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Literatur und Schriften
Agamen The origin of Saharan biodiversity is poorly understood, in part because the geological and paleoclimatic events that presumably shaped species diversity are still controversial, but also because few studies have explored causal explanations for the origin of Saharan diversity using a phylogenetic framework. Here, we use mtDNA (16S and ND4 genes) and nDNA (MC1R and CMOS genes) to infer the relationships and biogeographic history of North African agamas (genus Agama). Agamas are conspicuous, diverse and abundant African lizards that also occur in the Saharan xeric and mesic environments. Our results revealed the presence of three Agama lineages in North Africa: one Afrotropical, one Sahelo-Saharan, and one broadly distributed in North Africa and mainly Saharan. Southern Mauritania contains the highest known diversity, with all three lineages present. Results suggest that agamas colonized the Sahara twice, but only one lineage was able to radiate and diversify there. Species in the Saharan lineage are mostly allopatric, and their splitting, genetic diversity and distribution are greatly explained by mountain ranges. One species in this lineage has colonized the Mediterranean climatic zone (A. impalearis), and another one the Sahel savannah (A. boueti). The other lineage to colonize the Sahara corresponds to A. boulengeri, an eminently Sahelian species that also inhabits Saharan mountain ranges in Mauritania and Mali. Phylogenetic analyses indicate that allopatric montane populations within some currently recognized species are also genetically divergent. Our study therefore concludes that vicariant speciation is a leading motor of species diversification in the area: Inside the Sahara, associated to mountain-ranges isolated by dune seas and bare plains; outside, associated to less harsh climates to the North and South. Paleoclimatic oscillations are suggested as causal explanations of the vicariant distribution and origin of species. Agamas are thought to have colonized northern Africa during wet periods, with subsequent dry periods fragmenting species distribution and leading to allopatric populations associated to milder and wetter climates in the Mediterranean, Sahel, and in Saharan mountains, in an island-model fashion. Finally, our results support the synonymization of A. castroviejoi with A. boueti, the reciprocal monophyly of all other North African agamas, and suggest one candidate species within A. boulengeri. Agama aculeata aculeata, Agama planiceps planiceps. HENLE, K. (1995): A brief review of the origin and use of ‘stellio’ in herpetology and a comment on the nomenclature and taxonomy of agamids of the genus Agama (sensu lato). – Herpetozoa, Wien, 8 (1/2): 3-9. Zusammenfassung: JOGER, U. (1991): A molecular phylogeny of agamid lizards. – Copeia, 1991: 616-622. KLOSSOVSKY, B.N. & E.G. BALASHOV (1961): On different constructions of circulatory system of Agama brain. – Zool. Zh., 40: 251-257. LEACH, A.D., CHONG, R.A., PAPENFUSS, T.J., WAGNER, P., BÖHME, W., SCHMITZ, A., RÖDEL, M.-O., LEBRETON, M., INEICH, I., CHIRIO, L., BAUER, A., ENIANG, E.A. & S. BAHA EL DIN (2009): Phylogeny of the genus Agama based on mitochondrial DNA sequence data. – Bonner zoologische Beiträge, Bonn, 56 (4): 273-278. LEACHÉ, A.D., WAGNER, P., LINKEM, C.W., BÖHME, W., PAPENFUSS, T.J., CHONG, R.A., LAVIN, B.R., BAUER, A.M., NIELSEN, S.V., GREENBAUM, E., RÖDEL, M.-O., SCHMITZ, A., LeBRETON, M., INEICH, I., CHIRIO, L., OFORI-BOATENG, V., ENIANG, E.A., BAHA EL DIN, S., LEMMON, A.R. & F.T. BURBRINK (2014): A hybrid phylogenetic–phylogenomic approach for species tree estimation in African Agama lizards with applications to biogeography, character evolution, and diversification. – Molecular Phylogenetics and Evolution, 79: 215–230. Africa is renowned for its biodiversity and endemicity, yet little is known about the factors shaping them across the continent. African Agama lizards (45 species) have a pan-continental distribution, making them an ideal model for investigating biogeography. Many species have evolved conspicuous sexually dimorphic traits, including extravagant breeding coloration in adult males, large adult male body sizes, and variability in social systems among colorful versus drab species. We present a comprehensive time-calibrated species tree for Agama, and their close relatives, using a hybrid phylogenetic–phylogenomic approach that combines traditional Sanger sequence data from five loci for 57 species (146 samples) with anchored phylogenomic data from 215 nuclear genes for 23 species. The Sanger data are analyzed using coalescent-based species tree inference using *BEAST, and the resulting posterior distribution of species trees is attenuated using the phylogenomic tree as a backbone constraint. The result is a time-calibrated species tree for Agama that includes 95% of all species, multiple samples for most species, strong support for the major clades, and strong support for most of the initial divergence events. Diversification within Agama began approximately 23 million years ago (Ma), and separate radiations in Southern, East, West, and Northern Africa have been diversifying for >10 Myr. A suite of traits (morphological, coloration, and sociality) are tightly correlated and show a strong signal of high morphological disparity within clades, whereby the subsequent evolution of convergent phenotypes has accompanied diversification into new biogeographic areas. We conducted field studies in 15 West African countries and collected one thousand specimens of lizards of the genus Agama. Based on these collections, literature, molecular analysis of selected specimens, and examination of Linnean type-specimens of A. agama, we review the phylogeny, taxonomy, geographic distribution and ecology of the West African species of the genus Agama. Seventeen different species are recognized in the genus Agama in West Africa, northern Cameroon and Chad: A. africana, A. agama, A. boensis, A. boueti, A. boulengeri, A. castroviejoi, A. cristata, A. doriae benueensis, A. gracilimembris, A. insularis, A. lebretoni, A. paragama, A. sankaranica, A. weidholzi, and three new species. We design a lectotype for A. agama (Linnaeus, 1758) and attribute to A. wagneri, sp. nov., the populations from northern and central Cameroon of the A. agama complex. Agama parafricana, sp. nov., is described from wet savannah areas of Togo and Benin. Agama sylvanus from southern Ghana is a junior synonym of A. africana. Agama cf impalearis from northern Niger and Mali corresponds to an undescribed species. Agama boensis is resurrected from the synonymy of A. sankaranica. According to biogeographic areas, four species are Sahelian, seven species are Sudanian, four species are Guinean, and two species are ubiquitous. PETERS, G. (1971): Die Wirtelschwänze Zentralasiens (Agamidae: Agama). Ergebnisse der Mongolisch-Deutschen Biologischen Expeditionen seit 1962, Nr. 59. –Mitt. zool. Mus Berlin, 47: 357-381. Immature, adult males and females of the lizard, Agama agama (L.) collected at Ago-Iwoye, Nigeria during the second and third quarters of 1987, differed most in snout-vent lengths (SVL). They averaged 59 ± 3 mm, 125 ± 2 mm and 104 ± 3 mm SVL respectively and also differed in dimensions and weights of other body structures. These structures, including scales, eyes, tail and hind limb, correlated well with SVL and grow allometrically in relation to it. They can therefore be used to separate Agama agama into age-sex classes. Females are capable of breeding at 91 mm SVL and those in breeding condition differed from non-reproductive members in weights and dimensions of gonadal structures (P<0.001) but not SVL and body weight (P>0.10). Males in reproductive condition differed from non-reproductive ones in SVL, body weight and reproductive parameters (P<0.001).
Ground Agama HEIDEMAN, N.J.L. (1997): Comparative analysis of nematode infection in Agama aculeata and Agama planiceps, and its effect on body condition and fecundity. – Copeia, 1997 (4): 875-880. HEIDEMAN, N.J.L. (2002): A comparison of the breeding and non-breeding season diet of Agama aculeata and Agama planiceps (Reptilia: Agamidae) in Windhoek, Namibia. - Journal of Herpetology, 36 (3): 515-520. MERREM, B. (1820): Description of Agama aculeata. – In: “Versuch eines Systems der Amphibien I (Tentamen Systematis Amphibiorum)”. J. C. Kriegeri, Marburg, 191 pp. Agama aculeata aculeata MERREM, 1820 Ground Agama HEIDEMAN, N.J.L. (1993): Social organization and behaviour of Agama aculeata aculeata and Agama planiceps planiceps (Reptilia: Agamidae) during the breeding season. – J. Herpetol. Assoc. Afr., 42: 28-31. HEIDEMAN, N.J.L. (1994): Reproduction in Agama aculeata aculeata and Agama planiceps planiceps females from Windhoek, Namibia. – Amphibia-Reptilia, Leiden, 15: 351-361. (00.353) Agama aculeata aculeata and Agama planiceps planiceps females were reproductively active during summer, which is the rainy season in Namibia. Both species were iteroparous; clutch size and relative clutch mass in the smaller A. a. aculeata were significantly larger than in A. p. planiceps. Egg and newborn hatchling size of A. a. planiceps on the other hand were significantly greater than in A. a. aculeata. It is suggested that these differences may have evolved as a result of different strategies which the two species possibly employ to cope with predation pressure, namely, evasion through crypsis in A. a. aculeata and through speed of escape in A. p. planiceps. Ground Agama Analysis of stomach contents can provide insights into foraging mode, habitat use, and dietary specialization of animals. In this paper, we make observations on the poorly known diet of four southern African agamid species, Agama aculeata distanti (Eastern Ground Agama), Agama armata (Peter’s Ground Agama), Agama atra (Southern Rock Agama), and Acanthocercus atricollis (Southern Tree Agama). We examined the diet of 67 individuals by identifying and weighing prey items after stomach flushing lizards in the field. We found that these agama species fed on a broad spectrum of arthropods (11 orders). A high relative importance of ants was present for all agama species examined here, which suggests that ants are a major food source in the arid ecosystem. We found that active prey such as ants, beetles, and highly mobile flying insects like wasps and flies to be major components of the diet, indicating that these lizards are ambush predators. We also found that 43% of the stomachs contained herbaceous material and 39% contained sand particles. Agama atra had the most diverse dietary niche, eating fewer ants and more beetles, hemipterans, and dipterans than other species, whereas A. armata had a narrower dietary niche consisting mainly of ants. Lastly, although low in sample size, we found that juveniles qualitatively had a diet of functionally similar prey items, albeit with a narrower niche breadth, when compared to adults. We discuss how diet corresponds with differences in foraging behavior and habitat specialization.
Siedleragame / Rainbow Lizard / Common Agama ABULUDE, F.O., ADESANYA, W.O., OGUNKOYA, M.O., ONIBON, V.O. & E. AJAYI (2007): Compositional studies on tropical species of Agama agama lizards. – Int. J. Zool. Res., 3 (4): 218-222. ADEGOKE, J.A. (1988): Studies on the chromosomes of the rainbow lizard Agama agama agama (L.) with notes on polypoidy in the spermatocytes. - Cytologia 53: 233-239. ADEGOKE, J.A., IGHAVINI, B.O. & R.O. ONUIGBO (1991): Characteristic features of the sonicated DNA of Agama agama agama L. (Reptilia, Agamidae) on hydroxyapatite columns, using mouse DNA as a reference. – Genetica, 83: 171-180. Hydroxyapatite column chromatography has been used to study some properties of the extensively sheared DANN of the Rainbow lizard, Agama agama agama. Reassociation studies show that the genome has a Cot,,? of 370. Approximately 15% of the genome is highly repetitive in nature. This repetitive fraction is resolved into thermally stable and less stable fractions. The stable fraction has a base composition of 47% GC, higher than the 40.2% GC for the native DNA. This stable fraction is believed to be of recent origin. Chromatography of the total DNA of the lizard with linear gradients of phosphate buffer containing 1 M urea resolves it into two components which were shown by thermal fractionation, also in the presence of 1 M urea, to vary in base composition. This behaviour may be characteristic of reptilian genomes and may be used as a basis for studying the structural organisation of the reptilian genome.
