Account written by Axel Hernandez, author of Crocodile Newts: The Primitive Salamandridae of Asia (Genera Echinotriton and Tylototriton), 2016 Edition Chimaira and may contain excerpts from here. More on the author and book.
The North African Fire Salamander
BEDRIAGA 1883 by Axel Hernandez
Université Pasquale Paoli de Corse, faculté des Sciences et Techniques, Corte, 20250, Haute- Corse.
The Fire salamanders, genus Salamandra Garsault, 1764 are a group of terrestrial and tailed amphibians widely distributed across the western Palearctic, extending from the Iberian Peninsula and North Africa to Asia Minor, comprising seven species: S. salamandra, S. longirostris, S. algira, S. corsica, S. infraimmaculata, S. atra and S. lanzai (Dubois and Raffaëlli 2009; Raffaëlli 2013). The genus is particularly diversified in the Iberian Peninsula, possibly caused by several events of allopatric speciation (Steinfartz et al. 2000). In North Africa, Salamandra algira Bedriaga, 1883 is the sole representative species of the genus Salamandra (Bons and Geniez 1996; Schleich et al. 1996; Escoriza and Ben Hassine 2019).
The genus Salamandra most likely colonized the African continent during the Messinian Salinity Crisis via the Gibraltar land bridge (Escoriza et al. 2006; Beukema et al. 2010, 2013). Subsequent divergence was initiated approximately 3.6 mya on both sides of the Moulouya River in eastern Morocco, due to a period of cyclic fluctuations in climate and consequently vegetation in north-western Africa (Beukema et al. 2010, 2013; Ben Hassine et al. 2016). Climatic oscillations during the Upper Pliocene and Pleistocene have driven at least mitochondrial divergence in the hitherto recognized taxa (Beukema et al. 2010, 2013; Dinis et al. 2019). Salamandra algira shows a fragmented distribution pattern which is mostly confined to mountain ranges at an altitude of 0 - 2455 m above sea level (asl), with five subspecies recognized (Donaire-Barroso and Bogaerts 2003; Escoriza and Comas 2007; Beukema et al. 2013; Hernandez and Escoriza 2019): S. a. tingitana occupies the northwestern Rif, ranging from Ceuta to Chefchouen; S. a. splendens extends into the south-central Rif, from Chefchouen to Al Hocéima southwards; S. a. atlantica is found in the northern and central parts of the Middle Atlas; S. a. spelaea is restricted to a small range in the Beni Snassen massif, near the Morocco-Algeria border; and the nominal S. a. algira is restricted to Algeria, in the coastal mountainous regions of Annaba, Kabylia, Blida Atlas and Oranie. Suitable areas were also identified in northwest Tunisia, suggesting that, despite previously reported inability to detect S. algira in the region (Bogaerts et al. 2013), climatic conditions are suitable for the occurrence of this species and the possibility of its persistence in the area cannot be fully discarded (Escoriza and Ben Hassine 2019). The separation among allopatric populations over North African mountainous systems can be attributed to the Late Neogene aridification (Griffin 2002; Escoriza and Ben Hassine 2015). The split between both subspecies of the western group, S. a. tingitana and S. a. splendens, was estimated during the Pliocene, approximately 1.6 ‒ 3.8 million years ago (Mya) while between the eastern subspecies, S. algira algira and S. a. spelaea was estimated during the Plio-Pleistocene boundary, approximately 2.5 Mya (Beukema et al. 2010; Ben Hassine et al. 2016). Salamandra a. atlantica is completely isolated from other populations of S. algira by the arid Moulouya and the Saïss plains. These contemporaneous arid barriers precluded gene flow between Riffean and Middle Atlas populations and the Beni Snassen (Escoriza and Ben Hassine 2015; Hernandez and Escoriza 2019).
