by human-mediated factors like habitat disturbance (increased visitors attendance, water pollution, construc-
tion activities), especially in the Paghman area.
Key Words
Batrachuperus, conservation, Hindu Kush, Hynobiidae, occurrence, Paghman stream Salamander, Pakistan
The Afghanistan Mountain Salamander Paradactylodon
(Afghanodon) mustersi (Smith, 1940), described initially
as Batrachuperus mustersi Smith, 1940, is currently
understood to be endemic to Afghanistan, and is considered
to be one of the world’s least-known salamanders (Wagner
et al. 2016; Ahmadzadeh et al. 2020). However, the
possible presence of the species across a wider area of the
Hindu Kush range (see Wall 1911), which is zoologically
one of the least explored places on earth, would not be
unexpected. Paradactylodon mustersi is a member of the
family Hynobiidae, and was initially discovered at the
Paghman streams area in Kabul Province (Smith 1940).
The original habitat where the species was initially found
mainly comprises a four- kilometer-long, glacier-formed
valley (Smith 1940; Nawabi 1965; Böhme 1982; Reilly
1983; Böhme and Jablonski 2022). Overall, the species
is known from elevations between ~1800 to 3750 m
(Reilly 1983; Wagner et al. 2016). Its population is
estimated between 1000–2000 individuals (Papenfuss et
al. 2004), but recent data are missing, and it is assumed
that the populations of P. mustersi declined over the last
40 years of armed conicts and instability in Afghanistan
Herpetozoa 35: 133–139 (2022)
DOI 10.3897/herpetozoa.35.e86028
Copyright Ahmad Samim Ayobi et al. This is an open access article distributed under the terms of the Creative Commons Attribution
License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and
source are credited.
Ahmad Samim Ayobi et al.: Additional distribution data on the Paghman Salamander134
(Jablonski et al. 2021). Although the species is considered
as a Critically Endangered B2ab (iii) according to IUCN
criteria, and also listed amongst the most Evolutionarily
Distinct and Globally Endangered species (EDGE;, there is still very
limited knowledge about the distribution, morphology,
ecology, and current threats of this enigmatic amphibian
(see also Böhme and Jablonski 2022).
Despite P. mustersi being listed by the Afghan
Government as a legally protected species since 2009, it
remains unclear to what extent the species is threatened
due to continuing political changes and the still
complicated security situation in Afghanistan that may
aect its conservation (see Gaynor et al. 2016). Thus,
data that could inform conservation priorities for the
Afghanistan mountain salamanders and other Hindu Kush
biota are missing. Most of the information we currently
have regarding the presence of the species originate from
older published work, comprising studies of P. mustersi
in the wild or in captivity (Nawabi 1965; Mertens 1970;
Seufer 1974; Sparreboom 1977; Bohmë 1982; Reilly
1983; Böhme and Jablonski 2022). In summary, this
salamander is commonly found in fast-owing melted
glacier waters with temperatures ranging between 0 to 14
°C (Reilly 1983). The total recorded length of P. mustersi
ranges from 119 to 215 mm, with individuals showing
14 costal grooves, that extend into the tail. The snout is
broadly rounded and limbs are well-developed. The tail
is slightly shorter than the body, and the color of the
body is dark olive-brown to yellowish-olive and speckled
indistinctly with tiny pigmented dots (Reilly 1983). From
an evolutionary point of view, P. mustersi is a unique
species characterized by a long-term independent history
and an isolated distribution, when compared to other
members of the family Hynobiidae (Zheng et al. 2011;
Ahmadzadeh et al. 2020). According to Wagner et al.
(2016), there are around 200 specimens of the species in
eight museum collections worldwide, mostly housed in
the United States.
