×

Abstract The Equatorial African group of lacertids comprises nine species which share a number of derived features and appear to constitute a holophyletic assemblage. Revision of the group using morphological characters analyzed by parsimony and compatibility methods results in the following taxonomic changes: Lacerta jacksoni = Adolfus jacksoni, Lacerta echinata = Gastropholis echinata, Bedriagaia tropidopholis = Gastropholis tropidopholis, Bedriagaia moreavi = Gastropholis prasina, the latter species being validated and removed from the synonymy of Gastropholis vittata. Adolfus jacksoni has most features in common with the hypothetical ancestor of the group, while Holaspis guentheri and the species of Gastropholis are the most derived forms. Geographical variation in Adolfus alleni is described and what is known of the ecology of the Equatorial group summarized. Most are forest dwellers with the majority of the more primitive forms living on tree boles and among litter and timber on the forest floor (Adolfus jacksoni, A. africanus, A. vauereselli). Another, Holaspis guentheri, is confined to tree boles but glides between them, while the species of Gastropholis appear to be climbers in flimsy vegetation and Adolfus alleni is ground-dwelling above the tree line. Many of the morphological features characterizing these lizards appear to be functionally related to their particular habitats.

×

Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.

×

In this paper we deal on the ultraviolet color (invisible to us): where we can find it, the capability of animals to see it and the advantages that this color perception offers to them. As the simplest way to detect it is the photography, we describe and review how to photograph the UV, as a result of 15 years of amateur experience, searching and testing nearly in complete blindness due to the lack of practical information about “how to do it”. We describe the different kinds of photography (chemical and digital); the cameras and objectives suitable (both astronomically expensive ones and cheap options); what are the best characteristics that the objectives should have for this purpose; the films suitable for their use in chemical photography; the different filters (current or discontinued) manufactured along the years; and the subtle combinations among the different materials to obtain pure UV photographs. This kind of scientific photography is mainly used in forensics, forgery detection, art dermatology and less in Natural History, despite the fact that a great part of animals see this color and use it in important questions of their biology as the social behavior, mate choice or the food search.

×

The sclerotic ring consists of several bones that form in the sclera of many reptiles. This element has not been well studied in squamates, a diverse order of reptiles with a rich fossil record but debated phylogeny. Squamates inhabit many environments, display a range of behaviours, and have evolved several different body plans. Most importantly, many species have secondarily lost their sclerotic rings. This research investigates the presence of sclerotic rings in squamates and traces the lineage of these bones across evolutionary time. We compiled a database on the presence/absence of the sclerotic ring in extinct and extant squamates and investigated the evolutionary history of the sclerotic ring and how its presence/absence and morphology is correlated with environment and behaviour within this clade. Of the 400 extant species examined (59 families, 214 genera), 69% have a sclerotic ring. Those species that do not are within Serpentes, Amphisbaenia, and Dibamidae. We find that three independent losses of the sclerotic ring in squamates are supported when considering both evolutionary and developmental evidence. We also show that squamate species that lack, or have a reduced, sclerotic ring, are fossorial and headfirst burrowers. Our dataset is the largest squamate dataset with measurements of sclerotic rings, and supports previous findings that size of the ring is related to both environment occupied and behaviour. Specifically, scotopic species tend to have both larger inner and outer sclerotic ring apertures, resulting in a narrower ring of bone than those found in photopic species. Non-fossorial species also have a larger sclerotic ring than fossorial species. This research expands our knowledge of these fascinating bones; with further phylogenetic analyses scleral ossicles could become an extremely useful character trait for inferring the behaviour of fossil squamates.

×

Animals communicate via a variety of sensory channels and signals. Studies on acoustic and visual communication systems suggest that differences in the physical environment contribute to the variety of signalling behaviour, with species investing in those signals that are transmitted best under the local conditions. Whether or not environmental tuning also occurs in chemical communication systems has received much less attention. In the present study, we examined the effect of several aspects of the physical environment on the chemical communication system of lacertid lizards (family Lacertidae). The numbers of femoral pores are used as a proxy reflecting how much a particular species invests in and relies upon chemical signalling. Femoral pores are specialized epidermal structures that function as a secretion channel for the waxy substance produced by glands. In some lacertid species, the secretion carries infochemicals that play an important role in social communication. The number of femoral pores varies considerably among species. We have compiled data on femoral pore numbers for 162 species and tested for the effects of climate and substrate use. After correcting for body size and taking the phylogenetic relationships among the species into account, we found no effect of climate conditions or latitude on species pore numbers. Substrate use did affect pore numbers: shrub-climbing species tended to have fewer femoral pores than species inhabiting other substrates.