Pathogenicity studies were conducted using two bacteria (Proteus mirabilis and Pseudomonas aeruginosa) that were previously recovered from oral and cloacal swabs of red-headed rock agamas (Agama agama) living in poultry farms at Oyo State, Nigeria. Both bacteria were inoculated in two experiments having 60 broilers each. In each experiment, three groups of 15 broilers were given separately graded doses of P. mirabilis or P. aeruginosa. A group of 15 non-inoculated broilers were used in each experiment as negative controls. Clinico-haematological findings, and bacterial examination of cloacal swabs of the broilers were evaluated weekly for 5 weeks post-inoculation (PI). At necropsy, some tissues were harvested for bacteriological and histopathological examinations. Apart from the significant increase (p<0.05) recorded in the total white blood cell and heterophilic counts at week 3 and 4 PI in birds with high and medium doses of Proteus, all haematological values remain within reference intervals in Psedomonas groups. P. mirabilis and P. aeruginosa were isolated from cloacal swabs in all inoculated birds at the 3-4 weeks PI. The Proteus groups (25.0%; 15/60) showed moderate lung and liver congestion, and a few greyish white nodules on the myocardium while extensive myocardiac fiber degeneration, necrosis and loss of striation with numerous inflammatory cells were also observed. This is the first documented report of P. mirabilis induced nodular myocarditis in chickens. These findings also suggested that P. mirabilis could be pathogenic in adult broilers with possible oral transmission of this microorganism from lizards to poultry, thus, could represent a threat to human health. The diet composition of rainbow lizards (Agama agama complex) populations was studied by feces analysis at eight distant places across a mega-transect in the Gulf of Guinea (West Africa), covering three countries: Togo, Benin and Nigeria. The effects of geography (= linear distance between study sites) and local conditions (using the mean annual rainfall as a proxy of the site-specific conditions) on dietary similarity of rainbow lizards were tested. Rainbow lizards were mainly insectivorous at all sites. Multivariate analyses identified four main groups of localities in terms of diet diversity indexes, with populations inhabiting forest towns tending to have less prey taxa richness than conspecifics from more arid areas, which instead had higher dietary evenness. Food niche overlap between populations was high among populations (range 0.631-0.940, x = 0 . 839 ), and decreased with increases in the difference of mean annual rainfall between sites. There was no effect of the geographic distance on the similarity in diet composition between populations. A UPGMA dendrogram revealed a geographic trend in terms of presence/absence of the various prey types in the diets, with all the Nigerian study sites forming one cluster, whereas Lomé and Cotonou, two cities situated within the Dahomey Gap, being grouped apart. Overall, rainfall of the various sites seems to be more important than geographic distance for determining the taxonomic diet composition similarity of these lizards. Most species of lizard are either diurnal or nocturnal, and it is extremely rare to find species that operate both diurnally and nocturnally, or that may shift from diurnality to partial nocturnality when conditions allow. However, niche expansion from diurnal to nocturnal habits (often referred to as the night-light niche) has rarely been reported in lizards (mainly in Anoles), and mostly through anecdotal reports. In West Africa, the rainbow lizard Agama agama is a conspicuous species across the region but also lives in suburban areas of towns and villages. It is a diurnal sun-basker operating at relatively high body temperatures of 36 °C and higher. In this paper, we describe a night-light niche expansion, i.e. nocturnal foraging and thermoregulation, in a small number of A. agama populations living in suburban areas. These lizards utilised radiant heat from incandescent light bulbs situated on the walls of buildings to mostly achieve target body temperatures and forage for and fed on five different groups of invertebrates. Foraging lizards had significantly higher body temperatures than inactive lizards. However, variance in body temperature was significantly greater in foraging lizards than in inactive lizards probably due to the necessity to shuttle between the incandescent night lights and cooler foraging areas during activity, a known cost of thermoregulation. Regression analysis of body temperatures against time of night in foraging lizards supported the notion that the lizards were maintaining body temperatures by actively thermoregulating whilst in inactive non-basking resting lizards during the same time period body temperatures declined. Although our results indicate a potential thermoregulatory benefit from using the night-light shift, we cannot be certain that this benefit is the direct cause of the shift, rather than an additional advantage when foraging. The Eufilariinae of Agama agama from upper Volta Saurositus agamae hamoni n. s. sp. is differentiated from S. a. agamae Macfie, 1924 by small-sized microfilarias and large spicules ; it is occurring in savannah whereas the type species is presumably restricted to forests. At 24-25° its larval development is completed in seven and a half days in the wing muscles of A. stephensi; it exhibits original characters among Reptile Filarians such as a short and very stout sausage-shaped body, a 300 μ long infective stage with very long glandular oesophagus, which usually are only to be found in highly evoluted Filarians of Birds (Eufilariinae and Splendidofilariinae).
BEIER, U. (1974): Observations on the rainbow lizard. – Niger Fld., 39: 137-140. BRINGSØE, H. (1994): Live observations and egg laying of the lizard, Agama agama, in Nigeria. – Nordisk Herpetologisk Forening, 37 (5): 85-91. (in Dänisch) The distribution, population structure and behaviour of agamid: lizards at Nsukka during January and early February 1981 are described. The animals were almost always found near buildings, trees, rubble and fallen logs etc.: they tended to remain in the same place day after day. Territorial behaviour was most marked in adult males, but individuals of all sizes displayed by 'nodding'. Mating is described. The animals roosted at night in trees and shrubs and on buildings, Activity was greatly influenced by weather. Escape from enemies is described, and it is suggested that, in addition to indicating 'social space', 'nodding' may act as a protean anti-predator strategy. These lizards are omnivorous. We report the first documented observations of predation on Dorylus (Anomma) congolensis Santschi, 1910 by the agamid lizard Agama agama (Linnaeus, 1758), the only reptile known to predate on this aggressive army ant. We also report two other observations of predation by Gabonese Agama agama on Dorylus sp. These observations confirm the extreme dietary adaptability of Agama agama. Seasonal histological features of the testis and epididymis were studied in male A. agama agama rom July l990 to June, 1992 at Ile-Ife, Nigeria. Tests weights showed no signicant diference (P> 0.05) in the dry and rainy seasons, but were generally low from August to January. Whereas seminiferous tubule diameter and epithelia heights showed no seasonal variation (p > 0.05), the epididymal tubule diameter and epithelia heights varied seasonally (P < 0001). Although males in full breeding condition were caught all through the months, such were more prevalent from April toJuly. Females with eggs or enlarged ovarian follicles were caught all through the study period. Cases of multiple clutches were predominant fromFebruary to July. However vitellogenc activitiesdecreased from August to January thus coinciding with the observed decrease in spermatogenic activity in the male. We propose that individual male Agama lizards maintain peculiar breeding patterns and that reproduction in Agama seems to be influenced by food availability as well as microclimatic conditions at oviposition sites. Zusammenfassung: Abstract: HALSTEAD, L.B. (1970): Some observations of the rainbow lizard, Agama agama (L.), at the University of Ife. – Niger. Fld., 35: 86-89. HARRIS, V.A. (1963): The anatomy of the rainbow lizard Agama agama (L.). With a glossary of anatomical terms. – Hutchinson tropüical monographs (Anchor Press Ltd.). pp. 104 LEACHÉ, A.D., GRUMMER, J.A., MILLER, M., KRISHNAN, S., FUJITA, M.K., BÖHME, W., SCHMITZ, A., LeBRETON, M., INEICH, I., CHIRIO, L., OFORI-BOATENG, C., ENIANG, E.A., GREENBAUM, E., RÖDEL, M.O. & P. WAGNER (2017): Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). – Syst. Biodiv., 15 (3): 192-203. The savannah and tropical forest biomes of Africa have a long history of expansion and contraction, and the recent and rapid spread of dry savannah habitats has influenced the spatial and temporal diversification of vertebrate taxa across this region. We used a combination of species tree and phylogeographic methods to describe the spatio-temporal changes through time and across space (D species diffusion) in a clade of seven West African lizard species in the Agama agama species group. A Bayesian species tree diffusion approach was used to compare the relative rates at which species ranges changed across the landscape. We found that some species have high diffusion rates characterized by significant movement in their range location and minor changes to their overall range size, whereas other species show little movement in their range centre with an exponential increase in range size. This discrepancy between the rates that range locations shift versus change in their relative area could be linked to populations tracking their preferred habitats through time. A continuous Bayesian phylogeography approach using a relaxed random walk model was used to estimate the timing and rate of population size change and geographic diffusion in A. picticauda, the single species in the group with an extensive African distribution from Mauritania to Ethiopia. The mean dispersal rate of A. picticauda increased dramatically throughout the Pleistocene, and a Bayesian skyride analysis supports exponential population growth over this same time period. A comparison of genetic diversity across different loci and species suggests that A. lebretoni experienced a mitochondrial selective sweep that has caused a deficit of variation at this locus in relation to nuclear loci. Stomach flushing is one of the proposed techniques to study lizard diets. Apparently, it is ranged, together with direct observation and faecal analysis, as a non-harmful method for dietary studies. Some works explored the usefullness of stomach flushing, but we lack information about its effect on lizard's survival probabilities. In this paper we studied the effect of stomach flushing in an urban population of the rainbow lizard (Agama agama) from Calabar (Nigeria). During a period of five months of 2010, 147 lizards were noosed, sexed and individually marked. One group of lizards was stomach flushed only once, whereas the rest of lizards were not flushed. The flushed sample of lizards showed a lower survival than non-flushed lizards of all sex and age classes. In this study, the pictured diet from stomach flushing was very similar to results obtained with a faecal analysis of the same lizard population. Thus, both methods seem to be reliable to study the diet of the rainbow lizard. However, our results indicate that stomach flushing increases the probability of mortality (or at least emigration rates) in all age and sex classes, precluding its extensive use as a method to study lizard's diets.