Besides the area of Paghman (Kabul Province), the
species has also been recorded in provinces surrounding
Kabul, i.e. Ghazni, Parwan, and Wardak (Bohmë 1982;
Wagner et al. 2016; Ahmadzadeh et al. 2020 and Fig. 1
in this study). However, its current presence in localities
Figure 1. The distribution of Paradactylodon (Afghanodon) mustersi represented by published (red circles; for review see Wagner
et al. 2016; Ahmadzadeh et al. 2020 and Suppl. material 2) and new records (green circles; this study) from Afghanistan: 1. Paghman
area, Kabul Province; 2. Rakuľ, border of Wardak and Kabul provinces; 3. osprings of Paghman stream, Parwan Province; 4. San-
glakh, Wardak Province; 5, 6. Salang Pass, Parwan Province; 7. Gardan Diwal in the Koh-i-Baba Massif, Wardak Province; 8. Dasht-
i-Nawar, Ghazni Province; 9. Gardana Qalatak, Salang Valley, Parwan Province; 10. Shutul Valley, Panjsheer Province; 11. Qal‘ah-
ye Salim Khan, Kabul Province. The asterisk represents the presence of the species mentioned by locals from the Surkh-i-Parsa area,
Parwan Province (ca. 34.727°N, 68.740°E). The question mark represents the Wall (1911) record from the Chitral Valley, Pakistan.
Herpetozoa 35: 133–139 (2022)
beyond Kabul Province (Paghman area; see Jablonski et
al. 2020) has not been veried. Therefore, this study aims
to bring forth new information from recent eld surveys
carried out to map the occurrence of P. mustersi in
Afghanistan and to highlight areas of further investigation
and the possible threats the species is facing.
Materials and methods
Because the species is historically well-known from
three tributaries of the Paghman stream, four kilometers
above Paghman town (Kabul Province), we carried out
time-constrained, visual encounter surveys mostly in
this area. Besides, suitable habitats were investigated
for the presence of this species in Parwan and Panjsheer
provinces. As the presence and abundance of P. mustersi
may vary throughout the year (Reilly 1983), we made
targeted eld visits in four distinct seasons between 2017
and 2021. At the beginning of the study in Paghman,
semi-structured questionnaires in English (see Suppl.
material 1), translated into Dari Persian, and printed
photos of the species (available in the literature or
internet sources) were distributed among residents and
farmers so as to determine the possible localities of the
species. The information regarding the presence of the
species in Panjsheer and Parwan provinces was gathered
during our public awareness lectures in university and
school classes, later by distribution of the questionnaires
among local people. During eld surveys, basic
morphological data of encountered individuals, together
with geographic coordinates, elevation, and water and air
temperature, were recorded (if possible) from the places
where salamanders were observed. The total number of
eld trips was 12, each spanning over ve hours’ time-
constrained searching eorts in a day. Localities were
divided across the dierent areas as follows: eight in
the Paghman area (Kabul Province), two in the Gardana
Qalatak area of the Salang valley (Parwan Province),
and two in the Shutul valley (Panjsheer Province). The
Paghman area was surveyed on April 8, 2017; December
17, 2017, May 11, 2018; August 5 and 17, 2019;
September 29, 2019 and July 15 and 16, 2021. The Shutul
valley (Panjsheer Province) was surveyed on November
9, 2018 and January 17, 2019. The Salang district
(Parwan Province) was surveyed on December 26, 2018
and May 3, 2019. We searched for the species using a
combination of a visual encounter and randomized walks
along the streams. Several observed individuals and their
habitats were photographed. Due to security reasons,
we were not always able to spend enough time in the
studied places, take photographs or particular data, and
for the same reasons only the generalized state of the
habitat, including anthropogenic eects, were noted.
The distribution data (see Suppl. material 2: Table S1)
were reviewed with the literature, visualized using QGIS
Desktop 3.20.1 software (2021) and used for approximate
estimation of the species range.
We conrm the presence of Paradactylodon (Afghanodon)
mustersi in several localities of Kabul (Paghman area is
considered as one locality) and Parwan provinces, and
for the rst time, we report its occurrence in Panjsheer
Province (Figs 1–3).