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Salamandra a. algira
Salamandra a. atlantica
Salamandra a. tingitana
Salamandra a. spelaea
Salamandra a. splendens
Salamandra algira algira
The Algerian Fire Salamander
Diagnosis and taxonomy
The holotype is not known to exist; However, a neotype was later designated by Eiselt (1958), NHMW 9251 from the same locality as the original description of the holotype "Mt. Edough bei Bône, Algerien"(= Mts Edough, Algeria). Salamandra algira algira is a slender bodied salamander with a relatively long, laterally-flattened tail. The background is usually black, but can also be dark brown. The animals possess a few small yellow spots on dorsal parts, arranged irregularly. Red pigment is rarely present but can be found in juveniles and sometimes subadults. Ventral parts are totally black. Adults of both sexes attain a total length of about 230 mm, sometimes longer. The females are usually larger than males. The male cloaca is much more swollen than that of females. Red pigment is rarely present in the nominal subspecies but can be observed in some specimens, especially from the Blida Atlas region. In addition to the diagnostic characteristic, S. a. algira differs from all other relatives by presenting a combination of the following morphological characters (see Tab. 1): multiple small white spots dispersed in different parts of the body (belly, sides, legs and throat) and a high number of large red spots (type locality and Blida Atlas region). These red spots are surrounded by a ring of small white spots on the lower part of the body and sometimes on the legs, thus creating specific eyespots that are often aligned along the lower sides. According to recent phylogenetic studies, Algerian populations of Salamandra algira are genetically distinct from the Moroccan ones: S. a. algira is clustered with S. a. spelaea in a separate clade. The species is named after the country of its discovery, Algeria.
Salamandra a. algira shows a broad range of distribution from 297 m (Zitouna) to 1550 m asl. (Kabylie region) in the northern coastal mountains of Algeria including: Annaba, Kabylia, Blida Atlas and Oran regions. Its possible presence in northwestern Tunisia is discussed by Bogaerts et al. 2013 and Escoriza and Ben Hassine 2019). Specimens from Rarh el Maden, in the extreme northwest of Algeria, belong to S. a. spelaea.
Habitat, ethology and ecology
This subspecies remains poorly studied. Almost no data on the phylogeography, ecology, reproduction, and morphological variation are known. This nocturnal and crepuscular subspecies is found near wet places in oak or mixed forests mainly covered by Mediterranean maquis, Pinus and Quercus trees. It is only active during periods of relatively high rainfall in the winter. 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 or caves from the end of April to September. The Arabic common name for this species, Arous Chta, means "bride of the rain."
This subspecies is ovoviviparous. 10 to 20 larvae are deposited in tanks, reservoirs, and small springs, brooks and temporary ponds from early October to February - March.
Status, threats and conservation
This relict subspecies is considered as Vulnerable by IUCN (VU) due to its severely fragmented distribution and ongoing decline in the extent and quality of its forest habitat in Morocco and Algeria.
Salamandra algira atlantica
HERNANDEZ AND ESCORIZA 2019
Atlas Fire Salamander
Diagnosis and taxonomy
The holotype (MNCN 50499) is an adult male from Jbel Tazekka, Taza Province, Middle Atlas Mountains, northern Morocco collected at 810 m asl on 25 March 2013 by Daniel Escoriza. This subspecies is named after the word ‘Atlas’ from Greek Aτλας in reference to the Atlas Mountains (Morocco), where this subspecies is found. According to Hernandez and Escoriza (2019), Salamandra a. atlantica is a large subspecies of S. algira with a maximum total length of 246.21 mm showing a variable number (0 - 7) of yellow to golden yellow dorsal blotches rounded in shape. It’s the only known subspecies in which coral red coloration can exceed the proportion of black. It can be distinguished from the other subspecies by the following combination of characters (see Tab. 1):
There are intraspecific population variations regarding coloration pattern in adult specimens. Within the same population, yellow spots on the dorsum of the head can be divided into four semi- symmetric 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 color can be limited to the edge of the yellow spots or be the dominant background color (Fig. 3). The specimens can show a completely black ventral region, or show some reddish coloration 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 coloration, with five small yellow patches in the dorsum and seven on the tail.
- 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 a. algira and S. a. spelaea usually show only little red coloration, mostly around yellow blotches on the dorsum of head, limbs and tail and gular region. Salamandra a. tingitana typically lacks red coloration. Salamandra a. splendens never shows full red background coloration; red is 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. a. algira and S. a spelaea, 3 – 10). Salamandra a. tingitana can be completely black or with numerous fragmented little yellow spots or with few large blotches in an arrangement similar to S. a. atlantica. Salamandra a. splendens and S. a. atlantica have similar numbers of yellow blotches, but these can be more irregular in their shape in the former. Moreover in S. algira atlantica dorsal yellow blotches can fade progressively into white, composing a pattern that does not appear in S. a. splendens.
