Basic and Applied Herpetology 27 (2013): 23-50

Chapter 25 Amphibians of Morocco, including Western Sahara: a status report Ricardo Reques1,*, Juan M. Pleguezuelos2, Stephen D. Busack3, Philip de Pous4 1 Departamento de Ecología Evolutiva, Estación Biológica de Doñana, Sevilla, Spain. 2 Departamento de Zoología, Facultad de Ciencias, Universidad de Granada, Granada, Spain. 3 North Carolina Museum of Natural Sciences, Raleigh, North Carolina, USA. 4 Faculty of Life Sciences and Engineering, Departament de Producció Animal (Fauna Silvestre), Universitat de Lleida, Lleida, Spain.

* Correspondence: Departamento de Ecología Evolutiva, Estación Biológica de Doñana, Cl. Americo Vespucio, s/n, 41092, Isla de la Cartuja, E-41092 Sevilla, Spain. Phone: +34 616810551, Email: [email protected] Received: 10 January 2013; received in revised form: 30 September 2013; accepted: 21 October 2013.

Morocco has one of the highest rates (28.6%) of amphibian endemism among countries bordering the Mediterranean Sea and, while large areas of Morocco are crucial for conserving amphibian biodiversity, some areas are not afforded legal protection. We examine biodiversity, identify immediate anthropogenic threats, dis- cuss critical habitat for the conservation of amphibian diversity and the role of currently protected areas in meet- ing conservation goals within Morocco, Western Sahara included. The study area harbours 14 amphibian species, eight of which are assigned to the categories of Endangered (Pelobates varaldii), Vulnerable (Salamandra algira, Amietophrynus xeros, and Hoplobatrachus occipitalis) or Near Threatened (Pleurodeles waltl, Alytes maurus, Bufo spinosus, and Barbarophryne brongersmai) using IUCN criteria at the regional level of the study area. Habitat loss and degradation due to conversion of land for agriculture, urbanization, or industry are major threats, but infrastructure for tourism, freshwater pollution by chemicals, introduction of non-native species to aquatic ecosystems (Gambusia holbrooki), pathogens (Batrachochytrium dendrobatidis), road-kills, and natural disasters (drought), are also rapidly increasing threats. In addition, consequences from global warming must also be considered. The present Conservation Area Network (CAN) does not include distributional ranges of some amphibian species, and a more complete CAN in Atlantic and desert areas is suggested. The northwestern Atlantic, Rif-Middle Atlas, Central Atlantic, and Tiris regions should be considered priorities for conservation because of amphibian endemism and/or the existence of isolated amphibian populations.

Key words: amphibian decline; conservation planning; Morocco; survival threats; Western Sahara.

Los anfibios de Marruecos, incluyendo Sáhara Occidental: Informe sobre su situación. Marruecos posee una de las tasas más elevadas de anfibios endémicos (28.6%) entre los países de la cuenca mediterránea y, a pesar de que exten- sas áreas de Marruecos son fundamentales para la conservación de la biodiversidad de anfibios, otras zonas no gozan de protección legal. En este estudio examinamos la biodiversidad, identificamos las amenazas antropogénicas más inmediatas y discutimos el hábitat crítico para la conservación de la diversidad de anfibios y el papel de las áreas actualmente protegidas en el cumplimiento de los objetivos de conservación en Marruecos, Sáhara Occidental inclu- ido. El área de estudio alberga 14 especies de anfibios, ocho de las cuales se asignan a las categorías de En Peligro (Pelobates varaldii), Vulnerable (Salamandra algira, Amietophrynus xeros y Hoplobatrachus occipitalis) o Casi Amenazada (Pleurodeles waltl, Alytes maurus, Bufo spinosus y Barbarophryne brongersmai), utilizando criterios de la UICN a nivel regional en la zona de estudio. La pérdida de hábitat y la degradación debido a la conversión de ter- renos para la agricultura, urbanismo o industria son las principales amenazas, pero las infraestructuras relacionadas con el turismo, la contaminación de agua dulce por productos químicos, la introducción de especies exóticas en los ecosis- temas acuáticos (Gambusia holbrooki), la llegada de patógenos (Batrachochytrium dendrobatidis), los atropellos y los desastres naturales (sequía), también son amenazas en rápido aumento. Del mismo modo, también deben tenerse en cuenta las consecuencias del calentamiento global. La actual Red de Áreas de Conservación (CAN) no incluye ran-

This chapter should be cited as: Reques R., Pleguezuelos J.M., Busack S.D. & de Pous, P. (2013). Amphibians of Morocco, including Western Sahara: A Status Report. Chapter 25 in Part 2. Mauritania, Morocco, Algeria, Tunisia, Libya and Egypt in Vol. 11. Conservation and Decline of Amphibians: Eastern Hemisphere of the series Amphibian Biology. Basic and Applied Herpetology 27: 23-50. DOI: http://dx.doi.org/10.11160/bah.13003/ REQUES 24 ET AL. gos de distribución de algunas de las especies de anfibios, por lo que se sugiere una red de conservación más comple- ta incluyendo zonas del Atlántico y el desierto. Las regiones del Atlántico Noroeste, Rif-Atlas Medio, Atlántico Centro y Tiris deben ser consideradas prioridades de conservación debido a los endemismos presentes y/o la existen- cia de poblaciones aisladas de anfibios.

Key words: amenazas para la supervivencia; declive de anfibios; Marruecos; planificación de conservación, Sáhara Occidental.

