Alytidae Fitzinger

BEUKEMA, WOUTER, DE POUS, PHILIP, DONAIRE-BARROSO, DAVID, BOGAERTS, SERGÉ, GARCIA-PORTA, JOAN, ESCORIZA, DANIEL, ARRIBAS, OSCAR J., MOUDEN, EL HASSAN EL & CARRANZA, SALVADOR, 2013, 3661, Zootaxa 3661 (1), pp. 1-60 : 31-33

Family Alytidae Fitzinger

Moroccan Midwife Toad, Alytes (Baleaphryne) maurus Pasteur and Bons 1962

Figs. 11A–C .

Background information. A few decades after the initial discovery of the genus Alytes in Morocco by Galan (1931), Pasteur and Bons (1962) attributed the African populations to Alytes obstetricans maurus based on divergent tadpole morphology. In contrast, Arntzen and Szymura (1984) suggested Moroccan populations to be of anthropogenic origin and identical with Alytes obstetricans ( Laurenti 1768) , based on minimal recovered electrophoretic divergence in respect to Iberian and Western European populations. The position of A. maurus within Alytes would remain obscure for some time. Concurrently however, the discovery of extant Alytes muletensis ( Sanchíz & Adrover 1979 “1977”) on the Balearic island of Mallorca ( Mayol et al. 1980) led to a systematic rearrangement of the genus ( Dubois 1987 “1986”). While the subgenus Baleaphryne was initially proposed solely for A. muletensis, Donaire-Barroso and Bogaerts (2003b) elevated A. maurus to species level and placed the taxon in Baleaphryne Sanchíz & Adrover, 1979 “1977”. The validity of the subgenus, and confirmation of the correct attribution of its members, was subsequently provided by means of osteological, mitochondrial and nuclear evidence ( Martínez-Solano et al. 2004; Gonçalves et al. 2007), albeit not explicitly. According to the most recent phylogenetic analyses ( Gonçalves et al. 2007), A. maurus forms a trichotomy with the Iberian Alytes dickhilleni Arntzen and García-París 1995 and A. muletensis . A more recent study including several nuclear genes places A. maurus with high support as sister to A. dickhilleni , while A. mulentensis is shown to be sister to the A. maurus - A. dickhilleni clade (M. Vences pers. comm.). The collapse of the Gibraltar land bridge at the end of the Miocene has most likely led to the rapid diversification of the Baleaphryne members, separating the ancestor of A. maurus on the African continent ( Martínez-Solano et al. 2004). The current, fragmented distribution range of A. maurus is likely a result of increasing temperatures throughout the recent Quarternary, as a fossil record of a Baleaphryne sp. from the Upper Pleistocene in the arid Jbilets northwest of Marrakech ( Hossini 2001) suggest a much wider historical distribution.

Bioacoustics. Márquez et al. (2011) recently described the advertisement call of male individuals recorded in the Middle Atlas Mountains. Males were encountered calling at dusk and during rainy nights, located beneath cover objects or out in the open. Pulse duration ranges between 77–217 ms (mean 97.3 ms), while emitted at intervals of several seconds. To the ear, the call resembles a short tonal burst of sound with a very brief rise time and a longer fall time. While the advertisement call of A. maurus is significantly shorter in duration than that of A. dickhilleni , it falls within intraspecific variation of A. obstetricans .

Tadpole. Pasteur and Bons (1962) gave a comprehensive description of the tadpole of A. maurus ( Fig. 12A ). The following account is a summary of the original description. Fully developed tadpoles are characterized by their large size (up to 90 mm), midventral spiracle and spotted tail. Eyes positioned dorsally on head. Interorbital space at least twice as wide as the internarial space. Oral disc ventral, significantly wider than interorbital space. Two tooth rows on the upper labium and three on the lower. Upper supralabial row composed of two series, lower supralabial row and all infralabial rows composed of at least three series. Upper infralabial row with very narrow median gap. Spiracle midventral. Colour grey, brown or olive with conspicuous pattern of smaller and larger golden and dark patches and spots.