- Absence of white spots on the flanks of the body, which appear in the nominotypic subspecies and S. a. spelaea.
Salamandra a. atlantica is endemic to the northern and central Middle Atlas Mountains, northern Morocco including Tazekka National Park, Bou Iblane Massif and Jbel Sidi Ali.
Habitat, ethology and ecology
This subspecies is generally found between 600 – 2455 m asl near springs and streams in humid mesothermal forests of conifers (Abies pinsapo, Cedrus atlantica, Pinus halepensis) and oaks (Quercus ilex, Q. canariensis, Q. suber). It is a crepuscular and nocturnal species with surface activity from autumn to spring. There are troglophile populations at Grotte de Chaâra, Grotte d’Izora and Gouffre du Friouato which reproduce inside the caves at 400 m from the entrance (Hernandez 2018b,c). It is a crepuscular and nocturnal species having a surface activity from autumn to spring.at middle to high altitudes, from 600 - 2250 m asl in patch of mixed forests, alpine grasslands or sometimes into caves where they can breed. The southernmost record is Jbel Sidi Ali, southeastern Bou Anguer township, Midelt province, central Middle Atlas at 2206 m asl. One adult male was recently found in a ditch containing water seepage in this locality. The habitat was damp with high level of humidity in the air (76.8 %) and on the ground level (87.2 %). The temperature ranged from 3.4°C to 8.3°C during the daytime. The microhabitat was poorly vegetated and contained a fragmented patch of a Cedar forest (Cedrus atlantica) with some Quercus rotundifolia and Juniperus thurifera on a volcanic plateau at the vicinity of a large permanent and eutrophic lake, the Aguelmame Sidi Ali (Hernandez 2018a). The soil includes rocks of various size and perennial herbs. The habitat around the forest is similar to an alpine wetland in appearance with many ephemeral water ponds. Snow was present near the location at 2350 m asl. Mean annual precipitation in the area is 817 mm, most of which falls between November and April, when the mean minimum monthly temperature is – 6 °C. Snow may also persist into early summer. Occasional intense rainstorms account for much of the summer precipitation. The area under study belongs to the supra-Mediterranean sub-humid bioclimatic zone with cold winter variation. Individuals have incidentally been encountered in caves and suitable habitats were also observed in the same mountain ranges such as the Jbel Bouyzane, Jbel Inifif, Jbel Moulay Sadik, Jbel Myadir or the Jbel Boulkhitane through the southern Moulouya basin where the subspecies probably had a continuous distribution range. The only adult observed at Sidi Ali was a slender male with a rounded cloaca and a large size of 227 mm in total length (TL). It was characterized by having the following morphological combination: an elongated flat head, 1½ times longer than wide; long well separated limbs, fingers and tail; a general black coloration with few and irregular yellow markings. The red discoloration was almost absent, only indistinguishable red spots being present on the parotoid glands and below the throat. In the northern part of its range, five adults, one juvenile and 22 larvae of S. a. atlantica were observed in a large cave commonly known as Grotte de Chaâra located 1213 m asl. The adults and juvenile were walking on rocks. They were located from a few hundred meters up to 2 km into the cave and near a large permanent stream in a large chamber. Larvae were generally found inside the cave at 160-400 m from the entrance in five pools, about 68 - 185 cm in length and 32 - 55 cm deep. In all the pools containing larvae, water quality was acidic to neutral with a pH between 6.13 - 7.11, air temperature at midday was 13.1 - 16.2 °C and water temperature 11.1 - 12.6 °C. The habitat was characterized by various rocks and small to large pools in complete darkness. The external habitat comprised pine and oak forests with Pinus sp. and Quercus suber.
S. a. atlantica is ovoviparous and reproduces mainly during the winter period up to the beginning of the springtime. However, larvae can be found in caves throughout the whole year (pers. obs.). Larvae are characterized as follows: Head depressed, with relatively large eyes situated laterally. 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 coloration varies according to the stage of development, with the terminal phases showing 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; see Hernandez and Escoriza 2019).