Morocco (including Western Sahara), Morocco has only 14 of Africa’s 993 with an area of 705 850 km2, is biogeograph- amphibian species (Table 1) (IUCN, 2009). ically within the Maghreb. In the north, mild One of these species, Pelobates varaldii, is and wet winters alternate with long, hot and endemic but three other species, Discoglossus sco- increasingly dry summers as one progresses vazzi, Alytes maurus, and Barbarophryne southward and eastward. Annual rainfall averages brongersmai, may be considered quasi-endemic 950 mm in the north (Tangier), 430 mm along because most of their distributional area is with- the Atlantic Coast (Casablanca), and less in Morocco and only isolated populations are than 100 mm in most of the Western Sahara. known from within Algeria. Morocco, therefore, Four mountain ranges, the Rif (Jebel has almost exclusive responsibility for global Tidighine, 2456 m), the Middle Atlas (Jebel Table 1: Moroccan amphibian species. Bou Naceur, 3356 m), the High Atlas (Jebel Toubkal, the highest peak in North Africa at 4167 m) and the Anti-Atlas (Jebel Siroua, CAUDATA 3304 m) traverse the country from north to Salamandridae Pleurodeles waltl Michahelles, 1830 south. The Sahara Desert begins south of the Salamandra algira Bedriaga, 1883 Anti-Atlas and High Atlas and extends east for more than 800 km and south for more ANURA Alytidae than 1000 km to 21º N latitude. Alytes maurus Pasteur and Bons, 1962 Morocco probably has the most complete Discoglossus pictus Otth, 1837 dataset regarding amphibian distribution and Discoglossus scovazzi Camerano, 1878 Pelobatidae status across northern Africa, and BONS & Pelobates varaldii Pasteur and Bons, 1959 GENIEZ (1996), SCHLEICH et al. (1996), Hylidae MATEO et al. (2003), and GENIEZ et al. (2004) Hyla meridionalis Böttger, 1874 have been relied upon heavily for summaries Bufonidae of distribution and habitat preferences. While Amietophrynus mauritanicus (Schlegel, 1841) Amietophrynus xeros (Tandy, Tandy, Keith and this manuscript was in review BEUKEMA et al. Duff-Mackay, 1976) (2013) described a third subspecies within Bufotes boulengeri Lataste, 1879 Salamandra algira (S. a. splendens) and provid- Barbarophryne brongersmai (Hoogmoed, 1972) Bufo spinosus Daudin, 1803 ed natural history data separately for the three Dicroglossidae Moroccan subspecies; supplemental natural Hoplobatrachus occipitalis (Günther, 1858) history data for other amphibian taxa are also Ranidae provided throughout this paper. Pelophylax saharicus (Boulenger, 1913) STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 25 conservation of these four amphibians. Table 2: Red List status (IUCN categories) for Proximity to the Iberian Peninsula has provided amphibians of Morocco (PLEGUEZUELOS et al., 2010). CR=Critically Endangered; EN=Endangered; Morocco with the greatest European influence VU=Vulnerable; NT=Near Threatened; LC=Low on its amphibian fauna among all African Concern; NSp=number of species; End=number of countries; two species (Pleurodeles waltl and Bufo endemic species; for comparative purposes global spinosus) and some genera (Salamandra, data for the amphibians of the Mediterranean Basin Discoglossus, Alytes, Pelobates, Pelophylax) are of are included (COX et al., 2006). European origin. Because of greater habitat diversity, higher rainfall, and lower rates of evap- CR EN VU NT LC NSp End oration, amphibian species richness is highest in the north while large areas within the Sahara Morocco, Anura 1 2 3 6 12 1 Morocco, Caudata 1 1 2 0 lack amphibians (BAHA EL DIN et al., 2008). Morocco, Amphibia 1 3 4 6 14 1 The global conservation status of Moroccan Mediterranean, Amphibia 2 13 13 17 61 106 68 amphibians has been assessed quite recently (Table 2; see also COX et al., 2006) and while large areas of Morocco are crucial to conserving in northern wetlands and mountains, but these amphibian biodiversity (Table 3), many signifi- habitat types are not always well-represented in cant areas are not afforded legal protection the current Conservation Area Network (CAN, (RONDININI et al., 2006; DE POUS et al., 2011). MINISTÈRE DE L’AGRICULTURE, 1994). Threatened amphibian species and populations Currently, no amphibian species is legally pro- are found in southern desert and savannah, and tected in Morocco.

Table 3: Precipitation and attributes of the distributional ranges of amphibians in Morocco (including Western Sahara). See text for procedure.

Species Range within Percentage of global Precipitation Morocco (km2) range within Morocco Minimum Maximum Mean

Alytes maurus 5 500 ~95 700 > 2000 995.7 Amietophrynus mauritanicus 395 300 47.88 < 100 2000 466.1 Amietophrynus xeros 12 300 0.21 < 100 Barbarophryne brongersmai 95 100 ~ 95 250 600 293.7 Bufo spinosus 183 200 1.16 500 > 2000 541.9 Bufotes boulengeri 438 600 22.44 250 > 2000 441.9 Discoglossus pictus 5 300 2.22 150 600 270.6 Discoglossus scovazzi 190 600 ~ 95 350 2000 614.9 Hoplobatrachus occipitalis 525 0.01 < 100 Hyla meridionalis 240 900 31.67 300 2000 580.8 Pelobates varaldii 7 000 100 600 800 710.7 Pelophylax saharicus 398 300 28.21 < 100 2000 463.7 Pleurodeles waltl 60 500 14.11 500 2000 733.2 Salamandra algira 10 800 48.74 800 > 2000 943.9 All Species 441.8 Morocco 413.1 REQUES 26 ET AL.

Figure 1: Major habitats utilized by amphibians at the regional level (from PLEGUEZUELOS et al., 2010).

In the following pages amphibian biodiver- of uncultivated sandy soil next to temporary sity in Morocco is examined, the most impor- ponds; DE POUS et al. (2012) provided 134 geo- tant habitats (Fig. 1) and the role of protected referenced localities. Spawning sites are ephemer- areas are discussed, and immediate threats to al ponds and dayas (Fig. 3a); modified habitats this biodiversity are identified. It is hoped that are avoided, and it is the most stenoecious this information will assist environmental and Moroccan amphibian (BEUKEMA et al., 2013). conservation planning in Morocco by identify- The northernmost locality is a small, forested ing non-random threats to diversity in terms of area just south of the Tangier airport, easternmost species’ associations or ecological preferences. localities are Khemisset and south of Ouezzane, In addition, this treatment provides important and the southernmost locality is in the vicinity of information on a group of terrestrial verte- Oualidia. Suitable areas for this species along brates under-represented in the CAN of most most of the Moroccan Atlantic coastline, into the countries and frequently neglected in conserva- Souss Valley and southwards, and discontinuous- tion policy (MILNER-GULLAND et al., 2006). ly along the Mediterranean coast east to the Algerian border, potentially do exist. IUCN RED LIST STATUS OF MOROCCAN ESCORIZA & BEN HASSINE (2013) have AMPHIBIANS (2009) reported a newly-discovered population of Pelobates varaldii, represented by a tadpole Endangered from one of 21 ponds surveyed in March 2013, in the Ben Slimane region. They sug- Pelobates varaldii: The Moroccan spadefoot gest this population likely demonstrates the toad (Fig. 2a), an endemic, is locally distributed relictual nature currently found in a formerly within the northwestern coastal plain among continuous distribution through northern cork oak and other forested habitat and in areas Morocco’s Atlantic coastal plain. STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 27

a b c

d e f

Figure 2: Portraits of some Moroccan amphibians: (a) Pelobates varaldii (Mamora cork oak forest). (b) Salamandra algira (Yebel Musa). (c) Alytes maurus (Chefchaouen). (d) Discoglossus scovazzi (Beni Snassene). (e) Amietophrynus xeros (Atar, Mauritania). (f) Bufotes boulengeri (Aïn Leuh). Photo credits: (a) P. de Pous, (b) S. Yubero, (c) S.D. Busack, (d) R. Reques and (e & f) J.M. Pleguezuelos.

