Salamandra Algira Research Articles

First Mention of Cannibalism in Salamandra Algira (Caudata, Salamandridae) Larvae in Algeria with Some Ecological Notes.

Animal cannibalism is a practice well known to scientists, it consists of partially or entirely eating a living individual of the same species by necessity or by instinct. Although originally thought to be abnormal behavior, it has now been recorded in over 1300 species across a diverse group of taxa, ranging from protists and invertebrates to humans. This type of behavior exists in amphibians in different forms and it is observed in 12 families of toads, 7 families of salamanders and 1 family of gymnophionas (Lissamphibians) and this phenomenon is known in natural populations of Salamandridae larvae. The only representative of this family and genus Salamandra in North Africa is Salamandra algira with five subspecies, of which only one is reported for Algeria (Salamandra algira algira) and cannibalism among Salamandra algira larvae has been described for Morocco at Jbel Tazzeka in the Middle Atlas where gigantism has also been observed in certain larvae whose size can vary from 25 to 74 mm. In this study we report for the first time the practice of cannibalism in Algeria in S.A. algira and we also observed gigantism in some larvae. Regarding salamanders, there is still a lot to do because their distribution and ecology are not well known.

Multiple paternity in the pueriparous North African fire salamander, Salamandra algira, supports polyandry as a successful mating strategy in low fecundity Salamandra lineages

Abstract Multiple paternity is widespread in nature and despite costs, it has many associated benefits like increased genetic diversity and fertilization success. It has been described in many viviparous systems, suggesting the existence of some fitness advantages counteracting the inherent costs of viviparity, such as fecundity reduction and high parental investment. Reproductively polymorphic species, like the urodele Salamandra algira, which shows two types of viviparity: larviparity (i.e., delivering aquatic larvae), and pueriparity (i.e., delivering terrestrial metamorphosed juveniles), are suitable systems to study the relationship between reproductive modes and polygamous mating. Here, multiple paternity is confirmed in a pueriparous lineage of S. algira, as previously verified for the pueriparous lineages of the reproductively polymorphic species S. salamandra, suggesting polyandry is a successful mating strategy in pueriparous systems with reduced brood sizes. We discuss the potential benefits of polyandry in the context of viviparity evolution in urodeles.

Phylogenomic inference of species and subspecies diversity in the Palearctic salamander genus Salamandra

The salamander genus Salamandra is widespread across Europe, North Africa, and the Near East and is renowned for its conspicuous and polymorphic colouration and diversity of reproductive modes. The phylogenetic relationships within the genus, and especially in the highly polymorphic species S. salamandra, have been very challenging to elucidate, leaving its real evolutionary history and classification at species and subspecies levels a topic of debate and contention. However, the distribution of diversity and species delimitation within the genus are critically important for identifying evolutionarily significant units for conservation and management, especially in light of threats posed by the pathogenic chytrid fungus Batrachochytrium salamandrivorans that is causing massive declines of S. salamandra populations in central Europe. Here, we conducted a phylogenomic analysis from across the taxonomic and geographic breadth of the genus Salamandra in its entire range. Bayesian, maximum likelihood and network-based phylogenetic analyses of up to 4905 ddRADseq-loci (294,300 nucleotides of sequence) supported the distinctiveness of all currently recognised species (Salamandra algira, S. atra, S. corsica, S. infraimmaculata, S. lanzai, and S. salamandra), and all five species for which we have multiple exemplars were confirmed as monophyletic. Within S. salamandra, two main clades can be distinguished: one clade with the Apenninic subspecies S. s. gigliolii nested within the Iberian S. s. bernardezi/fastuosa; and a second clade comprising all other Iberian, Central and East European subspecies. Our analyses revealed that some of the currently recognized subspecies of S. salamandra are paraphyletic and may require taxonomic revision, with the Central- and Eastern-European subspecies all being poorly differentiated at the analysed genomic markers. Salamandra s. longirostris – sometimes considered a separate species – was nested within S. salamandra, consistent with its subspecies status. The relationships identified within and between Salamandra species provide valuable context for future systematic and biogeographic studies, and help elucidate critical evolutionary units for conservation and taxonomy.

