×

Morocco is one of the richest countries in the Mediterranean Basin, comprising 120 terrestrial amphibians and reptiles, of which 30 are endemic species (25%). Remarkably, the genus Chalcides hits top record with nine endemic species out of 16 existing species. Updated distribution maps were elaborated for each species. A new species was added to the taxonomic list of endemics of Morocco. Three main distribution patterns regarding the occupied area and number of localities were identified: 1/ large, extensive, or widespread, 2/ punctual, and 3/ restricted distribution. Three main distribution patterns regarding the occupied region were identified: 1/ mountain, 2/ Atlantic, and 3/ southern Morocco. The areas apparently accommodating most of the endemic species are located within the Tangitan Peninsula, Atlantic coast, Rif Massif, and High and Middle Atlas.

×

This study reports the observations of 54 species of amphibians and reptiles obtained during four field surveys to Morocco, including the southern and southeastern regions. Our records reveal a notable expansion of the current distribution range for several species especially in the eastern part of the country, highlighting the need for more intensive sampling within this region.

×

In the herpetological collection of ZFMK 528 scientific species group names are represented by type materi- al. Of these, 304 names are documented by primary type specimens (onomatophores) while for 224 further names sec- ondary type specimens (typoids) are available, ranging chronologically from 1801 to 2010. The list is a shortened pred- ecessor of a comprehensive type catalogue in progress. It lists name bearing types with their catalogue numbers includ- ing information on further type series members also in other institutions, while secondary types are listed only by pres- ence, both in ZFMK and other collections including holotype repositories. Geographic origin and currently valid names are also provided.

×

he systematics of the genus Acanthodactylus was classically based on external morphological traits, osteological characters and morphology of the hemipenes. Although the identification of sepcies complexes is relatively easy, the destinction within some groups is difficult due to a high variability of the external morphology. Partial mitochondrial (12S and 16S rRNA) sequences (371 and 499 base pairs, respectively) were analysed from 32 specimens of the A. pardalis group from northern Africa including the described species A. busacki, A. maculatus, A. mechriguensis and A. pardalis. Several highly distinct genetic units were resolved, but with little support for relationships between them. These units did not coincide with current taxonomic units, but showed geographic structuring. Although the A. pardalis group displays significant variation, the present taxonomy of the group must be considered unsatisfactory since it is not supported by genetic evidence. For some forms, such as A. mechriguensis there is no support and it is suggested that it should by synonymized with A. maculatus. More data are clearly needed for other forms. Complex microevolutionary patterns due to the recent contraction/expansion phases of the Sahara Desert probably are related with the phylogenetiv patterns observed.

×

The list of the amphibians and reptiles of Western Sahara includes 46 species (4 amphibians and 42 reptiles), while 13 further species unknown in this area could conceivably be discovered there. Among the important facts, we exclude Tarentola boehmei from the fauna of Western Sahara, and report on the effect of the Jbel Ouarkziz which acts as a biogeographical boundary, separating the herpetofaunal elements from Morocco (north of the Jbel Ouarkziz) and from Western Sahara. The herpetofauna of Western Sahara is composed as follows: 10 species can be considered as being of Mediterranean affinities, 2 as North Saharan, 19 as purely Saharan, 7 as being of Sahelian affinities, 4 are tropical relicts and 4 are limited to the oceanic fringe of the Sahara and can be considered as `continental Macaronesian endemics`. This confirms that the herpetofauna of Western Sahara is mainly of Saharan origin but that this area is, in spite of its apparently uniform landscape, a biogeographical cross-roads for Amphibians and Reptiles. This can be related to the wide latitudinal amplitude of Western Sahara and by the climate contrast between the oceanic fringe, comparatively humid and temperate, and the remaining of the country, subject to a dry Saharan climate with strongly contrasted temperature.