Current major threats to survival derive lophotus rather than P. clarkii [fide John from loss and degradation of habitat due to Cooper, North Carolina State Museum, in conversion of suitable land to pasture for, litt., October 2010]) in a pond east of and water pollution by, livestock. Survival is Larache (35.08122°N, 6.06908°W) signals contingent upon sandy soil, and industrial an additional threat to this and other farming is expanding into coastal areas (DE species. Invasive crayfish present an POUS et al., 2012). Populations are often increasingly serious threat to aquatic restricted to temporal ponds where hydro- organisms, including amphibian larvae, periods may not be sufficient for comple- wherever they become established (COOPER tion of larval development (developmental & ARMSTRONG, 2007). period is unstudied, but is dependent upon Batrachochytrium dendrobatidis, a globally water depth, temperature, and availability of distributed fungus responsible for chytrid- food; four to six months is likely). Tadpoles iomycosis and mass mortality among amphib- remaining in permanent bodies of water are ians worldwide (HEATWOLE, 2013), has disappearing because of introduced predato- recently been reported from P. varaldii in ry fish (Gambusia holbrooki; see also Morocco. One of ten larvae sampled in April SALVADOR, 1996 and STUART et al., 2008) 2009, from ± 20.5 km SE Larache and sudden draining during the larval peri- (35.038110°N, 6.029248°W) demonstrated a od. Additional discovery (November 2008, 0.4-genome equivalent infection intensity of this DE POUS et al., 2012) of an indeterminate pathogen (EL MOUDEN et al., 2011). This finding, species of Procambarus (photographic docu- while suggesting a low prevalence of infection, mentation is suggestive of P. [Ortmannicus] warrants additional investigation. Populations of 28 REQUES ET AL.

a b

c d

e f

g h STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 29 these amphibians are decreasing and there is warranted (STEINFARTZ et al., 2000; DUBOIS urgent need for conservation and manage- & RAFFAËLLI, 2009; BEUKEMA et al., 2010, ment; additional protected areas within its 2013). Salamandra algira populations occur- current range, coupled with management ring within the Rif and northern border of actions for conservation, are needed to miti- the Middle Atlas have been partitioned into gate continued population reduction. two subspecies, S. a. splendens Beukema, de The Evolutionary Distinct and Globally Pous, Donaire-Barroso, Bogaerts, Garcia- Endangered (EDGE) program at the Zoological Porta, Escoriza, Arribas, El Mouden & Society of London (http://www.edgeofexistence. Carranza 2013 and S. a. tangitana Donaire- org/) recently listed P. varaldii at number 36 on Barroso & Bogaerts 2003. Some populations their global amphibian top 100. of Salamandra are ovoviviparous (DONAIRE- BARROSO & BOGAERTS, 2003a; BEUKEMA et Vulnerable al., 2013). Viviparous populations from the Tingitane Peninsula have also been proposed Salamandra algira: The range of the for species status (as S. tingitana; DUBOIS & North African fire salamander (Fig. 2b) is RAFFAËLLI, 2009). The population in Beni widely fragmented throughout northern, Snassen is assigned to S. algira spelaea, which rather wet, mountain ranges in Morocco and is more related to the Algerian populations. Ceuta, Spain (MARTÍNEZ et al., 1997; ESCORIZA The species is locally common in the western et al., 2006; BOGAERTS et al., 2007; ESCORIZA and central Rif Mountains as well as in the & COMAS, 2007; BEUKEMA et al., 2010, Middle Atlas (FAHD et al., 2006; BEUKEMA et 2013). Recent morphological, genetic, and al., 2010, 2013), in habitat associated with ecological studies suggest that there are dif- humid, montane areas of Atlas Cedar ferent genotypes and phenotypes with parap- (Cedrus atlantica), Pyrenean Oak (Quercus atric distributions within Moroccan pyrenaica), and mixed forest (Abies, Cedrus, Salamandra, and further taxonomic study is Pinus, and Quercus) with an abundance of montane streams (MARTÍNEZ-MEDINA, 2001) (Fig. 3b), and has been cited in caves Figure 3: Habitats for some of Morocco’s amphibian (AELLEN, 1951; DONAIRE BARROSO & species. (a) Nufar pool, Kasr-el-Kebir; P. varaldii. (b) BOGAERTS, 2001). Salamandra algira occurs Stream, Talassemtane, Bab Taza; S. algira, A. maurus, between 100 and 2100 m above sea level. and B. spinosus. (c) Temporary pond, Asilah. Breeding habitat, P. waltl. (d) Urban habitat, Populations are threatened by habitat loss Chefchaouen, S. algira, A. maurus, D. scovazzi, due to the agriculture of Cannabis sativa in and P. saharicus. (e) Spur of the Anti-Atlas, west the Rif, deforestation, alteration and chan- of Agdz; B. brongersmai. (f) Lanasser pool, Rif nelization of water, overgrazing by livestock Mountains, H. meridionalis, A. mauritanicus, throughout its range and, locally, by road-kill and P. saharicus. (g) Fort Bou Cherif, B. boulengeri. (TAIQUI, 1997; TAIQUI & MARTÍN- (h) Dayet Sjri, Merzouga, Tafilalt, A. mauritanicus. CANTARINO, 1997). The pet trade, climatic Photo credits: (a) D. Donaire, (b & f) R. Reques, change, and the chytrid fungus are emerging (c & d) S.D. Busack, (e, g, & h) J.M. Pleguezuelos. concerns (BOGAERTS, 2007). REQUES 30 ET AL.