A new subspecies of African fire salamander Salamandra algira (Urodela, Salamandridae) from the Middle Atlas Mountains, Morocco

A new subspecies within the Salamandra algira species complex from north-western Africa is described. Previous molecular analysis showed that the populations of S. algira splendens from north-western Morocco consisted of two well supported clades: clade 1 distributed in the Rif Mountains, from Chefchaouen (type locality) to Al Hoceima, and clade 2, located southern from clade 1 being isolated in the northern and central Middle Atlas Mountains. Clade 2 is herein described as a distinct subspecies: Salamandra algira atlanticassp. nov. based on morphological data, allopatric range and molecular divergence. This new subspecies shows an uncorrected pairwise distance of 0.0265 from clade 1 based on cytochrome b DNA sequences. Salamandra algira atlanticassp. nov. is a slender and large sized salamander with a highly variable colouration pattern. It can be distinguished from S. algira splendens by the greater proportion of coral red in the background colouration, being the only known subspecies of S. algira in which coral red can exceed the proportion of black. Variable number (0–5) of yellow to golden yellow dorsal blotches, but usually in lower numbers than the nominotypical subspecies. Salamandra algira atlanticassp. nov. inhabits subhumid to humid forests and karstic systems at mid to high elevations. We briefly discuss the phylogenetic and taxonomic issues among the genus Salamandra which encompasses more valid species than currently recognised.

Allopatric diversification and evolutionary melting pot in a North African Palearctic relict: The biogeographic history of Salamandra algira

North Africa is a climatically and topographically complex region with unique biotic assemblages resulting from the combination of multiple biogeographic realms. Here, we assess the role of climate in promoting intra-specific diversification in a Palearctic relict, the North African fire salamander, Salamandra algira, using a combination of phylogenetic and population genetic analyses, paleoclimatic modelling and niche overlap tests. We used mitochondrial DNA (Cyt-b), 9838 ddRADseq loci, and 14 microsatellite loci to characterize patterns of genetic diversity and population structure. Phylogenetic analyses recover two major clades, each including several lineages with mito-nuclear discordances suggesting introgressive patterns between lineages in the Middle Atlas, associated with a melting pot of genetic diversity. Paleoclimatic modelling identified putative climatic refugia, largely matching areas of high genetic diversity, and supports the role of aridity in promoting allopatric diversification associated with ecological niche conservatism. Overall, our results highlight the role of climatic microrefugia as drivers of populations’ persistence and diversification in the face of climatic oscillations in North Africa, and stress the importance of accounting for different genomic regions when reconstructing biogeographic processes from molecular markers.

First report of Euryhelmis parasites (Trematoda, Heterophyidae) in Africa: conservation implications for endemic amphibians.

In this study, we report, through molecular identification, the first African records of a digenean trematode parasite of the genus Euryhelmis. We recovered metacercariae encysted in an anuran, the endemic Moroccan painted frog (Discoglossus scovazzi), and a vulnerable caudate, the North African fire salamander (Salamandra algira), from four localities in North Africa (Morocco). Our records go back to the past century and have been confirmed in successive fieldwork seasons thereafter. Metacercarial stages of these parasites require amphibians as the last intermediate host, but the exact identity of the primary hosts and predators of the infected animals in Africa remain unknown. Our searches with basic local alignment search tool (BLAST) from Genbank revealed that hosts were infected by parasites of Euryhelmis costaricensis, which showed almost the same genetic identity (with only one substitution) to previous reports from Costa Rica and Japan, suggesting a recent introduction in Morocco. We proceed to discuss the likely role of introduced mustelids as the potential definitive hosts of trematode adults. Under this assumption, we conclude that the infestation of Discoglossus scovazzi and Salamandra algira might pose a risk to these threatened species.

