The Agamidae is a widespread Old World lizard family. Apart from the New World, it is naturally lacking only in the Malagasy subregion of the Afrotropical region (Wermuth 1967; Moody, 1980). So far, there are only two species of agamids recorded that have established populations outside their natural distribution ranges. Calotes versicolor sensu lato, a widespread Oriental tree-dwelling species complex has successfully conquered the western Indian Ocean islands Mauritius, La Réunion and Rodriguez (Vinson and Vinson 1969, Henkel and Schmidt 1995) and Florida in the south-eastern United States (Enge and Krysko 2004). A similar successful invader is Agama agama sensu strictu, a western Afrotropical species complex distributed from the southwestern margin of the Sahara desert in the north down to Angola in the south and Uganda in the east. It has colonised parts of Florida in the United States and has succeeded in establishing even some reproducing populations (e.g. Wilson and Porras, 1983, Bartlett and Bartlett 1999, Enge et al. 2004) which are generally believed to be escapers from the pet trade. Bartlett and Bartlett (1999) observed Agama agama in Miami Dade County close to pet shops. Enge et al. (2004) reported about 40 escaped specimens from a reptile store in

Sanford. According to the description given by Harris (1964), they were assigned to the subspecies A. agama africana from West Africa. From the Mediterranean island of Malta two introduced individuals from Agama agama were recorded (Schembri and Schembri 1984), which, however, did not establish a population. One of the specimens was found and captured on crates of imported beer. Also from Palermo in Sicily, another Mediterranean island, Agama agama is known as introduced by a transport from Africa, but also in this case a population was not established (Lo Valvo and Longo 2001, Sindaco et al. 2006). In the western Indian Ocean region, A. agama has been recorded from the Mascarene island La Réunion (Guillermet et al. 1998, Probst 1999). Very recently, the species was also recognized as a single voucher from Santo Antão, Cape Verde Islands (Vasconcelos et al. 2009). In the present note, we record Agama agama for the first time from Madagascar and discuss its identity on the Comoro Islands.

The first two specimens suggesting occurrence of Agama agama in Madagascar were found in the herpetological collection of the Zoologisches Museum der Universität Hamburg (ZMH). According to the jar label, one female and one juvenile (ZMH R07734-35), catalogued in 1953, were collected by Fritz Engelmann from the ‘Aquarium Hamburg’, but were labelled only with “Madagaskar”, without more precise locality data. However, it can not be excluded that this record is due to erronneous label data, especially since the provided data is very general and the putative collector is not known as a collector of any other amphibians or reptile species in Madagascar.

Herpetology Notes, volume 2: 73-77 (published online on 3 June 2009)

Studies on African Agama IV: First record of Agama agama (Sauria: Agamidae) from Madagascar

and identity of the alien population on Grande Comore Island

Philipp Wagner1, Frank Glaw2, Kathrin Glaw3, Wolfgang Böhme1

1Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. Email: philipp.wagner.zfmk@uni-bonn.de, w.boehme.zfmk@uni-bonn.de

2Zoologische Staatssammlung München, Münchhausenstr. 21, 81247 München, Germany. Email: Frank.Glaw@zsm.mwn.de

3LMU Biozentrum, Department Biologie II, Tierhaltung, Grosshaderner Strasse 2, 82152 Planegg-Martinsried, Germany. Email: KathrinGlaw@zi.biologie.uni-muenchen.de

Abstract. Invasions of allochthonous species are encouraged through international traffic and trade. Here we present first distribution records of the highly invasive species Agama agama for Madagascar and discuss its identity and introduction on the island Grande Comore.

Key words. Agama agama, new distribution record, Madagascar, Comoro Islands.

Philipp Wagner et al.74

In January 2004, the British ornithologist and Kenyan resident Brian Finch took photographs of a large Agama basking on a building wall of Antananarivo Airport (pers. comm. to WB). This lizard, an adult male (Fig. 1), shows the typical colouration characters of Central and West African A. agama sensu stricto particularly a tricoloured tail, and therefore has certainly not been introduced from the opposite East African coast. [East African populations of the A. agama complex, which were previously believed to represent various subspecies (e. g. A. agama dodomae, A. a. elgonis, A. a. lionotus, A. a. usambarae), have proved to be specifically distinct from typical A. agama (Böhme et al. 2005) or synonyms (Wagner 2007)]. Although there is no doubt about the reliability of this record, attempts in 2006 and 2008 by FG to confirm the presence of A. agama around the airport and the nearby crocodile farm failed, suggesting that no population of this eye-catching species might

have established so far. According to the colouration of the specimen (Figs. 1-2) it is tentatively attributed to Agama agama sensu stricto.

The first record of Agama agama from the Comoro Islands was published by Meirte (2004) who stated 1998 as the year of introduction of A. agama to the island Grande Comore. Two of us (FG and KG) found many individuals of this species in February 2000 at several localities in Moroni, the capital of Grande Comore. Voucher specimens were deposited in the Zoologische Staatssammlung München (ZSM 704-705/2000 and ZSM 322-323/2002). All individuals were recorded in the northern areas of Moroni including the coastal road and the hotel „La Grillade“. The staff of this hotel told us that the lizards were living at the hotel compound already since several years. This information, as well as the relatively large range and high abundance

Figure 1. Agama agama from Antananarivo Airport (Madagascar). Photo by B. Finch.