Near Threatened genetic analysis shows that A. maurus pre- sents low levels of mtDNA variability with Pleurodeles waltl: The Sharp-ribbed newt is no clear geographical structuring (DE POUS distributed mainly throughout the coastal plain et al., 2013). Its current, fragmented range is of northwestern Morocco (GARCÍA-PARIS et al., likely a result of increasing temperatures 2004; BEUKEMA et al., 2013) within a roughly throughout the Quaternary, as a fossil record triangular area connecting the Tingitane likely attributable to A. maurus suggests a Peninsula, Souk Jemaa des Oulad Abbou, and much wider historical distribution (BEUKEMA Anosseur, but three recorded localities et al., 2013; DE POUS et al., 2013). (Talamrhecht, Safi, Île d'Mogador) fall outside Populations in the Chefchaouen district this area. In general, P. waltl inhabits ponds, are threatened by pollution due to human lakes, ditches, and slow-moving streams with activity (Fig. 3d) and by introduction of the non-permanent water (Fig. 3c) and adjusts invasive Eastern mosquito fish (Gambusia to habitat modified by cultivation (BEUKEMA holbrooki) (DONAIRE-BARROSO et al., 2009a). et al., 2013); populations are quite fragmented In the Middle Atlas some breeding localities and are declining throughout the country in the Jebel Bou Iblane region are within pro- (BEJA et al., 2009). tected areas (site of bio-ecological interest, This species is Near Threatened because priority 1) but pressure from cattle grazing, of a general decline in population numbers deforestation, canalization of mountain- (almost 30% over ten years) and widespread water, and soil erosion are increasing threats loss of habitat (BEJA et al., 2009). Main outside of protected areas (DONAIRE- threats to survival are agrochemical pollution BARROSO et al., 2006). Additional threats to and eutrophication caused by livestock, loss this species include the potential spread of of aquatic habitat through drainage, and loss recently detected chytrid fungus (EL and fragmentation of terrestrial habitats MOUDEN et al., 2011) as well as the effects of (BAHA EL DIN et al., 2008). climatic change. Due to its reduced distribu- tion (about 30 known localities in an area of Alytes maurus: The Moroccan midwife less than 5000 km2), the species has been list- toad (Fig. 2c) is often found in association ed as Near Threatened (DONAIRE-BARROSO with larvae-bearing populations of S. algira et al., 2009a). (DONAIRE-BARROSO et al., 2006; BEUKEMA et al., 2013). It has a discontinuous distribution Barbarophryne brongersmai: The genus throughout humid areas in montane-karst Barbarophryne was recently published and forested areas (mainly Q. pyrenaica) close (BEUKEMA et al., 2013) to reduce polyphyly to water sources in the western and central within the “green toad group” by removing Rif Mountains (Fig. 3b) between 200 and this species from the genera Bufo, Bufotes, or 2050 m above sea level (Jebel Tazekka) (see Pseudepidalea to which it has been variously DE POUS et al., 2013). Its distribution in the assigned. Brongersma’s toad is found from 5 m Middle Atlas is poorly known (LIBIS, 1985; to 1000 m above sea level in suitable habitat DONAIRE-BARROSO et al., 2006). A recent south of Casablanca (HOOGMOED, 1972; STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 31

GENIEZ et al., 2000; BEUKEMA et al., 2013) 2001; ZANGARI et al., 2006). Abundant in where it inhabits semiarid areas of Argania sub-humid and humid bioclimatic zones, but spinosa, Euphorbia, grass-like vegetation, and also in semiarid zones around Casablanca, ploughed fields. Temporary ponds where it this frog inhabits temporal and ephemeral breeds are generally located in rocky areas ponds of fresh or slightly brackish water and (Fig. 3e), and the species has been observed montane streams (BEUKEMA et al., 2013). in artificial bodies of water (GARCÍA-MUÑOZ Localized loss of breeding sites through agri- et al., 2009). It is threatened by increased cultural development in inland areas and aridity and pollution and drainage of breed- increasing salinity in lagoons are the main ing habitat throughout much of its range. threats (SALVADOR et al., 2009); it is pre- Although present in the Parc National de sumed that the species can resist light modi- Souss-Massa, the species is probably in fication of its habitat by deforestation. decline, and at a loss rate of almost 30% over Chytrid fungus, B. dendrobatidis, recently has ten years makes it close to qualifying for been reported from D. scovazzi at two locali- Vulnerable status (SALVADOR et al., 2006). ties in Tétouan Préfecture. One metamorphic Populations in the Souss Valley, Ifni, and individual sampled in November, 2006, from Low Draa and Tekna regions, and those in Agnane (near Tétouan at 35.535881°N, northern Western Sahara, are being nega- 5.386177°W) presented an infection intensi- tively affected by the proliferation of cisterns ty of 29.9 genome equivalents and one adult being built to water cattle. Toads, attracted sampled in February, 2007, approximately by the humidity, fall into these underground 18.5 km SE Larache (35.043940°N, cisterns and die when these watering struc- 6.046156°W) presented an infection intensity tures dry up (GARCÍA-MUÑOZ et al., 2009). of 60.3 genome equivalents (EL MOUDEN et al., One-meter-deep decantation chambers linked 2011). Additional research regarding the to these cisterns are frequently used by actual extent of the distribution of this fun- breeding toads and, while post-metamorphic gus is of paramount importance. toads can probably climb the vertical walls, adults are less likely to escape in this Discoglossus pictus: The western limit of manner (L. García-Cardenete, personal distribution for the common painted frog is communication). unclear. In Morocco apparently it is limited to a strip from the Cap des Trois Fourches Least Concern Peninsula (Melilla included) to Morocco’s northern border with Algeria and possibly Discoglossus scovazzi: The white-bellied extending westward beyond the Moulouya painted frog (Fig. 2d) inhabits suitable habitat Basin (BEUKEMA et al., 2013). It breeds in in Morocco and the Spanish territory of most types of still water, including temporal Ceuta at elevations from near sea level to ponds, marshes, and brackish water. Habitat 2650 m above sea level in the High Atlas alteration, including changes in traditional Mountains (see Fig 9 in BEUKEMA et al., land-use and urbanization (BOSCH et al., 2013) (DUBOIS, 1982; MARTÍNEZ-MEDINA, 2009), is its principal threat. REQUES 32 ET AL.