Status, threats and conservation
In terms of conservation, the southernmost populations of S. a. atlantica seem to be very scarce, based on the paucity of specimens found during the last decades. This subspecies is classified as Vulnerable (VU) under IUCN criteria. It is threatened by anthropogenic pressure including water pollution near the lake and habitat destruction. Climate change may be also another major threat for the coming years especially in the Mediterranean region where a high biodiversity of unique salamander species and other endemic amphibians occurs. A re-assessment of the outdated Vulnerable status (VU) of S. algira is needed, which will reflect the current taxonomic revisions and the increasing threats from the international pet trade and habitat loss, which have taken place over the last decade.
Salamandra algira tingitana
DONAIRE-BARROSO & BOGAERTS 2003
Tanger’s Fire Salamander
Diagnosis and taxonomy
The holotype (MNCN 41037) was designed from: "500 m altitude on Jabal Muse (= Jebel Musa) north Morocco". The specimen is an adult female collected on 7 December 1997 by David Donaire- Barroso and Francisco Javier Martinez-Medina (See Beukema et al. 2013 for justification of validity as an electronic publication). According to Donaire-Barroso and Bogaerts (2003), this subspecies is characterized by the following morphological characters: Dorsal ground color is jet black with ten daffodil- yellow spots arrayed in a more or less longitudinal pattern; four spots on the head (one on each eye pad and paratoid), one small one in the neck, four in the dorsum and three connected ones in the sacrum. The spot on the left eye pad is threefold shaped, whereas the spot on the right eye pad is similarly shaped but with a greater reduction in yellow coloration. Four small dots on the right lateral side and none in the left lateral one, of which two are in the trunk and two in the sacral side; the tail is mainly black but with two dots at the base on the left side and one dot at the base of the right side. There is one spot on the left forearm, one at the base of the right arm, one at the base of the left leg and two very small vague ones on the right leg. Ventral side coloration, including tail and throat regions, is dark reddish-brown with no yellow spots. Males of S. a. tingitana may develop conspicuous glandular skin protuberances on the whole dorsum and head during mating season. 'Nuptial pads' were never observed in other populations. General coloration differs from other populations: they lack the vivid black-yellow coloration that is common in other Salamandra and the ground color is dark brown instead of black. There is also a high tendency towards hypoluteism and even melanism. Red coloration is totally absent. According to Donaire-Barroso and S. Bogaerts (2003), about 30 animals were photographed and partly measured in the field. From photographs they found that 15 animals (50 %) have four spots on the head (paratoids and eye pads), while the other 15 animals (50 %) have fewer or no spots at all on their heads. Thirteen (43.3 %) of these 30 animals, especially juveniles, show blue whitish coloration on the eye pads. Two animals (6.6%) were completely melanistic, whereas 5 (16.6%) showed very strong reduction in the yellow coloration (hypoluteism). Fifteen (50%), mostly adult, animals showed signs of reduction in the yellow pattern and the remaining eight (26.6%) had clear full colored spots. Red coloration was absent in all of the photographed animals (100%) and all others that were observed. Ground color within this subspecies varies from completely jet-black to reddish-dark brown. The yellow coloration varied from orange-yellow to daffodil yellow in 10 animals (33.3 %), which had a dark ground color, whereas it varied from daffodil-yellow to vague whitish-yellow in 13 animals (43.3%) that showed a brownish ground color. Juveniles, and sometimes adults, can have white-bluish or greenish coloration of the spots above the eyes, instead of yellow. Dorsal yellow spots array in a single longitudinal or “zigzag” pattern. They never occur in two rows nor do they form longitudinal lines; however ontogenetic changes can give the illusion of two rows of spots. Moreover, Salamandra a. tingitana is divided into two main forms with two various reproduction modes (pueriparous and larviparous) while mtDNA studies showed the taxon to consist of three different clades which diverged during the Pleistocene (see Beukema et al., 2010). Populations from northwestern Morocco consistently lack red discoloration while often showing white specks or rosettes on the underside, they are pueriparous and larviparous (Clade 2); The northernmost populations were shown to possess a high tendency towards hypoluteism and melanism and are mainly pueriparous (Clade 1) while southern populations shows similar morphology as S. a. algira and are mainly larviparous (Clade 3). The boundary between these is approximately located near the Oued Martil, with larvae occasionally having been found west of Tetouan, the southernmost population bearing fully- developed juveniles. The viviparous form of S. a. tingitana, which is only distributed in the extreme northern parts of the Tingitana peninsula, was previously considered as a full valid species by Dubois and Raffaëlli (2009). This consideration is still under debate, despite unanimous recognition of two very divergent phenotypes and genotypes. According to Hernandez and Escoriza (2019), S. a. tingitana may warrant taxonomic revision including new phylogenetic and morphological studies.