Phylogeny, age structure, growth dynamics and colour pattern of the Salamandra algira algira population in the Edough Massif, northeastern Algeria

The distribution of the Fire Salamander in North Africa is discontinuous and the Edough Peninsula, Algeria, is considered as the eastern edge of the distribution area. In the current study, we establish a description of the Salamandra algira algira population in its type locality. In this context, an analysis of the mitochondrial DNA D-loop of 47 sequences comes to confirm the phylogenetic status of our population with regard to the other Algerian and Moroccan populations. Also, we used the skeletochronological method for establishing the age structure of the population. Maximum longevity reached 18 years, with a high frequency of young adults, which suggests a good survival of the juveniles. The growth of males is faster than that of the females, although the maximum size of the males is 180 mm, while that of the females is 210 mm. The Edough’s salamander’s phenotype is characterized by multiple small white spots dispersed in different parts of the body (belly, sides, legs and throat) and a high number of large red spots. 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.

Phylogeographic relationships and shallow mitochondrial divergence of Algerian populations of Salamandra algira

The North African fire salamander, Salamandra algira, is distributed in Algeria, Morocco and Ceuta (Spanish territory located on the north coast of Africa), but until now rather limited information has been available on the populations across the Algerian part of its range. We here provide a first analysis of the phylogeography of this species in Algeria, based on 44 samples from populations distributed across 15 localities in Central Algeria. We sequenced three segments of mitochondrial DNA, covering 12S rRNA, cytochrome b (Cytb) and the D-loop. The mtDNA sequences of the Algerian samples were strongly different from the Moroccan populations occurring west of the Moulouya River (corresponding to the subspecies S. a. tingitana and S. a. splendens) but sister to the genetically rather similar population from the Beni Snassen Massif in eastern Morocco (subspecies S. algira spelaea). Among the Algerian specimens studied, those from the westernmost site, Chrea Massif, were the sister clade to the remaining populations, and the overall genetic divergence was low, with a maximum of five mutational steps in a 295 bp fragment of cytochrome b.

Sexual dimorphism and age of Mediterranean salamanders

We analysed sexual size dimorphism (SSD) for two Mediterranean species of the “true” salamander clade possessing distinct life histories (Salamandra algira and Mertensiella caucasica) and equilibrated the morphometric approach to individual age by using skeletochronology. For species that have a short breeding season and live at high altitudes, such as Mediterranean amphibians, the fecundity advantage hypothesis predicts female-biased SSD to maximise reproductive success. Our results showed no SSD in either species; however, morphometric data indicated a male-biased dimorphism in limb (arm and leg) dimensions in both species when compared to body size. Limb dimorphisms are likely related to the particular mating system, which involves an amplexus during spermatophore transfer. Arm length appeared sexually dimorphic during ontogeny both in viviparous S. algira and oviparous M. caucasica. A review on SSD indicated monomorphy of body size as a common lineage-specific pattern among the “true” salamander clade, but also the common presence of other traits such as sexually dimorphic limb proportions.

Microclimatic variation in multiple Salamandra algira populations along an altitudinal gradient: phenology and reproductive strategies

Salamandra algira is one of the southernmost species of the genus , and most of its ecology remains poorly known. We studied the microhabitat conditions of the sites occupied by several populations of S. algira along an altitudinal gradient, and the use of water bodies for reproduction. The microclimate conditions were analysed at six sites in northern Morocco: one site in Beni Snassen massif ( S. algira spelaea ), two in the Middle Atlas and central Rif mountains ( S. algira splendens ), and three in the western Rif mountains and Peninsula Tingitana ( S. algira tingitana ), where a viviparous population also occurs. The microclimate was characterized using temperature and relative humidity data loggers for a period of two years. We also measured the surface area and depth of the water bodies where we found S. algira larvae. Our results showed an autumn-winter reproductive period for all ovoviviparous populations studied . In most of the aquatic habitats examined, larvae appeared between November and March, although this period could extend to May at higher altitudes. Larval abundance and their size variability did not correlate with water body size or microclimate conditions. The decrease in the number of larvae per water body coincided with the existence of suitable conditions for post-metamorphic dispersal. Salamandra algira occurred in regions with moist conditions (annual average relative humidity greater than 64 %) and with mean annual temperatures of 13.6-18.6 °C, but populations were largely segregated along a gradient of humidity, with some showing higher and more constant values than others. The viviparous population occurs in a region with maritime influence and greater microclimate stability than the other sites studied.

Review of the systematics, distribution, biogeography and natural history of Moroccan amphibians.