Hyla meridionalis: Mediterranean treefrogs tus based on mitochondrial DNA (see STÖCK are widely distributed throughout the western et al., 2006, 2008; BEUKEMA et al., 2013). Mediterranean (TEJEDO & REQUES, 2002); Inhabiting forested areas, shrubland, dry northern Mediterranean locations appear to grassland, semi-desert, and desert at eleva- have been colonized recently from Africa tions from near sea level to 2670 m above sea (RECUERO et al., 2007; but see STÖCK et al., level (Fig. 3g), Boulenger’s toad is one of 2012). Widespread in Morocco from coastal the most widespread amphibians in wetlands to montane habitats, it inhabits most Morocco. The main threat facing the species areas adjacent to still or moving water (Fig. 3f); appears to be loss of breeding habitat through current gaps in its apparent range in many drainage of wetlands, management of natural parts of northeastern Morocco and the water sources, and water pollution (IUCN, Middle-Atlas and High Atlas are likely due to 2006). In the southern belt of its Moroccan a lack of distributional research (BEUKEMA et al., distribution B. boulengeri is the species most 2013). Loss of terrestrial and aquatic habitats likely to die inside modern underground cis- is the major threat, but populations remain terns built for watering cattle (L. García- locally abundant. One of ten H. meridionalis Cardenete, personal communication). larvae sampled in April, 2009, approximately 18.5 km SE Larache (Tétouan Préfecture; Bufo spinosus: The common toad is a 35.043940°N, 6.046156°W) demonstrated a European species with genetically distinct 395.9 genome equivalent infection intensity relictual populations in Morocco (GARCÍA- of the fungus B. dendrobatidis (EL MOUDEN PORTA et al., 2012). Isolated populations are et al., 2011). found along Mediterranean mountains and in other mountains except the Anti-Atlas Amietophrynus xeros: The desert toad (Fig. 2e) (FAHD et al., 2006; BEUKEMA et al., 2013). It has been recorded from the extreme south of inhabits very humid areas near permanent Western Sahara. The population found in water, mostly in mountainous regions (Fig. 3b), temporal pools at Aouadi, a well close to Wadi and attains an elevation of 2750 m on Jebel Aïn Ascaf, is probably relictual and adversely Tinergouet in the High Atlas. Populations impacted by regional drought; in areas near might be locally impacted by deforestation, and within Adrar Atar (Mauritania) the species water pollution, and draining of traditional is always present around natural, temporal, breeding places (IUCN, 2006). While not small reservoirs. Listed as Least Concern at threatened at the global level (AGASYAN et al., the global level, at the regional level of 2009), at the regional level of Morocco it Morocco it could be considered Vulnerable can be considered Near Threatened (PLEGUEZUELOS et al., 2010). (PLEGUEZUELOS et al., 2010).

Bufotes boulengeri: Previously considered a Amietophrynus mauritanicus: The Mauritanian member of the Palearctic green toad complex toad is a Maghrebian endemic with a wide as Bufo viridis, this toad (Fig. 2f) has been distribution from the northern Sahara Desert recently reassigned independent species sta- to the Mediterranean coast, but its presence STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 33 in Western Sahara is unconfirmed (DONAIRE- alteration of its habitat (Figs. 3d, 3g), this BARROSO et al., 2009b). It frequents nearly species is presumed to have no major threats any water source (Figs. 3f, 3h), including to survival. Survival, in fact, appears assisted temporary ponds used for crops or cattle at in some regions by artificial ponds installed elevations from sea level to 2650 m above sea for agricultural purposes. level in the High Atlas (DUBOIS, 1982). This may be the most abundant and widely dis- ECOLOGICAL FACTORS AFFECTING tributed amphibian in Morocco and it cur- DISTRIBUTION rently does not require conservation action. Some populations, however, are affected by Understanding environmental factors and habitat loss and casualties due to motor vehi- ecological requirements for survival of cles (FAHD et al., 2002) and, attracted by amphibian populations is key to developing humidity, some individuals die inside under- management strategies for conservation ground cisterns in the southern limit of its (SEMLITSCH & ROTHERMEL, 2003). Moroccan distribution (J.M. Pleguezuelos, Amphibian species richness is influenced by personal observation). latitude within Africa, with maximum values at the Equator and minimum values at north- Hoplobatrachus occipitalis: The groove- ern and southern continental extremes crowned bullfrog, widely distributed further (RONDININI et al., 2006). Other factors south in Africa, recently has been found in accounting for African amphibian richness are Western Sahara. Known only from the south- the presence of suitable breeding places, rain- western margin of Western Sahara (Gleïb fall, and terrestrial habitats used during non- Ladjir) it survives in temporary ponds of breeding periods; below we review these fac- fresh water in regions with a Sahelian climate. tors within Morocco. The principal threat to Western Saharan pop- With the exception of the ovoviviparous ulations is its dependence on water; at the S. a. tingitana in mountains north and west of regional level of Morocco it could be consid- Tétouan (DONAIRE BARROSO & BOGAERTS, ered Vulnerable (PLEGUEZUELOS et al., 2010). 2001; BEUKEMA et al., 2010), the presence of every native species is contingent upon aquatic Pelophylax saharicus: Populations of the habitat for larval development. Mediterranean Saharan green frog are generally widespread wetlands and other suitable breeding areas are in Morocco from sea level to 2670 m above highly temporal and unpredictable, and sea level in the Middle Atlas but fragmented Morocco has suffered considerable degradation in the desert because of patchy availability of and loss of wetlands (RAMDANI et al., 2009); habitat (oases). Locally abundant where wet- ecological monitoring of 24 wetlands and adja- land habitat exists, P. saharicus is one of the cent habitat (4529 ha) recorded a shrinkage in most commonly recorded amphibians in area of 25% from 1978 to 1999 (MORGAN, Morocco. An aquatic species that inhabits 1982; BIRKS et al., 2001; RAMDANI et al., 2001; irrigation ditches, ponds, springs, rivers, and GREEN et al., 2002). Oligohaline ponds (along temporary pools, and with high tolerance to the Atlantic coast), montane lakes (in the REQUES 34 ET AL.