Coll. No. Locality SVL TL HL HW HH SR SL IL OS SD IN TS

ZMH R07734 Madagascar 92.0 145.3 23.3 16.6 10.7 -- 11 11 11 23 6 8

ZMH R07735 Madagascar 67.8 -- 19.0 15.6 8.4 76 10 10 10 21 5 9

ZSM 704/2000 Grande Comore 92.2 -- 23.0 16.7 10.5 74 11 11 12 25 7 10

ZSM 705/2000 Grande Comore 60.8 112.7 16.5 12.9 7.5 66 9 9 9 20 7 8

ZSM 322/2002 Grande Comore 75.8 123.1 19.3 14.8 9.4 70 10 11 8 23 6 8

ZSM 323/2002 Grande Comore 96.1 136.4 24.7 17.1 10.8 -- 10 10 10 21 6 8

SVL= snout-vent length; TL= tail length; HL= head length; HW= head width; HH= head height; SR= scale rows around midbody; SL=Supralabialia; IL= Infralabialia; OS= occipital scales; SD= lamellae under the right 4th toe; IN= internasal scales; TS= temporal scales.

Table 1. Measurements of the collected vouchers of Agama agama (all measurements in mm).

Agama in Madagascar and Grande Comore 75

within Moroni strongly suggest that A. agama has been introduced at least a few years before 1998. The observed populations contained many juveniles indicating successful reproduction, but additional observations in 2002 did not reveal any significant range extension. Inquiries concerning the origin of A. agama resulted in the information that the first alien specimens have reached Grande Comore in a parcel obtained by the Coca Cola repository at Moroni which was also populated by A. agama. This information seems consistent with the apparent absence of A. agama at the waterfront of Moroni in 2000, which makes a passive introduction by boat traffic unlikely. Another invasion scenario was reported by Yahaya Ibrahim (pers. comm. in Hawlitschek 2008). According to him, a South African diplomat of the African Union in Moroni kept some Agama agama as a pets at his home close to the Coca Cola repository, but released them when he left the island. The origin of these specimens is unclear, but due to the nuptial colouration of the observed males (including one collected voucher), these agamas again can not have originated from the African opposite coast of the Mozambique Channel, because they can be identified as typical Agama agama s. str. from West/Central Africa. This makes their introduction by a South African less likely, where one would expect Agama atra or A. planiceps rather than A. agama as pet from this region.

Although we can attribute both the Malagasy specimen and the Comoroan population to Agama agama sensu stricto their exact geographic origin within Central or West Africa of cannot be identified at present. First, scale counts from Madagascar are only available from one single juvenile (ZMH R07735), the accompanying female being too damaged for taking scale counts. Second, the African populations of the entire Agama agama complex are extremely understudied on a continental scale, and the few scalation data published so far (e.g. Thys van den Audenaerde 1963, Grandison 1968) do not allow for a geographic assignment of the Madagascan and Comoran specimens (see Tab. 1 for scale counts). However, the Comoran specimens are possibly peculiar in terms of their small size. A sexually mature, i. e. fully coloured male (ZSM 323/2002) measures only 96.1 mm snout-vent length and 136.42 mm tail length. This is relatively small as compared with typical A. agama. However, it is not clear whether such a small size is actually characteristic of the Comoran population.

Invasive species can become a serious threat to local herpetofaunas, and island faunas with a high degree of endemism are particularly endangered (see e.g. Rodda and Fritts 1992, Fritts and Rodda 1998). It is therefore possible that the invasion of this expansive and anthropophilic agamid on Grande Comore could threat the rare iguanid Oplurus cuvieri comorensis which is restricted to very small areas on this island (Meirte 1992, 2004). Although genetically deeply nested within O. cuvieri populations from Madagascar, chromatic and ecological pecularities suggest that this species colonized Grande Comore by natural oversea dispersal (Münchenberg et al. 2008) and therefore merits conservation. Thus the range expansion of Agama agama on Grande Comore and the area around the Antananarivo Airport should be carefully monitored in order to prevent damage from the endemic potential competitors.

Figure 1. Agama agama from Antananarivo Airport (Mada-gascar). Photo by B. Finch.

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Acknowledgements. We are grateful to the Ministère de la pro-duction et de l’environnement, Direction Générale de l’environ-nement, Iles des Comores, in particular to the General Director Monsieur Mohamed Youssouf Oumour, Madame Foudzia, et Monsieur Dossar for the necessary permissions. We also thank Dr. Jakob Hallermann, Hamburg, who loaned us specimens under his care, and Dr. Birgit Blosat, Jünkerath, who helped to obtain some literature. The research of FG was granted by the Deutsche Forschungsgemeinschaft (DFG: GL 314/1).

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Figure 3. Agama agama male from Moroni. Photo by K. Glaw.

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Agama in Madagascar and Grande Comore

Figure 3. Agama agama female from Moroni. Photo by K. Glaw.

Accepted by Miguel Vences; Managing Editor: Miguel Vences