Description

The North African Salamander (Salamandra algira) is a striking amphibian endemic to North Africa. It measures between 15-25 cm in total length, with a robust body and rounded snout. The most distinctive feature of this salamander is its dramatic coloration, which consists of a glossy black background adorned with bright yellow or orange-red patterns that form irregular spots, stripes, or blotches along the body and tail^[1].

This species exhibits considerable variation in color pattern across its range, with populations displaying different proportions of yellow and red markings depending on their subspecies and geographical location^[2]. Sexual dimorphism is present, with males typically smaller than females and having a more swollen cloaca during the breeding season^[3].

4 out of the 5 subspecies can be found in Morocco, the nominal form being endemic to Algeria.

Salamandra algira spelaea

The Salamandra algira spelaea subspecies is more constant in coloration compared to other subspecies. Yellow markings are usually small and numerous. White dots can sometimes be found on the body, particularly in the lateral, gular, and ventral regions^[5]. This subspecies is only found in the Beni Snassen massif in northeastern Morocco, where it has adapted to life in or near cave systems^[6].

Salamandra algira splendens

The Salamandra algira splendens subspecies shows high variability in coloration, ranging from mostly black individuals with few red markings and abundant yellow dots, to surprisingly colorful individuals with very high amounts of red^[7]. Individuals from the eastern-most part of the range display a darker shade of red, almost grape purple colored. This subspecies is characterized by a longer tail and a more elongated form. Red pigmentation is particularly prominent in Rif populations, especially around the parotoid glands, throat, and between limbs^[8].

Salamandra algira tingitana

The Salamandra algira tingitana subspecies is composed of both viviparous and ovoviviparous forms that inhabit the Tingitana Peninsula region, and be split in 3 distinct genetical clades. The viviparous form (Clade 1) is characterized by a brown-black to completely black coloration (melanism) with few scattered yellow spots, and is uniquely localized to the north of Oued Martil^[9]. The ovoviviparous form has more pronounced yellow patterning on the dorsal surface and resembles the nominotypical subspecies from Algeria. Males of this subspecies develop swollen forearms during the breeding season^[9].

Natural History

Salamandra algira is a primarily terrestrial species that is most active during cool, humid conditions. These salamanders are nocturnal, spending daylight hours concealed under rocks, logs, or in burrows, emerging at night to forage. Their diet consists mainly of invertebrates, including insects, spiders, worms, and slugs^[10].

Like other salamander species, Salamandra algira possesses skin glands that secrete toxins as a defense against predators. When threatened, they may arch their back and lift their tail to display their warning coloration more prominently, a behavior known as the unken reflex. The bright colors serve as aposematic signals, warning potential predators of their toxicity^[11].

Reproduction in Salamandra algira is quite variable across its range. Some populations are ovoviviparous, with females giving birth to fully developed larvae that are deposited in water, while others (particularly at higher elevations and in the viviparous form of S. algira tingitana) are fully viviparous, giving birth to fully metamorphosed juveniles^[9][12]. Mating typically occurs on land during rainy periods in autumn or spring. Depending on the subspecies and local conditions, females may produce 10-40 offspring per reproductive cycle. Larvae typically develop in cool, oxygen-rich streams or springs for 3-5 months before metamorphosing into terrestrial juveniles^[13].

These salamanders can live up to 15-20 years in the wild and are adapted to the Mediterranean climate of North Africa, becoming dormant during hot, dry summer months (estivation) and sometimes during the coldest winter periods^[14].

Distribution

The North African Salamander is endemic to the mountainous regions of northwestern Africa. Its distribution is highly fragmented due to climate and topographical barriers, with populations isolated in different mountain ranges^[15].

In Morocco, different subspecies occupy distinct ranges:

• S. algira tingitana is found in the northwestern Rif Mountains (Tingitana Peninsula), ranging from Ceuta to Jbel Sougna^[16].
• S. algira splendens inhabits the central and eastern Rif Mountains, from Chefchaouen to Al Hoceima, including recently discovered populations in the Bokkoyas massif which represent its easternmost occurrence in the Rif^[17].
• S. algira spelaea is restricted to the Beni Snassen massif in northeastern Morocco, where it is confined to a small area of approximately 45 km²^[5].
• S. algira atlantica is endemic to the northern and central Middle Atlas Mountains, particularly around Jbel Tazekka and Bou Iblane Massif^[4].

