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Studies on African Agama VII Bonner zoologische Beiträge Band 56 Heft 4 Seiten 285–297 Bonn, November 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 Philipp WAGNER*1, Michael F. BAREJ1 & Andreas SCHMITZ2 1 Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany. 2 Muséum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland. * Corresponding author: [email protected]. Abstract. 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. Key words. Reptilia: Sauria, Agamidae, Agama sp. n., Agama mehelyi, Africa, Cameroon, Gabon, morphology, taxon- omy. INTRODUCTION After research activities on the genus Agama Daudin, 1802 who restricted the erroneous type locality ‘America’ of in the last few years many species were described, syn- Agama agama to Cameroon but failed to fix this decision, onymised or revalidated (PADIAL 2005, WAGNER 2007, according to article 76 of the Internatio-nal Code of Zo- WAGNER et al 2008a, WAGNER et al. 2008b). Currently, the ological Nomenclature (ICZN 1999) by designating a lec- genus contains 35 recognised species, but preliminary re- to- or neotype. Now, after the proper de-finition of a neo- sults (especially from the West African clade of the genus) type (WAGNER et al. 2009b), a taxo-nomic revision of this show several cryptic taxa within the Agama agama (Lin- species group is possible and the already recognised new naeus, 1758) complex. Therefore, many populations are species can be described. under investigation and especially vouchers from Cameroon were of special interest to the authors, because CHIRIO & LEBRETON (2007) listed seven species of Aga- the type locality of Agama agama has been restricted to ma for Cameroon: Agama a. agama (Linnaeus, 1758); this country. Agama doriae benueensis Monard, 1951; Agama gracil- imembris Chabanaud, 1918; Agama mehelyi Tornier, 1902; However, until recently it was not possible to character- Agama paragama Grandison, 1968; Agama sankaranica ize the ‘true’ Lacerta agama Linnaeus, 1758, because the Chabanaud, 1918 and Agama sylvanus MacDonald, syntypes illustrated in SEBA (1734) are not available and 1981. In the adjoining countries of Cameroon, no other the proposed type material in the Museum Adolphi Fri- than the above mentioned Agama species occur (WAGN- derici collection of the Naturhistoriska Riksmuseet (= The ER, unpubl. data) but CHIRIO & LEBRETON (2007) also Swedish Museum of Natural History) is not identical with identified four probable new Agama species from the illustrated specimens. WAGNER et al. (2009b) clarified Cameroon and one of them (Agama sp. 1) turned out to this situation by designating a neotype (ZFMK 15222) be identical with the new species recognized by us before, from Cameroon. The authors followed MERTENS (1938) and we describe it herein. 286 Philipp WAGNER et al.: Studies on African Agama VII. New species of the Agama agama-group MATERIAL & METHODS plify a portion of the mitochondrial 16S ribosomal RNA gene. PCR cycling procedure was as described in The type material of the new species is deposited in the SCHMITZ et al. (2005). PCR products were purified using Zoologisches Forschungsmuseum A. Koenig, Bonn, Zo- Quiaquick purification kits (Quiagen). Sequences were ob- ologisches Museum für Naturkunde, Berlin and the tained using an automatic sequencer (ABI 377). Sequences Muséum d’histoire naturelle, Geneva. It was compared were aligned using ClustalX (THOMPSON et al. 1997; de- with vouchers from Cameroon and other African countries fault parameters) and manually checked using the origi- housed in those collection (see appendix) and with data nal chromatograph data in the program BioEdit (HALL from literature (GRANDISON 1968, CHIRIO & LEBRETON 1999); this resulted in a total of 505 bp for the chosen sec- 2007). tion of the 16S gene. Phylogenetic trees were calculated in the program Paup* 4.0b10 (SWOFFORD 2002) using the Measurements were taken with a digital calliper to the neighbor-joining algorithm (NJ) and bootstrapping with nearest of 0.1 mm. Measurements and scale counts were 20000 pseudo-replicates to estimate node support as well done following GRANDISON (1968), MOODY (1980) and as with a maximum parsimony approach (heuristic search MOODY & BÖHME (1984). with TBR branch swapping, stepwise random addition with 100 replicates, bootstrapping with 2000 pseudo-repli- Molecular data were collected to examine sequence vari- cates). Paup* 4.0b10 was also used to compute the