Amphibians of North Africa

Amphibians of North Africa

Daniel Escoriza

University of Girona, Girona, Spain

Jihène Ben Hassine

University of Tunis El Manar, Tunis, Tunisia

Table of Contents

Cover image

Title page

Copyright

Dedication

Preface and Acknowledgments

Chapter 1. Introduction

Abstract

1 Amphibian evolution and diversity

2 Amphibian natural history

3 Amphibian global decline

Chapter 2. Amphibians of north-west Africa

Abstract

1 Environment of north-west Africa

2 Tectonics and zoogeography

3 Fossil record

4 Life history

5 List of the north-west African amphibians

6 Insular species

7 Dubious and unconfirmed records

8 Conservation

Chapter 3. Ecology of amphibians in north-west Africa

Abstract

1 The vegetation and the phytogeographical regions

2 Batrachocenosis

3 Amphibians in the ecosystems

Chapter 4. Amphibian surveys

Abstract

1 Survey methods

Chapter 5. History of batrachology in North Africa

Abstract

Chapter 6. Identification keys

Abstract

1 Keys to the orders of amphibians in North Africa

2 Key to the genera of salamanders (Fig. 6.1)

3 Key to the species of Pleurodeles

4 Key to the genera of frogs and toads (Figs. 6.2 and 6.3)

5 Key to the family Bufonidae species (Fig. 6.4)

6 Key to the family Ptychadenae species

7 Keys to the orders of amphibians in North Africa: larvae

8 Key to the genera of salamanders: larvae

9 Key to the genera of frogs and toads: larvae

10 Key to the genera of amphibians: eggs

Chapter 7. Species descriptions

Abstract

1 Methods

Chapter 8. North-western Africa amphibians

Abstract

1 Caudata, Salamandridae

2 Pleurodeles nebulosus Algerian newt

3 Pleurodeles poireti Edough newt

4 Pleurodeles nebulosus-Pleurodeles poireti hybrids

5 Pleurodeles waltl Spanish newt

6 Salamandra algira Bedriaga, 1883 North African fire salamander

7 Anura

8 Alytes maurus Moroccan midwife toad

9 Discoglossus pictus Painted frog

10 Discoglossus scovazzi Moroccan painted frog

11 Bufonidae

12 Barbarophryne brongersmai Tiznit toad

13 Bufo spinosus Spiny toad/Mediterranean common toad

14 Bufotes boulengeri African green toad

15 Sclerophrys mauritanica Berber toad

16 Sclerophrys xeros Subdesert toad/Desert toad/Savanna toad

17 Dicroglossidae

18 Hoplobatrachus occipitalis African groove-crowned frog/Giant swamp frog/African tigrine frog

19 Hylidae

20 Hyla meridionalis Mediterranean tree frog

21 Pelobatidae

22 Pelobates varaldii Moroccan spadefoot toad

23 Ranidae

24 Pelophylax perezi Perez’s frog/Iberian green frog

25 Pelophylax saharicus (Boulenger, 1913) North African green frog

Chapter 9. Amphibians of Mauritania and Egypt

Abstract

1 Bufonidae

2 Sclerophrys dodsoni (Boulenger, 1895) Dodson’s toad

3 Sclerophrys kassasii Nile Valley toad

4 Sclerophrys pentoni Penton’s toad

5 Sclerophrys regularis Square-marked toad/Egyptian toad/Reuss’s toad/Common African toad

6 Dicroglossidae

7 Hylidae

8 Hyla savignyi Audouin, 1827 Savigny’s tree frog/Lemon-yellow tree frog

9 Hyperoliidae

10 Kassina senegalensis Senegal running frog/Senegal Kassina/Bubbling Kassina

11 Phrynobatrachidae

12 Phrynobatrachus natalensis Natal puddle frog

13 Ptychadenidae

14 Ptychadena bibroni Broad-banded grass frog

15 Ptychadena nilotica Nile grass frog/Nile ridged frog

16 Ptychadena schillukorum Sudan grassland frog/Schilluk ridged frog

17 Ptychadena trinodis Dakar grassland frog

18 Pyxicephalidae

19 Pyxicephalus edulis Lesser bullfrog/Edible bullfrog

20 Tomopterna milletihorsini Mali screeching frog/Mali sand frog

21 Ranidae

22 Pelophylax bedriagae Levant green frog/Levant water frog

References

Index

Copyright

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Dedication

We dedicate this book to our daughter, Lydia.