A total of more than 260 individuals were observed
during eld trips to the Paghman area (Kabul Province;
several sites around 34.6155°N, 68.9125°E; Figs 2A–C,
3A, B) at an elevation of about 2,600 meters above sea
level in a stream of approximately 4.5 km length. The
individuals were found under rocks and near vegetation
where the stream ow was low, with a shallow water
depth, and with water temperatures ranging between 3
to 16 °C. During the last visit (16 July 2021), the air
temperature was 22 °C. The water tilt was low, and sal-
amanders were mostly hidden under rocks and the shade
of shrubs and bushes in shallow water. In addition to
salamanders, we observed two other amphibian species
from families Dicroglossidae and Bufonidae; Chrys-
opaa sternosignata (Murray, 1885) and Bufotes pseudo-
raddei (Mertens, 1971), both species in adult, subadult,
juvenile as well as tadpole stages. During the Septem-
ber 2019 survey, we observed the highest number of
salamander individuals, i.e., 241, with a minimum of
three at a single spot to a maximum of 17. Against this,
however, we only observed 16 individuals during our
July 2021 survey. One individual has been observed in
a human-made pool used as a rainwater reservoir (15
July 2021; 34.6142°N, 68.9117°E; 2,634 m a.s.l. Fig.
2C). During December investigations, we observed
three individuals with air temperatures of -4 °C. From
a conservation point of view, we noted the increase in
the tourist inux and construction activities (including
the construction of man-made pools) inside the stream
in the Paghman area.
Two eld surveys in Shutul Valley, Panjsheer Prov-
ince (35.1994°N, 69.2611°E, 2,170 m, Fig. 1, loc. 10;
Fig. 2D, E) resulted in the sighting of ve individuals
(Fig. 3C, D), three during the 2018 November survey
and two in January 2019. One individual was found
under rocks among the shade of shrubs in snow-melt-
ed fed streams. Two individuals were found in a hu-
man-made pool used as rainwater reservoir. At the
same site, two individuals were found in January. They
were moving and active. The environmental character-
istics of the area were similar to that of the Paghman
stream, but the stream ow was very low. The stream
dries up during the late summer and autumn seasons,
during which the salamanders probably live under
rocks near springs that produce water throughout the
year (this information emanates from local people).
The vegetation cover of the area was very low, limited
to the vicinity of water bodies. The common vegetation
is Nasturtiumocinale, which provides shelter to the
species. This represents the rst record of the species in
Panjsheer Province.
Ahmad Samim Ayobi et al.: Additional distribution data on the Paghman Salamander136
Two eld surveys in Gardana Qalatak (35.2317°N,
69.2086°E, 2,009 m; Fig. 1, loc. 9; Fig. 2F) of
the Salang Valley in Parwan Province resulted in
observations of three individuals (Fig. 3E). These were
found under rocks in shallow water, where the stream
ow was relatively low, as well as near springs that
produced cooler water (10.8 °C) than that of the stream.
One individual was found during the 2018 December
investigation in running water and was fully active.
Vegetation that covered streams included N.ocinale,
Schoenoplectus lacustris, and Cynodon dactylon. This
locality conrmed the historical records of the species
in the Salang area.
Additionally (24 July 2021), we recorded the species
from Qal‘ah-ye Salim Khan (34.7749°N, 69.0059°E; eleva-
tion between 2,200 to 2,400 m, Fig. 1, loc. 11), situated near-
by Qal‘ah-ye Mīrzā, Farza District, Kabul Province. One
adult individual (Fig. 3F) was observed in a local mountain
stream. This record represents a new locality in Kabul Prov-
ince, north of the Paghman area, connecting localities from
Parwan and Panjsheer provinces. The specimen is currently
stored in the Pakistan Museum of Natural History, Islam-
abad, under the voucher number PMNH 2263.
We took basic morphological measurements from 11 in-
dividuals (three from Paghman, ve from the Shutul Valley,
and three from the Salang Valley) that we sampled during
Figure 2. Localities and habitats in Afghanistan where Paradactylodon (Afghanodon) mustersi were observed. A, B. Paghman
area, Kabul Province (three individuals at April 8, 2017); C. Paghman area, Kabul Province, a human-made pool (one individuals,
July 15, 2021); D, E. Shutul Valley, Panjsheer Province (ve individuals, November 9, 2018); F. Gardana Qalatak area, Salang,
Parwan Province (three individuals, May 3, 2019).