The nominotypical S. algira algira is found in Algeria, across the pericoastal mountains of Annaba, Collo, Kabylia, and the Blida Atlas^[19]. These populations are separated from Moroccan populations by approximately 800 km^[3].

The fragmented distribution of Salamandra algira is primarily due to its requirement for humid habitats in predominantly arid regions. The main barrier between eastern and western populations is the arid Moulouya River basin, which creates a significant biogeographical boundary^[15].

Habitat

In Morocco, Salamandra algira inhabits predominantly humid, forested mountain regions with abundant rainfall. These salamanders are typically found between elevations of 30-2,455 m above sea level, occupying a variety of Mediterranean forest types and microhabitats depending on the subspecies^[15].

The four Moroccan subspecies have adapted to slightly different habitat types:

• S. algira atlantica is found in subhumid to humid forests and karstic systems at mid to high elevations (600-1,300 m), typically in forests dominated by Quercetalia ilicis vegetation including cork oak, holm oak, and cedar forests^[4].
• S. algira spelaea inhabits rocky environments and caves in the Beni Snassen massif, at elevations of 600-1,300 m. This troglophilic subspecies can be found deep within caves, as well as in the Mediterranean forest outside^[6].
• S. algira splendens occurs in diverse habitats from 280-2,200 m elevation in the Rif Mountains and Middle Atlas, preferring calcareous soils in pine, cedar, and oak forests. It is typically found near springs and small streams^[7]. Recently discovered populations in the region of Al Hoceima have been found at lower elevations (around 220 m) in mesic mixed forest with dense scrub^[17].
• S. algira tingitana occupies varied ecological niches in karstic regions at elevations of 390-1,274 m. The vegetation is predominantly Mediterranean maquis with small shrubs, herbs, and forests of pine and cork oak^[7].

The species requires access to clean, flowing water bodies such as mountain springs, streams, and seeps for larval development, typically avoiding still waters. Populations from caves and certain high-elevation areas have adapted to viviparous reproduction, eliminating the need for aquatic habitats^[12].

Conservation

Salamandra algira is listed as Vulnerable on the IUCN Red List, with populations showing a declining trend. The species faces numerous threats throughout its limited range in North Africa, resulting in increasingly fragmented populations and local extinctions^[20].

Major threats to this species in Morocco include:

• Habitat loss due to deforestation, agricultural expansion, and infrastructure development^[21].
• Climate change and increasing drought frequency, reducing habitat suitability and breeding opportunities^[22].
• Water pollution and modification of stream habitats, affecting larval development^[21].
• Over-collection for the pet trade, particularly of the more colorful subspecies^[23].
• Introduced predatory fish that prey on salamander larvae^[24].
• Emerging fungal diseases, including chytridiomycosis, which threatens amphibian populations globally^[25].

Conservation efforts for Salamandra algira in Morocco include protection within several national parks and nature reserves, such as Talassemtane National Park in the Rif Mountains and the Parc National d'Al-Hoceima. The species is legally protected in Morocco, though enforcement is inconsistent^[26]. Research and monitoring programs have been established to better understand population dynamics and genetic diversity across the species' range.

The cave-dwelling subspecies S. a. spelaea is of particular conservation concern due to its extremely restricted range and specialized habitat requirements. This subspecies may warrant elevation to Critically Endangered status in future assessments^[5].