The goal of studying the North African Amphibians determined the course of most family holidays. Having her around, looking to the pond with interest, asking about every animal observed or captured, carrying her pink dipnet with unconcealed enthusiasm and infinite curiosity made her an invaluable driving strength-giving and motivating companion in all our field trips….

Preface and Acknowledgments

Over the past 12 years, we have published our findings in a variety of papers, but as almost every field trip yielded new data and significant new distributional or biological data, we postponed the preparation of a more broadly based review of the batrachofauna as a whole. We feel that the time has come to offer a general synthesis of the actual state of the North African amphibians in the form of this book.

We conceived this book to meet a major need in northern Africa: it is the first book dedicated only to the amphibians of North Africa and that describe all the amphibian species and map their distributions as known at present. This book is intended for the nature lovers but also meant to be a source of information for more expert readers and that will be of value to both in the field and laboratory. It is the aim of this book to provide a guide that will enable the user to identify any amphibian’s species at any development stages encountered in North Africa. Along with the keys and the species descriptions, the drawing and photographs should allow readers to identify all amphibian species occurring in the region. In addition to its use as an identification guide, we hope that the book will serve to direct future studies by highlighting gaps in our knowledge of this batrachofauna and that this will ultimately translate into greater conservation concern for the biota of this region.

Many people have assisted us in the various phases of our work in North Africa. To all we owe a sincere debt of gratitude. Collecting permits were provided by Le Haut Commissaire aux Eaux et Forêts et à la Lutte Contre la Désertification in Morocco and by Le Ministre de l’Agriculture, du Développement Rural et de la Pêche Direction Général des Forêts in Tunisia. Valuable assistance with these permits was provided by Pr. Soumia Fahd and Pr. Saïd Nouira.

The authors owe much to the following people for providing field assistance, answering various letters of inquiries, providing literature and permission to use photographs, and for many other kinds of help. We arrange them alphabetically as every contribution was valuable and we thank them all:

Alberto Sánchez-Vialas, Arnaud Jamin, Axel Hernandez, Bayram Göçmen, Dani Boix, Frank Deschandol, François Maillet, Iñigo Martinez-Solano, Jean Lescure, Jean Rafaëlli, Jorge Gutiérrez-Rodríguez, Keith Wilson, Mabrouk Chetoui, Mark Oliver Rödel, Maximilian Dehling, Mehdi Chetibi, Mohamed Wassim Hizem, Philippe Geniez, Ridha el Ouni, Roberto López Ortiz, Salah Eddine Selmi, Stephano Doglio, and Victoire Koyamba.

Finally, we wish to thank our families for their understanding and support from our formative years onward.

Chapter 1

Introduction

Abstract

Amphibians are a group of vertebrates that originated in the Paleozoic Era. North Africa has few species, but these belong to very divergent lineages. Most of the species belong to Eurasian groups that colonized the region through land bridges in the Miocene. No mass extinctions have been detected in the region, but some marginal populations became extinct recently. The prolonged droughts could affect the relict populations of tropical amphibians in the central Sahara.

