Déficit de connaissances de la biodiversité et biologie de la conservation : le cas de l’herpétofaune d’Algérie

Algeria is a case study of biodiversity ignorance. To date, there are neither complete inventories for any taxa nor atlas across the country! Yet, Algeria is of great importance for global biodiversity. The coastal area of Algeria is part of the global biodiversity hostpot which is the Mediterranean perimeter and has several regional red spots of biodiversity. Moreover, the Saharan part encompasses a diversity of endemic organisms adapted to the strong conditions of drought. The Algerian authorities are deploying an ambitious project for conservation aiming to reach 50% of the country's surface as protected areas! However, the location of these protected areas and their management don’t obey to a scientific evidence based, but are rather based on experts opinion. The main aim of this thesis is the study of the biodiversity ignorance distribution in Algeria by focusing on the "Linnean, Wallacean, and Darwinian Shortfalls" and to contribute to reduce them. The first chapter aims at reducing the Linnean Shortfall by proposing 1) a first checklist of reptiles and amphibians of Algeria which is the result of an accurate review of scientific publications on these taxa in North Africa. 2) a first pre-assessment of the conservation status of reptiles and amphibians of Algeria for the production of the first national red list. The second part of this manuscript deals with the ignorance of the geographical distribution of species. The purpose of this part is to map the ignorance which is defined as the difference between the expected specific richness diversity and the observed specific richness. The predicted species richness was modelled with two approaches: 1 ecological niche modeling with the maximum entropy method (MaxEnt) to predict the suitable habitats for each species and then add the different binary layers of species presence to calculate the specific richness. 2) Minimum convex polygon method was used to create range maps of each species and were auditioned to obtain predicted species richness. Both approaches have shown the same trend at a national scale: the concentration of the richest areas along the Saharan Atlas and the high plateaux which are the transition zone between the Sahara and the the Mediterranean part of Algeria. The opposition of the Saharan with a low species richness and the northern part with a higher species richness. Finally, the Saharan massifs of the Hoggar and Tassili form a very distinct zone with a much greater wealth compared to the rest of the Sahara. In terms of gaps, in the northern part, Kabylie, El Kala National Park and the Oran region are fairly well surveyed. In the Sahara, only a few points are clearly well prospected as Biskra, Bechar and some areas of the Hoggar and Tassili. The third part deals with the distribution of genetic llineages in the Maghreb. The aim of this section is to locate the phylogeographic breaks between the divergent eastern and western populations of the Maghreb and to try to understand the mechanisms that led to this pattern of genetic diversity. To answer these questions, I carried out a comparative phylogeography on 11 species of reptiles and amphibians with continuous and wide distribution in the Maghreb. Two suture zones have been identified: one zone on the border of Algeria and Morocco, the other in Kabylia west of the Soummam valley. The divergences between the eastern clades and the western clades occurred between the pre-Messinian periods up to the plio-pleistocene boundary and were maintained in climatic refugia in the east and west of the Maghreb.L’Algérie est un cas d’école en matière d’ignorance en biodiversité. A ce jour, on ne dispose d'aucun inventaire complet pour aucun taxon ni aucun atlas à l’échelle du pays ! Pourtant, l’Algérie est d’une grande importance pour la biodiversité mondiale. La façade littorale de l’Algérie fait partie du hostpot de biodiversité mondial qui est le pourtour méditerranéen et compte plusieurs points rouges de biodiversité régionaux. Par ailleurs, la partie saharienne contient une diversité d’organismes endémiques adaptés aux fortes conditions de sécheresse. Les autorités algériennes déploient un projet ambitieux pour la conservation en fixant 50 % de la surface du pays comme objectif pour les aires protégées ! Mais, l’emplacement de ces aires protégées et leur gestion n’obéit pas à des critères basés sur une bonne connaissance de la diversité, mais sont plutôt panifiés à dire d’expert. L’objectif général de cette thèse c’est l’étude de la distribution de l’ignorance en biodiversité en Algérie en se concentrant sur les « Linnean, Wallacean et Darwinian Shortfalls » et de contribuer à les réduire. J’ai consacré un chapitre qui a pour but de réduire le Linnean Shortfall en proposant 1) une première checklist des reptiles et amphibiens d’Algérie qui est le fruit d’un examen précis des publications scientifiques sur ces taxons en Afrique du Nord. 2) une première pré-évaluation des statuts de conservations des reptiles et amphibiens d’Algérie pour la production de la première liste rouge nationale. La deuxième partie de ce manuscrit traite la question de l’ignorance de la distribution géographique des espèces. Le but de cette partie est de cartographier l’ignorance qui est la différence entre la diversité la richesse spécifique attendue et la richesse spécifique observée. La richesse spécifique attendue a été modélisée avec deux approches : 1 l’utilisation des modèles de niches avec la méthode de maximum d’entropie (MaxEnt) pour prédire les habitats favorables pour chaque espèce puis additionner les différentes couches binaires de présence des espèces pour calculer la richesse spécifique. 