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New records of Echis leucogaster in Morocco

Authors:
  • Zoopark Zájezd
The white-bellied carpet (or saw-scaled) viper Echis
leucogaster ROMAN, 1972 is considered one of the
rarest snakes in Morocco (Spawls and Branch, 1995;
Bons and Geniez, 1996; Schleich et al., 1996). Recent
genetic studies have subdivided the genus Echis into four
main clades consisting of the E. ocellatus, E. coloratus,
E. pyramidum and E. carinatus groups (Arnold et al.,
2009; Pook et al., 2009). Within the E. pyramidum
clade, E. leucogaster inhabits the western Sahel region,
with possibly isolated populations in the Algerian
Hoggar (or Ahaggar) massif, in southern Morocco
and in Tunisia (Pook et al., 2009; Geniez, 2015). Due
to low genetic variability, it has been proposed that E.
leucogaster should be regarded as a subspecies of E.
pyramidum (Arnold et al., 2009; Sindaco et al., 2013;
Geniez 2015).
Until less than two decades ago, E. leucogaster was
known in Morocco from only a single locality near
Aouinet Torkoz (presently Aouinet Lahna), where
it was discovered in 1963 (Bons and Dakka, 1963).
However, starting from 1999 observations were made
at other locations, resulting in a fragmented distribution
with three major known populations: in Aouinet Lahna,
which has been confirmed by several observations over
the years (Herrmann et al., 2000; Aymerich et al., 2004;
personal observations by M. Berroneau, T. Charlton, S.
Clerc, D. Herrero, B. Rebollo, M. Sassoe, G.J. Verspui
and M. Wilson); another one in a region located between
Agdz and Bou Azzer and extending south to Allougoum,
which comprises the northernmost record of the species
in Morocco (Maran and Geniez, 1999; Aymerich et
al., 2004; Escoriza et al., 2009; Pook et al., 2009); and
in the Zemmour region near Amgala, which is known
from only one dead individual and may be contiguous
with populations in northern Mauritania (Aymerich
Herpetology Notes, volume 11: 655-657 (2018) (published online on 24 August 2018)
New records of Echis leucogaster in Morocco
Daniel Koleska1, Jaroslav Karhánek2, Gabriel Martinez del Marmol Marin3, and Marco Sassoè-Pognetto4,*
1 Department of Zoology and Fisheries, Czech University of
Life Sciences Prague (CULS), Kamýcká 129, 165 00 Praha
6, Czech Republic.
2 Nopova 56, 61500 Brno, Czech Republic.
3 Cl. Pedro Antonio de Alarcon 34, 18002 Granada, Spain.
4 Department of Neuroscience “Rita Levi Montalcini”,
University of Torino, C.so Massimo d’Azeglio 52, 10126
Torino, Italy.
* Corresponding author. E-mail: marco.sassoe@unito.it
Figure 1. Photographs of the two Echis individuals found near
Assa. (A) juvenile. (B) male.
Daniel Koleska et al.
656
et al., 2004). More recently, a juvenile E. leucogaster
has been found near Tata, suggesting that there could
be a continuous distribution between the population of
Aouinet Lahna and that of Agdz along the Drȃa valley
(Martinez del Marmol and Rebollo Fernandez, 2012).
In support of this idea, we report here two novel
observations of the species near the town of Assa. A
juvenile specimen (approximately 20 cm in length) of
E. leucogaster was located at the southern exit of Assa
(28.58°N 9.41°W) on May 5, 2017 (Fig. 1A). The snake
was found active at night in a dry river bed with sparse
vegetation forming potential shelter (Fig. 2). Another
individual, a male of approximately 45 cm in total length,
was spotted in the same area inside a small bush on the
night of September 28, 2017 (Fig. 1B). The habitat in
which the vipers were found is bordered by plantations,
which corresponds with previous records of this species
(see Martinez del Marmol and Rebollo Fernandez,
2012). Other reptiles and amphibians observed in the
surroundings include Chalcides ocellatus, Ptyodactylus
oudrii, Stenodactylus mauritanicus, Tarentola boehmei,
and Pelophylax saharicus and Sclerophrys mauritanica
in a nearby water pond.