Keywords

Evolution; fossil record; Batrachia hypothesis; Eurasian lineage; functional guilds; global decline

1 Amphibian evolution and diversity

Amphibians are a class within Chordata whose origin dates back to the Paleozoic Era. In the Late Devonian, crossopterygian fishes (lobe-fin fishes) developed anatomical traits which allowed them to temporarily use the terrestrial habitats that surrounded rivers and streams. These features included the development of lungs and four limbs (robust and modified fins, but without fingers), which support their body weights on land. Transitional forms such as Elpistostege watsoni and Tiktaalik roseae (found in Canada) and Elginerpeton pancheni (found in Scotland) were recovered in deposits from 375 million years ago (Ma) (Ahlberg, 1995; Downs et al., 2008; Schoch, 2014). More derived forms appear 365 Ma, which show more developed limbs with seven or eight fingers (Acanthostega gunnari, Ichthyostega; Carroll, 1988). Ichthyostega could possibly inhabit streams and use the contiguous terrestrial habitats, whereas A. gunnari could have been exclusively aquatic (Ahlberg et al., 2005; Clack, 2002).

In the Early Carboniferous period (359–318 Ma), several Tetrapodomorpha showed a mosaic anatomy, combining traits typical of amphibians along with those found in other Tetrapoda (e.g., Crassigyrinus, Eucritta melanolimnetes; Panchen and Smithson, 1990; Clack, 2001). During the Late Carboniferous/Permian (318–270 Ma), the first true amphibians appear (Marjanović and Laurin, 2008; San Mauro, 2010). The monophyly of modern amphibians (subclass Lissamphibia) is supported by shared synapomorphies: monospondylous vertebrae, short ribs which do not encircle the body cavity, pedicellate teeth, presence of granular and mucous glands in the skin, fat bodies associated with the gonads, and skull structure (Duellman and Trueb, 1986). However, there is still no consensus on whether modern amphibians have evolved from a temnospondyl or lepospondyl ancestor and about the phylogenetic relationships of extant groups of amphibians: Procera (salamanders and caecilians are monophyletic) versus Batrachia (salamanders and frogs are monophyletic) hypotheses (Ruta and Coates, 2007; Fong et al., 2012; Maddin et al., 2012).

Based on the Batrachia hypothesis, the separation between frogs and salamanders occurred during the Late Permian/Triassic (270–240 Ma; Marjanović and Laurin, 2013). The first transitional amphibians that shared some characters with Anura appeared in the Early Triassic in Madagascar and Europe (250 Ma), Triadobatrachus massinoti and Czatkobatrachus polonicus (Sigurdsen et al., 2012). These species retained several plesiomorphic characters, like a high number of vertebrae, but possibly they were capable of hopping or performing short jumps (Shubin and Jenkins, 1995; Evans and Borsuk-Bialynicka, 1998; Sigurdsen et al., 2012). Stem-frogs were recovered in Lower and Upper Jurassic deposits (200–190 Ma) in Argentina and Arizona: Vieraella herbsti and Prosalirus bitis (Notobatrachidae) (Gao and Wang, 2001). The first crown-group frogs Callobatrachus sanyanensis (Discoglossoidea), Eodiscoglossus oxoniensis (Discoglossoidea), and Rhadinosteus parvus (Pipoidea, Rhinophrynidae) were found in younger deposits in China, England, and North America (Upper Jurassic/Lower Cretacic, 168–125 Ma; Evans et al., 1990; Henrici, 1998; Gao and Wang, 2001).

Salamanders possibly evolved in Eurasia during the Middle Jurassic (165 Ma), although there is an older candidate (Triassurus sixtelae; 240 Ma, Kirghizistan). The remains of T. sixtelae were small in size and low ossified, possibly a larval specimen and not allowing definitive conclusions (Skutschas, 2013). During the Middle Jurassic, stem-group salamanders are known to occur in Central Asia (Karaurus sharovi, Kokartus honorarius, and Urupia monstrosa) and England (Marmorerpeton kermacki; Gao and Shubin, 2003). The first crown-group Caudata, the genera Chunerpeton (Cryptobranchoidea, Middle Jurassic) and Beiyanerpeton and Qinglongtriton (Salamandroidea, Late Jurassic) are known from exceptionally well-preserved fossils found in China (Gao et al., 2013; Jia and Gao, 2016).