2) l’addition des couches des aires d’occurrences des espèces construite par la méthode du minimum convex polygon pour produire la carte de distribution de la richesse spécifique. Les deux approches ont montré la même tendance à l’échelle nationale, c’est-à-dire la concentration des zones les plus riches sur le long de l’Atlas Saharien et des hauts plateaux qui sont la zone de transition entre le Sahara et la partie méditerranéenne de l’Algérie. L’opposition de la partie saharienne globalement pauvre en espèce et la moitié nord plus riche. En fin, les massifs sahariens du Hoggar et Tassili forment une zone très distincte avec une richesse nettement supérieure par rapport au reste du Sahara. Pour ce qui est des lacunes, dans la Partie nord, la Kabylie, le parc national d’El Kala et la région d’Oran sont assez bien prospectées. Dans le Sahara, seuls quelques points sont assez bien prospectés comme Biskra, Béchar et quelques zones du Hoggar et Tassili. En fin, la troisième partie porte sur la distribution des lignées génétiques dans le Maghreb. Les objectif de cette partie est localiser les zones de sutures entre les lignées génétiques divergentes des populations de l’est et de l’ouest du Maghreb et d’essayer de comprendre les mécanismes qui ont conduit à ce patron de diversité génétique. Pour répondre à ces questions, j’ai effectué une phylogéographie comparées sur 11 espèces de reptiles et amphibiens à distribution continue et large dans le Maghreb. Deux zones de sutures ont été identifiées : une zone à la frontière de l’Algérie et du Maroc, l’autre EN Kabylie à l’ouest de la vallée de la Soummam. Les divergences entre les clades est et les clades ouest ont eu lieu entre la période pré-messinienne jusqu’au à la limite plio-pleistocène et se seraient maintenues dans des refuges climatiques à l’est et à l’ouest du Maghreb

Biodiversity shortfalls and biodiversity conservation : the case of Algerian herpetofauna

L’Algérie est un cas d’école en matière d’ignorance en biodiversité. A ce jour, on ne dispose d'aucun inventaire complet pour aucun taxon ni aucun atlas à l’échelle du pays ! Pourtant, l’Algérie est d’une grande importance pour la biodiversité mondiale. La façade littorale de l’Algérie fait partie du hostpot de biodiversité mondial qui est le pourtour méditerranéen et compte plusieurs points rouges de biodiversité régionaux. Par ailleurs, la partie saharienne contient une diversité d’organismes endémiques adaptés aux fortes conditions de sécheresse. Les autorités algériennes déploient un projet ambitieux pour la conservation en fixant 50 % de la surface du pays comme objectif pour les aires protégées ! Mais, l’emplacement de ces aires protégées et leur gestion n’obéit pas à des critères basés sur une bonne connaissance de la diversité, mais sont plutôt panifiés à dire d’expert. L’objectif général de cette thèse c’est l’étude de la distribution de l’ignorance en biodiversité en Algérie en se concentrant sur les « Linnean, Wallacean et Darwinian Shortfalls » et de contribuer à les réduire. J’ai consacré un chapitre qui a pour but de réduire le Linnean Shortfall en proposant 1) une première checklist des reptiles et amphibiens d’Algérie qui est le fruit d’un examen précis des publications scientifiques sur ces taxons en Afrique du Nord. 2) une première pré-évaluation des statuts de conservations des reptiles et amphibiens d’Algérie pour la production de la première liste rouge nationale. La deuxième partie de ce manuscrit traite la question de l’ignorance de la distribution géographique des espèces. Le but de cette partie est de cartographier l’ignorance qui est la différence entre la diversité la richesse spécifique attendue et la richesse spécifique observée. La richesse spécifique attendue a été modélisée avec deux approches : 1 l’utilisation des modèles de niches avec la méthode de maximum d’entropie (MaxEnt) pour prédire les habitats favorables pour chaque espèce puis additionner les différentes couches binaires de présence des espèces pour calculer la richesse spécifique. 2) l’addition des couches des aires d’occurrences des espèces construite par la méthode du minimum convex polygon pour produire la carte de distribution de la richesse spécifique. Les deux approches ont montré la même tendance à l’échelle nationale, c’est-à-dire la concentration des zones les plus riches sur le long de l’Atlas Saharien et des hauts plateaux qui sont la zone de transition entre le Sahara et la partie méditerranéenne de l’Algérie. L’opposition de la partie saharienne globalement pauvre en espèce et la moitié nord plus riche. En fin, les massifs sahariens du Hoggar et Tassili forment une zone très distincte avec une richesse nettement supérieure par rapport au reste du Sahara. Pour ce qui est des lacunes, dans la Partie nord, la Kabylie, le parc national d’El Kala et la région d’Oran sont assez bien prospectées. Dans le Sahara, seuls quelques points sont assez bien prospectés comme Biskra, Béchar et quelques zones du Hoggar et Tassili. En fin, la troisième partie porte sur la distribution des lignées génétiques dans le Maghreb. Les objectif de cette partie est localiser les zones de sutures entre les lignées génétiques divergentes des populations de l’est et de l’ouest du Maghreb et d’essayer de comprendre les mécanismes qui ont conduit à ce patron de diversité génétique. Pour répondre à ces questions, j’ai effectué une phylogéographie comparées sur 11 espèces de reptiles et amphibiens à distribution continue et large dans le Maghreb. Deux zones de sutures ont été identifiées : une zone à la frontière de l’Algérie et du Maroc, l’autre EN Kabylie à l’ouest de la vallée de la Soummam. Les divergences entre les clades est et les clades ouest ont eu lieu entre la période pré-messinienne jusqu’au à la limite plio-pleistocène et se seraient maintenues dans des refuges climatiques à l’est et à l’ouest du Maghreb.Algeria is a case study of biodiversity ignorance. To date, there are neither complete inventories for any taxa nor atlas across the country! Yet, Algeria is of great importance for global biodiversity. The coastal area of Algeria is part of the global biodiversity hostpot which is the Mediterranean perimeter and has several regional red spots of biodiversity. Moreover, the Saharan part encompasses a diversity of endemic organisms adapted to the strong conditions of drought. The Algerian authorities are deploying an ambitious project for conservation aiming to reach 50% of the country's surface as protected areas! However, the location of these protected areas and their management don’t obey to a scientific evidence based, but are rather based on experts opinion. The main aim of this thesis is the study of the biodiversity ignorance distribution in Algeria by focusing on the "Linnean, Wallacean, and Darwinian Shortfalls" and to contribute to reduce them. The first chapter aims at reducing the Linnean Shortfall by proposing 1) a first checklist of reptiles and amphibians of Algeria which is the result of an accurate review of scientific publications on these taxa in North Africa. 