Middle Atlas) and temporal wetlands (both in and tadpoles (STUART et al., 2004), and desic- Saharan and coastal areas), habitats considered cation of temporal pools results in loss of eggs most threatened, are precisely the habitats most and tadpoles (PECHMANN et al., 1991) in important for amphibian reproduction. amphibian populations in arid regions Degradation resulting from hydrological alter- (SEMLITSCH et al., 1996; BEJA & ALCAZAR, ation (in Atlantic plains), siltation (in some 2003). In general, all habitat modification eastern semiarid areas), pollution by cattle and reducing hydroperiod in breeding areas affects domestic sources (mainly in Atlantic plains) via survival of amphibian populations. processes very similar to those which have Annual rainfall influences distribution and occurred in southern Spain (GREEN et al., 2002; abundance (BORKIN, 1999) and in Morocco REQUES, 2009) are common, yet only 10 of 47 annual average rainfall is approximately 400 wetlands with areas greater than two hectares mm for most of the country, increasing to surveyed between 1997 and 1999 by GREEN et al. between 1500 mm and 2000 mm in northern (2002) have been provided legal protection. mountains and decreasing to about 50 mm in Alteration or loss of pools, streams, and the Sahara. Data on rainfall were obtained gullies suitable for amphibian reproduction from GLOBCOVER (2008; full resolution frequently goes unnoticed because of the small mode, spatial resolution of 300 m), amphib- size and temporal character of these areas ian distributional data were obtained from the (OERTLI et al., 2005; REQUES, 2009) yet tem- Global Amphibian Assessment (IUCN, porary wetlands with extended hydroperiods 2006) and both datasets were entered into facilitate breeding success for most amphibians ArcGIS 9.2® (ENVIRONMENTAL SYSTEMS (PECHMANN et al., 1989; ROWE & DUNSON, RESEARCH INSTITUTE, 2006) to obtain precip- 1995; WEYRAUCH & GRUBB, 2004). itation ranges and means for distributional Permanent wetland habitat is threatened by areas of the 14 species in Morocco. As the frequent introduction of non-native species boundaries between D. pictus and D. scovazzi (mainly fish) that feed upon amphibian eggs are currently not clear (see Fig. 9 in BEUKEMA et al. 2013), we deal with this uncertainty by considering all populations of painted frogs from Cap des Trois Fourches Peninsula east to Algeria (including the Beni Snassen) as D. pic- tus. Most species are distributed throughout northern and western Morocco in localities where annual precipitation ranges between 400 mm and 1000 mm. Only two species, H. occipitalis and A. xeros in the extreme south, are found in localities with an annual rainfall of less than 150 mm. Together with A. mauritan- Figure 4: Amphibian species richness in relation icus, B. boulengeri, B. brongersmai, and P. sahar- to rainfall classes. See text for data collection pro- icus, these species represent the only Moroccan tocol. species inhabiting the desert. At the opposite STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 35 extreme, S. algira and A. maurus inhabit areas with an annual precipitation greater than 600 mm (DONAIRE-BARROSO & BOGAERTS, 2003b) (Table 3, Fig. 4). With the exception of P. saharicus and most populations of P. waltl, Moroccan amphibians can be considered terrestrial, and that is significant for conservation. Little is known about microhabitat use by most Moroccan amphibians (but see EL HAMOUMI et al., 2007; DE POUS et al., 2012; BEUKEMA Figure 5: Correspondence between amphibian dis- et al., 2013) and any analysis must be per- tribution and habitat type. Rc = Rainfed croplands; formed within the framework of major habi- Ba =Bare areas; Mo1 = Mosaic cropland, 50-70% tats. Landscape ecology can highlight rela- vegetation/20-50% grassland-shrubland-forest; tionships among species and landscapes, fol- Mo2 = Mosaic vegetation, 50-70% grassland- low shifts in relationships through time, and shrubland-forest/20-50% cropland; Co: Closed to open, >15% broadleaved or needle-leaved evergreen monitor the influence of anthropogenic or deciduous, <5m shrubland; Sp = Sparse, <15% changes in the landscape (NAUGLE et al., vegetation; Ot= Other. Data from GlobCover 2005). Data on the distributional area of dataset (GLOBCOVER, 2008). Alymau = Alytes major habitat types within the range of each maurus; Amimau = Amietophrynus mauritanicus; species were obtained from GLOBCOVER Amixe = Amietophrynus xeros; Barbro = Barbarophryne (2008; see above), and the frequency of habi- brongersmai; Bufspi = Bufo spinosus; Bufbou = Bufotes boulengeri; Dispic = Discoglossus pictus; Dissco = tat use for each amphibian species analyzed Discoglossus scovazzi; Hopocc = Hoplobatrachus occip- by correspondence analysis (STATSOFT, italis; Hylmer = Hyla meridionalis; Pelvar = Pelobates 2001), which allowed assessment of relation- varaldii; Pelsah = Pelophylax saharicus; Plewal = Pleurodeles ships between landscape variables and waltl; Salalg = Salamandra algira. amphibian species (PÉREZ-LÓPEZ, 2005) (Fig. 5). Hoplobatrachus occipitalis and A. xeros, associated with barren areas typical of desert PROSPECTS FOR CONSERVATION regions, were again identified as isolated from the remaining species. At the opposite Synthesis of Threats to Amphibian Assemblages extreme were A. maurus and S. algira, both of which occur in forested or shrubland Survival problems faced by Moroccan areas of northern Morocco. Other associa- amphibians can be attributed to environmen- tions of conservation interest were those of tal change of anthropogenic origin (IUCN, P. varaldii with P. w a l t l , and of D. scovazzi 2009); habitat loss is the major threat (Fig. 6). and/or D. pictus with H. meridionalis. The negative effect of human replacement of Anurans of the genera Bufo, Bufotes, natural habitats with agricultural, urban, or Barbarophryne, and Pelophylax utilized the industrial sites involves loss both of suitable landscape in an eclectic manner. habitat for breeding and of connectivity REQUES 36 ET AL.