Origin of north-west African amphibians

North-west African amphibians, despite the low number of species, represent very divergent evolutionary lineages (Fig. 1.1). Most of the species belong to Eurasian groups which migrated to north-west Africa from the Mediterranean peninsulas, using the land bridges established intermittently during the Miocene (Veith et al., 2006; Gibert et al., 2013). The exceptions are the Sahelian species (Hoplobatrachus occipitalis, Sclerophrys xeros) and Sclerophrys mauritanica, the only bufonid belonging to a tropical lineage present in the mesic belt of north-west Africa (Pramuk et al., 2008).

Figure 1.1 Evolutionary relationships (measured as divergence times, in Ma) of north-west African amphibians, based on published molecular analyses (Pyron and Wiens, 2011; Pyron, 2014).

The Salamandridae family evolved in Europe, as independently indicated by the fossil record and phylogenetic analyses (Zhang et al., 2008). The oldest salamandrid (Koalliella genzeli, a stem-group pleurodeline) has been recovered from paleocenic deposits (57 Ma) in France and Germany, but the origin of the family was older, possibly between 86 and 58 Ma (Roček, 1994a; Marjanović and Witzmann, 2015). In north-western Africa, the older salamandrid remains (Pleurodeles) were recovered in Early Pleistocene sites (Bailon et al., 2011). Alytidae are a basal group of anurans which include three extant genera (Alytes, Discoglossus, and Latonia; Pyron and Wiens, 2011; Biton et al., 2016). Fossils of alytids were recovered in Europe and Israel (Latonia; 22–0.12 Ma) and in Europe (Discoglossus, 23–0.12 Ma; Alytes 2.5–0.12 Ma; López-Martínez and Ruiz Bustos, 1977; Sanchiz and Adrover, 1977; Roček, 1994b; Rodríguez et al., 2011; Gardner and Rage, 2016). In north-western Africa, alytids (Latonia and/or Latoglossus) were found since the Miocene (Gardner and Rage, 2016). The family Pelobatidae comprises a single extant genus (Pelobates). Fossils of possible crown-group Pelobatoids were recovered in Cretaceous and Paleogene deposits in central Asia (Chen et al., 2016), Gobiates spinari (87 Ma) and Macropelobates osborni (30 Ma). In Europe, pelobatids are known since the Oligocene, Eopelobates (34–12 Ma) and Pelobates (28–0.12 Ma). Pelobates decheni found in Germany (25 Ma) could be the sister lineage of all the extant Pelobates (Chen et al., 2016). In north-western Africa, pelobatids were found since the Late Pleistocene (Stoetzel et al., 2010). The family Hylidae originated in America and invaded Eurasia in two separate migratory events, during the Paleogene and the Miocene (Smith et al., 2005). Hylids were recovered since the Pliocene in north-west Africa (Raynal et al., 1990). True toads (Bufonidae) originated in America (Baez and Nicoli, 2004), and migrated to Eurasia through the Beringian land bridge during the Paleocene (Pramuk et al., 2008). Fossils of bufonids were found in north-west Africa since the Miocene (Gardner and Rage, 2016). The older crown-group ranoids were found in Cretaceous deposits from Sudan, supporting the hypothesis of an African origin (Bossuyt et al., 2006). The Raninae group migrated to Eurasia about 65 Ma (Bossuyt et al., 2006). The genus Pelophylax evolved in Eurasia and migrated back to northern Africa during the Miocene (Lymberakis et al., 2007). Fossils of Pelophylax frogs were found in north-west Africa since the Early Pleistocene (Hossini, 2001).

Amphibian diversity

At this moment, 6973 species of Anura and 722 species of Caudata are recognized (Frost, 2018). Gymnophiona (209 species) are not treated in this section because it has no extant representatives in the study region (Frost, 2018). Caudata are mostly confined to the northern hemisphere (Wells, 2007). Shared-derived characters of Caudata are the presence of a developed tail, the similar length of all their limbs, presence of gills in aquatic larvae, presence of teeth on both jaws, the absence of tympanic ears, and presence of ribs (Duellman and Trueb, 1986). Caudata show the longest genomes among vertebrates (e.g., the genomes of Ambystoma mexicanum and Amphiuma means are 10 and 30 times larger than that of humans, respectively; Hartl and Ruvolo, 2011; Keinath et al., 2015). Salamanders are also unique among vertebrates by their early development of autopodial structures during the skeletogenesis (Fröbisch and Shubin, 2011).

