Philip de Pous

Area prioritization and performance evaluation of the conservation area network for the Moroccan herpetofauna: a preliminary assessment

The integration of spatial area prioritization algorithms and species distribution modelling has shown great promise in conservation planning in recent years. However, despite the fact that reptiles and amphibians have the highest threat status of all terrestrial vertebrates, these species are often under-represented in conservation planning. The Kingdom of Morocco possesses the richest and most varied herpetofauna in the Maghreb and the western Mediterranean, and is characterized by high species richness, endemism and number of European relict species. Despite the fact that Moroccan reptiles and amphibians have been the subject of numerous studies by a large number of international herpetologists since the beginning of the 20th century, few or none of these concerned their conservation. This study had three main objectives: (1) to identify those areas that harbour the highest species richness; (2) to evaluate the existing and proposed future ‘important biological and ecological sites’ (SIBES) conservation area network (CAN) with respect to their ability to protect the herpetofauna adequately; and (3) to identify priority areas into which the existing protected areas can be augmented. We used maximum-entropy species distribution modelling to run distribution models for 11 amphibian and 86 reptile species (27.6% endemics and 12.4% threatened) for which we had 2,170 single geographic records. A total of 97 models were used to create a richness map of the Moroccan herpetofauna and thereby detect both areas of high species richness and the distribution patterns of individual species. This map was subsequently used as a basis for performance evaluation of the CAN and area prioritization using the ConsNet conservation planning software initialized by “Rarity” first, while using representation targets of 5% and 10%. Additionally, the proposed future Moroccan CAN (SIBES) was evaluated in terms of its overlay and proximity with ConsNet solutions using visual interpretation and distance measurements in a GIS. Our results show that Moroccan herpetofauna is poorly protected under the existing and future CAN. Prioritization of areas shows that a major increase in conservation area is required to guarantee the persistence of individual herpetofauna species even with a global minimum representation target of only 10%. An increase of the existing CAN is especially needed along parts of the Atlantic coast, in the north-western Mediterranean region, on the north-eastern Moroccan coast, as well as in several areas in the Sahara, notably vast proportions of the Vallée du Haut and Bas Drâa.

Elusive but widespread? The potential distribution and genetic variation of Hyalosaurus koellikeri (Günther, 1873) in the Maghreb

The genetic variability and the potential distribution under past (Last Glacial Maximum; LGM (MIROC and CCSM simulations)) and present conditions were studied for the anguid Hyalosaurus koellikeri, using analyses of two mitochondrial (ND1 and ND2) and one nuclear (PRLR) gene and species distribution modelling (SDM) including 19 geographical coordinates, covering most of its distribution range. Unexpectedly, the genetic results show that H. koellikeri presents a very low level of variability both in the mitochondrial and nuclear genes studied. The present predicted distribution of H. koellikeri revealed a large potential distribution in both north and eastwards directions, with suitable areas predicted in places where the species has never been reported before, as for instance the Rif Mountains in Morocco, as well as into most parts of northern Algeria and Tunisia. The LGM distribution is even larger compared to the present, with a continuous predicted distribution from Morocco to Tunisia, and even into Libya under the MIROC simulation. The results of the genetic and SDM analyses suggest that the now isolated populations from Debdou and Tlemcen have probably been in contact during the LGM, but its absence from both present and past predicted suitable areas is still a mystery. Hyalosaurus koellikeri depends mainly on closed deciduous forests (typically Cedrus atlantica and Quercus sp.) and open deciduous shrubland with high amounts of annual rainfall. The results of this study and the absence of recent sightings of the species outside the core distribution might indicate a regression of the species. Hence, a reevaluation of the conservation status of the species seems warranted.

Age structure and growth in a population of Pelobates varaldii (Anura, Pelobatidae) from northwestern Morocco

Age structure and growth in the Moroccan spadefoot toad, Pelobates varaldii were estimated by skeletochronology and reported for the first time for a population living in NW Morocco. Snout vent length and body mass did not significantly differ between the sexes, although females appeared larger and heavier than males. Age ranged 2-7 years in males (mean age±SD: 4.5±1.2, n = 66) and 2-10 years in females (4.7±2.4, n = 20). The difference in age was not significant between the sexes. The modal age was 5 years for males and 3 years for females. The age distributions significantly differed between the sexes. Von Bertalanffy growth curves showed a similar profile between the sexes although growth coefficient was higher in males than in females. Longevity and growth rates of P. varaldii were compared with those of other species of the genus Pelobates.