ments that modify natural water sources. Since construction of the Ouarzazate reservoir on the Oued Drâa, regular flooding ceased and oases that formerly provided suitable breeding places for A. mauritanicus, B. brongersmai, H. merid- ionalis, and B. boulengeri disappeared. Due to the high number of species affected, four of them threatened, intrinsic factors of the species should be considered the second major threat Figure 6: Major threats to Moroccan amphibians (from PLEGUEZUELOS et al., 2010). to Moroccan amphibians (Fig. 6). Restricted distributional area and low population densi- between patches of suitable habitat ty place Moroccan populations of S. algira, (LAURANCE, 2001; STUART et al., 2008); the P. waltl, A. maurus, P. varaldii, A. xeros, B. spinosus, last is an increasing threat in Morocco, and H. occipitalis, all species with reduced disper- because of the spreading network of paved sal capacity, in very sensitive situations. roads. These threats are particularly harmful Amphibians cannot escape natural effects to species with restricted distributions or resulting from climatic change (PARMESAN et ecological requirements, such as S. algira, al., 1999; CAREY & ALEXANDER, 2003) as P. varaldii, and A. maurus. Some species, changes in temperature and humidity affect however, do benefit from anthropogenic physiology and phenology of the reproductive change; deforestation increases open habitat process (WALTHER et al., 2002; ROOT et al., that may be rapidly colonized by generalist 2003; READING, 2007) in addition to pro- species such as A. mauritanicus and H. meridionalis. moting loss and fragmentation of habitat Anurans also adapt well to agricultural land- (TEWKSBURY et al., 2008). Earlier estimates scapes if habitat change is not extreme; new that at least four Endangered species and 13 and modified habitats can provide artificial, Vulnerable species within Africa, most inhab- but suitable, breeding habitat in the form of iting the northern half of the continent, irrigation ponds for P. saharicus and A. mau- would be affected by climatic change (IPCC, ritanicus (STUART et al., 2004). Currently 2007) have been reinforced by MARTÍNEZ- only four amphibian species, A. mauritanicus, FREIRÍA et al. (2013) who predict a reduction B. boulengeri, P. saharicus and, to a lesser in suitable area for 50% of Moroccan reptil- extent, H. meridionalis (because of overall ian species in the future, suggesting a serious resilience) exhibit relatively high tolerance to potential threat to amphibians as well. human-induced change to the landscape Natural disasters like the current drought (deforestation and agriculture in particular). affecting Western Sahara (BENASSI, 2008) Other recent threats include loss of suitable could hamper survival of A. xeros and H. occipi- coastal habitat due to increasing development talis, species living in desert habitats at the for tourism (affecting mainly P. varaldii along southern extreme of Western Sahara. the Atlantic and Discoglossus species along the Batrachochytrium dendrobatidis, the glob- Mediterranean), and spread of other develop- ally distributed fungus responsible for STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 37 chytridiomycosis and mass mortality among ative (BOONE & BRIDGES, 2003), different amphibians worldwide (HEATWOLE, 2013), species do respond differently (BRIDGES & has recently been reported in populations of SEMLITSCH, 2000). Low pollution levels can P. varaldii, D. scovazzi, and H. meridionalis increase trophic resources (BOONE & JAMES, from Tétouan Préfecture. While opportunis- 2005), and the relative abundance of some tically sampling 51 sites on the Gharb plain generalist species (P. saharicus, A. mauritanicus, and in the Rif and Middle Atlas mountains and H. meridionalis) residing on the out- between 2005 and 2009, EL MOUDEN et al. skirts of urban areas, such as Chefchaouen, (2011) examined 203 amphibian specimens Ifrane, Goulimine, and Ouarzazate (FAHD for presence of chytrid fungus. Three sites et al., 2006, 2007) may be artificially high (6%; all north of latitude 35.038°N) proved because of pollution. Species such as S. algira, positive for chytrid fungus. While sample however, cannot tolerate this kind of pollution sizes for individuals determined to carry the (DONAIRE-BARROSO & BOGAERTS, 2003a). fungus were low (one P. varaldii tadpole of Non-native species in aquatic ecosystems 10 sampled, one adult H. meridionalis of can pose serious threats to native amphibian four sampled, and two D. scovazzi [a meta- populations (MOUSLIH, 1987; KATS & morph and an adult] from each of two sites; FERRER, 2003; KIESECKER, 2003). Mosquito see IUCN Red List Status, above) and infec- fish (G. holbrooki), for example, are known to tion rates cannot be accurately determined be detrimental to amphibian breeding success from the data obtained, the presence of this (SEGEV et al., 2009) and recent introductions fungus in northern Morocco introduces of mosquito fish into numerous montane springs another serious threat to the survival of may compromise populations of A. maurus and native amphibians. other species (DONAIRE-BARROSO & BOGAERTS, When ranked by number of species affected 2003b; BEUKEMA et al., 2013). Discovery of by each major threat to Moroccan amphibians, New World crayfish Procambarus cf lophotus pollution placed third (Fig. 6). Chemical pol- (DE POUS et al., 2012) suggests impending lution of freshwater has been recognized as a threats to aquatic amphibians in this area as serious global threat to amphibian popula- well. “Once established, no methods exist by tions (BRIDGES & SEMLITSCH, 2000; which a non-indigenous crayfish can be SPARLING et al., 2000) but while there are exterminated without unacceptable harm to data concerning the effect of some pesticides native crayfishes and other organisms” on the Northern Bald Ibis (Geronticus eremita; (LODGE et al., 2000). ARMESTO et al., 2006) in Morocco, data are Road-kills may cause local extirpation of lacking on the effect of pesticides upon populations when traffic levels in proximity amphibians. Pollutants in favourable breed- to breeding areas are high (DODD & SMITH, ing habitat in Morocco are generally nitroge- 2003; HOULAHAN et al., 2006). There are no nous derivatives from agricultural fertilizer, data on such an impact on Moroccan but urban and cattle waste also contribute. amphibians, but the threat from traffic is pre- Although the relationship between pollutant sumed to be increasing because the number level and amphibian richness is generally neg- of roads and vehicles continues to increase. REQUES 38 ET AL.

An increasing threat to amphibian assem- blages in arid areas of the south (Souss Valley, Ifni, Low Draa, Tekna) is the building of more and more reinforced concrete cisterns by shep- herds. These deep, underground and covered structures with one or two uncovered decanta- tion chambers positioned at the bottom of lightly sloping valleys accumulate humidity and attract amphibians. Individuals fall into decantation chambers or cisterns, both of which serve as death-traps (e.g. GARCÍA-MUÑOZ et al., 2009). Although some B. brongersmai may Figure 7: Priority areas (shaded) for amphibian con- breed in decantation chambers, others are sub- servation (see text). jected to predation or to dehydration as these structures become empty (L. García-Cardenete, Protection of places of interest, with amphibians personal communication). This increasing acting as umbrella species, would conserve other threat could rapidly deplete Saharan popula- vertebrates as well (RONDININI & BOITANI, tions of A. mauritanicus, B. brongersmai and, 2006). Using models for 11 amphibian and 86 particularly, B. boulengeri, in regions where they reptilian species, DE POUS et al. (2011) defined currently are scarce because of arid conditions. priority areas into which the existing protected This hazard can be corrected easily by installa- areas could be extended. tion of evacuation slides. Amphibians occur in most major habitats in Morocco, and threatened species are notably Priority Areas present in wetlands as expected, but also in shrublands, savannahs, and deserts (Fig. 1). The Mediterranean Basin is considered a Threatened Moroccan amphibians have very global hotspot for amphibian diversity and restricted ranges, but ranges of the most threat- endemism (STUART et al., 2008). ened species include portions of the ranges of Biogeographically, this eco-region includes four most other remaining species. Well-designed mountain chains and the northern half of the conservation area networks (CAN) will include Moroccan lowlands and, as with other most threatened species and much of Mediterranean areas, Moroccan habitats are Morocco’s amphibian species richness (Fig. 7; highly degraded and scarcely protected (BROOKS see also DE POUS et al., 2011). In northern et al. 2004; WILSON et al., 2007; DE POUS et al., Morocco, for example, protection of the entire 2011). Gap analysis of diversity and distribution range of A. maurus will protect eight addition- of amphibians, mammals, and protected areas in al species; DE POUS et al. (2011) showed that Africa illustrated the convenience of broadening only about 7% of the distribution of A. maurus protected areas around the western Rif, western falls inside protected areas (e.g. National and Middle Atlas, and some localities along the Natural Parks of Jebel Bou Hachem, Bou Atlantic coast (RONDININI et al., 2005, 2006). Iblane, Talassemtane, Koudiat Tidighine, Al STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 39