×

A complete catalogue is provided for the type specimens of lacertid lizards in the herpetological collection of the Zoological Museum, Berlin, as of December, 1993. The collection contains a total of 514 type specimens, representing 63 taxa, of which 42 are currently regarded as valid at the specific or subspecific level. Types representing an additional four taxa appear to have been lost from the collection. The collections are especially rich in African lacertid types and in historically significant specimens collected early in the Nineteenth Century by Pallas, Eversmann, and Hemprich & Ehrenberg. Also present are the type series of many subspecies of Podarcis lilfordi and P. pityusensis described by Eisentraut.

×

Acanthodactylus arabicus, Acanthodactylus aureus, Acanthodactylus bedriagai, Acanthodactylus blanci, Acanthodactylus blanfordii, Acanthodactylus boskianus boskianus, Acanthodactylus boskianus asper, Acanthodactylus boskianus euphraticus, Acanthodactylus boueti, Acanthodactylus busacki, Acanthodactylus cantoris, Acanthodactylus dumerili dumerili, Acanthodactylus dumerili exiguus, Acanthodactylus erythrurus erythrurus, Acanthodactylus erythrurus belli, Acanthodactylus erythrurus lineomaculatus, Acanthodactylus felicis, Acanthodactylus gongrorhynchatus, Acanthodactylus grandis, Acanthodactylus guineensis, Acanthodactylus haasi, Acanthodactylus longipes, Acanthodactylus maculatus, Acanthodactylus masirae, Acanthodactylus micropholis, Acanthodactylus opheodurus, Acanthodactylus orientalis, Acanthodactylus pardalis, Acanthodactylus robustus, Acanthodactylus savignyi, Acanthodactylus schmidti, Acanthodactylus schreiberi schreiberi, Acanthodactylus schreiberi syriacus, Acanthodactylus scutellatus scutellatus, Acanthodactylus scutellatus hardyi, Acanthodactylus spinicauda, Acanthodactylus tilburyi, Acanthodactylus tristrami, Acanthodactylus yemenicus, Adolfus africanus, Adolfus alleni, Adolfus jacksoni, Adolfus vauereselli, Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus nigropunctatus, Algyroides nigropunctatus kephallithacius, Australolacerta australis, Australolacerta rupicola

×

A recent herpetological collection in Cameroon yielded some 28 anuran, 16 lizard and 10 snake species. The species are listed and commented on under faunistical, taxonomical, ecological and zoogeographical aspects. Notes and corrections as to 7 other species listed in previous surveys are included. Specimens of Agama sankaranica, Mabuya buettneri, Panaspis aloysiisabaudiae and Lycophidion irroratum are first records from Came- roon. Noteworthy species are also Bufo villiersi, Adolfus africanus and Prosymna ambi- gua. The two last named are first records from the Adamaoua highlands, as are Holaspis guentheri, Hylarana albolabhs and Cardioglossa gratiosa. The Adamaoua population of C. gratiosa is described as a new subspecies C. g. peternageli.

×

In indoor terraria Adolfus jacksoni lays 3 - 5 clutches yearly, each batch consisting of 3 - 4 eggs. Oviposition takes place in the period January - September. Averaged over three years most clutches were found in January and May - July.

×

The amphibian fauna of northwestern Zambia was quite well known, but no systematic collection of reptiles had ever been made. This was rectified when I spent the period 23 September to 12 October 1990 based at Sakeji School near Ikelenge and collected 258 reptiles and 77 amphibians. Five reptile species are new for Zambia, i.e. Adolfus africanus, Causus lichtensteinii, Limnophis bicolor, Thrasops j. jacksonii and Rhamnophis aethiopissa ituriensis.

×

A review of the reptiles recorded from southeast Katanga Province in the Democratic Republic of Congo, employing the evolutionary species concept, indicates that there are at least 119 species in the area (7 chelonians, 37 lizards, 4 amphisbaenians, 69 snakes and 2 crocodilians), 15 of which are endemic (12.6 %). Analysis of zoogeographical affinities shows that this reptile fauna is derived from all directions, with forest forms entering from the north accounting for 14.4 % and savanna forms entering from the east contributing 22 %.