×

The evolutionary diversification of many terrestrial vertebrate groups is strongly linked to climatic events in the Cenozoic, the period from 65 Million years ago to today when modern animals first appeared. I investigated the effects of Cenozoic climate change on the taxonomic and morphological diversification of the Old World lizard family Lacertidae, with particular emphasis on the African radiation. African lacertids exhibit an unusual pattern of diversification, in which their highest species richness occurs in deserts north and south of the equator, despite being spread throughout the continent. This disparity is particularly surprising given that desert lacertids are thought to be evolutionarily younger than their mesic-dwelling relatives, suggesting increased diversification rates in arid habitats. To identify the evolutionary factors underlying this pattern, I use a combination of phylogenetic, morphological and ecological techniques. In Chapter 1, I apply Bayesian methods and fossil-based calibrations to molecular sequence data to construct a time-calibrated phylogeny for Lacertidae. I estimate that the family arose in the early Cenozoic, with the majority of their African radiation occurring in the Eocene and Oligocene. In Chapter 2, I describe changes in lacertid body shape across biomes and substrates, and find widespread morphological convergence in similar habitat types. I suggest that in addition to foraging demands, fluctuating and extreme climatic conditions, largely driven by precipitation and temperature, contribute to morphological convergence across independent arid-dwelling clades. Finally, I test if ancestral transitions in ecology, morphology, and rates of diversification temporally coincide with paleoclimatic events in the Cenozoic. I use High Resolution X-ray Computed Tomography to characterize changes in the skull related to life in arid habitats, and apply maximum likelihood methods to test if the origins of those traits temporally coincide with significant shifts in habitat, diversification rates and climatic changes. My results show that African lacertids experienced three major peaks in diversification, accompanied by the evolution of suites of arid-adapted morphological traits. These changes coincide with climatic shifts in Africa, including the transition from closed forests to open grasslands and savanna in the late Oligocene, prior to the peak temperatures of the mid-Miocene Climatic Optimum, and following the formation of the Benguela current leading to hyper-aridity in southern Africa. I conclude that deserts are important centers for reptile evolution, but that expected changes in climate due to global warming may outpace the ability of arid-dwelling species to adapt and persist in the future.

×

Aim To map and assess the richness patterns of reptiles (and included groups: amphisbaenians, crocodiles, lizards, snakes and turtles) in Africa, quantify the overlap in species richness of reptiles (and included groups) with the other terrestrial vertebrate classes, investigate the environmental correlates underlying these patterns, and evaluate the role of range size on richness patterns. Location Africa. Methods We assembled a data set of distributions of all African reptile species. We tested the spatial congruence of reptile richness with that of amphibians, birds and mammals. We further tested the relative importance of temperature, precipitation, elevation range and net primary productivity for species richness over two spatial scales (ecoregions and 1° grids). We arranged reptile and vertebrate groups into range-size quartiles in order to evaluate the role of range size in producing richness patterns. Results Reptile, amphibian, bird and mammal richness are largely congruent (r = 0.79–0.86) and respond similarly to environmental variables (mainly productivity and precipitation). Ecoregion size accounts for more variation in the richness of reptiles than in that of other groups. Lizard distributions are distinct with several areas of high species richness where other vertebrate groups (including snakes) are species-poor, especially in arid ecoregions. Habitat heterogeneity is the best predictor of narrow-ranging species, but remains relatively important in explaining lizard richness even for species with large range sizes. Main conclusions Reptile richness varies with similar environmental variables as the other vertebrates in Africa, reflecting the disproportionate influence of snakes on reptile richness, a result of their large ranges. Richness gradients of narrow-ranged vertebrates differ from those of widespread taxa, which may demonstrate different centres of endemism for reptile subclades in Africa. Lizard richness varies mostly with habitat heterogeneity independent of range size, which suggests that the difference in response of lizards is due to their ecological characteristics. These results, over two spatial scales and multiple range-size quartiles, allow us to reliably interpret the influence of environmental variables on patterns of reptile richness and congruency.

×

The albumins from 41 species of the family Lacertidae representing a substantial part of genera as well as infrageneric groups of the collective genus Lacerta s.l. were investigated by means of the MCF technique. The data allow a chronological estimation of the most important radiation events and their correlation with paleogeographical facts. Our results are in remarkable contradiction to the phylogeny presented by ARNOLD (1989a) based mainly on morphological characters. The most important phylogenetic relationships are: 1. The genera Gallona and Psammodromus are already isolated from the other taxa since the Oligocene. We assess these main branches as subfamilies Gallotiinae and Lacertinae. 2. Lacerta s.str. and Lacerta subgenus Zootoca form the sister group of the remaining Lacertinae. 3. A colonization of Africa in the Lower Miocene has led to a separation in two lines, with mainly Eurasian and African members respectively. 4. ARNOLD`S (I.e.) `Ethiopian and advanced Saharo-Eurasian clade` proved to be a paraphyletic unit, one group of genera being more closely related to European taxa. All techniques applied to the systematics of Lacertidae hitherto are critically discussed. An area-time- hypothesis concerning the phylogeny of Lacertidae is presented. Key words: Lacertidae, phylogeny of; Gallotiinae subfam. nov.; Lacertinae subfam. nov.; micro complement fixation; albumin evolution.