Natural history. Available natural history information on this species has been presented and summarized by Donaire-Barroso and Bogaerts (2003b) and Donaire-Barroso et al. (2006). Alytes maurus is commonly found in open forests, agricultural terraces bordered by stone walls and open, rocky valleys in the vicinity of streams or water sources. Dominant vegetation consists mainly of Quercus sp. , Juniperus sp. and Olea europaea . In in the Middle Atlas populations are additionally encountered in Cedrus atlantica forest ( Donaire-Barroso et al. 2006). Occurrence in high-altitude mountain meadows has occasionally been noted ( Harris et al. 2008). Habitats are often dominated by an undergrowth of maquis and high abundance of rocks and (karstic limestone) rocky outcrops. Near Chefchaouen in the western Rif Mountains advertisement calls have been heard from February until April, while larvae close to metamorphosis were observed in August ( Donaire-Barroso & Bogaerts 2003b). In the Middle Atlas (Taza), advanced-staged larvae were encountered in August and September (D. Donaire-Barroso pers. obs.). It is not uncommon for larvae to grow to large sizes and hibernate in permanent streams. Indeed, larvae have also been observed during January and February ( Donaire-Barroso & Bogaerts 2003b). Populations have been encountered from 200 up to 2142 m a.s.l. ( Donaire-Barroso et al. 2006), although most occur at intermediate altitudes (Donaire- Barroso & Bogaerts 2003b). Sympatric occurrence with larvae-bearing populations of S. a. tingitana and S. a. splendens ssp. nov. is common.

Distribution. Donaire-Barroso and Bogaerts (2003b) and Donaire-Barroso et al. (2006) reviewed the distribution of this species in the Rif Mountains and Middle Atlas Mountains, respectively. Alytes maurus is commonly found in brook valleys of the western- and central Rif Mountains and several small peripheral ranges. Presence in the Middle Atlas Mountains seems to be limited to the Tazekka and Bou Iblane Massifs. Despite prior mentions of A. maurus north of the Oued Martil (e.g. in Ceuta, Bons & Geniez 1996), such records have not been confirmed by subsequent inventories of the area (e.g. Martínez-Medina 2001; Donaire-Barroso & Bogaerts 2003b) and are consequently not displayed on the map. The distribution map ( Fig. 9C ) includes records from Bons and Geniez (1996), Donaire-Barroso and Bogaerts (2003b), Donaire-Barroso et al. (2006), Fahd and Mediani (2007), Fahd et al. (2007) and Harris et al. (2008). Newly discovered localities largely fill in gaps within the known distribution range. The niche model ( Fig. 9C ) shows potential suitable areas in the Middle Atlas and east of the Moulouya Basin, which are unoccupied due to major geographical barriers separating these from the realized distribution.

National Red List Status. Near Threatened.

Figures

FIGURE 9. Niche models and distribution records of Salamandra algira (A); Pleurodeles waltl (B); Alytes maurus (C); and Discoglossus spp. (D) in Morocco. Warmer colours indicate higher climate suitability. Dark spots indicate records derived from literature, white spots show new records.

FIGURE 11. Members of the Alytidae and Discoglossidae families native to Morocco. A-C: Alytes maurus; A–B: Tazekka (Gabriel Martínez del Mármol); C: Chefchaouen (SB). D-G: Discoglossus scovazzi; D–E: Jbel Musa (WB); F: Oualidia (WB). I-J: Discoglossus pictus; G: Beni Snassen (DE); H: Saidia (SB); I: Beni Snassen (SB).

FIGURE 12. Tadpoles of Moroccan anurans. A: Alytes maurus, Ketama. B: Discoglossus scovazzi, Safi. C: Discoglossus pictus, Tafuyalt (Algeria). D: Amietophrynus mauritanicus, Mohammedia. E: Barbarophryne brongersmai, Smimou. F: Bufo bufo, Tazekka. G: Bufotes boulengeri, Dayet Aoua. H: Hyla meridionalis, Mohammedia. I: Pelobates varaldii, Kenitra. J: Pelophylax saharicus, Sidi Ifni. All drawings made by DE.