S. a. tingitana is distributed in the Tingitana peninsula across the Oued Martil up to southern Chefchaouen. Populations of this subspecies are separated with S. algira splendens by a natural barrier, the river Oued Laou.
Habitat, ethology and ecology
Salamandra a. tingitana occupies distinct habitats along its distribution, which have been suggested to be associated with distinct reproductive strategies. Pueriparous populations were originally described from Pinus ssp. and Quercus ssp. forests and in barely vegetated limestone formations up to 390 m asl, in the Mediterranean-influenced northeast Tingitana peninsula, while a distinct habitat has been described for the southernmost larviparous populations of this subspecies, consisting of mountainous or hilly terrain up to at least 1,274 m asl near water sources and in half-open forests, agricultural terraces and limestone outcrops (Donaire-Barroso and Bogaerts 2003; Beukema et al. 2013). Ecological niche-based models have identified a weaker association with vegetation abundance and increased reliance on karstic substrates among the pueriparous populations (Beukema et al. 2010). 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 (Donaire-Barroso and Bogaerts 2003). Low temperatures (less than 10ºC) and presence of considerable wind does not have a negative influence on activity in general (Beukema et al. 2013).
Most of the S. a. tingitana populations are larviparous. However, pueriparous populations have been described north of Oued Martil, and this has been connected with a distinct ecological niche for the larviparous populations of this subspecies (Beukema et al. 2010, 2013; Escoriza and Ben Hassine 2014).
Status, threats and conservation
Vulnerable (as part of S. algira). The northernmost populations are highly threatened by rock extractions (pers. obs.).
Salamandra algira splendens
BEUKEMA, DE POUS, DONAIRE-BARROSO, BOGAERTS, GARCIA-PORTA, ESCORIZA, ARRIBAS, EL MOUDEN, & CARRANZA 2013
Splendid Moroccan Salamander
Diagnosis and taxonomy
The holotype (RMNH 40173) was designed from: "Aïn Tissimilan, Jebel el Kelaâ, Chefchaouen, western Rif Mountains, Morocco (N 35°10.5, W 5°14.6, 700 m asl)". The holotype is an adult male collected at 700 m asl in November 1996 by David Donaire-Barroso and César Barrio Amoros. 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. The subspecies is diagnosed by the following combination of characters: 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. Red pigment is less present on the body of this subspecies but can be visible mainly on parotoid glands and, in some specimens on body and tail especially in Ketama and Al Hocéima populations. In addition to the diagnostic characteristics already mentioned above, S. a. splendens differs from its relatives by a combination of color pattern and distribution (see Tab. 1). This is a very slender African fire salamander, showing a thin and elongated body and tail. This subspecies also lacks small white spots on the flanks and has little or no red spots. Yellow patches are without exception present on the eyes, parotoids and at the bases of limbs. Salamandra a. splendens has a similar number of yellow blotches as S. a. atlantica but these are more irregular in their shape (see Hernandez and Escoriza 2019).
The occurrence of S. a. splendens is restricted to the western- and central Rif Mountains from Chefchaouen to Imassinen, Jebel Aâloul, Jebel Rhelem up to the Bokkoyas Massif, Al Hocéima (Beukema et al. 2013; Hernandez and Escoriza 2017).
Habitat, ethology and ecology
Salamandra a. splendens has been found between 280 – 1700 m asl in the 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. 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 (Hernandez 2017, 2018b,c). The type locality 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 seems to allow survival of S. a. splendens at this location. The activity pattern of S. a. splendens ranges from late autumn to early spring, which coincides with periods of high rainfall. During a rainy night on the 15 – 16th of November 1996 near Chefchaouen, mating behavior 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 (Beukema et al. 2003).