Jabha Tazekka), although remaining popula- does not occur in any protected area (DE tions are in areas so rugged that alteration is not POUS et al., 2011, 2012). The Mamora cork imminent (DONAIRE-BARROSO & BOGAERTS, oak forest and surrounding area is the only 2003b). In western Morocco, protection of the locality where it is still common, and here it range of P. varaldii would include partial ranges is highly threatened by overgrazing, logging, of six additional species. This CAN would not exotic tree plantations, and industrial devel- include B. brongersmai, A. xeros, or H. occipitalis; opment. Reduction of these negative effects conservation of the latter two species requires and avoidance of further fragmentation management action in the southern extreme of (DE POUS et al., 2012) is essential, while Western Sahara, areas currently suffering establishing protected areas within this part extreme drought and political problems. of the range is urgent. Such action would also Expansion of areas under protection benefit P. waltl, D. scovazzi, H. meridionalis, would benefit A. maurus and other species B. boulengeri, A. mauritanicus, and P. saharicus. of concern in Morocco, e.g. S. algira and The CAN in Morocco does not currently B. spinosus. Because P. varaldii has not been include all amphibian species, and a more com- confirmed within limits of the Merja Zerga plete network including remaining species is Biological Reservation recently, it apparently advisable (DE POUS et al., 2011). Some threat-

Table 4: Characteristics of regions having priority for amphibian conservation. For approximate regional areas, currently-protected areas, and percentages of major habitats within regions, see GLOBCOVER (2008) and text for details. Cost.A.R. = Coastal Atlantic Region; R.A.M.R. = Rif-Middle Atlas Mountain Region; Cent.A.R. = Central Atlantic Region; T.R. = Tiris Region.

Cost.A.R. R.A.M.R. Cent.A.R. T.R.

Approximate regional area (km2) 5782 15 608 15 416 > 211 576 Currently-Protected areas Merja Zerga, Talassantane, Souss-Massa None Sidi Boughaba Tazekka Rainfed cropland 15.8 6.9 13.1 0 Mosaic cropland (50-70%) / other vegetation (grass- 36.3 24.0 16.0 0 land / shrubland / forest) (20-50%) Mosaic vegetation (grassland / shrubland / forest) 25.7 23.0 26.2 0 (50-70%) / cropland (20-50%) Closed broadleafed deciduous forest (> 40% of the 0.7 6.0 0 0 area with trees > 5m in height) Mosaic forest or shrubland (50-70%) / grassland 4.4 10.6 8.4 0 (20-50%) Closed to open (> 15%) (broad-leafed or needle-leafed, 11.7 19.3 4.6 0 evergreen or deciduous) / shrubland (< 5 m) Sparse (< 15% cover) vegetation 2.4 4.3 20.7 100 Bare areas 0.6 2.2 9.4 0 REQUES 40 ET AL. ened species are found in desert and savannah abandoned quarries, or deforested areas for but neither habitat type is well represented in example, may become suitable habitat for some the current CAN. Areas of high priority for amphibian species. protection are those in which historical eco- Besides foreseeable change in land use and logical processes continue to function and in pollutant levels, consequences from climatic which climatic changes forecasted for the change must also be considered. The future may be mitigated (PARRISH et al., 2003; Intergovernmental Panel on Climate Change LOVEJOY, 2006; DE POUS et al., 2011). It is (IPCC) predicts Morocco will be severely proposed here that the northwestern affected by an increase in temperature and a Atlantic, Rif-Middle Atlas, central Atlantic, decrease in rainfall in the near future (HULME et and Tiris regions be considered as priorities al., 2001; IPCC, 2007; KLAUSMEYER & SHAW, for amphibian conservation (Fig. 7; see also 2009; LOARIE et al., 2009; MARTÍNEZ-FREIRÍA RONDININI et al., 2005, 2006; CARVALHO et al., et al., 2013). A 15% decrease in average annual 2011; DE POUS et al., 2011). Currently pro- rainfall has been recorded in Morocco during tected areas in the first three regions should the past 30 years, with periods of drought be enlarged (Table 4) and protection should increasing in duration and intensity (BENASSI, be established for the Tiris region. Each of 2008). Following this scenario, Mediterranean these regions contains habitat meeting eco- wetlands will face altered hydroperiods, and logical requirements (GLOBCOVER, 2008; see wetlands currently permanent will become sea- above) for the survival of Moroccan amphib- sonal while wetlands currently temporal will ians (Table 4). disappear (REQUES, 2005; IPCC, 2007). Shortening of the hydroperiod would affect NEW ECOLOGICAL STAGES AFFECTING species like P. varaldii, a threatened species with MOROCCAN AMPHIBIANS a relatively long larval period. It would also have negative consequences for other species through Morocco will achieve significant human population and community effects population increase and economic ameliora- (SCHNEIDER, 1997; MOREY, 1998). Scarcity of tion in the near future (AFRICAN DEVELOPMENT freshwater would hardly affect availability of BANK, 2012), both of which likely will have water for humans (IPCC, 2007); competition negative effects on natural ecosystems. for available water will increase between Predicting how these changes will affect the humans and amphibians and the loser in this amphibian fauna is difficult. One cannot rely competition will always be the non-humans. upon modelling because time-series data for Many aspects of the biology and ecology of these effects on amphibian populations are not Moroccan amphibians remain unknown available, and one can only hypothesize future (BEUKEMA et al., 2013); to foresee climatic and relationships between environmental change anthropogenic effects of global change upon and amphibian populations. Knowledge of the amphibians it is necessary to understand the ability of Moroccan amphibians to adapt to, natural history and tolerance of each species to and survive, human-induced environmental environmental change. Amphibian decline change is also lacking; some degraded habitats, occurring on the southern Iberian Peninsula STATUS OF MOROCCAN AND WESTERN SAHARAN AMPHIBIANS 41 because of increasing aridity (unpublished REFERENCES data) is probably very similar to that occurring in Morocco. Effective measures in Spain AELLEN, V. (1951). 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