Herpetozoa 35: 133–139 (2022)
our visits. All individuals were adults (sex was not deter-
mined) and each of them had 14 coastal grooves extended to
their tails. Their total length ranged from 92.0 to 160.0 mm,
a tail length between 45.2–75.6 mm, head length 14.8–20.0
mm, and abdomen width 12.7–18.3 mm. The coloration of
the body was dark brown to yellowish olive, in some indi-
viduals indistinctly speckled with tiny dots (Fig. 3C, D). The
tail is oval at the base but attening at the end.
The present study provides rare insight into the distribution
of P. mustersi from Afghanistan after an almost 40-
year hiatus and provides ecological and conservation
notes. Although Jablonski et al. (2020) provided a short
contribution concerning predation on this species and
conrmed recent occurrence in the Paghman area in Kabul
Province (the type locality), detailed data on this critically
endangered species has been unavailable for a long time
(Böhme 1982; Reilly 1983; Stuart et al. 2008). The last,
detailed, eld-based study on the species by Reilly (1983)
stated that “Batrachuperus mustersi occurs only in the
Paghman Mountains of Afghanistan in a single stream
– a statement which was probably based on information
of Nawabi (1965). However, Reilly (1983) did not
survey the distribution extent of this species, as Mertens
(1970), and mostly Böhme (1982), had already published
Figure 3. Observed individuals of Paradactylodon (Afghanodon) mustersi from: A, B. Paghman stream, Kabul Province, May 11,
2018 and April 8, 2017, respectively; C, D. Shutul Valley, Panjsheer Province, November 9, 2018; E. Salang Valley, Parwan Prov-
ince, May 5, 2019; F. Qal‘ah-ye Salim Khan, Farza District, Kabul Province, July 24, 2021 (PMNH 2263).
Ahmad Samim Ayobi et al.: Additional distribution data on the Paghman Salamander138
several localities beyond Kabul Province that showed a
geographic expansion of about 200 km between the two
most distanced localities (Fig. 1). Based on published and
new distribution data, we expected that the species range
covers about 4,200 km2, limited to mountain streams of
the Hindu Kush range. A record of the salamander from
the Mastuj area, Chitral valley in Pakistan (Wall 1911),
found in the stomach of the dice snake (N. tessellata,
Natricidae), also suggests that the species could be
distributed in the wider area of the Hindu Kush, which
could ultimately extend the species range much further.
This needs further investigations in Pakistan. We, for the
rst time, provide the record of this species for Panjsheer
Province in Afghanistan. This suggests that provinces with
the species presence mentioned in the literature (Böhme
1982; Wagner et al. 2016), as well as suitable habitats of
other provinces of Afghanistan (and Pakistan), need to
be examined. We highly anticipate the presence of this
species in dierent places of already recorded provinces
(e.g. Ghourband Valley in Parwan Province where local
people mentioned observation in Surkh-i-Parsa area, ca.
34.727°N, 68.740°E), as well as in Baghlan, Kapisa and
Laghman provinces of Afghanistan where the species
has never been recorded so far (Fig. 1). Unfortunately, at
the time of our investigation, many of these places were
not possible to visit due to security reasons. If the record
from the Chitral valley in Pakistan is correct (Wall 1911),
we can also expect that the species could be present in
Afghan provinces bordering Pakistan (e.g. Badakhshan,
Kunar, Nuristan) and thus in more new localities in eastern
Afghanistan. In this context, we obtained interesting
information provided by local shepherds and nomads
about the presence of urodelan amphibian from the
Wakhan corridor in Badakhshan Province of Afghanistan.
