Salamandra algira atlantica, Hernandez, Axel & Escoriza, Daniel, 2019

Hernandez, Axel & Escoriza, Daniel, 2019, A new subspecies of African fire salamander Salamandra algira (Urodela, Salamandridae) from the Middle Atlas Mountains, Morocco, ZooKeys 893, pp. 143-158 : 143

Salamandra algira atlantica ssp. nov. Figs 2 , 3 , 4 Atlas fire salamander

Holotype.

MNCN 50499 ( Fig. 2 ), an adult male preserved in 70% ethyl alcohol from Jbel Tazekka, Taza Province, Middle Atlas Mountains, northern Morocco (34.15N, 4.00W) at 810 m a.s.l collected on 25 March 2013 by Daniel Escoriza.

Paratypes.

MZB 2010-0128 and MZB 2010-0129, two adult males preserved in 70% ethyl alcohol collected in the Jbel Tazekka, Taza Province, Middle Atlas Mountains, northern Morocco in December 2009 by Daniel Escoriza and Félix Amat.

Etymology.

The new taxon is named after the word ' Atlas ' from Greek Aτλας in reference to the Atlas Mountains (Morocco), where this subspecies is found.

English name.

Atlas fire salamander.

Diagnosis.

A large subspecies of S. algira with a maximum total length of 246.21 mm. Distinguished from the other subspecies by the following combination of characters (Table 2 ):

Background dorsal pattern variates from full grey-black to full coral red, being the only subspecies of S. algira in which the coral red can exceed the proportion of grey-black. Salamandra algira algira and S. algira spelaea usually show only few and little red colouration, mostly around yellow blotches on the dorsum of head, limbs and tail and gular region. Salamandra algira tingitana typically lacks red colouration. Salamandra algira splendens never shows full red background colouration, being mainly limited to the edges of the yellow blotches, on the dorsum of the head, limbs, tail and gular region.

Variable number (0-7, usually 2-4) of yellow to golden yellow dorsal blotches, but in lower numbers than the eastern subspecies ( S. algira algira and S. algira spelaea , 3-10). Salamandra algira tingitana can be completely black or with numerous fragmented little yellow spots or with few large blotches in an arrangement similar to S. algira atlantica ssp. nov. Salamandra algira splendens has a similar number of yellow blotches than S. algira atlantica ssp. nov., but these can be more irregular in their shape in the former. Moreover in S. algira atlantica ssp. nov. dorsal yellow blotches can fade progressively into white, composing a pattern that does not appear in S. algira splendens ( Fig. 3D ).

Absence of white spots in the flanks of the body, which appear in the nominotypic subspecies and S. algira spelaea .

Description of holotype.

Snout rounded in a dorsal aspect and in lateral profile. Head large and well differentiated from the body. Nostrils oriented dorsolaterally, situated close to the snout tip. Large-sized parotoids, elongatedly ovoid, slightly divergent anteriorly, showing contrasted glandular pores. Prominent eyes situated laterally, with barely visible vertical oval pupils. Gular fold present. Costal grooves 10. Fingers short and slightly rounded, with a relative length I <IV <II <III. Toes slightly rounded, with a relative length I <V <II <III <IV. Cloaca ovoid. Subcylindrical tail, ended in a blunt tip. Dorsal skin weakly granular. Two pairs of glandular pores disposed in parallel, along the vertebral axis.

Colour of holotype.

In life the background dorsal colouration is black with four elongated golden yellow spots. In the head appear two golden yellow spots on the orbits and two on the parotoids. Four yellow spots at the base of the limbs and five on the dorsum of the tail. Two yellow spots in lateral parts of the body, close to the insertion of the hindlimbs. One small yellow spot showing red colouration in lateral part of the head, located posterior and ventral to the orbit. Uniform greyish black ventral colouration. Eyes dark brown, almost black.

Measurements of holotype (in mm).

TL 175.19 mm, SVL 99.16 mm, HL 20.82 mm, PAL 9.89 mm, HW 14.57 mm, IOR 6.17 mm, ED 5.05, END 4.81 mm, LHU 8.01 mm, FAL 11.52 mm, HAL 13.07 mm, THL 6.53 mm, TIL 11.93 mm, FL 14.8 mm.

Variation.