×

A phylogenetic-comparative approach was used to assess and refine existing secondary structure models for a frequently studied region of the mitochondrial encoded large subunit (16S) rRNA in two large lizard lineages within the Scincomorpha, namely the Scincidae and the Lacertidae. Potential pairings and mutual information were analyzed to identify site interactions present within each lineage and provide consensus secondary structures. Many of the interactions proposed by previous models were supported, but several refinements were possible. The consensus structures allowed a detailed analysis of rRNA sequence evolution. Phylogenetic trees were inferred from Bayesian analyses of all sites, and the topologies used for maximum likelihood estimation of sequence evolution parameters. Assigning gamma-distributed relative rate categories to all interacting sites that were homologous between lineages revealed substantial differences between helices. In both lineages, sites within helix G2 were mostly conserved, while those within helix E18 evolved rapidly. Clear evidence of substantial site-specific rate variation (covarion-like evolution) was also detected, although this was not strongly associated with specific helices. This study, in conjunction with comparable findings on different, higher-level taxa, supports the ubiquitous nature of site-specific rate variation in this gene and justifies the incorporation of covarion models in phylogenetic inference.

×

Some experience with Jacksons forest lizard (Adolfus jacksoni) The author describes his over all experiences in keeping the African Jacksons forest lizard (Adolfus jacksoni). The article gives some information on distribution, husbandry conditions, feeding and (mating) behaviour. Adolfus is typified as a lizard with an attitude, similar to what one can see in European Podarcis- species. Therefore a male and female combination is recommended. In captivity Adolfus proved to be a sturdy species. The author also refers to some interesting references on Adolfus jacksoni.

×

The tongue shape and its epidermal structure was examined in a stock of palearctic and paleotropical lizards from Europe, Africa, the Middle East and south eastern Asia. The results of this preliminary report clearly point out the systematic importance of this morphological feature in providing valuable generic and specific characters. The fundamental pattern described for the genera Podarcis, Lacerta sensu stricto and Gallotia, e.g., is very distinctive despite several features in common, likely due to ancient phyletic relationships. A number of similar morphological trends stress the reciprocal affinities between Lacerta (Archaeolacerta) and the widely spread genus Podarcis which is still undergoing speciation. Another case is the minor but significant affinities linking the genera Lacerta sensu stricto and Gallotia, the latter also undergoing more complicated speciation due to its insular isolation. The interspecific differences in the taxa belonging to the so-called `species groups` such as Podarcis, are scanty or insignificant. Instead there are striking differences in the generic patterns of the tongue morphology in Afro-Asian lacertid lizards. However, a similar pattern was found in the African genera Adolfus and Holaspis or Centromastix (= Gastropholis according to Arnold, 1989) and Ichnotropis.

×

A large number of reptiles from the Central African Republic (CAR) were collected during recent surveys conducted over six years (October 1990 to June 1996) and deposited at the Paris Natural History Museum (MNHN). This large collection of 4873 specimens comprises 86 terrapins and tortois- es, five crocodiles, 1814 lizards, 38 amphisbaenids and 2930 snakes, totalling 183 species from 78 local- ities within the CAR. A total of 62 taxa were recorded for the first time in the CAR, the occurrence of numerous others was confirmed, and the known distribution of several taxa is greatly extended. Based on this material and an additional six species known to occur in, or immediately adjacent to, the coun- try from other sources, we present a biogeographical analysis of the 189 species of reptiles in the CAR.

×

Climatic conditions changing over time and space shape the evolution of organisms at multiple levels, including temperate lizards in the family Lacertidae. Here we reconstruct a dated phylogenetic tree of 262 lacertid species based on a supermatrix relying on novel phylogenomic datasets and fossil calibrations. Diversification of lacertids was accompanied by an increasing disparity among occupied bioclimatic niches, especially in the last 10 Ma, during a period of progressive global cooling. Temperate species also underwent a genome- wide slowdown in molecular substitution rates compared to tropical and desert-adapted lacertids. Evaporative water loss and preferred temperature are correlated with bioclimatic parameters, indicating physiological adaptations to climate. Tropical, but also some popu- lations of cool-adapted species experience maximum temperatures close to their preferred temperatures. We hypothesize these species-specific physiological preferences may con- stitute a handicap to prevail under rapid global warming, and contribute to explaining local lizard extinctions in cool and humid climates.

×

Jackson’s Forest Lizard (Adolfus jacksoni) is widespread throughout the highlands of the Albertine Rift, southern Uganda, western and central Kenya, and northern Tanzania. To understand the population genetics and phylogenetic relationships of this widespread taxon, we sequenced two mitochondrial (16S and cyt b) and two nuclear (c-mos and RAG1) genes from multiple populations. Population genetics analyses suggested a high degree of genetic differentiation among A. jacksoni populations, reflecting the high-elevation montane ‘‘islands’’ that they inhabit. Populations connected by a network of mountain ranges generally showed lower levels of genetic partitioning than those isolated by lowelevation habitat. Results from phylogenetic analyses and additional morphological data indicated that Adolfus jacksoni occurs throughout the Albertine Rift, likely from the Kabobo Plateau to the Lendu Plateau of Democratic Republic of the Congo, as well as southern Uganda, Mt. Elgon, and the highlands of western Kenya on the western side of the Kenyan Rift. Adolfus kibonotensis is removed from the synonymy of the latter taxon, elevated to full species, and recognized from the central Kenyan highlands to northern Tanzania on the eastern side of the Kenyan Rift. A new Adolfus species is described from the Mathews Range in central Kenya.