References

1. Hernandez, A. & Escoriza, D. (2017). Easternmost record of Salamandra algira splendens in Morocco. Boletín de la Asociación Herpetológica Española, 28(1): 60-61.
2. Beukema, W., de Pous, P., Donaire-Barroso, D., Bogaerts, S., Toxopeus, A.G., De Bie, C.A.J.M., Roca, J. & Carranza, S. (2010). Biogeography and contemporary climatic differentiation among Moroccan Salamandra algira. Biological Journal of the Linnean Society, 101: 626-631.
3. Escoriza, D. & Ben Hassine, J. (2019). Amphibians of North Africa. Academic Press, London, 350 pp.
4. 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.
5. Escoriza, D. & Comas, M.M. (2007). Description of a new subspecies of Salamandra algira Bedriaga, 1883 (Amphibia: Salamandridae) from the Beni Snassen massif (Northeast Morocco). Salamandra, 43: 77-90.
6. Hernandez, A. (2018). Salamandra algira splendens and Pleurodeles waltl in Moroccan caves; new distributional records. Herpetological Bulletin, 144: 36-38.
7. Hernandez, A. (2017). Les Urodèles du Maroc: Pleurodeles waltl, Salamandra algira tingitana, S. algira splendens et S. algira spelaea (Caudata: Salamandridae). Histoire naturelle, taxinomie, écologie et perspectives conservatoires. Situla Revue d'Herpétoculture Francophone, 29: 30-48.
8. Beukema, W., de Pous, P., Donaire-Barroso, D., Bogaerts, S., García-Porta, J., Escoriza, D., Arribas, O.J., El Mouden, E.H. & Carranza, S. (2013). Review of the systematics, distribution, biogeography and natural history of Moroccan amphibians. Zootaxa, 3661: 1-60.
9. Donaire-Barroso, D. & Bogaerts, S. (2003). A new subspecies of Salamandra algira Bedriaga, 1883 from northern Morocco. Podarcis, 4: 84-100.
10. 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: 116-126.
11. Raffaëlli, J. (2013). Les Urodèles du monde. 2ème édition. Penclen editions, 480 pp.
12. Donaire-Barroso, D. & Bogaerts, S. (2016). Observations on viviparity of Salamandra algira in Morocco and Algeria. Butlletí de la Societat Catalana d'Herpetologia, 23: 7-14.
13. Escoriza, D. & Ben Hassine, J. (2015). Niche partitioning at local and regional scale in the North African Salamandridae. Journal of Herpetology, 49(2): 276-283.
14. Schleich, H.H., Kästle, W. & Kabisch, K. (1996). Amphibians and Reptiles of North Africa. Koeltz Scientific Books, Koenigstein.
15. Escoriza, D., Comas, M.M., 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.
16. 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.
17. Hernandez, A. & Escoriza, D. (2017). Easternmost record of Salamandra algira splendens in Morocco. Boletín de la Asociación Herpetológica Española, 28(1): 60-61.
18. Hernandez, A. (2018). 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: 19-22.
19. Dinis, M., Merabet, K., Martínez-Freiría, F., Steinfartz, S., Vences, M., Burgon, J.D., Elmer, K.R., 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.
20. IUCN SSC Amphibian Specialist Group. (2013). Salamandra algira. The IUCN Red List of Threatened Species 2013.
21. Fahd, S., Martínez-Medina, F.J., Mateo, J.A. & Pleguezuelos, J.M. (2005). Anfibios y reptiles en los territorios transfretanos (Ceuta, Melilla e islotes en el norte de África). In: Pleguezuelos, J.M., Márquez, R. & Lizana, M. (eds.): Atlas y Libro Rojo de los Anfibios y Reptiles de España. Dirección General de la Conservación de la Naturaleza - Asociación Herpetológica Española, Madrid: 385-436.
22. Escoriza, D. & Ben Hassine, J. (2014). Microclimatic variation in multiple Salamandra algira populations along an altitudinal gradient: phenology and reproductive strategies. Acta Herpetologica, 9(1): 33-41.
23. Bogaerts, S. & Donaire-Barroso, D. (2005). Rival combats in the North Moroccan fire salamander Salamandra algira tingitana. Zeitschrift für Feldherpetologie, 12: 137-141.
24. Martínez-Solano, I., Bosch, J. & García-París, M. (2003). Demographic trends and community stability in a montane amphibian assemblage. Conservation Biology, 17: 238-244.
25. Bosch, J., Martínez-Solano, I. & García-París, M. (2001). Evidence of a chytrid fungus infection involved in the decline of the common midwife toad (Alytes obstetricans) in protected areas of central Spain. Biological Conservation, 97: 331-337.
26. Reques R, Pleguezuelos JM, Busack SD, et al. (2013). Amphibians of Morocco, including Western Sahara: a status report. Basic and Applied Herpetology, 27: 23-50.
27. Bogaerts, S., Valkenburg, K., Donaire-Barroso, D. & Espallargas, G. (2007). New localities of the north African fire salamander (Salamandra algira) at the southern limit of its distribution in Morocco. Zeitschrift für Feldherpetologie, 14: 238-241.