2) a first pre-assessment of the conservation status of reptiles and amphibians of Algeria for the production of the first national red list. The second part of this manuscript deals with the ignorance of the geographical distribution of species. The purpose of this part is to map the ignorance which is defined as the difference between the expected specific richness diversity and the observed specific richness. The predicted species richness was modelled with two approaches: 1 ecological niche modeling with the maximum entropy method (MaxEnt) to predict the suitable habitats for each species and then add the different binary layers of species presence to calculate the specific richness. 2) Minimum convex polygon method was used to create range maps of each species and were auditioned to obtain predicted species richness. Both approaches have shown the same trend at a national scale: the concentration of the richest areas along the Saharan Atlas and the high plateaux which are the transition zone between the Sahara and the the Mediterranean part of Algeria. The opposition of the Saharan with a low species richness and the northern part with a higher species richness. Finally, the Saharan massifs of the Hoggar and Tassili form a very distinct zone with a much greater wealth compared to the rest of the Sahara. In terms of gaps, in the northern part, Kabylie, El Kala National Park and the Oran region are fairly well surveyed. In the Sahara, only a few points are clearly well prospected as Biskra, Bechar and some areas of the Hoggar and Tassili. The third part deals with the distribution of genetic llineages in the Maghreb. The aim of this section is to locate the phylogeographic breaks between the divergent eastern and western populations of the Maghreb and to try to understand the mechanisms that led to this pattern of genetic diversity. To answer these questions, I carried out a comparative phylogeography on 11 species of reptiles and amphibians with continuous and wide distribution in the Maghreb. Two suture zones have been identified: one zone on the border of Algeria and Morocco, the other in Kabylia west of the Soummam valley. The divergences between the eastern clades and the western clades occurred between the pre-Messinian periods up to the plio-pleistocene boundary and were maintained in climatic refugia in the east and west of the Maghreb

Génétique de la population des Vipères péliades (Vipera berus - Linnaeus, 1758 ; Serpentes : Viperidae) dans la région du Hauts-de-France - apport de la génétique à la conservation locale de l'espèce

Genetics of the population of the common European adder ( Vipera berus - Linnaeus, 1758; Serpentes: Viperidae) in the French region of "Hauts -de- France" - contribution of genetics in the local preservation of the species. Genetic tools are frequently used for the protection and conservation of species. We used genetic markers to investigate the genetic diversity and genetic differentiation between five populations of the adder ( Vipera berus ) in the French Hauts-de-France region. Results demonstrated a limited genetic diversity especially within one population; this one seems genetically isolated, whereas three of the five studied populations displayed a limited but significant genetic differentiation. Historically they must have been connected and exchanged genes. The population with a limited genetic diversity should be specifically monitored, in order to determine if the lack of genetic diversity has an impact on the population dynamic. Résumé L'outil génétique est actuellement d'une grande aide pour la protection et la conservation des espèces. Il a été utilisé pour étudier la diversité et la différentiation génétiques entre cinq populations de Vipères péliades ( Vipera berus Linnaeus, 1758) dans le Pas-de-Calais. Les résultats montrent une diversité assez faible, surtout dans une population ; cette population semble d'ailleurs génétiquement isolée, alors que trois des cinq populations ont une différenciation génétique significative, mais peu marquée ; elles devaient donc historiquement échanger des gènes. Les résultats indiquent qu'une population est à suivre particulièrement, afin de voir si la faible diversité génétique se traduit dans une dynamique limitée au sein de cette population

Comparative phylogeography of amphibians and reptiles in Algeria suggests common causes for the east-west phylogeographic breaks in the Maghreb

International audienceA series of phylogeographic studies in the Maghreb identified a repeated pattern of deep genetic divergence between an eastern (Tunisia) and western (Morocco) lineage for several taxa but lack of sampling in Algeria made it difficult to know if the range limits between the eastern and western lineages were shared among taxa or not. To address this question, we designed a comparative phylogeographic study using 8 reptile and 3 amphibian species with wide distribution in the Maghreb as models. We selected species where previous studies had identified an East-West phylogeographic divide and collected sampled in Algeria to 1) examine whether the simple East-West divergence pattern still holds after filling the sampling gap in Algeria or if more complex diversity patterns emerge; 2) if the E-W pattern still holds, test whether the limits between the E and W clades are shared between species, suggesting that common historical process caused the E-W divergences; 3) if E-W limits are shared between species, use information on the age of the divergence to identify possible geological or climatic events that could have triggered these E-W differentiations. We found that the E-W pattern was generally maintained after additional sampling in Algeria and identified two common disjunction areas, one around the Algeria-Morocco border, the other one in Kabylia (central Algeria), suggesting that common historical mechanisms caused the E-W divergences in the Maghreb. Our estimates for the times to most common recent ancestors to the E and W clades span a wide range between the Messinian salinity crisis and the Plio-Pleistocene limit (except for one older split), suggesting different origins for the initial divergences and subsequent preservation of the E and W lineages in common climatic refugia in the west and the east of the Maghreb

Acanthodactylus montanus Miralles & Geniez & Beddek & Aranda & Brito & Leblois & Crochet 2020, sp. nov.