MADSEN, T. & J. LOMAN (1987): On the role of colour display in the social and spatial organization of male Rainbow lizards (Agama agama). – Amphibia-Reptilia, Leiden, 8: 365-372. (02.254) Abstract: We conducted field studies in 15 West African countries and collected one thousand specimens of lizards of the genus Agama. Based on these collections, literature, molecular analysis of selected specimens, and examination of Linnean type-specimens of A. agama, we review the phylogeny, taxonomy, geographic distribution and ecology of the West African species of the genus Agama. Seventeen different species are recognized in the genus Agama in West Africa, northern Cameroon and Chad: A. africana, A. agama, A. boensis, A. boueti, A. boulengeri, A. castroviejoi, A. cristata, A. doriae benueensis, A. gracilimembris, A. insularis, A. lebretoni, A. paragama, A. sankaranica, A. weidholzi, and three new species. We design a lectotype for A. agama (Linnaeus, 1758) and attribute to A. wagneri, sp. nov., the populations from northern and central Cameroon of the A. agama complex. Agama parafricana, sp. nov., is described from wet savanna areas of Togo and Benin. Agama sylvanus from southern Ghana is a junior synonym of A. africana. Agama cf. impalearis from northern Niger and Mali corresponds to an nondescribed species. Agama boensis is resurrected from the synonymy of A. sankaranica. According to biogeographic areas, four species are Sahelian, seven species are Sudanian, four species are Guinean, and two species are ubiquitous. PAUWELS, O.S.G., BURGER, M., GUIMONDOU, S. & W.R. BRANCH (2004): Agama agama (Linnaeus, 1758). Nocturnal activity. – Afr. Herp News, 37: 20-21. PAUWELS, O.S.G., GRAMENTZ, D. & A. KAMDEM TOHAM (2004): Natural history notes: Agama agama (Red-headed Rock Agama). Nocturnal activity. – Herpetol. Rev., 35 (2): 164-165. PROBST, J.-M. (1999): Nouvelles observations sur le comportement et les colorations variables de l’Agame des colons Agama agama (Linnaeus). - Bulletin Phaethon, 9: 11-12. RABIU, S. (2020): Dimensions of home range structure of Agama agama in the savanna region of Nigeria. – Herpetol. Cons. Biol., 15 (2): 318-324. Home Range (HR) evaluations provide data related to animal searches for food, mates, and other resources. In grasslands of Nigeria where populations of the semi-arboreal African Rainbow Lizard (Agama agama) are dense, I measured three dimensions of HR: home range area (HRA), home range length (HRL), and vertical height (VH). I surveyed 3–4 h in each of four sites between 0600–1300 and 1600–1800 over 6 d each month (June 2015 to May 2016). I recorded sighting grid coordinates and vertical positions of the lizards. After correction to 95% resolution, I programed the coordinates to calculate HRL (and VH) by Adjusted HRL method, and calculated HRA by Minimum Convex Polygon (MCP) method. Highest daily mean HRA (618 m2), HRL (63 m), and VH (262 cm) were for mature males during the dry season. Mature females, juvenile males, and juvenile females, in decreasing order, had lower means for all three dimensions of HR. Age, sex and age-sex classes, as well as seasonal changes had signiicant inluences on mean values of HRA, HRL, and VH. Regression analysis indicated that HRL is a signiicant predictor of HRA. There were age-sex class frequency overlays in all HR dimensions. I recommend a multi-dimensional approach to HR investigations for A. agama (and other semi-arboreal lizards) because of its possible link to age-sex class resource partitioning among other habitat-use interactions.
SCHALL, J.J., BROMWICH, C.R., WERNER, Y.L. & J. MIDLEGE (1989): Clubbed regenerated tails in Agama agama and their possible use in social interaction. – Journal of Herpetology, 23: 303-305. Agama agama is a successful invader that was recently introduced to Santo Antão Island of the Cape Verde archipelago. The species was then noted to be present also in São Vicente and Santiago Islands but the geographic origin of the introductions remains undetermined. Given its wide range and taxonomic identification problems, we compared new 16S RNA sequences with available Agama agama sequences in order to determine if some geographical structure was present that allowed identification of the country of provenance and number of introduction events on the Cape Verde Islands. Some geographical structure of genetic diversity and reasonably large within-group average divergence were revealed, especially in the countries around the Gulf of Guinea. Results support a single origin for this introduction, as a single 16S haplotype is found among the three Cape Verde Islands, and suggest Liberia as the country with the smallest genetic distance to the individuals found in the Cape Verde archipelago. We suggest that local authorities take action against this exotic species as it is spreading and reproducing and predation on endemic species of reptile has been confirmed.
BLUNDEN, T.K. & K.L. KRYSKO (2007): Natural history notes: Agama agama africana (African Rainbow Lizard). Reproduction. – Herpetol. Rev., 38 (1): 73. CAMPBELL, T.S., GODLEY, S., SANDMANN, D. & J. HOROWITZ (2008): Geographic distribution: Agama agama africana (African Rainbow Lizard). USA: Florida. – Herpetol. Rev., 39 (4): 482. DANIEL, P.M. (1960): Growth and cyclic behavior in the West African lizard Agama agama africana. – Copeia, 1960: 94-97. DANIEL, P.M. (1961): Notes on the life-history of Agama agama africana (Hallowell) in Liberia. – Spec. Publ. Ohio Herpet. Soc. 3: 1-5. ENGE, K.M., HUBBUCH, C. & D. HOFFER (2010): Geographic distribution: Agama agama africana (African Rainbow Lizard). – Herpetol. Rev., 41 (1): 106. ENGE, K.M., KRYSKO, K.L. & B.L. TALLEY (2004): Distribution and ecology of the introduced African rainbow lizard, Agama agama africana (Sauria; Agamidae), in Florida. – Florida Scientist, Gainsville, 67: 303-310. We document populations of the introduced African rainbow lizard (Agama agama africana) in Homestead, Miami-Dade County; Hollywood, Broward County; Palm City, Martin County; Punta Gorda, Charlotte County; and Sanford, Seminole County. The Homestead and Punta Gorda populations have been established for over 10 yr and have expanded at least 0.5 km from the point of introduction. The Palm City population has been established since 1999 and the Sanford population since 2000. All agamas were observed in urban or suburban situations perched on walls, rooftops, bridges, rocks, sidewalks, curbstones, or trees. We collected 33 voucher specimens from five populations 28 March 2002–11 March 2004. Maximum clutch size and maximum snout-vent length (SVL) of male and female A. a. africana in Florida exceeded those in native Nigerian populations. All adult females (> 94 mm SVL) collected May–August contained 5–18 vitellogenic follicles or oviductal eggs, but a female collected on 19 September was not gravid. Monitoring should be conducted to determine whether the species might eventually invade natural habitats and its potential impacts on native wildlife species.
Anchieta´s Agama BOCAGE, J.V. B. du (1896): Sur deux Agames d’Angola a ecaillure Heterogene. – Journal de Sciencias Mathematicas, Physicas e Naturaes, Lisboa, 4: 127-130. BUISWALELO, B.N. (2018): A description of reproduction, diet, nematode infection and sexual dimorphism in Agama anchietae and Pedioplanis undata undata (Reptilia: Lacertilia) in Namibia. - Master Thesis, University of Namibia. 31 pp. The objectives of this study were to describe reproduction, diet, nematode infection and sexual dimorphism in Agama anchietae and Pedioplanis undata undata, two Namibian lizard species with wide geographic distributions for which such baseline ecological information is still lacking. The specimens used in the study came from the preserved Herpetological Collection of the National Museum of Namibia. Agama anchietae occupies rocky areas and the findings were as follows, reproduction: (i) it reproduces in spring and summer, the warm and rainy season of Namibia when insect food abounds, presumably to optimize the survival of its offspring, (ii) the gonads were inactive during autumn and winter, (iii) the mass of its abdominal fat bodies and liver decreased during the reproductive period, presumably because they serve as sources of energy during that time; diet: (i) prey items consumed by the sexes belonged to the orders Hymenoptera, Coleoptera, Lepidoptera and Mantodea, (ii) in addition to these an item belonging to the Diptera was also found in males, (iii) no significant numerical differences were found between the sexes with respect to the items consumed, (iv) the fact that hymenopterans occurred in substantially higher numbers suggest that the species relies mainly on sit-and-wait foraging; nematode infection: (i) in both sexes a larger number of nematodes occurred in the stomach than in the intestine, (ii) no significant numerical differences in infection were found between the sexes; sexual dimorphism: (i) males were larger than females in absolute body size, (ii) relative hind-limb but not forelimb length of males was significantly longer than that of females; (iii) relative head breath, length and width of males were all significantly greater than that of females. Pedioplanis undata undata occurs in flat open areas and the findings were as follows, reproduction: (i) a similar pattern with respect to the activity of the gonads as reported for A. anchietae above was also found in this species, (ii) the same also applied to the mass fluctuations of the abdominal fat bodies and the liver; diet: (i) males consumed items belonging to the orders Hymenoptera, Isoptera, Coleoptera, Lepidoptera, Blatoidea, Solifugida as well as larvae, (ii) in females the prey items belonged to the Hymenoptera, Orthoptera, Coleoptera, Hemiptera and Aranea, (iii) no significant numerical differences were found between the sexes for similar prey items, (iv) the low numbers of hymenopterans but fairly high diversity of prey types suggest that this species is a wide forager; nematode infection: (i) in both the stomach and intestine nematode infection was low with that of the stomach marginally higher than that of the intestine, (ii) no significant numerical differences were found between males and females; sexual dimorphism: (i) males and females were of similar body size, (ii) males had significantly longer fore-and hind-limbs than females, and may therefore be able to run faster, (iii) relative head breath, length and width of males were all significantly greater than that of females. Agama anchietae is one of eight species of agama found in Namibia, its distribution range in the upper half of the country covers desert, Karoo and savannah type biomes and overlaps with that of some of its congeners. Here, we describe its sexual dimorphism, reproductive traits, predation, diet and nematode infection, and compare and contrast each aspect among the three biomes as well as to published findings for three other Namibian congenerics, Agama etoshae, Agama aculeata aculeata and Agama planiceps planiceps. Interesting similarities and differences were found among the biomes as well as with the three congenerics. Our hypothesis that the aspects studied in A. anchietae would be more in line with those of A. a. aculeata and A. etoshae, with which it shares similarities in body colouration, social organisation and microhabitat utilisation, was only partially confirmed. This cautions against using morphological and ecological similarities between agamas as proxies for making lifehistory strategy inferences. In the course of recent taxonomic studies in the African Agamidae the West African species were examined and new species have been identified. In this publication a new species of the genus Agama Daudin, 1802 is described from Cameroon and Gabon. Males of the new species differ from all other known Agama in the unique combination of the reticulate colouration of the throat and the blue tip of tail. It is compared with the other Cameroonian members of the genus and several other Agama species from Africa. Additionally, the status of Agama mehelyi Tornier, 1902, only known from its holotype, is discussed.
Bodenagame DIECKMANN, M. (2005): Das Agamenportrait. Bodenagame Agama armata PETERS, 1854. – Iguana-Rundschreiben, 18 (1): 4-8. Zusammenfassung: Analysis of stomach contents can provide insights into foraging mode, habitat use, and dietary specialization of animals. In this paper, we make observations on the poorly known diet of four southern African agamid species, Agama aculeata distanti (Eastern Ground Agama), Agama armata (Peter’s Ground Agama), Agama atra (Southern Rock Agama), and Acanthocercus atricollis (Southern Tree Agama). We examined the diet of 67 individuals by identifying and weighing prey items after stomach flushing lizards in the field. We found that these agama species fed on a broad spectrum of arthropods (11 orders). A high relative importance of ants was present for all agama species examined here, which suggests that ants are a major food source in the arid ecosystem. We found that active prey such as ants, beetles, and highly mobile flying insects like wasps and flies to be major components of the diet, indicating that these lizards are ambush predators. We also found that 43% of the stomachs contained herbaceous material and 39% contained sand particles. Agama atra had the most diverse dietary niche, eating fewer ants and more beetles, hemipterans, and dipterans than other species, whereas A. armata had a narrower dietary niche consisting mainly of ants. Lastly, although low in sample size, we found that juveniles qualitatively had a diet of functionally similar prey items, albeit with a narrower niche breadth, when compared to adults. We discuss how diet corresponds with differences in foraging behavior and habitat specialization.