uncor- ation between the new species and its morphologically rected pairwise distances for all sequences. closest relatives. A portion of the mitochondrial 16S rRNA gene of the holotype of the new species (ZFMK 87698; The following measurements and scale counts were part- GenBank accession number: GU133316), a series of oth- ly used to compare the different species (see table 1): er specimens of the new species (no voucher; GenBank Snout-vent length (SVL): measured from mental scale to accession number: GU133315; ZFMK 73241; GenBank cloaca; tail length (TL): measured from cloaca to tip of accession number: GU133317; ZFMK 75376; GenBank tail; head length (HL): measured from jugale to rostral accession number: GU133318; ZFMK 83762; GenBank scale; head height (HH): measured at the jugal-postorbital accession number: GU133319; ZFMK 83766; GenBank region; head width (HW): measured across the jugal-pos- accession number: GU133320), as well as the proposed torbital region just anterior to the external auditory mea- neotype of Lacerta agama (ZFMK 15222; GenBank ac- tus; Midbody scales (MS): scale rows around midbody; cession number: GU133323) from northern Cameroon, vertebral scales (VS): number of midline vertebral scales, Agama sankaranica (ZFMK 84992; GenBank accession counted from midpoint of pectoral region to midpoint of number: GU133327) from an unknown locality, Agama pelvic region; dorsal scales (DS): number of midline lon- paragama (ZFMK 15244; GenBank accession number: gitudinal dorsal scales, counted from midpoint of pectoral GU133321) from Cameroon, Waza, Logone et Chari, Aga- region to cloaca; cloacal pores (CP). ma agama africana (ZFMK 73845, ZFMK 73846; Gen- Bank accession numbers: GU133311, GU133312) both from Senegal, Dakar-Bel Air, Agama agama ssp. (MH- RESULTS & DISCUSSION NG 2689.53; GenBank accession number: GU133310) from Benin, Agama boueti (ZFMK 80057) from Mauri- Species of the genus Agama, unlike chameleons, the sis- tania, 30km NW of Rosso, and Agama finchi (ZFMK ter taxon of the Agamidae (e.g. MACEY et al. 2000), are 83652; GenBank accession number: GU133314) from very conservative in body form, without developing or- Kenya, Malaba (type locality) were sequenced. Sequences naments like e.g. horns or ear laps. Only in some species for Agama planiceps (GenBank accession number: body ornamentations like enlarged nuchal or tail crest are AF355476) and Agama castroviejoi (GenBank accession present. If one takes into consideration the results from number: AY522929) were added to the dataset from al- the chameleons (ZIEGLER & BÖHME 1997; BÖHME & ready published sequences (MATTHEE & FLEMMING 2002, ZIEGLER 2008), where morphologically similar species PADIAL 2005). Acanthocercus atricollis (ZFMK 41748; show high variations in hemipenis structures, a similar sit- GenBank accession number: GU133322 Botswana, uation in the Agamidae could be expected. In contrast, pre- Gaborone) was chosen as outgroup. liminary results (BÖHME 1988; WAGNER et al., unpubl. da- ta) show a high level of homoplasy and conservative mor- DNA was extracted using QuiAmp tissue extraction kits phology in the hemipenis structures in the genus Agama. (Quiagen) or a modified Chelex-Protocol (WALSH et al. Consequently, beside from body form and hemipenis 1991, SCHMITZ 2003). The primers 16sar-L (light chain; structure, there must be another mechanism of species 5’ – CGC CTG TTT ATC AAAAAC AT – 3’) and 16sbr- recognition. A pre-mating mechanism was identified by H (heavy chain; 5’ – CCG GTC TGA ACT CAG ATC several authors (LOVERIDGE 1933, THYS VAN DEN AUDE- ACG T – 3’) of PALUMBI et al. (1991) were used to am- NAERDE 1963, MCLACHLAN 1981, BÖHME et al. 2005, Bonner zoologische Beiträge 56 287 Fig. 1. Phylogenetic tree based on 505 bp of the mitochondrial 16S rRNA gene. Values above the nodes represent neighbour-joi- ning bootstrap (20000 pseudo-replicates) values in percent while the values blow the nodes are the corresponding maximum par- simony supports. Significantly supported values are in bold. Values below 50% are not shown. The branch length from the out- group (Acanthocercus) to the ingroup (Agama) has been shortened for visual purposes. 288 Philipp WAGNER et al.: Studies on African Agama VII. New species of the Agama agama-group 4 3 2 1 Table 1. Table 5 8 7 6 9 17 16 15 14 13 12 11 10 WAGNER 2007, WAGNER et al. 2008a, 2008b) as the Agama agama africana Agama agama Agama agama atricollis
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