<i>Acanthodactylus montanus</i> sp. nov. <p>(Appendixes I, III; Figs. 6, 8, 9)</p> <p> <b>Holotype.</b> Muséum national d’Histoire naturelle (Paris) MNHN-RA-2018.0026 (formerly BEV.11794, tissue sample in the BEV tissue collection, code T5771), adult male, collected on May 12 th 2012, 800 m after the Tizi n’Tichka pass towards Marrakech, southwestern High Atlas, Morocco, 31.2879°N, 7.3824°W (WGS84), 2,176 m a.s.l. by Pierre-André Crochet and Raphaël Leblois (Fig. 8A).</p> <p> <b>Paratypes.</b> Nine individuals, BEV.11791 to 11793, 11795 to 11800 (tissue sample codes T5768–5770, 5772– 5777), collected at the type locality at the same date by the same collectors.</p> <p> <b>Additional material.</b> Eight specimens from El Azib n-Iriri, Jbel Siroua, 30.7470°N, 7.6093°W, 2,319 m a.s.l., collected on April 23 rd 2005 by José C. Brito: BEV.10459–10460 (field number JCB.A544 & A547, tissue samples T11829 & T11834) and tissue samples + photos T11826 (JCB.A541), T11827 (A542), T11828 (A543), T11830 (A545), T11831 (A546) and T11833 (A458).</p> <p> We identified the following specimens as <i>A. montanus</i> on the basis of morphology, distribution and ecology: BEV.4973 from 5 km south of Askaoun, Jbel Siroua, 30.69°N, 7.77°W, collected on October 9 th 1974 by Gérard Délye; BEV.4974 from 3 km west of Askaoun, Jbel Siroua, 30.74°N, 7.81°W, collected on October 8 th 1974 by Gérard Délye; BEV.4975 from Tisfaldat mountain, Jbel Siroua, 30.68°N, 7.58°W, 2,460 m a.s.l., collected on March 13 th 1994 by Tahar Slimani.</p> <p> Lastly, a series of specimens exhibit morphological features of <i>A. montanus</i> but were apparently collected at a lower elevation than other specimens (around 1,800 m a.s.l.) and slightly south of the main massif of the Jbel Siroua. They are tentatively identified as <i>A. montanus</i> here (white square in Fig. 10) but this identification will require further data and possibly genetic confirmation before it can be firmly established: BEV.4932–4941 and 14898 from 40 km past Tazenakht along road to Taroudant, approx. 30.455°N, 7.558°W, around 1,800 m a.s.l., collected on April 5 th 1960 by Jacques Bons, Georges Pasteur and Bernard Girot.</p> <p> <b>Etymology.</b> The specific epithet <i>montanus</i> is a Latin adjective meaning “of mountains, belonging to mountains”, the high mountains of the Western High Atlas being the unique habitat where this species has been found until now.</p> <p> <b>Diagnosis.</b> A new species of the <i>Acanthodactylus erythrurus</i> species-group (small flat or carinated dorsal scales; three series of scales around the fingers; three entire supraoculars; 8–10, sometimes 12 straight longitudinal row of ventrals; slightly pectinate toes; undertail and underside of the hind limbs red or reddish in juveniles, subadults and young adults) from the south-western part of the High Atlas and from the Jbel Siroua above 2,000 m a.s.l., characterized by the combination of the following characters: (1) head scalation usually of the <i>“ bellii ”</i> type (subocular in large contact with the upper lip, wedged between the 4 th and 5 th supralabials), one internasal plate, no scales inserted between the prefrontals, but BEV.T11828 has no contact between the left supralabial and the lip; (2) a low number of dorsal scales (47–59 longitudinal rows around the body, mean 54.0), femoral pores (17–23, mean 19.9) on each side and subdigital lamellae (19–22, mean 19.9) on the 4 th toe; (3) one or one and a half row of supraciliary granules on each side with a very reduced number (15–46, mean 25.7) of scales and granules around the 2 nd and 3 rd supraoculars on each side; (4) temporals smooth or slightly keeled; (5) dorsal scales smooth or slightly keeled on the neck and progressively turning to distinctly tectiform or sometimes keeled on the back; (6) stocky proportions, especially in adults, with a relatively short tail (57.9–65.9% of the total length, mean 60.8%, almost always longer for all the other <i>Acanthodactylus</i> of the <i>erythrurus</i> group except <i>lacrymae</i>); (7) base of the tail of adult males very thick; (8) in juveniles, underparts of the tail orange red to intense coral red, slightly paler under the base of the tail, even paler under the rear of the thighs, this colour usually not reaching the rows of femoral pores; (9) some marginal ventral plates yellow of yellowish in adult and subadult males and females, sometimes in juveniles; (10) a series of small round ocelli on the flanks (each one containing 4–9 brightly coloured scales), usually yellow greenish in adults, which tend to disappear completely in adult females.