This novel locality of E. leucogaster is located 40 km
east of the well-known population of Aouinet Lahna.
The species has been recorded further down the Drȃa
valley, south of Tiglit (Aymerich et al., 2004), suggesting
that its distribution embraces the lower course of the
Drȃa river. This region is found within an area of high
occurrence probability as predicted by niche modelling
analysis (Escoriza et al., 2009; Brito et al., 2011). North
of Assa, the closest locality in which the species has
been recorded is Tata at 200 km (Martinez del Marmol
and Rebollo Fernandez, 2012), which is halfway from
the populations situated west of Agdz (Fig. 2). Further
exploration is needed to understand whether the lack
of data in intermediate areas between these localities is
due to poor sampling or rather reflects a fragmentary
Figure 2. Summary of the known distribution of Echis leucogaster in Morocco. Black dots indicate known localities, based on
published reports (see text). The site of the new observations near Assa is marked with a white dot. The inset shows the habitat
where the two individuals were found.
distribution of the species in isolated populations.
Similarly, it will be interesting to establish whether the
Moroccan populations of E. leucogaster are connected
to those of the Sahel or whether they should be regarded
as relict populations. Genetic analyses have indicated
that Moroccan Echis are very closely related to those
found in southern Mauritania, suggesting that these
populations either have a continuous distribution or have
been separated in recent times (Escoriza et al., 2009). A
better knowledge of the distribution of E. leucogaster
in Morocco is required for predicting factors that may
negatively impact the survival of its populations under
scenarios of climatic and environmental changes (Brito
et al., 2011).
One factor that may influence the distribution of E.
leucogaster in Morocco is the interaction with other
viperids (see for example Herrmann et al., 2000). The
southwestern part of Morocco in which Echis has been
found is an environmental transition zone with potential
sympatry of five distinct species of vipers (Brito et al.,
2011). Indeed, Cerastes cerastes has been found (e.g.
two adult individuals observed by M. Sassoe in May
2017) at sites near Aouinet Lahna where Echis has
also been recorded. Moreover, coexistence between
C. cerastes and Daboia mauritanica has been reported
in the vicinity of Aouinet Lahna (Martínez-Freiría et
al., 2016), suggesting that three species of vipers with
distinct biogeographical affinities occur in sympatry in
the same region. Fieldwork is needed to determine how
these species partition habitat resources in this contact
zone.
Acknowledgements. We would like to thank S. Borský, D.
Hegner, J. Lehký and V. Víta for their help during the fieldwork
and to Fernando Martínez-Freiría for his comments to the
manuscript. This work was supported by the Internal Grant
Agency of the Czech University of Life Sciences Prague (CIGA)
through Project No. 20172002, by Turin University (grant
RILO 17) and by Ministero dell’Istruzione, dell’Università e
della Ricerca – MIUR project “Dipartimenti di Eccellenza 2018
– 2022” to Dept. of Neuroscience “Rita Levi Montalcini”.
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New records of Echis leucogaster in Morocco 657
Accepted by Hendrik Müller
... The finding of Echis leucogaster in a cracked mud habitat lacking stones in Oued Drâa was novel as within this region, until this point, this snake was only known from rocky habitats in two main locations near the towns of Assa and Aouinet Lahna (Koleska et al., 2018). In poorly-explored arid regions, site accessibility can play a key role in our understanding of species' distributions (Kane et al., 2017) as more accessible locations receive greater survey attention. ...
... Garcia-Cardenete et al., 2014). Considering the records of E. leucogaster in the present work along with that reported by Hermann and Hermann (1999), combined with the rapid increase of known locations for this species in recent years (Koleska et al., 2018), it seems likely that this species may be found along much of the length of the Oued Drâa where it passes through suitable climatic zones (Brito et al., 2011). With future field work in this region a greater area of occurrence than is presently understood in Morocco may well be revealed. ...