Most of Caudata have small sizes, typically between 100 and 200 mm in length, although there are giant species, such as Andrias davidianus that reach lengths of more than 1000 mm (Wells, 2007; Browne et al., 2013). Caudata have different reproductive strategies. Most of the species oviposit in water, from which emerge larvae that metamorphose after an aquatic phase (e.g., Ambystoma, Triturus); more exceptionally they oviposit in land, emerging already metamorphosed individuals (e.g., Hydromantes, Bolitoglossa; Wake and Hanken, 2004; Wells, 2007). Some species show lecitrophic viviparity (the larvae feed on the oocyte); the females deliver larvae that can require an aquatic phase (e.g., Salamandra salamandra) or not (e.g., Lycisalamandra; Buckley et al., 2007). Matrotrophic viviparity (the larvae feed on the oocyte, but the mother provides additional nutrition) is only found in two species of alpine salamanders, Salamandra atra and Salamandra lanzai (Veith et al., 1998). Numerous Caudata species show very contrasting dorsal or ventral coloration (e.g., Paramesotriton, Salamandra, Taricha). These colorful patterns act as a warning to predators, because these species secrete toxic substances through cutaneous glands (Brodie et al., 1974). Toxic secretions are not present in the larvae, which have a deceptive coloration (Mebs and Pogoda, 2005; Altig and McDiarmid, 2015).

The highest species richness of salamanders appears in hyperhumid warm temperate-subtropical regions of North America and Asia, particularly in the Apalaches (north-western USA), the sino-Himalayan ranges (south-eastern Asia), and the Mexican ranges (Sierra Madre Occidental and Oriental and the Trans-Mexican Volcanic Belt; Frost, 2018). The countries with the highest species richness are the United States (196 species), Mexico (145 species), China (72 species), Guatemala (65 species), Japan (33 species), and Costa Rica (32 species; Frost, 2018). Only four species of the Salamandridae family occur in the north-west of Africa, being confined to the northern pericoastal mesic belt (Fig. 1.2).

Figure 1.2 Map showing the patterns of species richness of north-west Africa Caudata, based on vector polygons provided by the IUCN Red List of Threatened Species. Darker colors indicate higher species richness.

Anurans are distributed over most of the continents, except in Antarctica, regions with extreme climates (polar and hyperarid) and some oceanic islands (Wells, 2007). Shared-derived characters among adult anurans include the absence of tail, the hind legs relatively longer than the forelimbs, limbs with radius, the ulna, tibia, and fibula being fused, and a low number of vertebrae (9–4), with the last fused vertebrae forming the urostyle (Duellman and Trueb, 1986). Anuran larvae have a streamlined body shape, with keratinous oral structures and external respiratory orifices (spiracles) (Duellman and Trueb, 1986). Anuran larvae have the capacity to adapt their body shape (and feeding apparatus in some species) to the conditions of aquatic habitats; in some cases two phenotypes of the same species can coexist using distinct resources (Pfennig and Murphy, 2002).

Most anurans are small-sized (between 10 and 70 mm), with the largest species reaching between 230 and 320 mm (e.g., Conraua goliath, Pyxicephalus adspersus, Rhaebo blombergi, Rhinella marina; Duellman and Trueb, 1986; Wells, 2007). Most of the anurans oviposit in water or close to the water, and the larvae metamorphose after an aquatic or fossorial phase (McDiarmid and Altig, 1999; Senevirathne et al., 2016), but some genera have direct development (e.g., Eleutherodactylus, Myobatrachus; Callery et al., 2001; Anstis et al., 2007). There are also a few lecitrophic and matrotrophic viviparous species (Eleutherodactylus jasperi, Nectophrynoides, and Nimbaphrynoides; Drewry and Jones, 1976; Sandberger-Loua et al., 2017).