</p> <p> Body coloration similar to the other members of the <i>Acanthodactylus erythrurus</i> group. Juveniles are very dark (black or blackish) with three pale continuous lines on each side of the body (one on lower flanks, one at the junction between the dorsum and the flanks and one on the dorsum joining with the opposite one on the tail base) and a dark vertebral area dissected by a paler narrow vertebral area, frequently split on the nape by dark spots or a dark line and disappearing before the base of the tail; one or two series of pale ocelli are found in the dark interspaces between these stripes (juveniles of the other lineages are usually paler and lack the bifurcation of the pale central vertebral area on the nape). Adults are medium brown with faint pale lines on the dorsum and upper flanks (corresponding to the juvenile pattern) and a better marked line on the lower flanks; irregular dark marks are present on the back between the pale lines, sometimes invading the area on the centre of the dorsum. The flanks typically appear darker due to the large dark areas surrounding each pale ocellus. The pileus presents a complex pattern of pale and dark reticulations in juvenile; in adults it is of the same colour as the back with isolated irregular dark spots.</p> <p> Very similar to the allopatric <i>Acanthodactylus lacrymae</i> and single individuals are not always possible to separate but differs on average by the following characters: (1) in juveniles, parietals are less clearly striped but exhibit a more reticulated pattern; (2) orange red coloration on the undertail fading less toward tail base and more extensive around the cloacal slit; (3) supralabials usually with diffuse dark vertical stripes (uniformly pale creamy, even in juveniles, in <i>A. lacrymae</i>); (4) contrast between the lower border of the flanks and tail side and under surface of the belly and tail usually less marked in adults, especially along the sides of the tail; (5) temporal less flat and dorsal scales slightly killed in adults (smooth in <i>A. lacrymae</i> except sometimes on the rear of the back); (6) some specimens have one and a half row of supraciliary granules on each side (always one row in <i>A. lacrymae</i>) and with a larger number of scales and granules around the 2 nd and 3 rd supraoculars (maximum 46 against maximum 29 in <i>A. lacrymae</i>).</p> <p> <b>Description of the holotype.</b> An adult male (Fig. 8A) measuring 60.0 mm of snout-vent length, 14.3 mm of pileus length, 7.7 mm of pileus width, 8.4 mm of head height, 19.5 mm of forelimb length, 31.0 mm of hindlimb length, with missing tail tip, and having the following scalation features: 57 longitudinal rows of dorsal scales at mid-body which are smooth on the neck but carinated on the back, 25 gular scales along a line from the contact between the fourth pair of maxillary scales to the collar, 26 transversal rows of ventral plates from the collar to the anal plate, 21 and 20 femoral pores on the left and right sides respectively, 20 subdigital lamellae beneath the fourth toe, one and a half row of supraciliary granules on each side, with 41 (left) and 46 (right) scales and granules around the 2 nd and 3 rd supraoculars, one entire internasal, no scale inserted between the two prefrontals, two supralabials in contact with the subocular, the latter one in large contact with the upper lip on each side. Base of the tail very thick.</p> <p>Coloration in life: general ground colour light rufous brown with 3 longitudinal light grey stripes on each side; a series of irregular transverse black marks on the sides of the dorsum between the dorsolateral and dorsal pale lines and dark marks on the edge of the vertebral area between the two dorsal pale lines; a series of 7–8 small pale greenish-yellow ocelli, each surrounded a large black spot that are connected by black coloration, are present along the flanks; pileus roughly of the same colour as the dorsum but more intensely rufous, with very few dark marks; underparts pure white except for the tail and thighs (whitish with pale cream/salmon tinge) and some grey marks on the outer ventrals, 5 spots of yellow-green scales on the outer ventrals/lower dorsals; a poorly contrasted and fragmented dark longitudinal stripe runs along each side of the tail; tail base invaded by yellow-green spots of the same colour as the ones on the flanks and outer belly.</p> <p> <b>Distribution.</b> Known only from high altitudes (between 2,000 m and 2,500 m a.s.l. but precise elevation known for a very small number of localities only) in the southwestern High Atlas (Tizi n’Tichka) and in the Jbel Siroua massif, 60–70 km southwest of the Tzin-n-Tichka area (Fig. 10). Based on observations of animals with a <i>bellii</i> -like subocular scalation in the western High Atlas (Bons and Geniez 1995) and on habitat availability, the distribution probably extends rather continuously from just east of Tizi n’Tichka to Jbel Siroua but maybe not much further east. For example, no member of the <i>Acanthodactylus erythrurus</i> complex has ever been found at Oukaimeden in spite of the availability of similar habitats and of numerous visits by many herpetologists. If specimens of doubtful identity from between Tazenakht and Taroundant prove to be <i>A. montanus</i> (white square in Fig. 10) the distribution would extend to the northernmost parts of the eastern Anti-Atlas.