Southern Rock Agama BOULENGER, G.A. & J.H. POWER (1921): A revision of the South African agamas allied to Agama hispida and Agama atra. – Transact. Roy. Soc. S. Africa, Caope Town, 9: 229-287. BRUTON, M.N. (1977): Feeding, social behaviour and temperature preferences in Agama atra Daudin (Reptilia, Agamidae). – Zoologica, afr., 12 (1): 183-199. BURRAGE, B.R. (1974): Population structure in Agama atra and Cordylus cordylus cordylus in the vicinity of De Kelders, Cape Province. – Annals S. Afr.Mus., 66 (1): 1-23. FLEMMING, A.F. & P. le F.N. MOUTON (2000): Geographic variation in sexual size dimorphism in the rock agama, Agama atra (Sauria: Agamidae). – Afr. Zool., 35 (2): 233-249. Significant sexual dimorphism in overall size occurs in the rock agama, Agama atra (Sauria; Agamidae), with males growing larger than females. Geographic variation in the degree of sexual size dimorphism also exists, males growing significantly larger than females in Namaqualand and Namibia compared to populations in other areas. Sexual differences in scaling of head, limb and tail dimensions were mainly the result of differential asymptotic sizes reached by the sexes. Head size was also influenced by a faster increase in head dimensions with increasing snout to vent length in males compared to females, probably as a result of sexual selection. In females, scaling of limb and tail dimensions was decreased compared to males, possibly a result of differential energy allocation to reproduction. Aim Vicariance has played a major role in the evolution of the southern rock agama, Agama atra (Reptilia: Agamidae), and it is hypothesized that habitat shifts will affect small-scale patterns of gene flow. The Cape Floristic Region (CFR) is known for high levels of diversity and endemism; thus we set out to investigate whether genetic structuring of CFR populations of A. atra corresponds to regional environmental shifts. Analysis of stomach contents can provide insights into foraging mode, habitat use, and dietary specialization of animals. In this paper, we make observations on the poorly known diet of four southern African agamid species, Agama aculeata distanti (Eastern Ground Agama), Agama armata (Peter’s Ground Agama), Agama atra (Southern Rock Agama), and Acanthocercus atricollis (Southern Tree Agama). We examined the diet of 67 individuals by identifying and weighing prey items after stomach flushing lizards in the field. We found that these agama species fed on a broad spectrum of arthropods (11 orders). A high relative importance of ants was present for all agama species examined here, which suggests that ants are a major food source in the arid ecosystem. We found that active prey such as ants, beetles, and highly mobile flying insects like wasps and flies to be major components of the diet, indicating that these lizards are ambush predators. We also found that 43% of the stomachs contained herbaceous material and 39% contained sand particles. Agama atra had the most diverse dietary niche, eating fewer ants and more beetles, hemipterans, and dipterans than other species, whereas A. armata had a narrower dietary niche consisting mainly of ants. Lastly, although low in sample size, we found that juveniles qualitatively had a diet of functionally similar prey items, albeit with a narrower niche breadth, when compared to adults. We discuss how diet corresponds with differences in foraging behavior and habitat specialization. An understanding of the phylogeography and evolutionary processes involved in speciation is essential for the conservation and management of any particular species. To investigate the phylogeographic patterns in Agama atra from the Cape Fold Mountains (CFM), 98 individuals from 38 geographically close localities were analysed. In addition, to understand the phylogeographic associations between the CFM populations and the rest of Southern Africa, 18 specimens from 12 localities outside the CFM were also included. A total of 988 characters derived from two mitochondrial DNA fragments (control region and ND2) revealed 59 distinct haplotypes in the CFM. Parsimony, Bayesian and maximum likelihood analyses revealed four distinct clades associated with geography within the CFM. These clades were supported by a haplotype network and were defined as the Cape Peninsula clade, the Limietberg clade, the northern CFM clade and the central CFM clade. Analysis of molecular variance confirmed the high degree of genetic structure within the CFM, with more than 75% of genetic variation found among the geographic areas. SAMOVA and nested clade analysis (NCA) suggest that the central CFM clade may be more diverse than detected by the networks and the phylogenetic analyses. The processes that caused the four distinct genetic groups in the CFM are not yet clear. Using a speculative molecular clock estimate, the main cladogenesis of A. atra within the CFM took place, approximately ~6.5 - 9 MYA. This dating coincides well with the documented Miocene-Pliocene climate fluctuations which might have contributed towards the isolation among lineages. The genetic structure found in A. atra is also markedly congruent with what has been found in other taxa such as Mesamphisopus spesies, Potamonautes brincki, and Pedioplanis burchelli and this would further support vicariance as a main isolating factor here.
Atlasagame BOETTGER, O. (1874): Description of Agama impalearis. – In: “Reptilien von Marocco und von den canarischen Inseln”. - Abh. senckenb. naturf. Ges. (Frankfurt), 9: 121-191 [1873]. BONS, J. (1963): Note preliminaire sur l´oriéntation de l´embryon dans l´oeuf chez le lizard Agama bibroni Dum. – Bull. Soc. Sci. nat. Maroc, 43: 39-47. BONS, J. (1968): Comportement d’Agama bibroni A. Dum. 1851 (Sauria) durant la periode de reproduction. – Bull. Soc. Sci. nat. phys. Maroc, 48 (3): 93-99. A study of wild Agama bibroni indicated that adults fed largely on Orthoptera with an active selection for species in the middle to large size range. The juvenile diet was mainly of Hymenoptera, Formicidae. This dietary difference led to a study of jaw mechanisms suggesting that due to a differential growth of the cranial bones, particularly those of the lower jaw, the mechanical advantages of the adult and juvenile jaws differed considerably. This allowed the rapid bite of the adult to be two and a half times more powerful, per unit area, than that of the juvenile. During slow close there was almost no difference in efficiency. From this it is suggested that the differing diets are not only the result of different habitats but also due to skeletal arrangements. This is the first description of the embryonic development of an agamid lizard from North Africa, Agama impalearis. The developmental stages of only one species of the same family, Calotes versicolor, had been described previously. Two complementary studies had been performed on this Asian species, one dealing with intra-oviductal development, the other with extra-oviductal development. Among other iguanians, several cameleonid species and only two American iguanid species had apparently been examined. The other previous studies dealt with some species among the Lacertidae (complete or partial embryonic development), Scincidae, Cordylidae and Anguidae (partial embryonic development). In the present table, 42 stages have been distinguished during the embryonic development of A. impalearis. Stages 1 to 27 correspond to intra-oviductal development including egg segmentation, gastrulation, neurulation and the beginning of organogenesis. Stages 28 to 42 describe morphological changes during extra-oviductal development, i.e. between egg-laying and hatching, and correspond to the continuation of the organogenesis. Although data in other groups are still incomplete, a comparison of the A. impalearis table with that of other lizards reveals a great homogeneity of embryonic development in these reptiles, except for the external ornamentation which is specific. Patterns of growth were studied between 1993 and 1996 in a population of Bibron's agama Agama impalearis from an arid area in the central Jbilet Mountains, western Morocco. Non-linear regressions were used to model snout±vent lengths (SVL) and body masses of individuals caught during 1993±94 against skeletochronology age estimates. The overlap between male and female asymptotic SVLs (A) was negligible demonstrating sexual dimorphism, although there was no evidence of differences in characteristic growth rates (k) between sexes (males: A= 119.43. 2.29 mm and k= 0.849. 0.09 [month-1], females: A= 108.59 . 3.19 mm and k= 1.079 . 0.162 [month-1]). Asymptotic body mass differed substantially between sexes (males: 71.2. 3.1 g, females: 45.8. 4.2 g), but corresponding characteristic growth rates were similar (0.753. 0.339 and 0.789 . 0.623 [month-1], respectively). The mark±recapture method provided generally higher and more reliable asymptotic size estimates (using the logistic-by-length nonlinear regression model) than the skeletochronology-based estimates. Growth parameters were also estimated on an individual year basis. Only male characteristic growth rate showed a signi®cant year-toyear variation (0.010. 0.006 to 0.024 . 0.007 [day-1]), although this may have been due to a lack of statistical power. Annual variation in the absolute growth rate was detected only in hatchlings (0.087. 0.018 to 0.273 . 0.132 mm.day-1) and adult males (0.089. 0.030 to 0.206 . 0.100 mm.day-1). The aim of the present work is to investigate the female reproductive cycle of the North African agamid, Agama impalearis. Histological changes of the whole ovarian cycle were described. Folliculogenesis and oocyte growth were comparable to the general lizard pattern. Asingle germinal bed per ovary was found. Only previtellogenic follicles were present during the non reproductive period and the onset of vitellogenesis was observed just after emergence from winter dormancy. Two vitellogenic cycles which probably produce two successive clutches per breeding season were observed. The number of vitellogenic follicles can reach 12 per ovarian vitellogenic cycle. The oocyte growth was accompagnied by changes in the theca, granulosa, zona pellucida and zona radiata layers. The postovulatory follicles (corpora lutea) were observed until the onset of the second vitellogenic cycle and disappeared rapidly after them. Two kinds of atretic follicles, hydratation stage and vitellogenic stage atretic follicles, were described. Atresia was less frequent in Agama impalearis and particularly concerned with larger vitellogenic follicles. The effects of temperature on incubation time, embryo survival, sex ratio , embryo growth and size at hatching were investigated in the north African Agamid lizard, Agama impalearis. Seven constant temperature treatments (spanning 20-36°C) were employed and a split clutch design was used to assign eggs from the same clutch to the different treatments. Incubation time varied significantly with temperature treatments. Embryos incubated at 32 °C, 34 °C and 36 °C hatched between 41 and 46 days, whereas embryos incubated at 26 °C and 28 °C hatched at 83 and 67 days respectively. Hatching success was higher at 28 °C, 30 °C, 32 °C and 34 °C, but much lower at 26 °C and 36 °C; hatching did not occur at 20 °C. Eggs incubated at 26 °C and 36 °C produced only females. At 28 °C, 30 °C, 32 °C and 34 °C, the percentages of males were 9%, 5 3 . 5%, 32%, and 58% respectively. These sex ratios can be explained by a temperature-dependent mechanism of sex determination. The relative growth rates are highest early in incubation and lower for several days prior to hatching. The relationship between snout-to-vent length and age of embryos seems to be best described by a polynomial fitted regression. Growth rates at 26 °C were much lower than those at 34 °C. Constant incubation temperatures affected both snout-to-vent length and body mass at hatching, with max i mum body size occurring at intermediate constant incubation temperatures (30 °C, 32 °C and 28 °C). According to this study, the optimal temperatures of embryonic development probably lies within the range 28-34 °C. The possible adaptive significance of incubation temperature effects on some life history characteristics of A. impalearis is discussed. FRITZ, P. & A. HOLLERBAUM (2015): F²-Nachzucht der Atlasagame Agama impalearis BOETTGER, 1874 im Terrarium. – Sauria, Berlin, 37 (3): 35-36. The social and sexual behaviour of a marked group of Agama impalearis was observed under natural conditions in an arid area of the central Jbilet mountains (Western Morocco) during spring 1 995 . Supplementary observations were made either in Iaboratory or in the field on unmarked individuals at different periods of the year from 1 993 to 1 996. We examined the behavioural repertoire of adults. A quantitative analysis of field data allowed to assess relative time proportions and frequencies of the main behavioural categories. The diurnal emergence of animais from their refuges appears to be greatly influenced by temperature as agamas were observed to emerge in the field at different times of the day but at closely similar thermal conditions. The Iizards are highly heliophilous and predominantly sit-and-wait foragers. Feeding behaviour, escape from ennemies, courtship, oviposition and other behaviours were described. Adult males are strongly territorial and have specifie display action-patterns of aggressive behaviour. The displays involve postural change, pushups and head nodds. They were able to rapidly change their colours. The two sexes showed differences in nuptial colours and behavioural attitudes. Contrary to males, females exhibited neither pushup nor head nodds but a specifie display with mouth-gaping when males are courting them. Monogamy is the predominant mating system and there is no real social hierarchy. Home ranges were larger in males and showed low overlaps. Basking and alert postures are predominant through activity time and pushups are the most frequent display. The diet of adult and juvenile Agama impalearis was studied over a 1-year period at an arid area in western Morocco. Formicidae, Isoptera and Coleoptera constituted most of their food, but various other arthropods and plant materials were also taken. There was marked seasonality in prey size and volumetric proportions of the major food items. The principal seasonal dietary change was the increase of ants and termites during summer in juveniles and adults, respectively, while plants were consumed much less. Differences in prey size selection and proportions of eaten prey between adults and juveniles could be attributed not only to differences in their body size but also to their habitat and activity patterns. Comparisons with other lizard populations from different environments suggested that the importance of Formicidae and/or Isoptera as prey, and probably also the importance of plants during cold months, would be a common feature of the agamid food habits in arid hot ecosystems. A population study of Agama impalearis was conducted during two consecutive years (1993-1994) in a Juj ube bush site located in an arid area at the Jbilet Mountains (Western Morocco) . Lizards of both sexes required approximatively one year to reach sexual maturity. One to two clutches were produced per breeding season (May-August) and clutch size was significantly correlated with maternai body size. The annual reproductive output was in average of 20 eggs per female per year and did not exhibit a significant difference between the two years of study. Population size showed important seasonal changes throughout the study period and absolute spring density estimates were of 15 and 11 indlha, respectively in 1993 and 1994. In the post-reproductive period (September) the population density (excluding hatchlings) markedly declined and was only of 4-5 indlha. After the completion of hatching period (July to early November), the densities of hatchlings were 11 and 13 indlha respectively in 1993 and 1994 compensing for the population losses . The annual turnover rate of the population did not exhibit a significant yearly change and was of 77 %. Adult survivorship after the reproductive activity varied between years and j uvenile survival during the first year was estimated to be 35 %. Maximum longevity assessed by means of squelettochronology was 5 years.
