Taxonomic Notes
Based on traditional morphological characters Vipera monticola was recognized - first as a subspecies, and subsequently as a full species - as a distinct form occupying the High Atlas Mountains, with remaining North African members of the group retained within V. latastei.
Following research that found this arrangement to be inconsistent with genetic results (Freitas et al. 2020), Martínez-Freiría et al. (2021) revised the taxonomy of this group using an integrative analysis based on phylogenetic, morphological and ecological evidence. The resulting species concepts restricted Vipera latastei to the Iberian Peninsula, with all North African lineages referred to V. monticola, and this arrangement was supported with later genomic evidence (Dufresnes et al. 2024).
The resulting concept comprises three subspecies: V. m. monticola (central Moroccan High Atlas); V. m. atlantica (Western High Atlas); and V. m. saintgironsi in the remainder of the range (Martínez-Freiría et al. 2021).
Justification
This species is listed as Vulnerable under criterion B on the basis that this species has an area of occupancy calculated as 1,160 km² (including both historical and recent records), it is considered to occur as a severely fragmented population, and there is an observed continuing decline in the extent and quality of its habitat in both Morocco and Algeria driven by multiple processes, and a projected range-wide decline in the availability of suitable habitat as a result of climate change. It may now be extinct at the lowest elevations within its range, from which all records are historical and remaining habitat is scarce and degraded, and good subpopulations appear to remain only at high elevations. Based on recent records, the extant area of occupancy may be below 500 km², and further research may reveal that this species warrants listing in a higher category.
Geographic Range Information
This species occurs in several distribution centres in Mediterranean North Africa and the mountain ranges of the Maghreb (Martínez-Freiría et al. 2021), a distribution consistent with the hypothesis that the species is a Mediterranean relict (Freitas et al. 2018).
Most records are from Morocco, to which two of the three recognised subspecies are endemic, with a disjunct distribution centre in eastern Algeria (Tellian Atlas and the Aurès Mountains) (Martínez-Freiría et al. 2021). Records reported by Martínez-Freiría et al. (2023) - based on both field surveys and expert-confirmed citizen science data - increased the extent of its Moroccan range by 8.8% and confirmed its occurrence in Jbel Bou Naceur, where its occurrence was suggested as being likely by Brito et al. (2011). It is likely that surveys of the suitable high-elevation areas in the western High Atlas would reveal additional subpopulations (Martínez-Freiría et al. 2023). It is likely to be extinct in Salé, and a 1926 report from Ifrane is probably in error as the species has never been recorded again in this area (F. Martínez-Freiría pers. comm. 2024). Neither record is included on the accompanying map.
It is found at elevations up to 3,650 m asl (a reported maximum elevation of 3,900 m asl. in Martínez del Mármol et al. 2019 is an overestimate - F. Martínez-Freiría pers. comm. 2024). Most records in the last 15 years are from above 2,600 m asl, although it has been recorded as low as 450 m asl. in the Rif Mountains (Brito et al. 2011, Freitas et al. 2018). It formerly occurred as low as sea level, but at least in Morocco coastal subpopulations are believed to be extinct (Brito et al. 2011, Martínez-Freiría et al. 2021).
Records of Vipera latastei from Tunisia and from elsewhere in Algeria are believed to represent this species, but their identity requires genetic confirmation (Martínez-Freiría et al. 2021). Despite targeted and intensive searches in Tunisia it had not been recorded from this country in the 55 years prior to 2009 (J.M. Pleguezuelos pers. comm. 2008) and it has not since been recorded during surveys of historical Tunisian localities (Freitas et al. 2018).
Its area of occupancy is calculated as 1,160 km² (including both historical and recent records), although based on recent records, the extant area of occupancy may be below 500 km², but further research is needed to confirm this.
Population Information
It is thought to be a rare species. The low frequency of records even in relatively well-sampled areas suggest that it occurs at low population densities (Brito et al. 2011, Martínez-Freiría et al. 2023). Local extinction following tourist development has been proposed in the three known coastal subpopulations in Morocco, although Brito et al. (2011) caution that this is a secretive species and accurate determination of local extinction is therefore difficult. Subpopulations at mid-elevations are believed to be in decline as a result of high levels of habitat degradation (Freitas et al. 2018). It presumably occurs in "relatively good populations" at higher elevations in the Rif and Middle Atlas (above 1,500 m asl) (Freitas et al. 2018). In the High Atlas healthy subpopulations are restricted to areas above 2,100 m asl (F. Martínez-Freiría pers. comm. 2024). Of 136 records compiled by Freitas et al.(2018), more than 40% with associated dates were collected between 1950 and 2000 (mean 1.12 records / year) and 25.75% (mean two records / year) since, suggesting an increasing rate of detection. Surveys by these authors in around 55% of recorded localities found only five individuals, all in montane areas. The data compiled by Martínez-Freiría et al. (2023) consists of 45 records across five 10 x 10 km cells collected between 2017 and 2022, almost half of them from previously known localities in the Central High Atlas.