</p> <p> <b>Natural history.</b> At the Tizi n’Tichka locality, the habitat was very similar to the habitats occupied by <i>A. lacrymae</i> further east: earthy slopes with bare gravel ground or short grass interspaced with dense low bushes and some stones. Population density was rather high and the animals were relatively easy to catch (compared to many populations of the IM clade) due to their more confident behaviour and slower escape. The animals were active in sunny and calm weather and their behaviour was typical of the <i>A. erythrurus</i> complex, actively foraging in the open and retreating to the shelter of bushes when disturbed.</p>Published as part of <i>Miralles, Aurélien, Geniez, Philippe, Beddek, Menad, Aranda, Daniel Mendez, Brito, José Carlos, Leblois, Raphaël & Crochet, Pierre-André, 2020, Morphology and multilocus phylogeny of the Spiny-footed Lizard (Acanthodactylus erythrurus) complex reveal two new mountain species from the Moroccan Atlas, pp. 302-326 in Zootaxa 4747 (2)</i> on pages 316-322, DOI: 10.11646/zootaxa.4747.2.4, <a href="http://zenodo.org/record/3694824">http://zenodo.org/record/3694824</a&gt

Acanthodactylus lacrymae Miralles & Geniez & Beddek & Aranda & Brito & Leblois & Crochet 2020, sp. nov.

<i>Acanthodactylus lacrymae</i> sp. nov. <p>(Appendixes I, II; Figs. 6, 8, 9)</p> <p> <b>Holotype.</b> Muséum national d’Histoire naturelle (Paris), MNHN-RA-2018.0027 (formerly BEV.11991, tissue sample in the BEV tissue collection, code T5986), adult male, collected on May 29th 2012 by Philippe Geniez and Aurélien Miralles (Fig. 8B) along the north-eastern shore of the lake of Tislit, north-eastern High Atlas chain, north of Imilchil, Morocco, 32.1986°N, 5.6292°W (WGS84), 2,260 m a.s.l.</p> <p> <b>Paratypes.</b> Fifteen individuals, BEV.11980–11990, 11992–11995 (tissue sample codes T5975–5985, 5987– 5990), collected at the type locality at the same date by the same observers.</p> <p> <b>Additional material.</b> Twelve specimens: BEV.11996–12006 (tissue sample codes T5991–6001) from the western shore of the lake of Izely, Morocco, 32.2182°N, 5.5501°W, 2,276 m a.s.l, collected on May 29 th 2012 by P. Geniez and A. Miralles; photo PGe.1908, live subadult from the type locality.</p> <p> Moreover, we identified a number of specimens as <i>A. lacrymae</i> on the basis of morphology, distribution and ecology: BEV.4922–4923 from the Akkal Ouiad valley, north-west of Anefgou, 32.30°N, 5.42°W, 2,200 m a.s.l., collected on April 4 th 1961 by Georges Pasteur; BEV.4924 from Agoudim, south of Tounfite, 32.3763°N, 5.1797°W, 2,100 m a.s.l., collected on April 4 th 1961 by G. Pasteur; BEV.4927–4931 from a few km north of the Tizi n’Talremt, 32.61°N, 4.51°W, no elevation reported, collected on May 28 th 1962 by Jacques Bons; photo PGe.1541 from Jbel Aberdouz, 2,068 m a.s.l., 32.18222°N, 5.20532°W, photographed on July 7 th 2017 by Michel-Ange Bouchet; BEV.4921, from Aït-Moualil, near Bou-Drâa de l’Oudrhès, around 32.53°N, 5.19°W, at least 1,900 m a.s.l. (eastern High Atlas), collected on April 3 rd 1961 by K. Klemmer and G. Pasteur.</p> <p> In addition, two series of specimens exhibiting morphological features of <i>A. lacrymae</i> originate from outside the main distribution of the new species: we list them here but the identification is tentative only and they are represented by white diamonds in the map in Figure 10: BEV.4964–4966 from Itzer (presumably the area around Itzer), approx. 32.88°N, 5.05°W, no elevation reported, collected on June 5 th 1964 by Mr. Cauro; BEV.4920, from 14 km after Boulemane going towards Midelt, 33.25°N, 4.69°W, no elevation reported, collected on April 2 nd 1961 by G. Pasteur.</p> <p> <b>Etymology.</b> The specific epithet “ <i>lacrymae</i> ”, noun in apposition in the plural form (“tears” in English), comes from a local Berber legend, according to which two young people from different tribes fell in love, but were forbidden to see each other by their families. The grief led them to cry themselves to death, their tears being at the origin of the neighbouring lakes of Tislit (hers) and Isli (his).</p> <p> <b>Diagnosis.</b> A new species of the <i>Acanthodactylus erythrurus</i> species-group (small flat or carinated dorsal scales; three series of scales around the fingers; three entire supraoculars; 8–10, sometimes 12 straight longitudinal rows of ventrals; slightly pectinate toes; tail and underparts of the hind limbs red or reddish in juveniles, subadults and young adults) from the north-eastern part of the High Atlas Mountains of Morocco above 2,000 m a.s.l., characterized by the combination of the following characters: (1) head scalation always of the <i>“ bellii ”</i> type (subocular in large contact with the upper lip, wedged between the 4 th and 5 th supralabials), one internasal plate, no scales inserted between the prefrontals; (2) a low number of