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;
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
* Corresponding author. E-mail: email@example.com
Figure 1. Photographs of the two Echis individuals found near
Assa. (A) juvenile. (B) male.
Daniel Koleska et al.
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
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”.
Arnold, N., Robinson, M., Carranza, S. (2009): A preliminary
analysis of phylogenetic relationships and biogeography of
the dangerously venomous Carpet Vipers, Echis (Squamata,
Serpentes, Viperidae) based on mitochondrial DNA sequences.
Amphibia-Reptilia 30: 273–282.
Aymerich, M., Borof-Aymerich, E., Geniez, P. (2004): Neufunde
der seltenen Weißbäuchigen Sandrasselotter Echis leucogaster
Roman, 1972 in Marokko (Serpentes: Viperidae). Herpetozoa
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.
Bons, J., Geniez, P. (1996): Amphibiens et reptiles du Maroc
(Sahara Occidental compris). Atlas Biogéographique. Barcelona,
Asociación Herpetológica Española.
Brito, J.C., Fahd, S., Geniez, P., Martínez-Freiría, F., Pleguezuelos,
J.M., Trape, J.-F. (2011): Biogeography and conservation of
viperids from North-West Africa: An application of ecological
niche-based models and GIS. Journal of Arid Environments 75:
Escoriza, D., Metallinou, M., Donaire-Barroso, D., Amat, F.,
Carranza, S. (2009): Biogeography of the White-Bellied Carpet
Viper Echis leucogaster Roman, 1972 in Morocco, a study
combining mitochondrial DNA data and ecological niche
modeling. Butlletí de la Societat Catalana d‘Herpetologia 18:
Geniez, P. (2015): Serpents d’Europe, d’Afrique du Nord et du
Moyen-Orient. Paris, Delachaux et Niestlé.
Herrmann, H.-W., Herrmann, P.A., Geniez, P.H. (2000): Zur
Verbreitung von Echis leucogaster Roman, 1972 in Marokko.
Salamandra 36: 203–207.
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.
Martinez del Marmol Marin, G., Rebollo Fernandez, B. (2012): An
important new record of Echis leucogaster Roman, 1972 from
Morocco. Herpetology Notes 5: 229–231.
Martínez-Freiría, F., Stols, V.F., García-Cardenete, L. (2016):
Human-mediated syntopy between Cerastes cerastes and
Daboia mauritanica in the lower Drȃa Valley, Morocco. Boletín
de la Asociación Herpetológica Española 27: 2.
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.
Schleich, H.H., Kastle, W., Kabisch, K. (1996): Amphibians
and reptiles of North Africa. Koenigstein, Koelts Scientific
Sindaco, R., Venchi, A., Grieco, C. (2013): The Reptiles of the
Western Paleartic. Vol 2: Annotated checklist and distribution
atlas of the snakes of Europe, North Africa, Middle East and
Central Asia. Latina, Edizioni Belvedere.
Spawls, S., Branch, B. (1995): The dangerous snakes of Africa.
New records of Echis leucogaster in Morocco 657
Accepted by Hendrik Müller