Due to the combination of low dispersal abilities and its reliance on fragmented areas of largely unmodified habitat, this species is likely to experience severe climate change-linked declines resulting from reduced habitat suitability and these may already be underway (Brito et al. 2011, Martínez-Freiría et al. 2013). Even within suitable montane habitat where the species survives it species exhibits a patchy distribution, with records being isolated from one another, and it is likely to be extinct in intervening lowland areas (F. Martínez-Freiría pers. comm. 2024). As the species is expected to exhibit low dispersal ability, the population is considered to be severely fragmented (F. Martínez-Freiría pers. comm. 2024).
Habitat and Ecology Information
This species is largely restricted to Mediterranean humid and subhumid habitats (Brito et al. 2011). The three recognised subspecies appear to vary in their habitat preferences: the subspecies restricted to Morocco both occur in high-elevation xeric thorny scrub and grassland in rocky areas, and the more widespread subspecies Vipera monticola saintgironsi is also found in areas dominated by either evergreen (cedar or fir) or oak forest (Martínez-Freiría et al. 2021). The latter subspecies historically occurred in or around coastal sand dunes, but is thought likely to be extinct in these areas (Martínez-Freiría et al. 2021). Its habitats are characterised by moderate to high solar radiation and temperate - or extremely marginally subtropical - climates (with mean annual temperatures from 3-19 °C) (Martínez-Freiría et al. 2021). V. m. saintgironsi occurs across varied precipitation regimes, but the Moroccan endemic subspecies occur in areas with low or seasonal rainfall (Martínez-Freiría et al. 2021). Species distribution modelling indicates that the species is most likely to occupy areas with mean annual precipitation between 1,000 and 1,500 mm (Freitas et al. 2018).
High-elevation subpopulations are specialist feeders on lizards (lacertids, skinks, and diurnal geckos), although those at lower elevations may also feed on small mammals (Martínez del Mármol et al. 2019). These authors include amphibians in the diet, but there are no published observations of predation on amphibians (F. Martínez-Freiría pers. comm. 2024). It is viviparous, with litters typically containing two to eight, but sometimes only a single, young during April and May (Martínez del Mármol et al. 2019). The related V. latastei has a maximum recorded longevity in the wild of 11 (male) and 14 (female) years, with sexual maturity reached at four and five respectively (Brito and Rebelo 2003). Generation length in that species is estimated at nine years (F. Martínez-Freiría pers. comm. 2022).
Threats Information
This species exhibits traits that increase its sensitivity to local extinction as it is slow-growing, exhibits low reproductive frequency and dispersal capability, and is a dietary specialist in at least part of its range (Brito et al. 2011). Many recent records have been taken from montane areas where habitat loss has been limited and grazing was partially controlled, most often within protected areas (Brito et al. 2011, Martínez-Freiría et al. 2023). It is likely to have suffered historical and prehistoric local extinctions and fragmentation as a result of massive landscape transformation over several thousand years (Freitas et al. 2018). Outside protected areas, deforestation resulting from timber extraction, wildfires, cannabis cultivation, establishment of pine plantations (which are replacing oak woods in many mid-elevation localities; Freitas et al. 2018), and overgrazing by livestock threaten remaining habitats in both its known range and northwest Tunisia (summarised by Brito et al. 2011). Across the species' range more than 98% of 10 x 10 km cells containing suitable habitat contain human-modified habitats, mainly cropland, rangeland and villages (Freitas et al. 2018). Rangeland dominates the species' range in the Moroccan Atlas, while the Rif and Algerian Tell Atlas are most transformed by cropland and villages (Freitas et al. 2018). Potentially suitable habitats in the Rif, and within lowland areas of its historical range, suitable habitat is scarce and frequently degraded (Brito et al. 2011). In the Kabilye region, which contains the largest area of suitable habitat for this species in Algeria, an additional threat is the capture of springs at their source to support local villages (M. Beddek pers. comm. 2024). In the forested massif of Akfadou, more than 90% of springs have been captured (M. Beddek pers. comm. 2024). This practice results in mountain streams drying earlier in the year, affecting all amphibians and water-dependent snakes (M. Beddek pers. comm. 2024).