scales, especially for the dorsal scales (44–55 longitudinal rows around the body, mean 50.7), the femoral pores (16–23 or 17–23 on each side, means 19.5 and 19.4) and the subdigital lamellae (19–23, mean 19.8); (3) always less than two rows of supraciliary granules on each side and a reduced number of scales around the supraoculars (15–29 on each side, mean 22.05); (4) smooth temporals; (5) smooth dorsal scales on the neck and back, sometimes slightly but distinctly carinated (tectiform) on the back; (6) stocky proportions, especially in adults, with a relatively short tail (45.6–58.5% of the total length, mean 53.9%); (7) base of the tail extremely enlarged and thick in adult males; (8) in juveniles, undertail coral red turning paler toward the base, even paler on the rear of the underside of the thighs, where it does not reach the rows of femoral pores; undertail white in adults of both sexes; (9) some marginal ventral plates yellow or yellowish in adult and subadult males, sometimes in juveniles (but not clearly so in females); (10) a series of small rounded ocelli on each flank (3–10 brightly coloured scales), usually yellowish in adults, rarely grey greenish, which tend to disappear completely in old specimens.</p> <p> Body coloration is similar to the other members of the <i>Acanthodactylus erythrurus</i> group. Juveniles are dark (black or blackish) to brown with three pale continuous lines on each side of the body (one on lower flanks, one at the junction between the dorsum and the flanks continuing along the tail, and one on the dorsum joining with the opposite one on the tail base) and pale mid-vertebral area starting on the nape where it is often entirely divided by a dark line and often disappearing on mid-body or before the tail; one or two series of pale ocelli are found in the dark interspaces between these stripes. Adults are medium brown with faint pale lines on the dorsum and upper flanks (corresponding with the juvenile pattern) and a better marked line on the lower flanks; irregular dark marks are present on the back between the pale lines, often leaving the area on the centre of the dorsum unmarked. The flanks typically appear darker due to the large dark areas surrounding each pale ocelli. In many specimens, the pale dorso-lateral lines continue on the outer edge of the pileus to the back of the eye and continue in a faded way along the tail. The pileus is the same colour as the back, usually with dark spots.</p> <p> The combination of characters listed above separate most specimens from all other members of the <i>A. erythrurus</i> complex (except <i>A. montanus</i> sp. nov. which is extremely similar). Compared to <i>A. montanus</i>, <i>A. lacrymae</i> differs in (1) the parietals and sometimes supraoculars of juveniles with a more continuous and contrasting dark band along their outer edge, creating a more contrasted striped pattern with the pale dorsolateral band continuing through the supratemporals along the sides of the pileus (in <i>A. montanus</i>, parietals have a less distinctly striped pattern of dark marks and light reticulation, see Fig. 9); (2) the colour of the base of the undertail is paler and less extensive around the cloacal slit in juveniles (more intense orange red, more similar to the rest of the tail, and more extensive in juveniles <i>A. montanus</i>, Fig. 9); (3) the supralabials are unmarked, pale creamy even in juveniles (labials have diffuse dark stripes which persist in adults of <i>A. montanus</i>); (4) a sharper and better delimited border between the pale lower surface of the tail and a dark band running along the side of the tail, with a complex pattern in males (in <i>A. montanus</i> the pattern is less contrasted, especially in males).</p> <p> <b>Description of the holotype.</b> An adult male (Fig. 8B) measuring 54.0 mm of snout-vent length, 13.8 mm of pileus length, 7.9 mm of pileus width, 7.5 mm of head height, 16.5 mm of forelimb length, 25.6 mm of hindlimb length, with a relatively short non-regenerated tail (63 mm, representing only 54% of the total length), presenting the following scalation features: 54 longitudinal rows of smooth dorsal scales at mid-body, 23 gular scales counted along a line from the contact between the fourth pair of maxillary scales to the collar, 28 transversal rows of ventral plates from the collar to the anal plates, 21 femoral pores on each side, 20 subdigital lamellae beneath the fourth toe, 1 row of supraciliary granules on each side, with 29 (left) and 28 (right) granules, respectively, one entire internasal, no scale inserted between the two prefrontals, two supralabials in contact with the subocular, the latter in large contact with the upper lip on each side. Base of the tail extremely thick. Coloration in life: general ground colour light rufous brown with 6 longitudinal whitish to light grey stripes; the vertebral area between the two central light stripes is nearly uniform with only some vestigial black marks; a series of black transversal marks on the sides of the back; a series of 10 small light greenish ocelli, each surrounded by a large black spot, along the flanks; pileus of the same colour as the dorsum with some black and light marks corresponding approximately to the extension of the dorsal stripes; underparts pure white, a few marginal ventral plates pale yellow or with a black mark; colour delimitation between the side of the body and the underparts very strong and contrasted; a black longitudinal stripe runs along the side of the tail, the top of this stripe slightly wavy on the anterior part and straight on the rest of the side of the tail.