×

Currently, four species of the lacertid lizard genus Adolfus are known from Central and East Africa. We sequenced up to 2825 bp of two mitochondrial [16S and cytochrome b (cyt b)] and two nuclear [(c-mos (oocyte maturation factor) and RAG1 (recombination activating gene 1)] genes from 41 samples of Adolfus (representing every species), two species each of Gastropholis and Holaspis, and in separate analyses combined these data with GenBank sequences of all other Eremiadini genera and four Lacertini outgroups. Data from DNA sequences were analysed with maximum parsimony (PAUP), maximum-likelihood (RAxML) and Bayesian inference (MrBayes) criteria. Results demonstrated that Adolfus is not monophyletic: Adolfus africanus (type species), Adolfus alleni, and Adolfus jacksoni are sister taxa, whereas Adolfus vauereselli and a new species from the Itombwe Plateau of Democratic Republic of the Congo are in a separate lineage. Holaspis and Gastropholis were recovered in separate clades. Based on these molecular data, relatively substantial sequence divergence, and multiple morphological differences, we describe a new genus of lacertid for the lineage including A. vauereselli and the new Itombwe species. The recognition of this new, endemic genus underscores the conservation importance of the Albertine Rift, especially the Itombwe Plateau, a unique region that is severely threatened by unchecked deforestation, mining, and poaching.

×

DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.

×

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.

×

Background: Although current molecular clock methods offer greater flexibility in modelling evolutionary events, calibration of the clock with dates from the fossil record is still problematic for many groups. Here we implement several new approaches in molecular dating to estimate the evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and uncertain phylogeny. Four different models of rate variation are tested in a new program for Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by expressing multiple calibration dates as a range of probabilistic distributions. We also test the reliability of our proposed calibrations by exploring effects of individual priors on posterior estimates. Results: According to the most reliable model, as indicated by Bayes factor comparison, modern lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate much earlier origins for these clades than previously reported, and we discuss our results in light of paleogeographic trends during the Cenozoic. Conclusion: This study represents the first attempt to estimate evolutionary ages of a specific group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates from the fossil record as minimum node constraints. The bioinformatics program used here, TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with similar challenges.

×

While global variation in taxonomic diversity is strongly linked to latitude, the extent to which morphological disparity follows geographical gradients is less well known. We estimated patterns of lineage diversification, morphological disparity and rates of phenotypic evolution in the Old World lizard family Lacertidae, which displays a nearly inverse latitudinal diversity gradient with decreasing species richness towards the tropics. We found that lacertids exhibit relatively constant rates of lineage accumulation over time, although the majority of morphological variation appears to have originated during recent divergence events, resulting in increased partitioning of disparity within subclades. Among subclades, tropical arboreal taxa exhibited the fastest rates of shape change while temperate European taxa were the slowest, resulting in an inverse relationship between latitudinal diversity and rates of phenotypic evolution. This pattern demonstrates a compelling counterexample to the ecological opportunity theory of diversification, suggesting an uncoupling of the processes generating species diversity and morphological differentiation across spatial scales.

×

A lizard specimen recently collected from Gabon can not be assigned to any known species for the country. It is identified as Ichnotropis bivittata, a lacertid previouslnot reported for the country. The specimen is described in details and compared to related species. An older specimen examined by Boulenger (1921) was collected in the same area but not identified as from Gabon since the country was not always with the same borders and under the name `Congo français`.

×

the Lacertidae is one of the most diverse and widespread lizard families throughout Eurasia and Africa. Several studies so far have attempted to unravel the phylogeny of Lacertidae using morphological and mole- cular data. However, the intra-family relationships remain unclear. in an effort to explore the phylogenetic relationships within the family Lacertidae, a concatenated dataset of 5727 bp from six genes (two nuclear and four mitochondrial) and 40 genera was assembled based on GenBank database. Phylogenetic inference analy- ses were conducted using Maximum Parsimony (MP), Bayesian inference (Bi) and Maximum Likelihood (ML), revealing that even a combined dataset of both mitochondrial and nuclear genes is not able to resolve the phylogenetic relationships of the Lacertidae family under the tribe level.

