Salamandra algira splendens Beukema, de Pous, Donaire-Barroso, Bogaerts, Garcia-Porta, Escoriza, Arribas, El Mouden & Carranza

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 : 26-29

Salamandra algira splendens Beukema, de Pous, Donaire-Barroso, Bogaerts, Garcia-Porta, Escoriza, Arribas, El Mouden & Carranza , ssp. nov.

Figs. 8F–H

Salamandra algira ssp.—Steinfartz et al. 2000: p. 410.

Salamandra algira algira — Donaire-Barroso and Bogaerts 2003a: p. 97.

Salamandra sp. — Dubois and Raffaëlli 2009: p. 35.

Holotype. RMNH 40173 , an adult male collected at Aïn Tissimilan, Jebel el Kelaâ, Chefchaouen, western Rif Mountains , Morocco (N 35°10.5, W 5°14.6, 700 m a.s.l.) in November 1996 by David Donaire-Barroso and César Barrio ( Fig. 8F ).

Paratypes. EBD 29787, an adult female, collected at N 35°10.5, W 5°14.6, Aïn Tissimilan, Jebel el Kelaâ, Chefchaouen, western Rif Mountains in November 1996 by David Donaire-Barroso and César Barrio . MCNC 2010-0136 , a sub adult female collected at N 35º02.1, W 5º01.5, 1445 m a.s.l., Bab Berret , western Rif Mountains , Morocco in December 2009 by Daniel Escoriza and Félix Amat ( Fig. 8G ) . MCNC 2010-0128 , 2010-0129 , two adult males collected at N 34º85.7, W 2°21.0 and 1450 m a.s.l., Jebel Tazekka , Middle Atlas Mountains , Morocco in December 2009 by Daniel Escoriza and Félix Amat .

Etymology. The present participle splendens (Latin, from splendere = to shine) refers to the aesthetically pleasing appearance of this taxon, which is manifested in its pattern of bright yellow patches and red discolorations on a black background.

Diagnosis. A slender, large-sized (215 mm total length) North African member of the genus Salamandra having the combination of an elongated flat head, 1½ times longer than wide; long well separated limbs, fingers and tail; black coloration with rounded or elongated yellow markings and red discoloration; larvae-depositing. Salamandra algira splendens ssp. nov. can be distinguished from all other North African Salamandra taxa by a combination of colour pattern and distribution. Salamandra algira algira never possesses red discoloration and is restricted to the eastern Algerian Tell Atlas, comprising the Kabylies and the Annaba Peninsula ( Algeria). Salamandra algira tingitana similarly never possesses red discoloration and is found exclusively north of the Oued Laou. Northern populations of S. a. tingitana are characterized by hypoluteism ( Fig. 8B, C ) and bear fullydeveloped young, whereas larvae-bearing is the rule in S. a. splendens ssp. nov. Salamandra algira spelaea possesses only few and small red specks distributed across the head and body which do not extend to the degree observed in S. a. splendens ssp. nov. when present in the latter. Additionally, the former taxon is found exclusively in the Beni Snassen Massif east of the Moulouya Basin.

Description of the holotype. Habitus slender. Head flat, longer than wide. Snout short, slightly truncate in dorsal view, rounded in profile while exceeding beyond lower jaw. Parotoids large and elongated, with glandular pores, pointing slightly backwards. Double dorsolateral gland row from the posterior of the head to the basis of the hind legs. Skin smooth. Gular fold present. Limbs long and clearly separated from the body; fingers and toes long. Four fingers (fingers relative length 3>2>4>1) and five toes (toes relative length 4>3>2>5>1), all without webbing. Tail long, laterally slightly compressed, decreasing in width towards the tip. Measurements (in mm): SVL 94.0, LT 86.0, AL 31.0, PL 33.6, HL 25.0, HW 15.3 and PAL 13.2.

Colour of holotype in life. Body black with six elongated bright yellow patches on the dorsum. Parotoids and eyes covered by yellow patches, bordered by red discoloration. Yellow patches with red discolouration present at the base of each limb. Additional small red discolorations are present on the borders of the mouth, limbs, tail and gular region.

Colour of holotype in preservative. Body black. Yellow patches appear whitish. Red discoloration faded to dark brown, with smaller patches being difficult to distinguish.