The information was provided during a eld visit to
Wakhan in 2018 and locals conrmed (based on photos we
showed) the presence of salamanders in the area between
Qala Panja village (36.9500°N, 72.3185°E; ca 2,800 m a.
s. l.) and Baba Tungi mountain. Such information is also
interesting regarding another salamander, the enigmatic
Central Asiatic species Hynobius turkestanicus Nikolskii,
1910, and its possible presence in the mountains of Central
Asia, e.g. in the neighboring Pamir Mountains (cf. Borkin
and Litvinchuk 2011). However, without conrmation
based on photographic documentation in the eld such
information should be taken with caution. Nevertheless,
additional research is, therefore, needed for the critically
endangered, currently endemic species P. mustersi, as well
as for other poorly studied species and rather inaccessible
areas of Afghanistan (Jablonski et al. 2021).
Although our data are not robust and consistent (mostly
due to the sensitive security situation in the country during
surveys), we observed that P. mustersi is active through-
out the year (including December and January), and could
be found in streams even with snow cover around. On the
other hand, our observations suggest that the presence of
the species may be highly inuenced by humans (see also
Reilly [1983] mentioning the absence of salamanders in
places where streams are aected by human activities). In
the Paghman stream, the species was mostly observed in
areas where human activities were low. Unfortunately, cur-
rent habitat disturbance is very high, especially close to Ka-
bul city. This decreases the local distribution extent within
the Paghman stream area, which constitutes an easily ac-
cessible place for local people to seek rest and recreation.
Reilly (1983), Stuart et al. (2008), and Wagner et al. (2016)
indicate that irrigated cultivation, overgrazing, and physical
disturbance by pedestrians and livestock are major threats
to the species. This is exacerbated by environmental pollu-
tion (passenger transport, garbage), which we observed in
the Paghman area. Besides our observation, the movement
of vehicles around and even inside streams where the spe-
cies occurs and breeds is a common practice. The Paghman
area is close to Kabul capital city and receives hundreds of
residents from Kabul coming for recreation on a daily basis.
Moreover, the construction of swimming pools alongside
the streams or restaurants increases water pollution, which
could highly aect local populations of this endemic and
rare Afghan amphibian.
Threats to the species in localities of Parwan and Pan-
jsheer provinces were also documented. Due to an in-
crease in the local resident population, houses and com-
mercial areas are being built inside or nearby streams.
Water supply pipelines to Charikar city of Parwan Prov-
ince are additionally being constructed, taking water from
the streams for irrigation purposes. Overgrazing, recre-
ation pressure, and public unawareness constitute addi-
tional threats to the diminishing population of P. mustersi
in the Salang district of Parwan Province. In the Shutul
valley of Panjsheer Province, overgrazing and water dam
construction for the conservation of water for agriculture
purposes pose potential threats to P. mustersi.
Unfortunately, due to the long-term unstable situa-
tion in Afghanistan and the lack of biodiversity research
(Jablonski et al. 2021), our observations remain prelim-
inary. We, therefore, lack information about the current
presence of P. mustersi from other places mentioned in
the literature (see Böhme 1982), and thus we call for in-
tensive eld research to improve our knowledge of en-
dangered species of the Hindu Kush.
This study was supported by the Ruord Foundation
(project no. 21296-1). DJ was supported by the Slovak
Research and Development Agency under the contract
no. APVV-19-0076. We extend our profound thanks to
those who made the research possible, especially Meer-
wais Meerzaey, Ahmad Omran Khwajapoor, Ahmad Za-
hed Ayobi, Mohammad Rasoul Ashna, and Abdul Satar
Sarwary. We also thank the reviewer, Wouter Beukema,
for his valuable suggestions that improved the submitted
version of this manuscript. Finally, with heartfelt thanks,
we dedicate this study to a number of Afghan people who
provided information about the presence of this endan-
gered species and its possible habitats. Without them, this
study would not have been possible.
Herpetozoa 35: 133–139 (2022)
Ahmadzadeh F, Shahrokhi G, Saberi-Pirooz R, Oladi M, Taati M,
Poyarkov NA, Rödder D (2021) Alborz Heritage: geographic
distribution and genetic dierentiation of the Iranian Paradactylodon
(Amphibia: Hynobiidae). Amphibia-Reptilia 41: 519–534. https://
Böhme W (1982) Über dem Afghanischen Winkelzahnmolch,
Batrachuperus mustersi Smith, 1940. Elaphe 1982(3): 33–36.