Bocourt´s Agama ROCHEBRUNE, A.T. de (1884): Description of Agama bocourti. – In: “Faune de la Senegambie. Reptiles”. O. Doine, Paris, 1-221.
MONARD, A. (1940): Description of Agama boensis. – In:“Résultats de la mission du Dr. Monard en Guinée Portugaise 1937 –1938”. - Arq. Mus. Bocage, Lisbon, 11: 147-182.
Somali Agama BOULENGER, G.A. (1898): Description of Agama bottegi. – In: “Concluding report on the late Capt. Bottego’s collection of reptiles and batrachians from Somaliland and British East Africa”. - Annali del Museo Civico di Storia Naturale di Genova, (2) 18: 715-723 [1897].
Mali Agama CHABANAUD, P. (1917): Description of Agama boueti. – In: “Enumération des reptiles non encore étudiés de l'Afrique occidentale, appartenant aux collections du Muséum, avec la description des espèces nouvelles”. - Bull. Mus. nat. Hist. nat. Paris, 23: 83-105. MEDIANI, M. &F. CHEVALIER (2016): Agama boueti CHABANAUD, 1917: new to the herpetofauna of Morocco and the northwesternmost record of the species. – Herpetozoa, 28 (3/4): 187-191.
JOGER, U. (1979): Zur Ökologie und Verbreitung wenig bekannter Agamen Westafrikas (Reptilia: Sauria: Agamidae). – Salamandra, Frankfurt/Main, 15 (1): 31-52. (00.034) Zusammenfassung: KARNS, D.R. & M. CISSÈ (1975): Découverte d´Agama boueti CHABANAUD (Reptilia: Sauria: Agamidae) au Sénégal avec notes systématiques et écologiques. – Bull. Inst. Fondament. Afr. noire, sér. A, Dakar, 37 (4): 939-940. PADIAL, J.M. (2005): A new species of Agama (Sauria: Agamidae) from Mauritania. – Herpetological Journal, 15: 27-35. Abstract:
Boulenger´s Agama JOGER, U. (1979): Zur Ökologie und Verbreitung wenig bekannter Agamen Westafrikas (Reptilia: Sauria: Agamidae). – Salamandra, Frankfurt/Main, 15 (1): 31-52. (00.034) Zusammenfassung: Agama boulengeri is a West African endemic lizard. It occurs in arid rocky areas in the Mauritanian mountains and Kayes region of Mali. Data on the distribution of Agama boulengeri is however very coarse, and the contribution of climatic and habitat factors for population isolation are unknown. Using Maxent, GLM, and high-resolution data, we generated environmental niche models, and quantified suitable areas for species occurrence. Field observations and predicted suitable areas were used to evaluate the conservation status of Agama boulengeri. Results revealed the species occurs preferentially close to gueltas, bare areas, and rocky deserts and in areas of increasing rainfall. Suitable cells were mostly located in Mauritania, and four potentially fragmented subpopulations were identified. The conservation status of Agama boulengeri was determined to be of Least Concern. Patterns of biodiversity and evolutionary processes controlling them are still poorly studied in desert biomes. Finescale markers could help answer some of the pressing research questions for desert biomes and Sahara in particular. Such markers are available for some large mammals and crocodiles, but not for small vertebrates. Here we present a battery of microsatellite loci developed for Agama boulengeri, a promising model to study evolutionary and demographic processes in the Sahara-Sahel. Loci were selected by sequencing enriched DNA libraries with 454 pyrosequencing. A total of 23 polymorphic loci were successfully amplified in four multiplex reactions. Cross-amplification of the microsatellite loci in A. agama and A. boueti was partially successful. These markers are a promising tool for assessing genetic diversity, geneflow dynamics and demographic patterns in this group. Given the genus Agama is distributed throughout Africa, results presented here might also facilitate studies in other regions. Abstract:
Eimenteita Rock Agama BERTRAND, M. & D. MODRY (2004): The role of mite pocket-like structures on Agama caudospinosa (Agamidae) infested by Pterygosoma livingstonei sp. n. (Acari: Prostigmata: Pterygosomati-dae). – Folia Parasitologica, 51: 61-66. Abstract:
In the course of a recent taxonomic study on East African Agama species it became clear that a population of A. caudospinosa Meek, 1910 from Mt. Kenya is clearly distinct to specimens from the type locality at Lake Elmenteita (both in Kenya). Therefore, a new subspecies is described on the basis of morphological characters. This new subspecies shows a similar colouration in both sexes which is unique in the genus Agama.
Scortecci´s Agama MOCQUARD, F. (1905): Description de deux nouvelles espèces de Reptiles. – Bull. Mus. Hist. Nat., Paris 11: 288-290. TRAPE, J.-F. (2011): Agama cristata MOCQUARD, 1905 and Agama insularis CHABANAUD, 1918 (Squamata, Agamidae): two valid West African species. – Journal of Herpetology, 45 (3): 352-354. Agama cristata and Agama insularis are two poorly known West African species that are generally regarded either as dubious or synonyms. Agama cristata was described from a single specimen that was partially mutilated, thus preventing a precise count of midbody scale rows. There have been no further records for this species until it was considered as a senior synonym of A. insularis, a species that was until recently known only from Los Islands, off the coast of Guinea. A particularity of the type of A. cristata is the presence of two black spots on each side of the neck. During a field trip in southern Mali, I found a specimen with the same characteristics as the type of A. cristata. Further sampling provided 16 additional similar specimens. Comparison of pattern and meristic data indicates than A. cristata and A. insularis are two distinct species that can be distinguished easily both on pattern and midbody scale count, 71–90 for A. cristata versus 111–147 for A. insularis. Both species also differ by their geographic distribution and habitat. Abstract:
Ostafrikanische Siedleragame BURMANN, A. (2006): Phylogenie & Taxonomie der Agamen (Agama lionotus-Komplex) Ostafrikas: morphologische & genetische Untersuchungen. – Unpublished MSc. Thesis, University of Bonn. 1-244. Agama dodomae dodomae Ostafrikanische Siedleragame Ostafrikanische Siedleragame
Nigeria Agama BOULENGER, G.A. (1885): Description d’une espèce nouvelle d’Agame. – Ann. Mus. Civ. St. Nat. Genova (2) 2: 127-128. MOODY, S.M. & W. BÖHME (1984): Merkmalsvariation and taxonomische Stellung von Agama doriae Boulenger, 1885 und Agama benueensis Monard, 1951 (Reptilia: Agamidae) aus dem Sudangürtel Afrikas. – Bonner zoologische Beiträge, Bonn, 35 (1-3): 107-120. (01.175) 166 Exemplare der Artengruppe Agama doriae / A. benueensis wurden nach 35 morphologischen Merkmalen analysiert und statistisch ausgewertet. Ihre Verbreitung wird auf erläuterten Punktkarten ausführlich dargestellt. Die Ergebnisse zeigen, daß innerhalb beider Arten keine taxonomisch relevanten Gruppierungen abgrenzbar sind, und daß beide nominellen Arten als Subspeczies einer einzigen Art Agama doriae aufzufassen sind. Die nominellen Unterarten sennariensis und kordofanensis stellen Synonyme zur Nominat-Unterart dar. Nigeria Agama Nigeria Agama 166 Exemplare der Artengruppe Agama doriae / A. benueensis wurden nach 35 morphologischen Merkmalen analysiert und statistisch ausgewertet. Ihre Verbreitung wird auf erläuterten Punktkarten ausführlich dargestellt. Die Ergebnisse zeigen, daß innerhalb beider Arten keine taxonomisch relevanten Gruppierungen abgrenzbar sind, und daß beide nominellen Arten als Subspeczies einer einzigen Art Agama doriae aufzufassen sind. Die nominellen Unterarten sennariensis und kordofanensis stellen Synonyme zur Nominat-Unterart dar.