Range contraction and loss of genetic variation of the Pyrenean endemic newt Calotriton asper due to climate change

Many studies have identified climate warming to be among the most important threats to biodiversity. Climate change is expected to have stronger effects on species with low genetic diversity, ectothermic physiology, small ranges, low effective populations sizes, specific habitat requirements and limited dispersal capabilities. Despite an ever-increasing number of studies reporting climate change-induced range shifts, few of these have incorporated species’ specific dispersal constraints into their models. Moreover, the impacts of climate change on genetic variation within populations and species have rarely been assessed, while this is a promising direction for future research. Here we explore the effects of climate change on the potential distribution and genetic variation of the endemic Pyrenean newt Calotriton asper over the period 2020–2080. We use species distribution modelling in combination with high-resolution gridded climate data while subsequently applying four different dispersal scenarios. We furthermore use published data on genetic variation of both mtDNA and AFLP loci to test whether populations with high genetic diversity (nucleotide diversity and expected heterozygosity) or evolutionary history (unique haplotypes and K clusters) have an increased extinction risk from climate change. The present study indicates that climate change drastically reduces the potential distribution range of C. asper and reveals dispersal possibilities to be minimal under the most realistic dispersal scenarios. Despite the major loss in suitable climate, the models highlight relatively large stable areas throughout the species core distribution area indicating persistence of populations over time. The results, however, show a major loss of genetic diversity and evolutionary history. This highlights the importance of accounting for intraspecific genetic variation in climate change impact studies. Likewise, the integration of species’ specific dispersal constraints into projections of species distribution models is an important step to fully explore the effects of climate change on species potential distributions.

Phylogeny of the genus Rhynchocalamus (Reptilia; Colubridae) with a first record from the Sultanate of Oman.

The genus Rhynchocalamus comprises three species distributed in Southwest Asia. Little is known about them, most probably because of their secretive fossorial lifestyle. The poor knowledge of the genus is even underscored by the fact that its phylogenetic affinities remained unclear until very recently. The least known of the species, Rhynchocalamus arabicus, is known only from the holotype collected in Aden, Yemen, and it has not been observed since its description in 1933. Here we provide a second record for this species, which represents the first record of this genus for Oman. This extends its range in southern Arabia by more than 1000 km. The observed specimen was determined as R. arabicus on the basis of its similarity in size, color, and scalation with the holotype. Furthermore, we sequenced three mitochondrial (12S, 16S, cytb) and one nuclear (cmos) genes for R. arabicus and for two individuals of R. melanocephalus and one R. satunini and inferred the phylogenetic relationships of all currently recognized species of the genus for the first time. The results of our phylogenetic analyses indicate that Rhynchocalamus is a member of the Western Palearctic clade of Colubrinae and is sister to Lytorhynchus, with which it forms a very well supported clade and shares some morphological characters. As our results show, R. satunini is the basal lineage of the genus and R. melanocephalus is sister to R. arabicus.

Cryptic diversity in Ptyodactylus (Reptilia: Gekkonidae) from the northern Hajar Mountains of Oman and the United Arab Emirates uncovered by an integrative taxonomic approach

The Hajar Mountains of south-eastern Arabia form an isolated massif surrounded by the sea to the east and by a large desert to the west. As a result of their old geological origin, geographical isolation, complex topography and local climate, these mountains provide an important refuge for endemic and relict species of plants and animals. With 19 species restricted to the Hajar Mountains, reptiles are the vertebrate group with the highest level of endemicity, becoming an excellent model for understanding the patterns and processes that generate and shape diversity in this arid mountain range. The geckos of the Ptyodactylus hasselquistii species complex are the largest geckos in Arabia and are found widely distributed across the Arabian Mountains, constituting a very important component of the reptile mountain fauna. Preliminary analyses suggested that their diversity in the Hajar Mountains may be higher than expected and that their systematics should be revised. In order to tackle these questions, we inferred a nearly complete calibrated phylogeny of the genus Ptyodactylus to identify the origin of the Hajar Mountains lineages using information from two mitochondrial and four nuclear genes. Genetic variability within the Hajar Mountains was further investigated using 68 specimens of Ptyodactylus from 46 localities distributed across the entire mountain range and sequenced for the same genes as above. The molecular phylogenies and morphological analyses as well as niche comparisons indicate the presence of two very old sister cryptic species living in allopatry: one restricted to the extreme northern Hajar Mountains and described as a new species herein; the other distributed across the rest of the Hajar Mountains that can be confidently assigned to the species P. orlovi. Similar to recent findings in the geckos of the genus Asaccus, the results of the present study uncover more hidden diversity in the northern Hajar Mountains and stress once again the importance of this unique mountain range as a hot spot of biodiversity and a priority focal point for reptile conservation in Arabia.