The breeding season begins in October up to February which coincides with periods of high rainfall. Larvae are generally deposited from October to May in small water bodies, which vary from sources, small streams, temporary rain-filled ponds to small man-made concrete water reservoirs and irrigation channels. Metamorphosed juveniles can be found from beginning of January until the end of April, rarely up to summer or even past summer (Beukema et al. 2013; Raffaëlli 2013; Hernandez 2017). Some authors observed larvae and recently metamorphosed juveniles near Chefchaouen in late February (Beukema et al., 2013). They 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. Well-developed larvae were additionally observed during the end of April near Bab Berret (central Rif). Larvae of S. a. splendens can be found together with larvae and adults of Alytes maurus, Discoglossus scovazzi, Hyla meridionalis and Pelobates saharicus. Cannibalism in combination with a prolonged aquatic period and large growth (TL = 74 mm) has been observed. Metamorphosed juveniles can be found from beginning of January until the end of April, rarely up to summer or even past summer (Beukema et al. 2013).
Status, threats and conservation
This subspecies is considered as Vulnerable (as part of S. algira). The alteration and destruction of natural habitats are the main causes of population declines. The Rif region suffered from rampant deforestation.
Salamandra algira spelaea
ESCORIZA AND COMAS 2007
Beni Snassen’s Fire Salamander
Diagnosis and taxonomy
The holotype (MNCN 2005-05550) was designed from: "Ouartass, Beni Snassen massif, Northeast-Morocco at approximately 1300 m above sea level".
According to Escoriza and Comas (2007), this subspecies is characterized by having the following morphological characters: “a slender body and tail with moderate size (77 - 236 mm TL). Head flat, longer than wide, with prominent eyes having a black iris. Snout rounded, limbs large, tail large and cylindrical. Jet black ground coloration with scattered rounded or elongated yellow blotches, not arranged in bands, located at head, dorsum and tail; predominance of black over yellow. Large parotoid glands with reddish or some black glandular pores. Tiny red spots around parotoids, eyes, flanks, and on limbs and tail, sometimes as independent spots or fused to the yellow or white ones. Occasionally small white spots in lateral, gular and ventral region. Ventral and gular region black. Four fingers and five toes, all without membranes. Eight to eleven lateral tubercles. Tail a little shorter than head plus body. Ovoviviparous. Juveniles with similar color pattern as adults, with many white spots located on both sides of the body and without ventral spots”. Furthermore, according to various mitochondrial studies S. a. spelaea is the sister taxon of S. a. algira (Escoriza and Ben Hassine 2019).
The occurrence of S. a. splendens is restricted to the Beni Snassen Massif, Northeastern Morocco. The transition populations of S. a. algira recorded from Algeria in Rahr-el-Maden requires further studies to unravel their relationship with S. a. spelaea or the nominal subspecies.
Habitat, ethology and ecology
The subspecies has been found from 600 to 300 m asl occupying mixed forests that appear between these altitudinal ranges including Quercus, Pinus and Olea ssp. trees on limestone outcrops and granite soils near water bodies. Active individuals have been observed in November during humid or rainy weather, both in the late afternoon and night. During these observations, air temperature varied between 5.8 and 7 ºC while humidity ranges between 75 – 85%. Larvae have been found in November and January, while fully metamorphosed juveniles were observed in mid-February (Escoriza and Comas 2007).
The reproduction is poorly known for this subspecies. Due to the nature of the soil most water bodies used for reproduction are man-made, such as cattle watering troughs or springs, although small puddles and temporary streams are also used to deposit larvae. The onset of the activity period is initiated by the late autumn rains, and continues throughout the winter (Escoriza and Comas 2007; Beukema et al. 2013).
Status, threats and conservation
This subspecies is considered as Vulnerable (as part of S. algira). Salamandra a. spelaea was proposed to qualify as Endangered by Escoriza and Comas (2007). The entire range of S. a. spelaea is subject to intensive human impact by means of logging, construction of buildings, canalization of natural springs and construction of fountains which prevent access to the limited water bodies by pregnant females. These actions have led to considerable desertification, especially on the southern slopes of the massif (Escoriza and Comas 2007).