The amphibian fauna of the Kingdom of Morocco was traditionally regarded as poor and closely related to its European counterpart. However, an increase in research during the last decades revealed a considerable degree of endemism amongst Moroccan amphibians, as well as phenotypic and genotypic inter- and intraspecific divergence. Despite this increase in knowledge, a comprehensible overview is lacking while several systematic issues have remained unresolved. We herein present a contemporary overview of the distribution, taxonomy and biogeography of Moroccan amphibians. Fourteen fieldtrips were made by the authors and colleagues between 2000 and 2012, which produced a total of 292 new distribution records. Furthermore, based on the results of the present work, we (i) review the systematics of the genus Salamandra in Morocco, including the description of a new subspecies from the Rif- and Middle Atlas Mountains, Salamandra algira splendens ssp. nov.; (ii) present data on intraspecific morphological variability of Pelobates varaldiiand Pleurodeles waltl in Morocco; (iii) attempt to resolve the phylogenetic position of Bufo brongersmai and erect a new genus for this species, Barbarophryne gen. nov.; (iv) summarize and assess the availability of tadpole-specific characteristics and bioacoustical data, and (v) summarize natural history data.

Amphibians of Morocco, including Western Sahara: a status report

Morocco has one of the highest rates (28.6%) of amphibian endemism among countries bordering the Mediterranean Sea and, while large areas of Morocco are crucial for conserving amphibian biodiversity, some areas are not afforded legal protection. We examine biodiversity, identify immediate anthropogenic threats, discuss critical habitat for the conservation of amphibian diversity and the role of currently protected areas in meeting conservation goals within Morocco, Western Sahara included. The study area harbours 14 amphibian species, eight of which are assigned to the categories of Endangered (Pelobates varaldii), Vulnerable (Salamandra algira, Amietophrynus xeros, and Hoplobatrachus occipitalis) or Near Threatened (Pleurodeles waltl, Alytes maurus, Bufo spinosus, and Barbarophryne brongersmai) using IUCN criteria at the regional level of the study area. Habitat loss and degradation due to conversion of land for agriculture, urbanization, or industry are major threats, but infrastructure for tourism, freshwater pollution by chemicals, introduction of non-native species to aquatic ecosystems (Gambusia holbrooki), pathogens (Batrachochytrium dendrobatidis), road-kills, and natural disasters (drought), are also rapidly increasing threats. In addition, consequences from global warming must also be considered. The present Conservation Area Network (CAN) does not include distributional ranges of some amphibian species, and a more complete CAN in Atlantic and desert areas is suggested. The northwestern Atlantic, Rif-Middle Atlas, Central Atlantic, and Tiris regions should be considered priorities for conservation because of amphibian endemism and/or the existence of isolated amphibian populations.

A precipitous decline of the Algerian Newt Pleurodeles poireti Gervais, 1835 and other changes in the status of amphibians ofNumidia, North-Eastern Algeria

The worldwide erosion of biological diversity has not spared amphibians which are amongst the most affected taxa with numerous extinct or near-extinct species. Our knowledge of the status and ecology of amphibians of Algeria is still limited whereas local natural habitats are fast disappearing under a strong anthropogenic pressure. Eighty two localities were sampled between 1996 and 2010, to survey the amphibian community within Numidia, North-Eastern Algeria. Nine species, three Urodeles (Pleurodeles poireti, Pleurodeles nebulosus and Salamandra algira and six Anurans (Bufo bufo spinosus, Bufo mauritanicus, Bufo viridis, Discoglossus pictus, Hyla meridionalis and Pelophylax saharicus) were found within the region. Results suggest a marked, rapid and unexplained crash of the Algerian Newt Pleurodeles poireti, a species endemic to a small area in and around Djebel Edough (Annaba). Information on the status (abundance and distribution) of each recorded taxon is provided and conservation measures are discussed in the light of an apparent local decline of Anurans and the vulnerability of three species (S. algira, P. nebulosus and B. bufo spinosus).

Biogeography and contemporary climatic differentiation among Moroccan Salamandra algira

The mtDNA work was funded by grant CGL2009-11663/BOS from the Ministerio de Educacion y Ciencia, Spain. S.C. is a member of the Grup de Recerca Emergent of the Generalitat de Catalunya: 2009SGR1462.