Böhme W, Jablonski D (2022) Making forgotten information
available: An early study on the Afghanistan Mountain Salamander
Paradactylodon (Afghanodon) mustersi (Smith, 1940) (Caudata:
Hynobiidae). Bonn zoological Bulletin 71: 1–7.
Borkin LY, Litvinchuk SN (2011) On Turkestan salamander Hynobius
turkestanicus Nikolsky, 1910 and Vasily Nikolsky’s Pamirs travel
(1902). Modern Herpetology 11: 103–120. [In Russian]
Gaynor MK, Fiorella JK, Gregory GH, Kurz DJ, Seto KL, Withey LS,
Brashares JS (2016) War and wildlife: linking armed conicts to
conservation. Frontiers in Ecology and the Environment 14: 533–
Jablonski D, Farooqi J, Basit A, Masroor R (2020) Afghanodon mustersi
(Afghanistan Stream Salamander). Predation. Herpetological
Review 51: 287.
Jablonski D, Basit A, Farooqi J, Masroor R, Böhme W (2021)
Biodiversity research in a changing Afghanistan. Science 6549:
Mertens R (1970) Salamander aus Afghanistan: Batrachuperus
mustersi. Aquarien und Terrarien Zeitschrift 23(11): 346–348.
Nawabi S (1965) A rare amphibian from Afghanistan: Batrachuperus
mustersi. Science (Kabul) 8: 21–25. [in Farsi]
Papenfuss T, Anderson S, Kuzmin S (2004) Afghanodon mustersi. The
IUCN Red List of Threatened Species 2004: e.T2664A9465637.
[Downloaded on 08 June 2021]
QGIS Development Team (2021) QGIS Geographic Information System.
Open Source Geospatial Foundation Project.
Reilly SM (1983) The biology of the high altitude Salamander
Batrachuperus mustersi from Afghanistan. Journal of Herpetology
17: 1–9.
Sparreboom M (1977) Salamanders uit Afghanistan: Batrachuperus
mustersi. Lacerta 35: 125–128.
Wagner P, Bauer AM, Leviton AE, Wilms TM, Böhme W (2016) A
checklist of the Amphibians and Reptiles of Afghanistan, Exploring
Herpetodiversity Using Biodiversity Archives. Proceeding to the
California Academy of Science 63: 457–565.
Wall F (1911) Reptiles collected in Chitral. Journal of the Bombay Nat-
ural History Society 21: 132–145.
Seufer H (1974) Freilandbeobachtungen und Fang von Batrachuperus
mustersi. Das Aquarium 60: 269.
Smith MA (1940) Contributions to the herpetology of Afghanistan.
Annals and Magazine of Natural History 5: 382–384. https://doi.
Zheng Y, Peng R, Kuro-o M, Zeng X (2011) Exploring patterns and
extent of bias in estimating divergence time from mitochondrial
DNA sequences data in a particular lineage: a case study of
Salamanders (order Caudata). Molecular Biology and Evolution 28:
Supplementary material 1
Baseline survey questionnaire for Paghman
stream Salamander in Afghanistan
Authors: Ahmad Samim Ayobi, Rafaqat Masroor, Abdul Basit,
Daniel Jablonski
Data type: Adobe PDF le
Copyright notice: This dataset is made available under the Open
Database License (
odbl/1.0/). The Open Database License (ODbL) is a license
agreement intended to allow users to freely share, modify,
and use this Dataset while maintaining this same freedom
for others, provided that the original source and author(s)
are credited.
Supplementary material 2
Table S1. The distribution data of Paradactylodon
(Afghanodon) mustersi for the map vizualization
(Fig. 1)
Authors: Ahmad Samim Ayobi, Rafaqat Masroor, Abdul Basit,
Daniel Jablonski
Data type: excel le
Copyright notice: This dataset is made available under the Open
Database License (
odbl/1.0/). The Open Database License (ODbL) is a license
agreement intended to allow users to freely share, modify,
and use this Dataset while maintaining this same freedom
for others, provided that the original source and author(s)
are credited.