Variation. Measurements of the type series are summarized in Table 2. All paratypes show black body coloration. Yellow patches are without exception present on the eyes, parotoids and at the basis of each limb. Yellow patches on the dorsum and tail show random, variable patterns but are always rounded, occasionally elongated. Of the four paratypes, three possess red discoloration on the head, tail and gular region. In addition to the paratype series, a total of 12 adult individuals (8 males and 4 females) from National Park Tazekka were measured in the field during mid-December 2012 by S. Bogaerts and W. Beukema and subsequently released in order to document morphological variation. All of these individuals showed red discolorations on the head, surrounding the paratoids. Eleven individuals showed red discoloration on the gular region (see also Fig. 10C ), nine on the tails, eight on the legs and one on the ventral side (see also Fig. 10D ). Eight individuals show few (<10) tiny, irregularly-shaped whitish specks on the gular region; in two of these, the specks are present in a higher number while few are also present on the venter. Two males show glandular skin protuberances.

Natural history. Salamandra algira splendens ssp. nov. has been found between 600–2000 m a.s.l. in Middle Atlas Mountains, and between 280–1700 m a.s.l. in the western Rif Mountains. Within this range, the distribution is largely limited to forests characterized by Abies maroccana , Cedrus atlantica , Pinus sp. and Quercus sp. or open karstic limestone formations ( Donaire-Barroso & Bogaerts 2003b; Martínez-Medina 2007). At lower elevations populations can occur in shrub land chiefly composed of Pistacia sp. usually in the vicinity of brooks or springs. Individuals have incidentally been encountered in caves ( Aellen 1951). The terra typica consists of a rocky limestone mountain slope, where only Mediterranean maquis and garrigue is present due to human activities such as logging and the grazing of livestock. However, the combination of sufficient altitude, high annual rainfall and porous limestone outcrops characterized by many crevices seem to allow survival of S. a. splendens ssp. nov. at this location.

The activity pattern of S. a. splendens ssp. nov. ranges from late autumn to early spring, which coincides with periods of high rainfall. The earliest observation of an active individual was made on 9 October 2001, when four adult males were found along a stream south of Ras el Mâ (Taza). South of the latter locality in National Park Tazekka 12 active adult individuals were observed along a humid limestone cliff in a single night in December 2012, foraging and mating during temperatures around 5ºC. During a rainy night on the 15–16th of November 1996 near Chefchaouen, mating behaviour and deposit of larvae was observed amongst approximately 100 adult individuals. At higher altitudes the activity period is shifted towards late spring and early summer, as periods of frost and snow limit activity in the winter. Larvae are generally deposited from October to May in small water bodies, which vary from sources, streams, temporary rain-filled ponds to small man-made concrete water reservoirs and irrigation channels. Dorda (1984) observed larvae and recently metamorphosed juveniles near Chefchaouen in late February. We encountered different sized larvae on December 9th, 1998 near Chefchaouen, ranging from recently deposited individuals of 39 mm to larvae close to metamorphosis measuring 58 mm. On January 4th, 2012 we observed similar size differences (ranging from 40 mm up to 75 mm) among larvae on Jebel Tazekka, south of Ras el Ma (Middle Atlas). Well-developed larvae were additionally observed during the end of April near Bab Berret (central Rif). Larvae of S. a. splendens ssp. nov. can be found together with larvae and adults of A. maurus , D. scovazzi , H. meridionalis and P. saharicus . Cannibalism in combination with a prolonged aquatic period and large growth (TL = 74 mm) has been observed (Escoriza et al. 2006). Metamorphosed juveniles can be found from beginning of January until the end of April, rarely up to summer or even past summer.

Distribution. The occurrence of S. a. splendens ssp. nov. is restricted to the western- and central Rif Mountains (from Chefchaouen to Imassinen and Jebel Aâloul, Jebel Rhelem) and the north-eastern Middle Atlas Mountains (from Tazekka to the Bou Iblane Massif) as previously described by Bons and Geniez (1996), Bogaerts and Donaire-Barroso (2003), Fahd et al. (2005), Escoriza et al. (2006) and Bogaerts et al. (2007). Populations on the Rif- and Middle Atlas Mountains are separated by a large stretch of non-suitable lowland and have been separated from each other for approximately 0.7 my ( Beukema et al. 2010). The distribution map ( Fig. 9A ) is composed of records from Bons and Geniez (1996), Donaire-Barroso and Bogaerts (2003a), Bogaerts et al. (2007), Fahd and Mediani (2007) and Fahd et al. (2007). 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. 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. splendens ssp. nov. in relation to the other Moroccan Salamandra algira ssp.

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

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.