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... Morocco is one of the best sampled areas of the Maghreb, and the distribution of many of its amphibians and reptiles is relatively well known (see Bons & Geniez, 1996;Martínez del Mármol et al., 2019). However, some regions of the country have been, so far, poorly surveyed, and range extensions for certain species have been reported in recent times as a result of increased fieldwork in this area during the last few years (e.g., Barata et al., 2011;Koleska et al., 2018;Kane et al., 2019). ...
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In northwest Africa some species from Sahelian origin appear with relict populations and apparently isolated by the extreme aridity of the Sahara desert. However very tolerant to aridity species could maintain continuous populations as might be the case for Echis leucogaster as indicated by results from genetic analysis and bioclimatic models. Resum: Al nord-oest d'àfrica apareixen un grup d'espècies d'origen sahelià en poblacions relictes i aparentment aïllades pel desert del Sàhara. No obstant això espècies molt tolerants a l"aridesa podrien mantenir poblacions contínues, com podria ser el cas de Echis leucogaster segons indiquen els resultats de l'anàlisi genètica i els models bioclimàtics.
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Phylogenetic analysis of 1117 bp of mitochondrial DNA sequences (731 bp of cytochrome b and 386 bp of 16S rRNA) indicate that Echis consists of four main clades: E. ocellatus, and the E. coloratus, E. pyramidum, and E. carinatus groups. In the E. coloratus group, E. coloratus itself shows substantial genetic divergence from E. omanensis, corroborating their separate species status. In the E. pyramidum clade, E. pyramidum from Egypt and E. leucogaster from West Africa are genetically very similar, even though samples are separated by 4000 km. South Arabian populations of the E. pyramidum group are much better differentiated from these and two species may be present, animals from Dhofar, southern Oman probably being referable to E. khosatzkii. In the E. carinatus group, specimens of E. carinatus sochureki and E. multisquamatus are very similar in their DNA. The phylogeny indicates that the split between the main groups of Echis was followed by separation of African and Arabian members of the E. pyramidum group, and of E. coloratus and E. omanensis. The last disjunction probably took place at the lowlands that run southwest of the North Oman mountains, which are likely to have been intermittently covered by marine incursions; they also separate the E. pyramidum and E. carinatus groups and several sister taxa of other reptiles. The E. carinatus group may have spread quite recently from North Oman into its very extensive southwest Asian range, and there appears to have been similar expansion of E. pyramidum (including E. leucogaster) in North Africa. Both these events are likely to be associated with the marked climatic changes of the Pleistocene or late Pliocene. Similar dramatic expansions have also recently occurred in three snake species in Iberia.
Capture au Maroc de la vipère des Pyramides Echis carinatus (Schneider, 1801). C. R. Société des
  • J Bons
  • M Dakka
Bons, J., Dakka, M. (1963): Capture au Maroc de la vipère des Pyramides Echis carinatus (Schneider, 1801). C. R. Société des Sciences Naturelles et Physiques du Maroc, Rabat 29: 55-57.
Redécouverte de l'Echide à ventre blanc
  • J Maran
  • P H Geniez
Maran, J., Geniez, P.H. (1999): Nouvelles observations sur l'herpétofaune du Maroc, 7. Redécouverte de l'Echide à ventre blanc, Echis leucogaster ROMAN, 1972 (Reptilia, Serpentes, Viperidae) au Maroc. Bulletin de la Société Herpétologique de France, Paris 90: 63-64.
  • C E Pook
  • U Joger
  • N Stümpel
  • W Wüster
Pook, C.E., Joger, U., Stümpel, N., Wüster, W. (2009): When continents collide: phylogeny, historical biogeography and systematics of the medically important viper genus Echis (Squamata: Serpentes: Viperidae). Molecular Phylogenetics and Evolution 53: 792-807.