×

The lacertid lizard Adolfus africanus is recorded from Kakamega Forest, western Kenya, and Imatong Mountains, southern Sudan. Both localities represent first country records for the species. Kakamega Forest constitutes the easternmost locality for A. africanus which was also recorded from Cameroon. The known distribution of this rain forest species is reviewed and discussed in a biogeographical context.

×

The dangers of helminth infestations for lizards, particularly infestations with nematodes und cestodes, are reported. Infection routes, diagnoses, und treatments are described.

×

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.

×

This paper presents the composition of the lizard fauna of Maringa-Lopori-Wamba landscape in the western portion of the central Congo basin in the Democratic Republic of the Congo and its weak similarity with the lizard fauna of Gaboon and Ituri faunal areas. It also fills a gap of knowledge in a region where no previous research on Sauria has been undertaken and signals the presence of Leptosiaphos hylophilus and Lygodactylus n. sp. The paper mentions the finding of a subspecies of Gastropholis echinata. These data are important since collected for the first time in a spot where no herpetological research on lizards has previously been undertaken.

×

A commented lacertids list for Europe, Africa, the Middle East, including the Arabian Peninsula and Asia is given.

×

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 family Lacertidae encompasses more than 250 species distributed in the Palearctis, Ethiopis and Orientalis. Lacertids have been subjected in the past to several morphological and molecular studies to establish their phylogeny. However, the problems of convergent adaptation in morphology and of excessively variable molecular markers have hampered the establishment of well supported deeper phylogenetic relationships. Particularly the adaptations to xeric environments have often been used to establish a scenario for the origin and radiation of major lineages within lacertids. Here we present a molecular phylogenetic study based on two nuclear marker genes and representatives of 37 lacertid genera and distinct species groups (as in the case of the collective genus Lacerta). Roughly 1600 bp of the nuclear rag1 and c-mos genes were sequenced and analyzed. While the results provide good support to the hitherto suggested main subfamilies of Gallotiinae (Gallotia and Psammodromus), Eremiainae and Lacertinae [Harris, D.J., Arnold, E.N., Thomas, R.H., 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proc. R. Soc. Lond. B 265, 1939–1948], they also suggest unexpected relationships. In particular, the oriental genus Takydromus, previously considered the sister-group to the three subfamilies, is nested within Lacertinae. Moreover, the genera within the Eremiainae are further divided into two groups, roughly corresponding to their respective geographical distributions in the Ethiopian and the Saharo-Eurasian ranges. The results support an independent origin of adaptations to xeric conditions in different subfamilies. The relationships within the subfamily Lacertinae could not be resolved with the markers used. The species groups of the collective genus Lacerta show a bush-like topology in the inferred Bayesian tree, suggesting rapid radiation. The composition of the subfamilies Eremiainae and Lacertinae as well as their phylogeography are discussed.

×

Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk. Location World-wide. Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions. Results I obtained body size data for 99% of the world`s lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non-aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed. Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.

×

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.

×

Some observations in keeping the scarcley noticed Adolfus jacksoni are described. Showing only few remarkable specialities, breeding of the species was successful afte keeping it in an outdoor terrarium during summer. Some weeks old clutches that were found in November (soil temperature 5°C!) developed well. Growing of the hatchlings was without any problems.

×

Herein is presented a list of the reptiles and anurans from the Parc National des Volcans (PNV) (01°43’ S, 29°52’ W), an area in the west and north provinces of the Republic of Rwanda in the Albertine Rift region of Africa. Fieldwork was conducted between two and six days per week from June through August 2007 and 2008. We also conducted literature searches of all historical expeditions within the park for species records. Seventeen species of reptiles and anurans are recorded from the PNV. Nine of the species were anurans, distributed in five families: Arthroleptidae (3), Bufonidae (1), Hyperoliidae (3), Phrynobatrachidae (1), and Pipidae (1). Eight species of reptiles were recorded from five families: Chamaeleonidae (1), Lacertidae (2), Scincidae (2), Colubridae (2), and Viperidae (1). Eight of the seventeen species found in the PNV are endemic to the Albertine Rift.