Sampling in areas with recent records failed to record the species in localities where high levels of habitat degradation was observed (Martínez-Freiría et al. 2023). These areas were characterised by deforestation caused by wood extraction and wildfires, aridification resulting from water extraction to irrigate cannabis plantations and by overexploitation of waterbodies by domestic livestock, and overgrazing resulting from increases in stocking levels (Martínez-Freiría et al. 2017, Martínez-Freiría et al. 2023). All of these pressures are reported to be increasing by local shepherds, likely related to strong temperature rises and increasing droughts in the Mediterranean region (Martínez-Freiría et al. 2023). Persecution is also believed to be negatively impacting this species (Martínez-Freiría et al. 2021).
Brito et al. (2011) used maximum entropy modelling to predict habitat suitability for the subspecies Vipera monticola saintgironsiunder an ensemble of climate models, and predicted mean decreases in the extent of its core habitat (defined as areas where the probability of occurrence is greater than 50%) by 56.9% by 2050 and 89% by 2080, and corresponding increases in unsuitable habitat by 26.1 and 36.8%. Between 2011 and 2020 the subspecies was projected to have lost 18.1% of its core habitat, and of 32.2% in habitats where the probability of occurrence between 25 and 50% (Brito et al. 2011). A multi-algorithm correlative approach was used to infer climate change impacts on the potential distribution of endemic reptiles to Morocco by Martínez-Freiría et al. (2013) predicted a mean decrease in suitable habitats for the High Atlas subpopulations (i.e. V. m. monticola and V. m. atlantica) ranging from 31-35% in 2050 and 37-55% in 2080. None of these authors' models identified any areas of newly suitable habitat resulting from climate change. Subsequent research suggests however that both pieces of work underestimated the extent of the species' potential climatic niche (Freitas et al. 2018).
Records of two generalist, warm-adapted snakes, the Montpelier Snake (Malpolon monspessulanus) and Horseshoe Whipsnake (Hemorrhois hippocrepis), from high elevations in the central and High Atlas appear to be evidence of either human-mediated dispersal or range shifts resulting from climate change (Scaramuzzi et al. 2022). The former species has been observed feeding on young vipers of a related species elsewhere in its range, and the impacts of both species on the North African Atlas Dwarf Viper warrant special attention in future research (Scaramuzzi et al. 2022).
The species was reported to be less common in coastal and agricultural areas than in mountains by Brito et al. (2011), and Freitas et al. (2018) found that it tends to occupy areas with slopes between 12 and 20%. Brito et al. (2011) hypothesise that this is a consequence of habitat loss, which is most extensive in flatter areas. The extinction of known coastal subpopulations in Morocco may have occurred by the early 2000s, as intensive surveys reported from 2005 onwards have failed to record it and these areas have all been subject to urban development associated with tourism (summarised by Brito et al. 2011).
Use and Trade Information
There is no information on any use of or trade in this species.
Conservation Actions Information
This species is known from three large biosphere reserves in Morocco: the Arganeraie; Oasis Du Sud Marocain; and the northern Intercontinental Biosphere Reserve (Freitas et al. 2018). It is likely to be highly reliant on continued conservation in protected areas in mountains and cedar forests for its long-term persistence, as these are predicted to become climatic refugia (Cheddadi et al. 2009, Brito et al. 2011, Martínez-Freiría et al. 2013). Nevertheless, between 68 and 92% of suitable habitat across its range lies outside protected areas, and suitable habitat in Algeria is almost completely outside protected areas (Freitas et al. 2018). Population monitoring and surveys of areas where habitat suitability was predicted to decline by 2020 should be conducted to evaluate the accuracy of these predictions (Brito et al. 2011, Martínez-Freiría et al. 2013). If these authors' results are confirmed, strict protection should be enacted for all remaining subpopulations along with appropriate management actions including controls on overgrazing and potentially population translocation (Brito et al. 2011).
Further surveys are needed in areas with high predicted climatic and habitat suitability to clarify the status of subpopulations here (Freitas et al. 2018). Due to the disjunct nature of this species' subpopulations, which are largely isolated in individual mountain ranges, Martínez-Freiría et al. (2021) advocate local management of individual subpopulations or distribution centres at risk, potentially through identifying Evolutionarily Significant Units (ESUs), and wider genetic sampling across its range to delimit ESUs. Population monitoring is recommended in Jbel Oukaimeden, where the relatively high numbers of vipers recorded and ease of access are likely to make this feasible (Martínez-Freiría et al. 2023). This work should also collect data on ecological traits and life history data that can inform conservation actions to guarantee the viability of this and other subpopulations (Martínez-Freiría et al. 2023).