There are intraspecific population variations regarding colouration pattern in adult specimens ( Fig. 3 ). Within the same population, yellow spots on the dorsum of the head can be divided into four semisymmetric spots on the parotoids and the eyes (21.05% specimens) or they can be merged unilaterally (42.11% specimens) or bilaterally (36.84% specimens). The coral red colour can be limited to a thin colouration on the edge of the yellow spots or be the dominant background colour ( Fig. 3 ). The specimens can show a completely black ventral region, or show some reddish colouration in the throat and / or the cloaca region. More rarely, they can show small white spots in the gular region (10.53% specimens). The only known specimen from the southernmost population (Sidi Ali) showed a predominantly black dorsal colouration, with five small yellow patches in the dorsum and seven on the tail ( Fig. 3A ). The size (TL) of adults ranges from 125.93-246.21 mm in males (N = 15, mean = 190.65 mm ± 7.74 standard error, SE) and 133.43-205.81 mm in females (N = 4, mean = 171.54 mm ± 16.67 SE). The differences in TL between both sexes were not significant (Mann-Whitney U test: U = 23, P = 0.484). Males mean SVL 93.71 mm ± 2.82 SE, HL 21.57 mm ± 0.76, PAL 12.81 mm ± 0.53, HW 15.87 mm ± 0.45, IOR 6.73 mm ± 0.25, ED 4.04 mm ± 0.12, END 4.69 mm ± 0.14, LHU 7.81 mm ± 0.32, FAL 9.57 mm ± 0.36, HAL 12.18 mm ± 0.56, THL 7.49 mm ± 0.31, TIL 10.26 mm ± 0.38, FL 15.34 mm ± 0.59. Females mean SVL 90.72 mm ± 8.33 SE, HL 20.18 mm ± 1.90, PAL 12.82 mm ± 1.11, HW 16.28 mm ± 1.33, IOR 7.33 mm ± 0.57, ED 3.71 ± 0.18, END 3.95 mm ± 0.39, LHU 7.28 mm ± 0.67, FAL 8.40 mm ± 0.86, HAL 11.73 mm ± 1.27, THL 7.36 mm ± 1.06, TIL 9.64 mm ± 1.10, FL 15.15 mm ± 2.69. Males have an ovoid-shaped cloaca during the breeding season and females a flat cloaca. Larvae are characterised by having the following morphological characters: Head depressed, with relatively large eyes situated laterally ( Fig. 4 ). Snout rounded and semi-circular. Gills with three short rami and numerous fimbriae. Four fingers and five toes, narrow and pointed. Ten or eleven costal grooves. Tail equal in length to SVL or slightly smaller. Tail fin short, originating anterior to the pelvic girdle, ended in a bluntly pointed tip. Colour uniformly dark brown with a distinct pale spot at the base of the four limbs. The colouration varies according to the development and the terminal phases show diffuse yellow spots on the head and dorsum. SVL 29.0 mm, TAL 23.9 mm, HL 10.8, HW 6.2, TL 52.9 mm, MTH 5.6 mm (Taza, Morocco).

Distribution.

Salamandra algira atlantica ssp. nov. is endemic to the northern and central Middle Atlas Mountains, northern Morocco: Tazekka National Park, Bou Iblane Massif and Jbel Sidi Ali ( Fig. 5 ).

Natural history.

The new subspecies is found from 600-2455 m a.s.l. near springs and streams in humid mesothermal forests of conifers ( Abies pinsapo , Cedrus atlantica , Pinus halepensis ) and oaks ( Quercus ilex , Q. canariensis , Q. suber ; Fig. 6 A ). There are also troglophile populations at Grotte de Chaâra, Grotte d’Izora. and Gouffre du Friouato which reproduce inside the caves at 400 m from the entrance ( Fig. 6B ). It is a crepuscular and nocturnal species having a surface activity from autumn to spring.

Figures

Figure 2. Holotype of Salamandra algira atlantica ssp. nov. (MNCN 50499) in life. Photograph by Daniel Escoriza.

Figure 3. Variation in the colouration pattern of Salamandra algira atlantica ssp. nov.: A adult male from Jbel Sidi Ali, Midelt Province B adult female from Grotte de Chaara, Taza Province C, D adult males from Grotte de Chaara. Photographs by Axel Hernandez.

Figure 4. Larva of Salamandra algira atlantica ssp. nov. Photograph by Daniel Escoriza.

Figure 5. Map showing the distribution of the subspecies of Salamandra algira (red circles) and the localities included in the phylogenetic analyses (yellow crosses) in northern Morocco and Algeria.

Figure 6. A Habitat of Salamandra algira atlantica ssp. nov. at the type locality, Taza Province, Middle Atlas Mountains, northern Morocco B Cave habitat at Grotte de Chaara. Photographs by Axel Hernandez.