×

While the stability of sex chromosomes is widely accepted in viviparous mammals and birds, ectothermic vertebrates are still largely viewed as having frequent turnovers in sex determining systems. Frequent changes in sex determining systems in ectotherms could be problematic for field ecological studies as well as for breeding programs, as molecular sexing across a phylogenetically widespread spectrum of ectothermic vertebrates would not be possible. However, we recently documented that sex determining systems in three important reptile lineages (caenophidian snakes, iguanas, and lacertid lizards) are in fact highly conserved. 2.We applied a new molecular procedure to identify sex within each of these three lineages (encompassing altogether around 4,000 species, i.e. nearly 50% of the recent species of reptiles). This technique uses quantitative PCR (qPCR) to compare copy numbers of genes specific for their respective Z (in caenophidian snakes and lacertids) and X (in iguanas) chromosomes between male and female genomes. The DNA samples required can be collected relatively non-invasively. Unlike molecular sexing based on repetitive elements, this technique can be easily applied to previously unstudied species of these lineages, as the number of copies of protein coding genes linked to the differentiated sex chromosomes is evolutionary highly conserved in each. 3.We suggest that qPCR-based molecular sexing using the comparison of gene copy number is a practical choice for non-model species of caenophidian snakes, iguanas and lacertids. Morever, it should also soon be available for other reptile lineages with differentiated sex chromosomes.

×

Includes bibliographical references and index. Introduction. List of localities. New genus. List of new species and type localities. Preliminary note on the distribution of African lizards. Note on the distribution of African turtles -- Testudinata. Testudinae. Kinixys Bell. Pelomedusidae. Pelusios Wagler. Pelomedusa Wagler -- Loricata (Crocodilia). Crocodylidae. Crocodylus Laurenti. Osteoblepharon, new genus. Ecological notes on Congo crocodiles -- Squamata. Lacertilia. Geckonidae. Gonatodes Fitzinger. Hemidactylus Cuvier. Lygodactylus Gray. Agamidae. Agama Daudin. Varanidae. Varanus Merrem. Lacertidae. Lacerta Linnaeus. Bedriagaia Boulenger. Algiroides Fitzinger. Ichnotropis Peters. Eremias Wiegmann. Holaspis Gray. Gerrhosauridae. Gerrhosaurus Wiegmann. Scincidae. Mabuya Fitzinger. Lygosoma Gray. Ablepharus Fitzinger. Anelytropidae. Feylinia Gray. Chamaeleontidae. Chamaeleon Gronovius. Rhampholeon Günther -- Appendix A: Lists of the turtles, crocodiles, lizards, and chameleons of the rain forest and of the Sudanese subprovince -- Appendix B: Notes on the types of Hallowell`s West African species in the Academy of Natural Sciences of Philadelphia.

×

In this paper, the evolution and ecology of directional asymmetry (DA) during the developmental trajectory (DT) is compared with that of its product, morphological DA (MDA). DT and MDA are calculated for two bilateral morphological scale characters of lizards, the number of subdigital lamellae beneath the fourth toe in 10 agamid and 28 lacertid taxa, and the number of rows of ventral scales in 12 lacertid taxa. MDA, the subtraction between left and right sides (classical measure of DA), is functional in adult animals. Results confirm the hypodiesis that, in DT, the regression parameters a (constant) and b (regression slope) of counts on the right side with those on the left describe a developmental process. No phylogenetic or environmental effects were observed on a and b, but analyses considering both a and b together show non-random phyletic patterns. Independent analyses deduced the same ancestral DT in Agamidae and Lacertidae. In Lacertidae, distance between pairs of taxa in a + b (standardized values) correlates positively with die phylogenetic distance between taxa. Phyletic trends in MDA are indirect, and due to the link of MDA with a + b. The MDA of species is more dissimilar in sympatry than in allopatry. The phyletic trends suggest evolution of DT, while die association of MDA witii sympatry suggests that ecological pressures shape MDA in adult animals. Evolution of DT is independent from tiiat of its product, MDA–adaptive determinism defines the result of, but not the mechanistic process of, development. Deterministic environmental processes define MDA, and deterministic evolutionary processes define the interactive result of a and b, but not each separately. According to circumstances, different DTs produce similar or different MDA, and a particular DT can produce different MDAs.

×

This checklist records the 99 species of lizards known at present from Kenya, and which are divided amongst eight families: Gekkonidae 33 species, Agamidae seven, Chamaeleonidae 17, Scincidae 22, Lacertidae 12, Cordylidae five, Varanidae two, Amphisbaenidae one. Brief data on the distribution of all species is given, with some localities, details of habitat and (in some cases) status of subspecies. Some taxonomic notes on certain problematic species/genera are included, plus a brief discussion of the zoogeography of Kenya`s lizards, and a gazetteer of localities.