... Three species of Paradactylodon (Family: Hynobiidae) are found in the Middle East: P. mustersi, P. persicus, and P. gorganensis (Zhang et al., 2006). P. mustersi is endemic to the Hindu Kush mountain range of Afghanistan and is commonly found in fast-flowing melting glacier waters with temperatures ranging from 0°C to 14°C (Ayobi et al., 2022). Iranian Paradactylodons are endemic to the Hyrcanian forests of Iran, with P. persicus occupying the north and northwest regions and P. gorganensis occupying the northeast (Baloutch and Kami, 1995). ...
... Their conservation status is listed as Near Threatened for P. persicus (Papenfuss et al., 2009a), Critically Endangered for P. "gorganensis" (Papenfuss et al., 2009b), and P. mustersi (Papenfuss, Anderson and Kuzmin, 2004). Recent studies have investigated the distribution and conservation status (Ahmadzadeh and Kami, 2009;Ahmadzadeh et al., 2011;Hosseinian, 2021;Ayobi et al., 2022), genetic structure, Quaternary distribution dynamics, phylogeny (Yousefi Siahkalroodi, Khederzadeh and Ghadiri Abyaneh, 2015;Ahmadzadeh et al., 2020), skeletochronological evaluation (Zivari and Kami, 2017), reproductive biology, and histology (Rezapour et al., 2009;Heydari Nasrabadi, 2012) of the Paradactylodon species. However, it is unclear, if the climatic niches of Paradactylodon species had an effect on their diversification. ...
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The relict genus Paradactylodon is narrowly distributed in temperate forests throughout the Middle East region, including P. mustersi in Afghanistan, P. persicus, and P. gorganensis in northwestern and northeastern Iran. Recent phylogenetic analyses suggest that the Iranian Paradactylodon populations may belong to a single species, P. persicus. In this study, we addressed how climatic niche conservatism and/or niche divergence may have affected the evolution of all Paradactylodon species using ecological niche models and multivariate niche analyses to identify niche overlap and assess equivalency and similarity tests. We also used an ensemble of ten algorithms to predict Paradactylodon species distributions for recent (1970–2000) and future (2081–2100) climate conditions. The results show that future climate change may cause the loss of 80 to 97% of these species' suitable habitat, especially at lower elevations. The niche divergence hypothesis was supported by the results of niche equivalency tests on P. persicus vs. P. gorganensis (and vice versa) with moderate overlap (D = 0.23, I = 0.41) and P. persicus vs. P. mustersi (and vice versa) with little overlap (D = 0.05, I = 0.06). Meanwhile, the niche similarity test for the niche conservatism hypothesis revealed significant results for P. persicus vs. P. gorganensis, P. mustersi vs. P. persicus, and P. persicus + P. gorganensis (as a single species) vs. P. mustersi. Due to these complex evolutionary ecological patterns and allopatric distributions, we recommend that P. gorganensis be considered a valid subspecies.
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The first study after the official species description of the endemic Afghanistan Mountain Salamander Paradactylodon (Afghanodon) mustersi (Smith, 1940) was published in Kabul, Afghanistan, and only in Dari Persian. We, therefore, provide here an English translation of this paper, together with so far unpublished background information on this rare and endangered amphibian, and on the former scientific German-Afghan cooperation project from the 1960s in the framework of which this study had been performed.