BÖHME, W., WAGNER, P., MALONZA, P., LÖTTERS, S & J. KÖHLER (2005): A new species of the Agama agama group (Squamata: Agamidae) from western Kenya, East Africa, with comments on Agama lionotus BOULENGER, 1896. – Russian Journal of Herpetology, 12 (2): 143-150. HEIDEMAN, N.J.L. (1997): New distribution records for Agama etoshae (McLACHLAN 1981) in Northern Namibia. – Israel J. Zool., 43: 405-407. HEIDEMAN, N.J.L. (1998): Reproduction in Agama etoshae McLACHLAN 1981 from Namibia. – Amphibia-Reptilia, 19 (1): 99-103. HEIDEMAN, N.J.L. (1999): Summer diet of Agama etoshae MCLACHLAN 1981 from Namibia. – Russ. Jour. Herp., 6 (1): 45-47. HEIDEMAN, N.J.L. (2001): Sexual size dimorphism in Agama etoshae (McLachlan 1981). - African Journal of Ecology, 39: 303-305.
Finch’s agama Agama finchi finchi BÖHME WAGNER, MALONZA, LÖTTERS & KÖHLER, 2005 BÖHME, W., WAGNER, P., MALONZA, P., LÖTTERS, S & J. KÖHLER (2005): A new species of the Agama agama group (Squamata: Agamidae) from western Kenya, East Africa, with comments on Agama lionotus BOULENGER, 1896. – Russian Journal of Herpetology, 12: 143-150. Agama finchi leucerythrolaema WAGNER, FREUND, MODRÝ, SCHMITZ & BÖHME, 2011 WAGNER, P., FREUND, W., MODRÝ, D., SCHMITZ, A. & W. BÖHME (2011): Studies on African Agama IX. New insights into Agama finchi Böhme et al., 2005 (Sauria: Agamidae), with the description of a new subspecies. – Bonn zoological Bulletin, Bonn, 60 (1): 25-34. We present new information on the distribution and morphology of Agama finchi from eastern Africa. For the first time, material from three different populations (including the type locality) was available and the question of a possible subspecies in Uganda was positively answered. Based on the distribution pattern of Agama species groups, a general distribution pattern is discussed.
Benin Agama CHABANAUD, P. (1918): Étude complémentaire de deux Agama de l'Afrique occidentale et description de quatre espèces nouvelles de reptiles de la même région. - Bull. Mus. Natl. Hist. Nat. (1918) 24 (2): 104-112. GARTSHORE, M.E. (1985): Agama gracilimembris in Nigeria. – Herp. Jour., 1: 23-25. GRANDISON, A.G.C. (1969): Agama weidholzi (Sauria: Agamidae) of West Africa and its relationship to Agama gracilimembris. – Bull. Inst. Fundament. Afr. noire, sér. A, Dakar, 30 (2): 666-675.
Hartmann´s Agama PETERS, W.C.H. (1869): Eine Mittheilung über neue Gattungen und Arten von Eidechsen. - Monatsber. Königl. Preuss. Akad. Wissensch. Berlin, 1869: 57-66.
Common Spiny Agama BARRY, T.H. (1953): Contributions to the cranial morphology of Agama hispida (Linn.). – Ann. Univ. Stellenbosch, 29A (2): 55-77. BOULENGER, G.A. & J.H. POWER (1921): A revision of the South African agamas allied to Agama hispida and Agama atra. – Transact. Roy. Soc. S. Africa, Caope Town, 9: 229-287. CORNELIUS, A., MARAIS, A. & P. CUNNINGHAM (2012): Agama hispida (Linnaeus, 1754) Southern Spiny Agama. Geographical distribution. – Afr. Herp News, (56): 35-37. DOBIEY, M. (2005): Das Agamenporträt: Agama hispida (LINNAEUS, 1758. – Iguana-Rundschreiben, 18 (2): 15-19. GRAVENHORST, J.L.C. (1833): Über Phrynosoma orbicularis, Trapelus hispidus, Phrynocephalus helioscopus, Corythophanes cristatus und Chamaeleopsis hernandesii. - Acta Acad. Caes. Leop. Carol. Nat. Cur. (?) 16 (2): 909-958. KAUP, J. (1827): Description of Agama hispida. – In: “ Zoologische Monographien”. - Isis von Oken 20: 610—625. McLACHLAN, G.R. (1981): Taxonomy of Agama hispida (Sauria: Agamidae) in southern Africa. – Cimbebasia A 5: 219-227. Agama hispida hispida (KAUP, 1827) Common Spiny Agama Common Spiny Agama
Atlasagame WAGNER, P. (2014): A new cryptic species of the Agama lionotus complex from south of the Ngong Hills in Kenya. – Salamandra, 50 (4): 187-200. East Africa, especially if including the Horn of Africa, is a centre of diversity for African Agamid lizards and harbours the endemic lineage of the Agama lionotus complex, which currently comprises nine species. Species of the complex are mainly characterized by their throat pattern in adult males, which can be used for species identification. Among them, Agama lionotus and Agama dodomae show a very distinct colouration of a blue body and a white/blue annulated tail – a colour pattern that is otherwise only known from the southern African Agama kirkii. Within the complex, Agama lionotus is the most widely distributed taxon, ranging from Ethiopia to northern Tanzania and being replaced by Agama dodomae farther south in Tanzania. Other taxa of the complex are more restricted in their distribution. In this study, specimens from a larger area south of the Ngong Hills are examined and compared with other members of the complex, because they show an overall similarity to Agama lionotus, but are distinctly smaller. Examining the morphological (62 characters) and genetical (16S, ND4, CMOS) data indicates that these specimens represent a new species. Furthermore, phylogenetic analyses support the new taxon as not closely related to Agama lionotus itself, but as a member of the complex. The new species is especially characterized by its small size. Adult males have a vertebral stripe, a blue body colouration and an annulated white/blue tail. Further typical characters are the low number of scale rows around midbody, the pear-shaped and keeled nasal scale, the minute nuchal crest, and the feebly keeled vertebral scales, followed by dorsal and lateral keeled scales. The results of this study improve our understanding of the diversity of agamid lizards in East Africa and support the value of adult male throat coloration for the identification of species within the Agama lionotus complex.
Insular Agama CHABANAUD, P. (1918): Etude d’une collection de reptiles de l’Afrique occidentale française. - Bull. Mus. nation. Mist. nat. Paris, 24: 160-166. TRAPE, J.-F. (2011): Agama cristata MOCQUARD, 1905 and Agama insularis CHABANAUD, 1918 (Squamata, Agamidae): two valid West African species. – Journal of Herpetology, 45 (3): 352-354. Agama cristata and Agama insularis are two poorly known West African species that are generally regarded either as dubious or synonyms. Agama cristata was described from a single specimen that was partially mutilated, thus preventing a precise count of midbody scale rows. There have been no further records for this species until it was considered as a senior synonym of A. insularis, a species that was until recently known only from Los Islands, off the coast of Guinea. A particularity of the type of A. cristata is the presence of two black spots on each side of the neck. During a field trip in southern Mali, I found a specimen with the same characteristics as the type of A. cristata. Further sampling provided 16 additional similar specimens. Comparison of pattern and meristic data indicates than A. cristata and A. insularis are two distinct species that can be distinguished easily both on pattern and midbody scale count, 71–90 for A. cristata versus 111–147 for A. insularis. Both species also differ by their geographic distribution and habitat. Abstract:
LOVERIDGE, A. (1935): Description of Agama kaimosae. – In: “Scientific results of an expedition to rain forest regions in Eastern Africa – I. New reptiles and amphibians from East Africa”. – Bull. Mus. Comp. Zool. Harv. Coll., 79 (1): 3-19. WAGNER, P., BURMANN, A. & W. BÖHME (2008): Studies on African Agama II. Resurrection of Agama agama kaimosae LOVERIDGE, 1935 (Squamata: Agamidae) from synonymy and its elevation to species rank. – Russian Journal of Herpetology.
Kirk´s Rock Agama BOULENGER, G.A. (1885): Description of Agama kirkii. – In: “Catalogue of the Lizards in the British Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae”. London. 450 pp BROADLEY, D.G. (2002): Agama kirkii Boulenger 1885. Kirk’s Rock Agama. Geographical distribution. – Afr. Herp News, 35: 22-23. Agama kirkii kirkii BOULENGER, 1885 Kirk´s Rock Agama Kirk´s Rock Agama
WAGNER, P., LEACHÉ, A., MAZUCH, T. & W. BÖHME (2013): Additions to the lizard diversity of the Horn of Africa: Two new species in the Agama spinosa group. - Amphibia-Reptilia, 34: 363-387. The Horn of Africa is a center of diversity for African agamid lizards. Among the nine species of Agama occurring in the Horn of Africa, Agama spinosa is the most widely distributed. The A. spinosa group (sensu stricto, morphologically defined by possessing six clusters of spinose scales around the ear) contains two species: A. spinosa occurs from Egypt to Ethiopia and Somalia where it is replaced by the morphologically distinct and therefore sensu lato A. bottegi. Both species are only represented in museum collections by a small number of specimens from Ethiopia and Somalia, presumably the result of constant civil war that has plagued the region for decades and impeded field surveys. In this study, we examine species limits in the A. spinosa group using molecular genetic data (503 characters; mitochondrial 16S rRNA) and morphological data (67 characters). Deep divisions among populations of A. spinosa are supported by phylogenetic analyses and by multivariate analyses of morphometric data. Two new species from northern Somalia that differ from A. spinosa and A. bottegi are described. Furthermore, A. smithi, currently recognized as a synonymof A. agama, is re-assessed and recognized as a species of uncertain taxonomic position (i.e., incertae sedis). The results of this study improve our understanding of the evolution of agamid lizard diversity in the Horn of Africa, a significant biodiversity hotspot in Africa.
Ostafrikanische Siedleragame SANCHO, V. & O.S.G. PAUWELS (2015): An Accidental Importation of an Afrotropical Anthropophilic Lizard (Squamata: Agamidae: Agama lebretoni) into Spain. - Bull. Chicago Herp. Soc., 50 (12): 218-219. WAGNER, P. (2009): The arid corridor from Middle East to Africa – Insights from the Agamidae. - Abstracts of presentations hold on DeAGAMIS the 1st International Symposium on Agamid Lizards. Bonner Zoologische Beiträge, Bonn, 56 (4): 299. WAGNER, P., BAREJ, M.F. & A. SCHMITZ (2009): Studies on African Agama VII. A new species of the Agama agama-group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehelyi Tornier, 1902. – Bonner zoologische Beiträge, Bonn, 56 (4): 285-297.
