Salamandra algira tingitana Donaire-Barroso and Bogaerts 2003

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 : 24-25

Salamandra algira tingitana Donaire-Barroso and Bogaerts 2003

Figs. 8B–E .

Background information. Information about S. algira remained scarce for more than a century after the initial description of Bedriaga (1883) which was based on individuals from north-eastern Algeria. Both Bedriaga (1883)

and Pasteur and Bons (1959) reported S. algira to be larvae-bearing. Hence, observations of populations which deposit fully-developed juveniles among the limestone outcrops in extreme north-western Morocco (Donaire- Barroso & Bogaerts 2001; Donaire-Barroso et al. 2001) were unexpected. Subsequent research showed that populations from northwestern Morocco consistently lack red discoloration while often showing white specks or rosettes on the underside ( Fig. 10A&B ). The northernmost populations were shown to possess a high tendency towards hypoluteism and melanism (Bogaerts & Donaire-Barroso 2003). As a result of the aforementioned characteristics, Donaire-Barroso and Bogaerts (2003a) described S. a. tingitana (holotype: MNCN 41037; type locality: " 500 m altitude on Jabal Muse (= Jabal Mousa) north Morocco "). Recent studies have shown S. a. tingitana to possess a more extended distribution than initially thought, while mtDNA studies showed the taxon to consist of three clades which diverged during the Pleistocene ( Beukema et al. 2010). Only the northernmost clade deposits fully-developed juveniles, while populations located in the triangle formed by Tetouan, Ksar el-Kébir and Chefchaouen deposit larvae. The boundary between these is approximately located near the Oued Martil, with larvae occasionally having been found west of Tetouan (D. Donaire-Barroso pers. obs.), the southernmost population bearing fully-developed juveniles according to Beukema et al. (2010).

Natural history. Across the largely Mediterranean-influenced region located between the Strait of Gibraltar and Tetouan, S. a. tingitana inhabits chiefly limestone outcrops from sea level up to 390 m. Both barren eroded mountain summits (e.g. the type locality, Jbel Musa) as well as hilly terrain covered by maquis, Pinus and Quercus trees have been recorded as habitat ( Donaire-Barroso & Bogaerts 2003a). Populations recorded in this area deposit fully-developed juveniles, even though temporary surface water is available at multiple sites ( Donaire-Barroso & Bogaerts 2001; Donaire-Barroso et al. 2001; Donaire-Barroso & Bogaerts 2003a). On rare occasions, either a single or small number of larvae can be deposited in these water bodies during periods of abundant rainfall ( Martínez-Medina 2001; Donaire-Barroso & Bogaerts 2003a). The activity period ranges from October to March, throughout which individuals are exclusively nocturnal, and only emerge from their shelters during humid or rainy weather ( Martínez-Medina 2001; Donaire-Barroso & Bogaerts 2003a). Low temperatures (<10º C) and presence of considerable wind does not have a negative influence on activity in general, in contrast to most European Salamandra populations ( Thiesmeier & Günther 1996; Thiesmeier 2004; pers. obs. D. Donaire-Barroso & S. Bogaerts). During the day, individuals hide under superficial shelters such as rocks in humid weather throughout the activity period, while they retreat deep into karstic crevices from April to September ( Donaire-Barroso & Bogaerts 2003a). Up to 17 fully developed juveniles can be deposited between October to March (Donaire-Barroso et al. 2001).

Southwards, the occurrence is limited to mountainous or hilly terrain between Tetouan, Ksar el-Kébir and Chefchaouen up to at least 1274 m (Fôret de Bouachem), where populations generally occur close to springs or brooks in (half-open) forests, agricultural terraces and limestone outcrops (e.g. Donaire-Barroso & Bogaerts 2003b). General activity of these larvae-bearing populations is likely similar to that of the populations which deposit fully-developed juveniles, although the former might benefit from a more humid, Atlantic climate among the western mountain ranges of the Tingitana Peninsula ( Beukema et al. 2010). Larvae have been encountered in streams, springs and occasionally in temporary ponds. Donaire-Barroso and Bogaerts (2003b) encountered larvae at 700 m a.s.l. approximately 30 km south-west of Tetouan at the end of February, while D. Donaire-Barroso and W. Beukema (pers. obs.) observed recently metamorphosed juveniles with gill remnants at 400 m a.s.l. north-east of Ksar el-Kebir during the same period.

Distribution. The distribution map ( Fig. 9A ) is composed of records from Bons and Geniez (1996), Donaire- Barroso and Bogaerts (2003a) and Beukema et al. (2010). While most records fill in gaps within the continuous mountain ranges, several are located at the periphery of the distribution area, suggesting a wider range during the Pleistocene. An earlier suggested, isolated occurrence in the Ahl Serif northeast of Ksar-el-Kebir based on predictive species distribution modelling by Beukema et al. (2010) has been confirmed. As the niche model on Fig. 9A is composed of climatic data only (as opposed to that shown in Beukema et al. 2010), it is likely overpredicting the distribution of S. algira in Morocco. See Fig. 10 for the distribution of S. a. tingitana in relation to other Moroccan Salamandra algira ssp.

National Red List Status. Vulnerable (as part of S. algira ).

Taxonomic comment. The original description of S. a. tingitana by Donaire-Barroso and Bogaerts (2003a) appeared in Pod@rcis, an online bilingual herpetological journal which has irregularly appeared since 2000. As such, this publication appeared well before a recent amendment of the International Code of Zoological Nomenclature (ICZN) which permits electronic publication of new scientific names and nomenclatural acts (ICZN 2012). However, as detailed in the colophon of each Pod@rcis issue, copies of published papers were deposited on paper and CD-ROM at the PNMH, RMNH, Koninklijke Bibliotheek (The Hague, Netherlands), ZMA and ZMFK. Pod@rcis therefore meets the requirements of Article 8.6 of the ICZN, representing a published work.

Figures

FIGURE 8. Members of the Salamandridae family native to Morocco. A: Salamandra algira spelaea Beni Snassen (DE). B-D: Salamandra algira tingitana; B: Jbel Musa (PdP); C: Tagramt (SB); D: Cudia Adru (DD). E-H: Salamandra algira splendens ssp. nov.; E: Bou Iblane (DD); F: Chefchaouen (DD) (holotype specimen); G: Bab Berret (DE); H: Ketama (DD). I-K: Pleurodeles waltl; I: Forêt de Mamora (WB); J: Casablanca (Gabriel Martínez del Mármol); K: Moulay Abdeslam (SB).

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 10. Upper panel: distribution of Salamandra algira ssp. in Morocco, indicating identified geographical barriers. Lower panel: variation in ventral pattern and coloration among Moroccan Salamandra algira ssp. A: Salamandra algira tingitana, west of Tetouan; B: Salamandra algira tingitana, Moulay Abdeslam; C: Salamandra algira splendens ssp. nov., Bou Iblane; D: Salamandra algira splendens ssp. nov., Ketama.