×

The authors present an actualized and commented list of the amphibia and reptilia species noted in Morocco. The changes witch happened during the 16years after the last checklist established by BoNS(1972) are discussed. These are new additions (species recently discovered in Morocco and new taxa for Science, sorne of witch resulting from systematic modifications of other species or subspecies) and changes of names (new combinations) or status. Problems concerning the exact delimitation of several species are discussed, insisting particularly on the most conflictual groups (e.g. Chalcides, Aoenthodactylus). Thus we think that the number of species in the present checklist is probably surestimated. Taking in account these problems, we must consider this inventory as provisional, till clearing up the several questions still in suspense.

×

This is the first comprehensive review of the reptiles of Mauritania. It includes distributional information and comments. Mauritania harbors 86 species of reptiles belonging to 21 families. Among these families, Colubridae and Lacertidae are the most diverse, with 14 and 13 species respectively. Other families, such as Agamidae, Gekkonidae, Scincidae or Viperidae are also well represented. Among the 80 continental species, 47.5% are Saharan, 33.8% Afrotropical, 16.2% Sahelian and 2.5% Mediterranean. The marine turtles form another important group, with six species. Eight species are excluded from the country list because of old identification errors, there is not enough evidence of their presence or due to changes in political borders. Among the species expected to occur in Mauritania, at least nine may occur in Saharan environments, 13 in the Sahel savannah and two may have been introduced.

×

Many animals display complex colour patterns that comprise several adjacent, often contrasting colour patches. Combining patches of complementary colours increases the overall conspicuousness of the complex pattern, enhancing signal detection. Therefore, selection for conspicuousness may act not only on the design of single colour patches, but also on their combination. Contrasting long- and short-wavelength colour patches are located on the ventral and lateral surfaces of many lacertid lizards. As the combination of long- and short-wavelength-based colours generates local chromatic contrast, we hypothesized that selection may favour the co-occurrence of lateral and ventral contrasting patches, resulting in complex colour patterns that maximize the overall conspicuousness of the signal. To test this hypothesis we performed a comparative phylogenetic study using a categorical colour classification based on spectral data and descriptive information on lacertid coloration collected from the literature. Our results demonstrate that conspicuous ventral (long wavelength-based) and lateral (short wavelength-based) colour patches co-occur throughout the lacertid phylogeny more often than expected by chance, especially in the subfamily Lacertini. These results suggest that selection promotes the evolution of the complex pattern rather than the acquisition of a single conspicuous colour patch, possibly due to the increased conspicuousness caused by the combination of colours with contrasting spectral properties.

×

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 her- petofauna 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 Valle ́e du Haut and Bas Draˆa.

×

Die vorliegende Revision betrifft die Gattung Acanthodactylus (Damilie Lacertidae), die die Trocken- und Wüstengebiete des nördlichen Afrika und des südwestlichen Asiens bewohnt, mit Randpopulationen auf der Iberischen Halbinsel und auf Zypern. Mehr als 6000 Exemplare der Gattung, aus den bedeutendsten europäischen und amerikanischen Sammlungen, wurden untersucht. Zusätzlich wurden über 450 Röntgenaufnahmen für osteologische Untersuchungen ausgewertet. Die historische Übersicht behandelt in chronologischer Reihenfolge alle Arbeiten, die sich mit der Gattung Acanthodactylus beschäftigen, mit besonderer Berücksichtigung taxonomischer Studien. Ein beigegebener Bestimmungsschlüssel ermöglicht die Determination aller Arten der Gattung. Eine Reihe äußerer (Größe und Proportionen, Pholidose, Färbung und Zeichnung) und innerer (d.h. osteologischer) Merkmale wurde im Hinblick auf ihre Variabilität untersucht. Dann wurde ein Land ausgesucht, z.B. Ägypten, aus dem eine gute Materialbasis zur Verfügung stand. Für einen bestimmten Fundort in diesem Lande wurden sodann einige Kombinationen von Merkmalszuständen ermittelt, die sich aus dem Auftreten verschiedener Arten ergaben. Derselbe Vorgang wurde sodann auf den Rest Ägyptens übertragen, und die Merkmalskombinationen stellten sich als übereinstimmend heraus; sie erlaubten dadurch eine sichere Zuordnung von Tieren zu folgen Arten: A. boskianus, A. scutellatus und A. pardalis. Durch die Ausdehnung der Untersuchung nach Westen und Osten kamen weitere Taxa hinzu. Besonderer Aufmerksamkeit wurde diesen Merkmalskombinationen im Hinblick auf sympatrisches oder allopatrisches Auftreten der betroffenen Arten gezollt, mit dem Ergebnis, daß es in einigen Fällen möglich war, abzusichern, daß es sich um Populationen derselben Art handelt. Die Gattung Acanthodactylus wurde in mehrere Artengruppen gegliedert, die sich durch engere Beziehungen ihrer Arten untereinander auszeichnen. Die erste ist die micropholis-Gruppe mit der einzigen Art A. micropholis. Sie wird als die ursprünglichste der gesamten Gattung angesehen. Eine weitere Gruppe ist die boskianus-Gruppe, mit den Arten A. boskianus und A. schreiberi. Erstere wird ausführlich analysiert im Hinblick auf ihre außerordentlich große geographische Variabilität. Die yemenicus-Gruppe besiedelt die Arabische Halbinsel und besteht aus A. yemenicus sp.n., A. masirae, A. opheodurus und A. felicis. Eine weitere Gruppe ist die tristrami-Gruppe mit den Arten A. tristrami und A. robustus. Für erstere wird die Validität der Unterart orientalis verneint, die für iracensis dagegen bejaht. In die grandis-Gruppe wird hier nur A. grandis gestellt, während A. fraseri als dessen Synonym aufgefaßt wird. Die erythrurus-Gruppe enthält A. erythrurus, A. savignyi, A. blanci, A. boueti und A. guineensis. Letzterer rangierte bislang in der Gattung Eremias, gehört aber nach den hier vorgelegten Befunden in die erythrurus-Gruppe von Acanthodactylus. In der pardalis-Gruppe sind enthalten: A. pardalis, A. bedriagai, A. maculatus, A. spinicauda und A. busacki sp.n. aus Marokko. Die scutellatus-Gruppe besteht aus A. scutellatus, A. aureus, A. dumerili und A. longipes. Die letzte Gruppe ist die cantoris-Gruppe mit den Arten A. cantoris, A. arabicus, A. schmidti, A. blanfordi, A. gongrorhynchatus und A. haasi. Eine Bibliographie über die Gattung beschließt die Arbeit. Für jede Art werden Diagnosen, Verbreitungskarten, Synonymien, Beschreibungen und Materiallisten der untersuchten Tiere sowie auch Strichzeichnungen gegeben.