×

The staggering phenotypic variation among organisms and the striking fit between an organism`s morphology and its environment have fascinated naturalists for centuries. Darwin`s publication of `On the origin of species` was the first to offer an explanation for the apparently perfect form-function relationship. During the past century overly enthousiastic evolutionary biologists interpreted any direct link between an organism`s morphology and ecology as evidence for the theory of evolution by natural selection. However, for morphological variation to be considered adaptive it is necessary to explicitly test how this variation translates into the variation in ecologically relevant performance traits, which in turn translates into ecological (and fitness) variation. Nowadays, performance- based ecomorphological approaches are typically used to test the adaptive nature of phenotypic variation in order to provide a better understanding of the selective pressures operating on organisms. Using an ecomorphological approach, lacertid lizards have been used extensively to test whether and how they are adapted to their environment. The majority of these studies have mainly included European species, leaving African species underrepresented. This is suprising from an evolutionary viewpoint. Compared to European lacertids, African lacertid lizard clades are older and more speciose. Also, lacertid lizard communities on the African continent are more complex than the ones in Europe (i.e. comprising 7-8 species versus 2-3 species). In this talk, we will focus on the phenotypic divergence among African clades and the link with the variation in performance and ecology, and explicitly compare these results to results on European lacertids.

×

The aim of this study was to determine if divergence in habitat use among lacertid lizards is paralleled by morphological differentiation. For 35 lacertid species, we measured body, head and limb dimensions. Habitat use was inferred from the literature: ground-dwelling on open terrain, ground-dwelling in vegetated areas, shrub-climbing, tree-climbing, saxicolous (i.e. rock-climbing). Traditional (i.e. non-phylogenetic) statistical analyses suggest morphological differences among species groups with different habitat use. Ground-dwelling species from open habitats tend to have longer femurs, tibiae and humeri (relative to body length) than other groups. Cursorial (i.e. level-running) species have relatively high heads and trunks compared to climbing species. These differences follow biomechanical predictions and it is tempting to consider tham as adaptations to habitat use. However, phylogenetic analyses of the data fail to establish a clear relationship between habitat use and morphology in the data set considered. There is a weak indication that the differences in head and trunk height have evolved as an adaptation to different habitat use, but the differences in relative limb dimensions among species groups with different habitat use vanish. Either adaptation of limb dimensions to habitat use has not occurred in lacertid lizards, or our methods are unable to demonstrate such an adaptation. We show that uncertainties in the topology of the phylogenetic tree used are unlikely to influence the outcome of our study. We also address the fact that habitat use is often similar in different branches of the phylogenetic tree, and the consequences this may have for the power of our statistical analyses.

×

This study lists the amphibian and reptile species of Kibale National Park, Uganda, and discusses the natural history and biogeography of this unique herpetofauna. This herpetofaunal inventory was compiled based upon literature records and collections made during 17 mo fieldwork between 1995 and 1997, and includes 28 anuran, 15 lizard, and 32 snake species. Faunal comparisons with seven other tropical African forests show a high degree similarity between the Kibale herpetofauna and those of central and West Africa.

×

The amphibians and reptiles of Kibale National Park in western Uganda were inventoried over an 18-mo period in 1995 and 1996-97. A total of 75 species, including 28 amphibians and 47 reptiles, were collected or observed. Comparison with other equatorial African herpetofaunas confirms that the Kibale fauna is most similar to those of southwest Uganda and eastern Congo-Zaire, both hypothesized Pleistocene forest refugia. Comparison with a West Africa fauna also shows a fair degree of overlap, while almost no overlap was observed between Kibale and the forests of coastal East Africa. This confirms that the Kibale herpetofauna is an extension of the Guinea-Congolean forest faunas. Randomly placed 5 x 5 m plots were used to sample the herpetofauna of the forest leaflitter layer in unlogged forest, logged forest, and a neighboring exotic pine plantation. A total of 18 amphibian and reptile species were captured in the litter, a number similar to that observed in mid-elevation tropical forests in Central America and Southeast Asia. Density at Kibale was much lower than most previous studies. Analysis of the feeding ecology of the most abundant litter species showed that most diurnal litter frogs are active foragers of hard-bodied prey such as ants; sit-and-wait predators of larger soft-bodied prey are curiously absent. Plots sampled under fruiting Ficus natalensis trees showed significantly higher prey densities, but litter amphibians and reptiles did not seem to respond to this increase. Of the physical and biotic factors measured in each plot, seasonal changes in soil moisture were most closely correlated with the patterns of herpetofauna abundance observed in the forest. This is consistent with the fact that Kibale receives less rain than any site where the ecology of the litter herpetofauna has been studied, and that most of the species present in Kibale are believed to have evolved in the wetter forests of eastern Congo-Zaire.