Table 1. Identification key to Salamandra algira (derived and modified from Hernandez and Escoriza 2019)
| ||Salamandra algira algira ||Salamandra algira spelaea ||Salamandra algira tingitana ||Salamandra algira splendens ||Salamandra algira atlantica ssp. nov. |
|Total length (in mm) ||131 – 192 ♂ |
129 – 255 ♀
|177 – 194 ♂ |
206 – 236 ♀
|151 – 170 ♂ |
140 – 192 ♀
|140 – 178 ♂ |
161 – 261 ♀
|125 – 246 ♂ |
133 – 205 ♀
|Background coloration ||Grey-black ||Grey-black ||Grey-black ||Grey-black or coral red ||Grey-black or coral red |
|Number of dorsal yellow blotches ||3 – 10 |
|3 – 10 |
|0 – 7 |
|2 – 7 |
Irregular in shape
|0 – 7 |
Rounded in shape
|Red spots ||Usually small ||Small ||Absent ||Small to large ||Small to full |
|White spots on flanks ||Present ||Present ||Absent ||Absent ||Absent |
- Bedriaga J (1883) Beiträge zur Kenntniss der Amphibien und Reptilien der Fauna von Corsika. Archiv für Naturgeschichte, Berlin 49: 124–273.
- Ben Hassine J, Gutiérrez-Rodríguez J, Escoriza D, Martínez-Solano I (2016) Inferring the roles of vicariance, climate and topography in population differentiation in Salamandra algira (Caudata, Salamandridae). Journal of Zoological Systematics and Evolutionary Research 54(2): 116–126.
- Beukema W, De Pous P, Donaire D, Escoriza D, Bogaerts S, Toxopeus AG, De Bie CA, Roca J, Carranza S (2010) Biogeography and contemporary climatic differentiation among Moroccan Salamandra algira. Biological Journal of the Linnean Society 101(3): 626–641.
- Beukema W, De Pous P, Donaire-Barroso D, Bogaerts S, Garcia-Porta J, Escoriza D, Arribas O, El Mouden E, Carranza S (2013) Review of the systematics, distribution, biogeography and natural history of Moroccan amphibians. Zootaxa 3661: 1–60.
- Bogaerts S, Donaire-Barroso D, Pasmans F, Böhme W, Carranza S (2013) Do north African ﬁre salamanders, Salamandra algira, occur in Tunisia? Herpetological Notes 6, 301-306.
- Dinis M, Merabet K, Martínez-Freiría F, Steinfartz S, Vences M, Burgon JD, Elmer KR, Donaire D, Hinckley A, Fahd S, Joger U, Fawzi A, Slimani T, Velo-Antón G (2019) Allopatric diversification and evolutionary melting pot in a North African Palearctic relict: the biogeographic history of Salamandra algira. Molecular Phylogenetics and Evolution 130: 81–91.
- Donaire-Barroso D, Bogaerts S (2003) A new subspecies of Salamandra algira Bedriaga, 1883 from northern Morocco. Podarcis 4: 84–100.
- Dubois A, Raffaëlli J (2009) A new ergotaxonomy of the family Salamandridae Goldfuss, 1820 (Amphibia, Urodela). Alytes 26(1–4): 1–85.
- Eiselt J. (1958) Der Feuersalamander Salamandra salamandra (L.): Beiträge zu einer taxonomischen Synthese. Abhandlungen und Berichte für Naturkunde und Vorgeschichte Museum Magdeburg 10: 77–154.
- Escoriza D, Comas MM, Donaire D, Carranza S (2006) Rediscovery of Salamandra algira Bedriaga, 1883 from the Beni Snassen Massif (Morocco) and phylogenetic relationships of North African Salamandra. Amphibia-Reptilia 27: 448–455.
- Escoriza D, Comas MM (2007) Description of a new subspecies of Salamandra algira Bedriaga, 1883 (Amphibia: Salamandridae) from the Beni Snassen massif (Northeast Morocco). Salamandra 43: 77–90.
- Escoriza D, Ben Hassine J (2014) Microclimatic variation in multiple Salamandra algira populations along an altitudinal gradient: phenology and reproductive strategies. Acta Herpetolologica 9(1): 33–41.
- Escoriza D, Ben Hassine J (2015) Niche partitioning at local and regional scale in the north African Salamandridae. Journal of Herpetology 49: 276–283.
- Escoriza D, Ben Hassine J (2019) Amphibians of North Africa. Academic Press, London, 350 pp.
- Eiselt J (1958) Der Feuersalamander Salamandra salamandra (L.). Beiträge zu einer taxonomischen Synthese. Abhandlungen und Berichte für Naturkunde. Musem für Naturkunde Magdeburg 10: 77–154.