×

Acanthodactylus busacki ist ein relativ unbekannter Fransenfinger aus der Acanthodactylus-pardalis-Gruppe. Während mehrerer Aufenthalte in Marokko konnte ich diese stellenweise recht häufige Art beobachten.

×

Acanthodactylus lizards are among the most diverse and widespread diurnal reptiles in the arid regions spanning from North Africa across to western India. Acanthodactylus constitutes the most species-rich genus in the family Lacertidae, with over 40 recognized species inhabiting a wide variety of dry habitats. The genus has seldom undergone taxonomic revisions, and although there are a number of described species and species-groups, their boundaries as well as their interspecific relationships are largely unresolved. We constructed a multilocus phylogeny, combining data from two mitochondrial (12S, cytb) and three nuclear (MC1R, ACM4, c-mos) markers for 302 individuals belonging to 36 known species, providing the first large-scale time-calibrated molecular phylogeny of the genus. We evaluated phylogenetic relationships between and within species-groups, and assessed Acanthodactylus biogeography across its known range. Acanthodactylus cladogenesis is estimated to have originated in Africa due to vicariance and dispersal events from the Oligocene onwards. Radiation started with the separation into three clades: the Western and scutellatus clades largely distributed in North Africa, and the Eastern clade occurring mostly from Arabia to south-west Asia. Most Acanthodactylus species diverged during the Miocene, possibly as a result of regional geological instability and climatic changes. We support most of the current taxonomic classifications and phylogenetic relationships, and provide genetic validity for most species. We reveal a new distinct blanfordii species-group, suggest new phylogenetic positions (A. hardyi, A. masirae), and synonymize several species and subspecies (A. lineomaculatus, A. boskianus khattensis and A. b. nigeriensis) with their phylogenetically closely-related species. We recommend a thorough systematic revision of taxa exhibiting high levels of intraspecific variability as well as clear evidence of phylogenetic complexity such as A. guineensis, A. grandis, A. dumerilii, and A. senegalensis and the pardalis and erythrurus species-groups.

×

Recent molecular phylogenies of the Acanthodactylus pardalis species-group have revealed a deep genetic divergence within the nominal species A. busacki from north-west Africa. The species is phylogenetically separated into northern and southern lineages, which correspond to a previously observed morphological differentiation between the northern and southern populations of this species. Based on morphological comparisons of the type material and location of the type locality, the nomen Acanthodactylus busacki SALVADOR, 1982 is assigned here to the southern lineage, known from the northern Saharan Atlantic coastal desert. The northern lineage, described here as Acanthodactylus margaritae sp. nov., is prominently characterized by weakly keeled dorsal scales and a characteristic colour pattern. The new species is endemic to Morocco and confined to arid and semi-arid bioclimatic areas between the High Atlas and Anti-Atlas Mountains, from around Tamri in the north to Tiznit in the south and the Souss valley in the east.