Ostafrikanische Siedleragame BÖHME, W., WAGNER, P., MALONZA, P., LÖTTERS, S & J. KÖHLER (2005): A new species of the Agama agama group (Squamata: Agamidae) from western Kenya, East Africa, with comments on Agama lionotus BOULENGER, 1896. – Russian Journal of Herpetology, 12: 143-150. BREITFELD, M. (?): Kurze Bemerkungen zur Nahrung der Siedleragame. – Die Aquarien- und Terrarien-Zeitschrift, Stuttgart. 754. (00.356) BURMANN, A. (2006): Phylogenie & Taxonomie der Agamen (Agama lionotus-Komplex) Ostafrikas: morphologische & genetische Untersuchungen. – Unpublished MSc. Thesis, University of Bonn. 1-244. KNAACK, J. (1999): Beobachtungen einer Eiablage der ostafrikanischen Siedleragame (Agama lionotus) am Kilifi Creek, Kenia. – herpetofauna, Weinstadt, 21 (123): 10-11. The laying of eggs by the east african rainbow lizard (Agama lionotus) on the bank of the Kilifi Creek in kenya was observed and documented photographically. A gravid female inspected a clear sandy area of approx. 6 m², 1 m above the highest water level on 11 May 1996 at 10.25 am. From 10.32 am to 12.28 am the female dug a hole 140 mm deep with a Ø of 55-60 mm – stopping only for security pauses. The first and second deposits at 12.57 am and 1.12 pm were unseccesful. Between 1.45 pm and 4.12 pm further 7 eggs were deposited in the hole. At 4.35 pm the female began to cover the hole with her hindlegs – interrupted by a long pause. At 5.54 pm the cave with the 9 eggs was invisible; the female could be seen in the proximity of the breeding site for another four days. Ostafrikanische Siedleragame KNAACK, J. (1999): Beobachtungen einer Eiablage der ostafrikanischen Siedleragame (Agama lionotus) am Kilifi Creek, Kenia. – herpetofauna, Weinstadt, 21 (123): 10-11. (02.655) Abstract:
WAGNER, P. & A. BAUER (2011): A new dwarf Agama (Sauria: Agamidae) from Ethiopia. – Breviora, 527: 1-19. A new dwarf lizard of the genus Agama Daudin, 1802 (Sauria: Agamidae), is described from central Ethiopia. This dwarf agama (, 59 mm snout–vent length [SVL]) was compared with other dwarf congeners (, 65 mm SVL) in West, Central, and East Africa and to medium-sized agamas (, 92 mm SVL) occurring in the Horn of Africa. The new species is characterized by a homogeneous body scalation; keeled but nonmucronate dorsal scales; smooth ventral, gular, and upper head scales; and having the nasal scale on the canthus rostralis. It is morphologically well differentiated from all congeners to which it was compared in having a nuchal crest, few tufts of short spinose scales, and the nasal scale on the canthus rostralis and can further be identified by its large occipital scale, smooth gular and ventral scales, and keeled, but not strongly mucronate or spinose, dorsal scales. The new species may live in colonies, like many of the larger rock Agama species, and based on the presence of multiple age classes at the same time of year, it is probably not an annual species, as are some other dwarf Agama. The affinities of the new species are unclear but, on zoogeographic grounds, may lie with congeners to the west of the Rift Valley.
Montane Rock Agama BARBOUR, T. & A. LOVERIDGE (1928): A comparative study of the herpetological fauna of the Uluguru and Usambara mountains, Tanzania Territory with descriptions of new species. – Mem. Mus. Comp. Zool. Cambridge (Massachusetts), 50 (2): 85-265.
Mozambique Agama ANANJEVA, N.B. & G. PETERS (1981): On the validity of Agama pawlowskii, Cernov et Dubinin, 1946. – Trud. Zool. Inst. Akad. Nauk SSR, 101: 21-22. CERNOV, S.A. & V.B. Dubinin (1946): A new endemic from the mountains of Central Asia, Agama pawlowskii sp. nov. (Reptilia, Sauria). – C.R. Acad.Sci. Moscow N.S., 52 (8): 741-743.
Siedleragame / Common Agama HELMICH, W. (1957): Herpetologische Ergebnisse einer Forschungsreise in Angola. - Veröff. Zool. Staatssammlung München, 5: 1-91. BARTS, M. (2003): Agama mwanzae LOVERIDGE, 1923 Ostafrikanische Felsenagame oder Mwanza-Flachkopfagame. – Reptilia, Münster, 8 n(4): 51-54. BURMEISTER, E.G. (1989): Agama mwanzae erbeutet einen Finken. – Die Aquar. Terrar. Z., Stuttgart, 42 (10): 635. HOLTMANN, T. (2010): Die Fliederagame Agama mwanzae ist ein Traum. – TerralogNews No. 93: 15-17. LOVERIDGE, A. (1923): Notes on East African lizards colected 1920-1923 with the description of two new races of Agama lionotus Blgr. – Proceeding of the Zoological Society of London, 1923: 935-969. MENEGON, M., SPAWLS, S., WAGNER, P. & J. BERADUCCII (2014): Agama mwanzae. - The IUCN Red List of Threatened Species 2014: e.T170371A44810595. YARNELL, R.W. & B.H. JONES (2001): Notes on the behaviour and Morphology of Agama mwanzae in northern Tanzania. – Afr. Herp. News, 33: 4-9.
MEDIANNIKOV, O., TRAPÉ, S. & J.-F. TRAPÉ (2012): A molecular study of the genus Agama (Squamata: Agamidae) in West Africa, with description of two new species and a review of the Ttxonomy, geographic distribution, and ecology of currently recognized species. - Russ. J. Herpetol., 19 (2): 115-142. We conducted field studies in 15 West African countries and collected one thousand specimens of lizards of the genus Agama. Based on these collections, literature, molecular analysis of selected specimens, and examination of Linnean type-specimens of A. agama, we review the phylogeny, taxonomy, geographic distribution and ecology of the West African species of the genus Agama. Seventeen different species are recognized in the genus Agama in West Africa, northern Cameroon and Chad: A. africana, A. agama, A. boensis, A. boueti, A. boulengeri, A. castroviejoi, A. cristata, A. doriae benueensis, A. gracilimembris, A. insularis, A. lebretoni, A. paragama, A. sankaranica, A. weidholzi, and three new species. We design a lectotype for A. agama (Linnaeus, 1758) and attribute to A. wagneri, sp. nov., the populations from northern and central Cameroon of the A. agama complex. Agama parafricana, sp. nov., is described from wet savannah areas of Togo and Benin. Agama sylvanus from southern Ghana is a junior synonym of A. africana. Agama cf impalearis from northern Niger and Mali corresponds to an undescribed species. Agama boensis is resurrected from the synonymy of A. sankaranica. According to biogeographic areas, four species are Sahelian, seven species are Sudanian, four species are Guinean, and two species are ubiquitous.
False Agama GRANDISON, A.G.C. (1968): Nigerian lizards of the genus Agama (Sauria: Agamidae). – Bull. Br. Mus. nat. Hist. (Zool.), London, 17 (3): 67-90.
Similar Agama PARKER, H. W. (1942): The lizards of British Somaliland. - Bull. Mus. Comp. Zool. Harvard, 91: 1-101.
NUÑEZ, KRYSKO, K.L. & M.L. AVERY (2016): Confirmation of introduced Agama picticauda in Florida based on molecular analyses. – Bull. Florida Mus. Hat. Hist., 54 (9): 138-139. As of 2010, Florida had the largest number of introduced and established non-indigenous herpetofaunal species in the world. With the advent of molecular methodologies, researchers now are able to test hypotheses regarding introduction pathways, species identity, and native range origins. African Agamas, Agama Daudin 1802, found in Florida are hypothesized to be the African Rainbow Lizard, A. agama africana Hallowell 1844, based on color patterns and hypothesized native range origins (i.e., Benin, Ghana, and Togo) of imported specimens for the pet trade. However, recent systematic studies within the native range of the Agama complex have resulted in multiple taxonomic revisions, which calls into question the species identity of introduced populations in Florida. The purpose of this study is to determine the species identity of African agamas within Florida, as well as the native range origins of Florida populations. We conducted a comparative maximum likelihood analysis between individuals from Florida and individuals from the native range. Based on our results we determined that the species found in Florida is Agama picticauda from western Africa.
Namib Rock Agama BILLAWER, W.H. & N.J.L. HEIDEMAN (1996): A comparative analysis of diurnal behavioural activities in males of Agama aculeata aculeata and Agama planiceps planiceps in Windhoek, Namibia. – Journ. Herpet. Ass. Afr., 45 (2): 68-73. CARTER, A., GOLDIZEN, A.W. & R. HEINSOHN (2012): Personality and plasticity: temporal behavioural reaction norms in a lizard, the Namibian rock agama. – Anim. Behav., 84 (2): 471-477. Behavioural traits are often plastic and can allow animals important flexibility when environmental conditions vary. However, studies of animal personality suggest that behaviour can be ‘constrained’ to certain behavioural types, leading to consistent differences between individuals. Few studies have investigated the interaction between personality traits and the need for flexibility in individuals over different, naturally occurring situations in the wild. We investigated whether free-living Namibian rock agamas, Agama planiceps, were constrained by personality types or showed behavioural plasticity when their environmental conditions changed dramatically between the dry and rainy seasons. We examined three key behaviours: risk taking as measured by flight initiation distance, time budgets focusing on time spent conspicuous and rates of signalling directed at conspecifics. Risk-taking behaviour of males showed stable between-individual differences, but no between-individual differences in plasticity, supporting previous evidence that antipredator behaviour is determined by personality in this species. Time spent conspicuous showed a response to season, and an interaction between the individual and season, suggesting that behavioural plasticity itself may be a personality trait. In contrast, signalling behaviour changed in response to season but did not vary consistently between individuals. We suggest that where there are reliable environmental cues, individual plasticity will be favoured over consistency in the relevant behaviours. COWLEY, T. & P. CUNNINGHAM (2004): Agama planiceps PETERS, 1862 as prey item for Black Mongoose Galerella (sanguinea) nigrata [Short Note]. - Herpetozoa, 17 (1/2): 86. CUNNINGHAM, P. (2011): Agama planiceps (Peters, 1862) Namibian Rock Agama. Diet. – Afr. Herp News, (55): 19-20. GRUBERMANN, M.(2013): Einige fotografische Beobachtungen an Agamen in Kenia, Tansania, Malawi, Südafrika und Namibia. – Iguana, 26 (1): 23-33. HAMMERMANN, G. (1971): Belevenissen met enkele hagedissen. - Lacerta, 29 (4): 41-42. HEIDEMAN, N.J.L. (1990): Life History Note: Agama planiceps: Reproduction. – Jour. Herp. Ass. Afr., 37: 50. HEIDEMAN, N.J.L. (1992): Comparative reproductive biology, and aspects of behaviour and ecology of Agama aculeata aculeata and Agama planiceps planiceps (Reptilia: Agamidae) in the Windhoek area. – Unpublished Ph.D. dissertation, University of Stellenbosch. HEIDEMAN, N.J.L. (1993): Social organization and behaviour of Agama aculeata aculeata and Agama planiceps planiceps (Reptilia: Agamidae) during the breeding season. – J. Herpetol. Assoc. Afr., 42: 28-31. HEIDEMAN, N.J.L. (1994): Reproduction in Agama aculeata aculeata and Agama planiceps planiceps females from Windhoek, Namibia. – Amphibia-Reptilia, Leiden, 15: 351-361. (00.353) Agama aculeata aculeata and Agama planiceps planiceps females were reproductively active during summer, which is the rainy season in Namibia. Both species were iteroparous; clutch size and relative clutch mass in the smaller A. a. aculeata were significantly larger than in A. p. planiceps. Egg and newborn hatchling size of A. a. planiceps on the other hand were significantly greater than in A. a. aculeata. It is suggested that these differences may have evolved as a result of different strategies which the two species possibly employ to cope with predation pressure, namely, evasion through crypsis in A. a. aculeata and through speed of escape in A. p. planiceps. HIGHCOCK, L. & A.J. CARTER (2014): Intraindividual variability of boldness is repeatable across contexts in a wild lizard. - PLoS ONE, 9 (4): e95179. Animals do not behave in exactly the same way when repeatedly tested in the same context or situation, even once systematic variation, such as habituation, has been controlled for. This unpredictability is called intraindividual variability (IIV) and has been little studied in animals. Here we investigated how IIV in boldness (estimated by flight initiation distances) changed across two seasons—the dry, non-breeding season and the wet, breeding season—in a wild population of the Namibian rock agama, Agama planiceps. We found significant differences in IIV both between individuals and seasons, and IIV was higher in the wet season, suggesting plasticity in IIV. Further, IIV was highly repeatable (r = 0.61) between seasons and we found strong negative correlations between consistent individual differences in flight initiation distances, i.e. their boldness, and individuals’ IIVs. We suggest that to understand personality in animals, researchers should generate a personality ‘profile’ that includes not only the relative level of a trait (i.e. its personality), but also its plasticity and variability under natural conditions.