Phylogeny and biogeography of Arabian populations of the Persian Horned Viper Pseudocerastes persicus (Duméril, Bibron & Duméril, 1854)

The Persian Horned Viper (Pseudocerastes persicus) is distributed from northeast Iraq through the Iranian Plateau to western Pakistan with isolated populations in the Hajar Mountains of south-eastern Arabia. Like the other members of the genus Pseudocerastes, P. persicus is a sit-and-wait ambush feeder with low vagility, a characteristic that often results in high levels of population differentiation. In order to clarify the level of genetic variability, phylogenetic relationships, and biogeography of the Arabian populations of P. persicus we sequenced 597 base pairs of the mitochondrial cytochrome b of four individuals from the Hajar Mountains in south-eastern Arabia and inferred their phylogenetic relationships including 10 samples of P. persicus from Iran and Pakistan, four P. urarachnoides and one P. fieldi downloaded from GenBank. The four Arabian samples are genetically very similar in the gene fragment analysed and are phylogenetically very closely related to populations of P. persicus from coastal south Iran. Biogeographically, it appears that colonisation of the Hajar Mountains by P. persicus took place from Iran very recently, most probably during the last glaciation, when most of the Persian Gulf was above sea level and did not represent a barrier for dispersal.

Diversity, distribution and conservation of the terrestrial reptiles of Oman (Sauropsida, Squamata)

In the present work, we use an exceptional database including 5,359 records of 101 species of Oman’s terrestrial reptiles together with spatial tools to infer the spatial patterns of species richness and endemicity, to infer the habitat preference of each species and to better define conservation priorities, with especial focus on the effectiveness of the protected areas in preserving this unique arid fauna. Our results indicate that the sampling effort is not only remarkable from a taxonomic point of view, with multiple observations for most species, but also for the spatial coverage achieved. The observations are distributed almost continuously across the two-dimensional climatic space of Oman defined by the mean annual temperature and the total annual precipitation and across the Principal Component Analysis (PCA) of the multivariate climatic space and are well represented within 17 out of the 20 climatic clusters grouping 10% of the explained climatic variance defined by PC1 and PC2. Species richness is highest in the Hajar and Dhofar Mountains, two of the most biodiverse areas of the Arabian Peninsula, and endemic species richness is greatest in the Jebel Akhdar, the highest part of the Hajar Mountains. Oman’s 22 protected areas cover only 3.91% of the country, including within their limits 63.37% of terrestrial reptiles and 50% of all endemics. Our analyses show that large areas of the climatic space of Oman lie outside protected areas and that seven of the 20 climatic clusters are not protected at all. The results of the gap analysis indicate that most of the species are below the conservation target of 17% or even the less restrictive 12% of their total area within a protected area in order to be considered adequately protected. Therefore, an evaluation of the coverage of the current network of protected areas and the identification of priority protected areas for reptiles using reserve design algorithms are urgently needed. Our study also shows that more than half of the species are still pending of a definitive evaluation by the International Union for Conservation of Nature (IUCN).

NA2RE is reliable but aims for improvement: an answer to Vamberger and Fritz (2018)

A recent paper has suggested that NA2RE, the New Atlas of Amphibians and Reptiles of Europe, does not provide a reliable basis for ecological niche modelling studies due to errors flagging introductions and missing data for the native range of the pond turtle genus Emys. We point out that the original NA2RE paper already acknowledged that it was not aimed for fine-scale ecological distribution modelling and that it had the objective of stimulating research for improving the maps. New works now complement the Atlas in improving the coverage and providing new distribution maps for species within species complex. Moreover, we stress that the NA2RE web platform at present hosts only the distribution data compiled in 2014 from different sources, using the taxonomy adopted by the authors at the time. As with any large database, it is advisable that these data are carefully evaluated and quality-filtered before their use in scientific studies. We defend the reliability of the NA2RE web platform as the currently most comprehensive resource for the comparative chorological study of amphibians and reptiles in Europe, and encourage publication of updates and additions following the most recent taxonomic changes, to continuously improve this database and the Atlas.