The greatest richness of anuran species is found in the hyperhumid tropical and subtropical regions of America and Asia, particularly the eastern tropical Andes, the Amazonian basin, and the Brazilian Atlantic rainforest and in mainland south-eastern Asia, Sumatra, and Borneo (Frost, 2018). The countries with the highest species richness are Brazil (1011 species), Colombia (725 species), Perú (585 species), Ecuador (527 species), India (375 species), China (368 species), and Indonesia (341 species) (Frost, 2018). Only 13 species of anurans occur in the north-west of Africa and the peripheral archipelagos, with most of the species concentrated in the coastal plains and mountain ranges from the northern part of the region (Fig. 1.3; Schleich et al., 1996).

Figure 1.3 Map showing the patterns of species richness of north-west Africa Anura, based on vector polygons provided by the IUCN Red List of Threatened Species. Darker colors indicate higher species richness.

2 Amphibian natural history

The presence of amphibians in a given habitat is determined to a large extent by the vegetation cover, temperature, moisture, and the presence and typology of the aquatic habitats, and species respond differently to the environmental gradient (Hazell et al., 2001; Guerry and Hunter, 2002). Aquatic species show adaptations to move efficiently in this viscous environment, and terrestrial ones to reduce the risk of dehydration, facilitate on ground displacement, climbing, or the excavation of the substrate (Wells, 2007). These adaptations are reflected in the species external morphology (e.g., compressed tails, feet-webbing, toe-pads, spade-like metatarsal tubercles) and reproductive strategies (e.g., aquatic versus land ovipositing; direct versus indirect larval development); the combination of specific traits allows identify major functional guilds: fossorial, scansorial, terrestrial, or aquatic species (Wells, 2007; Hillman et al., 2009).

In Caudata, the species can be clustered in four guilds, according to Hillman et al. (2009): aquatic, semiaquatic, semiterrestrial, and terrestrial. Aquatic salamanders are those that retain external gills in the adult stage, but excluding facultative paedomorphics (Andreone and Dore, 1991). This guild includes salamanders with specialized eel-shaped bodies of the family Proteidae and Sirenidae and some stygobite Plethodontidae (Hillman et al., 2009). Semiaquatic salamanders spend a large part of their life cycle in aquatic habitats but adults do not retain external gills. These species can show specialized eel-shaped (Amphiumidae) or flattened bodies with laterally compressed tails (Cryptobranchidae, Rhyacotritonidae, and some rheophilic Salamandridae). Semiterrestrial salamanders spend a large part of their life cycle on land, but they return to aquatic habitats to breed. These species have relatively stout bodies, with laterally compressed or cylindrical tails, and in some cases, costal grooves, which facilitate the diffusion of water through the skin (Toledo and Jared, 1993). Examples of semiterrestrial salamanders are Ambystoma, Echinotriton, Salamandrina, Taricha, and Triturus. The north-western African salamanders are all included in this guild. Terrestrial salamanders do not return to aquatic habitats to oviposit or deliver the larvae. These species show costal grooves, slender bodies, and in the case of scansorial forms, thin and prehensile tails and interdigital webbing (Wells, 2007). Representatives of this guild include numerous species of plethodontids (e.g., Aneides, Hydromantes, Ensatina; Petranka, 1998) as well as some Salamandridae (S. lanzai, S. atra, Lycisalamandra). In north-west Africa, Gibraltar strait populations of Salamandra algira tingitana also belong to this guild (Mateo et al., 2003).