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Paradactylodon, a genus of Hynobidae, is distributed in mountainous regions of the Middle East. This genus comprises three species, Paradactylodon mustersi (Afghanistan), Paradactylodon persicus, and Paradactylodon gorganensis (Iran). Initially, P. persicus was reported from Talysh Mountains, and afterward, P. gorganensis was described from the eastern Alborz Mountains. Although these two Iranian Paradactylodon species were distinguished based on morphological features, there are arguments about their systematic status. In this contribution, 30 samples of hynobiid salamanders were collected from Talysh and Alborz Mountains and the phylogenetic relationship between the two species was assessed using two mtDNA markers (COI and 16S). Additionally, in order to survey the potential habitat suitability for the species, Species Distribution Models (SDMs) were performed and projected on climate scenarios that reflect current and past (6 ky and 21 ky before present) conditions. Our results indicated that P. gorganensis is nested within P. persicus. Overall, both SDM and molecular analyses signified that the Iranian Paradactylodon population was affected by the Quaternary glacial period, and according to haplotype networks, haplotype diversity was higher in the western part of the distribution range of the species. Given the low genetic distance among all samples, we suggest P. gorganensis be synonymized with P. persicus. Considering the conservational values and numerous threats that this endemic species is facing, we encourage a revision of the IUCN Red List category of the species immediately.
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Armed conflict throughout the world’s biodiversity hotspots poses a critical threat to conservation efforts. To date, research and policy have focused more on the ultimate outcomes of conflict for wildlife rather than on the ecological, social, and economic processes that create those outcomes. Yet the militarization that accom- panies armed conflict, as well as consequent changes in governance, economies, and human settlement, has diverse influences on wildlife populations and habitats. To better understand these complex dynamics, we summarized 144 case studies from around the world and identified 24 distinct pathways linking armed conflict to wildlife outcomes. The most commonly cited pathways reflect changes to institutional and socio- economic factors, rather than tactical aspects of conflict. Marked differences in the most salient pathways emerge across geographic regions and wildlife taxa. Our review demonstrates that mitigating the negative effects of conflict on biodiversity conservation requires a nuanced understanding of the ways in which conflict affects wildlife populations and communities.
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The study history of two salamanders (Hynobius turkestanicus Nikolsky, 1910 and Turanomolge mensbieri Nikolsky, 1918) collected by Vasily N. Nikolsky in Turkestan is considered in detail. Some comments on the nomenclature of these nominal taxa are made. New data on the collector V. N. Nikolsky and his Pamirs travel in 1902 are given. A possible distribution of Hynobius turkestanicus is suggested.
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In the practice of molecular dating, substitution saturation will bias the results if not properly modeled. Date estimates based on commonly used mitochondrial DNA sequences likely suffer from this problem because of their high substitution rate. Nevertheless, the patterns and extent of such expected bias remain unknown for many major evolutionary lineages, which often differ in ages, available calibrations, and substitution rates of their mitochondrial genome. In this case study of salamanders, we used estimates based on multiple nuclear exons to assess the effects of saturation on dating divergences using mitochondrial genome sequences on a timescale of ~200-300 My. The results indicated that, due to saturation for older divergences and in the absence of younger effective calibration points, dates derived from the mitochondrial data were considerably overestimated and systematically biased toward the calibration point for the ingroup root. The overestimate might be as great as 3-10 times (about 20 My) older than actual divergence dates for recent splitting events and 40 My older for events that are more ancient. For deep divergences, dates estimated were strongly compressed together. Furthermore, excluding the third codon positions of protein-coding genes or only using the RNA genes or second codon positions did not considerably improve the performance. In the order Caudata, slowly evolving markers such as nuclear exons are preferred for dating a phylogeny covering a relatively wide time span. Dates estimated from these markers can be used as secondary calibrations for dating recent events based on rapidly evolving markers for which mitochondrial DNA sequences are attractive candidates due to their short coalescent time. In other groups, similar evaluation should be performed to facilitate the choice of markers for molecular dating and making inferences from the results.
Batrachuperus mustersi occurs only in the Paghman Mountains of Afghanistan in a single stream. They inhabit the cold sources of the stream where they are active year-round, feeding on aquatic organisms. Morphological examination revealed many similarities to other members of the genus. Differences were found in the tail morphology, degree of skeletal ossification and gill development. The only adult sexual dimorphism is in the appearance of the cloaca. Redescriptions of the morphology of adults and larvae are given. Niche partitioning was found in habitat selection and prey size. Breeding coincides with the spring thaw when females lay at least two egg cases and attach them to submerged rocks or plants. Metamorphosis also takes place in the spring after two years of development.