Robecchi´s Agama BOULENGER, G.A. (1891): Description of Agama robecchii. – In: “On some Reptiles collected by Sig. L. Bricchetti Robecchi in Somaliland”. - Ann. Mus. St. nat. Genova (2) 12: 5-15. CHERCHI, M.A. (1958): Note su Agama robecchii Blgr. (Sauria). – Atti Soc. ital. Sci. nat., 97: 233-238.
Arboreal Agama VAILLANT, L. (1882): Description of Agama rueppelli. – In: “Reptiles et Batraciens. in: Révoil: Mission G. Révoil aux pays Çomalis, Faune et Flore. Faune et Flore des Pays Çomalis, Afr. orient”., 25 pp, 3 plates. Agama rueppelli rueppelli VAILLANT, 1882 Arboreal Agama Arboreal Agama Arboreal Agama
Senegal Agama CHABANAUD, P. (1918): Étude complémentaire de deux Agama de l'Afrique occidentale et description de quatre espèces nouvelles de reptiles de la même région. - Bull. Mus. nation. Hist. nat. Paris, 24: 104-112 JOGER, U. (1979): Zur Ökologie und Verbreitung wenig bekannter Agamen Westafrikas (Reptilia: Sauria: Agamidae). – Salamandra, Frankfurt/Main) 15 (1): 31-52. (00.609) Zusammenfassung:
WAGNER, P., LEACHÉ, A., MAZUCH, T. & W. BÖHME (2013): Additions to the lizard diversity of the Horn of Africa: Two new species in the Agama spinosa group. - Amphibia-Reptilia, 34: 363-387. The Horn of Africa is a center of diversity for African agamid lizards. Among the nine species of Agama occurring in the Horn of Africa, Agama spinosa is the most widely distributed. The A. spinosa group (sensu stricto, morphologically defined by possessing six clusters of spinose scales around the ear) contains two species: A. spinosa occurs from Egypt to Ethiopia and Somalia where it is replaced by the morphologically distinct and therefore sensu lato A. bottegi. Both species are only represented in museum collections by a small number of specimens from Ethiopia and Somalia, presumably the result of constant civil war that has plagued the region for decades and impeded field surveys. In this study, we examine species limits in the A. spinosa group using molecular genetic data (503 characters; mitochondrial 16S rRNA) and morphological data (67 characters). Deep divisions among populations of A. spinosa are supported by phylogenetic analyses and by multivariate analyses of morphometric data. Two new species from northern Somalia that differ from A. spinosa and A. bottegi are described. Furthermore, A. smithi, currently recognized as a synonymof A. agama, is re-assessed and recognized as a species of uncertain taxonomic position (i.e., incertae sedis). The results of this study improve our understanding of the evolution of agamid lizard diversity in the Horn of Africa, a significant biodiversity hotspot in Africa.
Lanza´s Spiny Agama HUSSEIN, H.K. & A.D.M. DARWISH (2000): Community structure, microhabitat use, sex ratio and sexual dimorphism in the agamid lizard, Agama agama spinosa. – Pakistan J. Biol. Sci., 3: 1700-1704. WAGNER, P., LEACHÉ, A., MAZUCH, T. & W. BÖHME (2013): Additions to the lizard diversity of the Horn of Africa: Two new species in the Agama spinosa group. - Amphibia-Reptilia, 34 (3): 363-387. The Horn of Africa is a center of diversity for African agamid lizards. Among the nine species of Agama occurring in the Horn of Africa, Agama spinosa is the most widely distributed. The A. spinosa group (sensu stricto, morphologically defined by possessing six clusters of spinose scales around the ear) contains two species: A. spinosa occurs from Egypt to Ethiopia and Somalia where it is replaced by the morphologically distinct and therefore sensu lato A. bottegi. Both species are only represented in museum collections by a small number of specimens from Ethiopia and Somalia, presumably the result of constant civil war that has plagued the region for decades and impeded field surveys. In this study, we examine species limits in the A. spinosa group using molecular genetic data (503 characters; mitochondrial 16S rRNA) and morphological data (67 characters). Deep divisions among populations of A. spinosa are supported by phylogenetic analyses and by multivariate analyses of morphometric data. Two new species from northern Somalia that differ from A. spinosa and A. bottegi are described. Furthermore, A. smithi, currently recognized as a synonymof A. agama, is re-assessed and recognized as a species of uncertain taxonomic position (i.e., incertae sedis). The results of this study improve our understanding of the evolution of agamid lizard diversity in the Horn of Africa, a significant biodiversity hotspot in Africa.
GENIEZ, P., PADIAL, J.M. & P.-A.CROCHET (2011): Systematics of north African Agama (Reptilia: Agamidae): a new species from the central Saharan mountains. – Zootaxa, 3098: 26-46. We studied the taxonomic status of the north African Agama species A. impalearis, A. castroviejoi, and A. boueti. The study of recently collected specimens and museum material, as well as phylogenetic analyses of a short 16S ribosomal RNA gene fragment, revealed the presence of an undescribed species in Adrar des Ifoghas (Mali), Aïr Mountains (Niger), Ahaggar Mountains (Algeria) and Tassili n’Ajjer (Algeria, Lybia), previously mistaken either as A. impalearis or A. agama. The new species, Agama tassiliensis n. sp., clearly belongs to the impalearis – boueti – spinosa species group but differs from these species, among other characters, by its red, reddish-orange or orange vertebral stripe in males, long and angular head, long limbs and toes (with 4th toe usually slightly longer than 3rd toe), long but low nuchal crest made of 10 to 15 spines (rarely 8–9 spines) and large number of supralabials (10–16, usually around 12). Interspecific uncorrected pdistances based on the 16S rDNA gene fragment are high, ranging from 3.9% between A. boueti and A. impalearis to 7.9% between A. spinosa and A. impalearis. The new species is sympatric with A. boueti at least in the Aïr Mountains and exhibits average mtDNA divergences of 6.2%, 7.4%, and 7.6% with A. spinosa, A. impalearis, and A. boueti, respectively. Agama boueti is paraphyletic relative to A. castroviejoi in the mtDNA tree, and mtDNA genetic divergences between populations of both species are lower than 1.0%, suggesting that A. castroviejoi is better treated as a junior synonym of A. boueti pending more detailed analyses. Scattered mountain ranges in the Sahara seem to have promoted lineage divergence and ultimately speciation in this group. Further work should be done to study species taxonomy and evolution in those areas.
WAGNER, P., KRAUSE, P. & W. BÖHME (2008): Studies on African Agama III. Resurrection of Agama agama turuensis LOVERIDGE, 1932 (Squamata: Agamidae) from synonymy and elevatin to species rank. – Salamandra, Rheinbach, 44 (1): 35-42. WAGNER, P., KRAUSE, P. & W. BÖHME (2008): Studien an afrikanischen Agama III.Revalidierung von Agama agama turuensis LOVERIDGE, 1932 (Squamata: Agamidae) und Erhebung in den Artrang. – Der Salamander, Rheinbach, 4 (1): 35-42. Zusammenfassung: Neues Material von Agama agama LINNAEUS, 1758, vom Mount Hanang, Tansania erweist sich als nicht zum Typenmaterial von Agama agama turuensis LOVERIDGE, 1932, unterscheidbar, einem Taxon, das bislang als Synonym von Agama lionotus elgonis LÖNNBERG, 1922, angesehen wurde. Unsere vergleichende morphologische Untersuchung zeigt, dass turuensis die größte Ähnlichkeit mit Agama mwanzae LOVERIDGE, 1923 und Agama kaimosae LOVERIDGE, 1935, aufweist und sich sowohl von A. agama als auch A. lionotus BOULENGER, 1896, unterscheidet. Agama turuensis kann gleichzeitig weder A. mwanzae noch A. kaimosae zugewiesen werden und muss daher als eigene Art betrachtet werden.
Marsabit Rock Agama MALONZA, P.K., SPAWLS, S., FINCH, B. & A.M. BAUER (2021): A new species of the Agama lionotus BOULENGER, 1896 complex (Squamata: Agamidae) from northern Kenya. – Zootaxa, 4920 (4): 543-553. Kenya has a high diversity of agamid lizards and the arid northern frontier area has the highest species richness. Among the Kenyan agama species, Agama lionotus has the widest distribution, occurring from sea level to inland areas in both dry and moist savanna as well as desert areas. This species mostly prefers rocky areas, both in granitic/metamorphic and volcanic rocks, although it also makes use of tree crevices as well as man-made structures. Recently in Marsabit, northern Kenya, a small-sized agama species, distinct from A. lionotus, was collected within a rocky lava desert area. This new species is characterized by its small size (mean SVL ~83 mm) as compared to typical A. lionotus (mean SVL ~120 mm). Past studies have shown the value of adult male throat coloration for the identification of species within the A. lionotus complex. Herein we also highlight female dorsal color pattern, which is a key character for distinguishing the new species from others in the group, including the similar A. hulbertorum. As in A. lionotus, displaying adult males have an orange to yellow head, a vertebral stripe, a bluish body coloration and an annulated white/blue tail. But the most diagnostic character is the coloration of females and non-displaying males, which exhibit a series of regular pairs of dark spots along the vertebrae as far posterior as the tail base. In addition, females have a pair of elongated orange or yellow marks on the shoulders and another on the dorsolateral margins of the abdomen. This study shows that more cryptic species in the Agama lionotus complex may still await discovery. The new species was found inhabiting dark desert lava rocks but should additionally be present in suitably similar sites in the northern frontier area. This underscores the need to re-examine populations of Agama lionotus from different microhabitats in this country.
Gambia Agama BÖHME, W. (2005): Vorkommen von Agama weidholzi WETTSTEIN, 1932 in Gambia, Westafrika. – Der Salamander, Rheinbach, 1 (3): 155-157. Zusammenfassung: Agama weidholzi WETTSTEIN, 1932 is for the first time recorded for The Gambia. The new locality, Bwiam on the southern shore of the Gambia River, is the westernmost locality of the species’ distribution area. A. weidholzi is one of the rare regional endemics of arid West Africa. The new record is based on an adult female collected in January (dry season), suggesting that former assumptions of a seasonal phenological displacement between age classes, or even an uniannual life cycle, in this species are unlikely. Zusammenfassung: |
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