Anura show greater ecological diversification than Caudata (Duellman and Trueb, 1986; Wells, 2007). Anuran species can be grouped into seven functional guilds, aquatic, semiaquatic, semiterrestrial, terrestrial, scansorial, fossorial, and semifossorial (Ernst et al., 2006; Hillman et al., 2009). Aquatic frogs show fully webbed feet, absence of tongue, and presence of lateral line, feeding mainly or exclusively on aquatic prey (Canedo et al., 2006). Pipids are the only representatives of this guild (Hillman et al., 2009). Semiaquatic frogs show fully webbed feet, smooth ventral skin, and relatively long hind legs, being strongly linked to aquatic habitats, but also using surrounding land (particularly juveniles) and including terrestrial prey in their diet. This guild includes species of Hylidae, Ranidae, and Dicroglossidae (Hillman et al., 2009). In north-west Africa, two species of the genus Pelophylax and H. occipitalis belong to this guild. Semiterrestrial frogs are associated with aquatic habitats mostly or exclusively during the breeding period. These frogs show incomplete feet webbing and rugosities in the ventral skin which facilitate water diffusion. In north-west Africa, most of the anurans are clustered within the semiterrestrial guild: Alytes, Discoglossus, Barbarophryne, Bufo, Bufotes, and Sclerophrys. Scansorial frogs are a specialized type of semiterrestrial frogs which show slender bodies and long legs with adhesive discs in the fingers and toes to facilitate climbing. Representatives of this guild are found in Centrolenidae, Hyperoliidae, Hylidae, and Phyllomedusidae families (Hillman et al., 2009). In north-west Africa, only one species belongs to this guild, Hyla meridionalis. Terrestrial frogs are morphologically similar but reproduce independently of water (Jennings and Hanken, 1998; Wells, 2007). Examples of this guild are found in Bufonidae, Ceratobatrachidae, Eleutherodactylidae, and Leiopelmatidae (Hillman et al., 2009). Fossorial and semifossorial frogs show compact skulls and bodies, short hind legs, and cornified structures on their feet. Fossorial frogs (e.g., Arenophryne) have direct development whereas semifossorial frogs use aquatic habitats to breed (Anstis et al., 2007; Wells, 2007). The fossorial and semifossorial guilds are composed by species inhabiting dry environments of Limnodynastidae, Myobatrachidae, Pelobatidae, and Scaphiopodidae (Hillman et al., 2009). In north-west Africa, this guild is represented by a single species, Pelobates varaldii.

3 Amphibian global decline

In the past four decades, amphibians have shown a worldwide decline, particularly affecting rheophilic tropical frogs (Houlahan et al., 2000; Stuart et al., 2004). The hypotheses explaining the global decline can be summarized in two groups, although their effect is possibly synergistic (Collins and Storfer, 2003; Beebee and Griffiths, 2005):

1. The decline attributable to well-known ecological processes which have been acting for an extended period time, such as the overexploitation of amphibians as a food resource and the habitat disturbance, including the spread of alien species and the action of chemical pollutants (particularly wind-borne pesticides; Davidson et al., 2002).

2. The decline linked to changes in climatosphere dynamics (e.g., global warming and changes in rainfall patterns; McMenamin et al., 2008) or the widespread and epidemic diseases (Ranavirus, Batrachochytrium dendrobatidis, Batrachochytrium salamandrivorans; Lips et al., 2008). The latter factors were repeatedly invoked to explain the decline of local amphibian populations, even in pristine environments (Becker and Zamudio, 2011; Parrott et al., 2017).

In north-west Africa, there are no data corroborating a general decline of amphibian populations, although the local extinction of marginal populations has been documented, in the ecological/distributional limits of a species or involving populations located in or close to cities (Ben Hassine and Nouira, 2012; Beukema et al., 2013; Ibrahim, 2014). The pathogenic fungus B. dendrobatidis was detected in northern Morocco, but it is unknown if it has any negative effects on the local amphibian populations (El Mouden et al., 2011). Irregular hydrological regimes in the Sahel (associated with fluctuations in the extension of the desert about 1.4 million km²; Tucker et al., 1991; Tucker and Nicholson, 1999) possibly have a very negative impact on the relic populations of amphibians that subsist in the extreme south of the region (Padial