</p> <p> <b>Distribution.</b> Until now, known only with certainty from a handful of localities in a narrow stretch of the eastern High Atlas of Morocco; at high elevations (above 2,000 m a.s.l. for all localities whose elevation is precisely known) between Tislit Lake to the west and the Tizi n’Talremt Lake to the east (Fig. 10). Based on the distribution of animals with the <i>bellii</i> -like subocular conformation in the eastern High Atlas, the potential distribution extends south to the upper Todra Valley (continuous white line in Fig. 10). If the identity of the specimens originating from the surroundings of Itzer and the area south of Boulemane (white diamonds in Fig. 10) is confirmed by further data, the distribution would extend to the southern edge of the Middle Atlas, potentially in sympatry with the local populations of the IM clade (doted white line in Fig. 10).</p> <p> <b>Natural history.</b> <i>Acanthodactylus lacrymae</i> lives on high plateaus above 2,000 m a.s.l. In both of the visited localities the habitat was similar: open, earthy or gravel ground with short grazed vegetation interspaced with barren patches, numerous stones and low shrubs/patches of taller plants (<i>Artemisia herba-alba</i> and <i>Hormathophylla spinosa</i>) that the lizards use as shelters. With its stocky proportions, <i>A. lacrymae</i> runs relatively slowly compared with other members of the <i>A. erythrurus</i> complex, which makes them easier to catch. Both visited populations were dense, with numerous individuals in activity (respectively 17 and 11 specimens caught and others seen but not captured in around 0.01 km ²). These high densities may be explained by the well-preserved habitats typical of high altitudes areas in the region and the absence of direct competitors (the only other observed Lacertid was the North- African wall lizard <i>Podarcis vaucheri,</i> which prefers rocky areas or dense vegetation).</p>Published as part of <i>Miralles, Aurélien, Geniez, Philippe, Beddek, Menad, Aranda, Daniel Mendez, Brito, José Carlos, Leblois, Raphaël & Crochet, Pierre-André, 2020, Morphology and multilocus phylogeny of the Spiny-footed Lizard (Acanthodactylus erythrurus) complex reveal two new mountain species from the Moroccan Atlas, pp. 302-326 in Zootaxa 4747 (2)</i> on pages 314-315, DOI: 10.11646/zootaxa.4747.2.4, <a href="http://zenodo.org/record/3694824">http://zenodo.org/record/3694824</a&gt

Morphology and multilocus phylogeny of the Spiny-footed Lizard (Acanthodactylus erythrurus) complex reveal two new mountain species from the Moroccan Atlas

International audienceWe generated an extensive morphological and multilocus molecular dataset to investigate the taxonomy of Acanthodactylus erythrurus, a widespread species across the Mediterranean and semiarid habitats of the Iberian Peninsula and the Maghreb. Our integrated analyses revealed the existence of at least five basal lineages: (i) an Ibero-Moroccan Glade widespread across Morocco and the Iberian Peninsula, from sea-level up to a maximal known altitude of 1.930 m, (ii) an Algero-Tunisian Glade, distributed in coastal and inland areas of eastern Algeria and Tunisia, (iii) a Central Algerian Glade, formed by two inland populations located in central Algeria (1,000-1,500 m a.s.l.), (iv) a western High Atlas Glade including two montane populations from Jbel Siroua and Tin n'Tichka (at 2,320 m and 2,176 m a.s.l.. respectively) and (v) an eastern High Atlas clade, including at least two montane populations from Isli and Tislit (both localities around 2,275 m a.s.l.). An integrated species delimitation approach combining molecular and multivariate morphological analyses demonstrated complete reproductive isolation and hence speciation between the Ibero-Moroccan Glade and the eastern High Atlas Glade in their contact zone. The divergence between all five Glades is broadly similar, supporting the existence of at least five species in the Acanthodactylus erythrurus complex. In the present work we describe the two well-differentiated endemic species from the Moroccan Atlas for which no name is available: Acanthodactylus lacrymae sp. nov. from Isli and Tislit and A. montanus sp. nov. from Jbel Siroua and Tizi n'Tichka Further work will be needed to fully resolve the taxonomy of this species complex