×

I compared species richness and habitat correlates of leaf-litter herpetofaunal abundance in undisturbed and selectively logged forests, and an abandoned pine plantation in Kibale National Park, Uganda. I sampled 50 randomly located 25 m2 litter plots in each area during the wet and dry seasons in 1997. Ten anuran, five lizard, and three snake species were captured in plots over the study. Assemblage composition was most similar at logged and unlogged sites. The logged forest herpetofauna had higher species richness and abundance than the unlogged forest, but diversity was greater in the unlogged forest due to greater evenness. In contrast, the pine plantation site had the highest richness, abundance, and evenness of the three study sites, but species composition was distinct from the other areas. Herpe- tofaunal densities were significantly lower in all three areas during the dry season than in the wet season. During the dry season, soil moisture, litter mass, topography, shrub cover, and number of fallen logs were significant positive predictors of herpetofaunal presence in litter plots, but only soil moisture was significant in the wet season. The interaction of moisture and topography appears to be important in determining seasonal patterns of litter herpetofaunal distribution. Comparison of litter herpetofaunal studies across the tropics have shown that mid-elevation faunas generally support fewer species than lowland faunas. Compared with other tropical mid-elevation litter faunas, Kibale supports an intermediate number of species, but at lower densities than observed at any other mid-elevation site reported in the literature.

×

Aim The Kakamega Forest, western Kenya, has been biogeographically assigned to both lowland and montane forest biomes, or has even been considered to be unique. Most frequently it has been linked with the Guineo-Congolian rain forest block. The present paper aims to test six alternative hypotheses of the zoogeographical relationships between this forest remnant and other African forests using reptiles as a model group. Reptiles are relatively slow dispersers, compared with flying organisms (Aves and Odonata) on which former hypotheses have been based, and may thus result in a more conservative biogeographical analysis. Location Kakamega Forest, Kenya, Sub-Saharan Africa. Methods The reptile diversity of Kakamega Forest was evaluated by field surveys and data from literature resources. Faunal comparisons of Kakamega Forest with 16 other African forests were conducted by the use of the ‘coefficient of biogeographic resemblance’ using the reptile communities as zoogeographic indicators. Parsimony Analysis of Endemism and Neighbour Joining Analysis of Endemism were used to generate relationship trees based on an occurrence matrix with paup*. Results The analysis clearly supports the hypothesis that the Kakamega Forest is the easternmost fragment of the Guineo-Congolian rain forest belt, and thus more closely related to the forests of that Central–West African complex than to any forest further east, such as the Kenyan coastal forests. Many Kenyan reptile species occur exclusively in the Kakamega Forest and its associated forest fragments. Main conclusions The Kakamega Forest is the only remnant of the Guineo- Congolian rain forest in the general area. We assume that the low degree of resemblance identified for the Guineo-Congolian forest and the East African coastal forest reflect the long history of isolation of the two forest types from each other. Kenyan coastal forests may have been historically connected through forest ‘bridges’ of the southern highlands with the Congo forest belt, allowing reptile species to migrate between them. The probability of a second ‘bridge’ located in the region of southern Tanzanian inselbergs is discussed. Although not particularly rich in reptile species, the area should be considered of high national priority for conservation measures.

×

We present an annotated checklist of the herpetofauna of Kakamega Forest with comments on the biology and systematics of the taxa. Twenty-five amphibian, one turtle, 22 lizard and 36 snake species are recorded from within the forest and its immediate environment. We discuss the generalized zoogeography of the forest and distribution pattern of the taxa comment on the protection of the forest. Analysis of the reptile species composition shows Kakamega Forest to be similar to the Guinea-Congolian rainforest and is considered the easternmost remnant of this forest block. Kakamega forest has a high diversity value for Kenya and represents a diversity hotspot on a national scale. Two species, Lrgodac- t}`lus gutturalis and Psammophis phillipsi. are recorded in Kenya for the first time. Several other first records and the description of a new species (Agamidae: Agama finchi) were published already separately.

×

The genus Adolfus Sternfeld, 1912 currently contains three species from Equatorial Africa. Two of these occur in widespread, low- to mid-elevation habitats, but Adolfus alleni is only known from four montane peaks (Aberdares, Mt. Kenya, Cherangani Hills, Mt. Elgon) in Kenya and Uganda. An integrative approach using 58 morphological characters and genetic analyses of mitochondrial (16S and cyt b) and nuclear (c-mos and RAG1) DNA sequence data revealed differences between these populations, and indicated that A. alleni is a complex of at least two cryptic species. Herein, we describe the populations from the Aberdares and Mount Elgon as a new species, and restrict A. alleni to Mount Kenya. This action underscores the importance of conservation strategies to protect these montane peaks, which may harbour additional, unique evolutionary lineages.

×

Lacertidae from page 84.