- García-París M, Alcobendas M, Buckley D, Wake DB (2003) Dispersal of viviparity across contact zones in Iberian populations of fire salamanders (Salamandra) inferred from discordance of genetic and morphological traits. Evolution 57: 129–143.
- Griffin DL (2002) Aridity and humidity: two aspects of the late Miocene climate of North Africa and the Mediterranean. Palaeogeography, Palaeoclimatology, Palaeoecology 182(1–2): 65–91.
- Hernandez A (2017) Les Urodèles du Maroc. Histoire naturelle, taxinomie, écologie et conservation. Situla Revue d’Herpétoculture Francophone, 29: 30–49.
- Hernandez A, Escoriza D (2017) Easternmost record of Salamandra algira splendens in Morocco. Boletín de la Asociación Herpetológica Española 28(1): 60–61.
- Hernandez A (2018a) New locality of Salamandra algira splendens Beukema, de Pous, Donaire-Barroso, Bogaerts, Garcia-Porta, Escoriza, Arribas, El Mouden & Carranza 2013 at the southern limit of its geographical distribution in Morocco. Bulletin de la Société Herpétologique de France 165: 18–22.
- Hernandez A (2018b) Salamandra algira splendens and Pleurodeles waltl in Moroccan caves; new distributional records. Herpetological Bulletin 144: 36–38.
- Hernandez A (2018c) When red embraces the darkness: first report of the most reddish specimen of Salamandra algira splendens Beukema, de Pous, Donaire-Barroso, Bogaerts, Garcia-Porta, Escoriza, Arribas, El Mouden & Carranza 2013 in a cave of northern Morocco. Bulletin de la Société Herpétologique de France 168: 30–34.
- Hernandez A, Escoriza D (2019) A New Subspecies of African Fire Salamander Salamandra algira (Urodela, Salamandridae) from the Middle Atlas Mountains, Morocco. ZooKeys 893: 143–158. DOI: 10.3897/zookeys.893.46649
- IUCN SSC Amphibian Specialist Group (2009) Salamandra algira. The IUCN Red List of Threatened Species 2009:e.T59464A11927380. http://dx.doi.org/10.2305/IUCN.UK.2009.RLTS.T59464A11927380.en. Downloaded on 07 August 2019.
- Joger U, Steinfartz S (1994) Zur subspezifischen Gleiderung der südiberischen Feuersalamander (Salamandra salamandra-complex). Abhandlungen und Berichte für Naturkunde. Musem für Naturkunde, Magdeburg 17: 83–98.
- Raffaëlli J (2013) Les Urodèles du monde.2ème edition. Penclen editions, Plumelec, 480 pp.
- Recuero E, Iraola A, Rubio X, Machordom A, García‐París M (2007) Mitochondrial differentiation and biogeography of Hyla meridionalis (Anura: Hylidae): an unusual phylogeographical pattern. Journal of Biogeography 34(7): 1207–1219.
- Seidel U, Gerhardt P (2016) The Genus Salamandra: History, Biology, Systematics, Captive Breeding. Edition Chimaira, Frankfurt am Main, Germany, 543 pp.
- Sparreboom M (2014) Salamanders of the Old World: the salamanders of Europe, Asia and northern Africa. Knnv Publishing, Zeist, Netherlands, 385 pp.
- Steinfartz S, Veith M, Tautz D (2000) Mitochondrial sequence analysis of Salamandra taxa suggests old splits of major lineages and postglacial recolonizations of Central Europe from distinct source populations of Salamandra salamandra. Molecular Ecology 9: 397–410.
- Thiesmeier B (2004) Der Feuersalmamander. Laurenti Verlag, Bielefeld, Germany, 192 pp.
- Thiesmeier B, Grossenbacher K (2004) Salamandra salamandra (Linnaeus, 1758) Feuersalamander. In: Thiesmeier B, Grossenbacher K (Eds) Handbuch der Reptilien und Amphibien Europas: Schwanzlurche IIB. Aula Verlag, Wiesbaden, 1059–1132.
- Veith M (1994) Morphological, molecular and life history variation in Salamandra salamandra. Mertensiella 4: 355–397.
- Veith M (1996) Are Salamandra atra and S. lanzai sister species?. Amphibia- Reptilia 17(2): 174–177.
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