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Annotated Checklist of the Fishes of Madagascar, Southwestern Indian Ocean, with 158 New Records

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Annotated Checklist of the Fishes of Madagascar, Southwestern Indian Ocean, with 158 New Records FishTaxa (2018) 3(1): 1-432 E-ISSN: 2458-942X Journal homepage: www.fishtaxa.com © 2018 FISHTAXA. All rights reserved Annotated checklist of the fishes of Madagascar, southwestern Indian Ocean, with 158 new records Ronald FRICKE*1, Jamal MAHAFINA2, Faustinato BEHIVOKE2, Henitsoa JAONALISON2, 3, Marc LÉOPOLD3, Dominique PONTON4 1Im Ramstal 76, 97922 Lauda-Königshofen, Germany. 2Institut Halieutique et des Sciences Marines, Jeune Équipe Associée à l’IRD ACOM, Université de Toliara, Route du Port Mahavatse II, Toliara 601, Madagascar. 3ENTROPIE, IRD-Université de La Réunion-CNRS, Laboratoire d’Excellence CORAIL, Institut Halieutique et des Sciences Marines, Route du Port Mahavatse II, Toliara 601, Madagascar . 4ENTROPIE, IRD-Université de La Réunion-CNRS, Laboratoire d’Excellence CORAIL, c/o Hydrô Réunion Station Marine, Port ouest, Magasin 10, 97420 Le Port, La Réunion, France. Corresponding author: *E-mail: [email protected] Abstract An annotated checklist of the fish species of the Madagascar EEZ (southwestern Indian Ocean) comprises a total of 1,798 species in 247 families. 158 species are recorded from Madagascar for the first time. The majority of the species is autochthonous; 28 species have been introduced, mainly in freshwater habitats. The fish fauna is mostly marine (95.4% of the total number of native fish species), with the Gobiidae, Labridae, Serranidae, Pomacentridae and Apogonidae being the families with most representatives; among the 90 native freshwater fish species (adults mainly occurring in freshwater), the Cichlidae are the dominating family, but there are also two endemic families, the Bedotiidae (16 species) and Anchariidae (6 species). The fish fauna at Madagascar is typical for offshore, high islands in the southwestern Indian Ocean. Zoogeographically, the main element of the marine fish fauna of Madagascar consists of widespread tropical Indo-Pacific species (978 species, 58.3% of the total native marine species). A total of 13 species (3.3%) are found worldwide, either circumtropical or circumtropical including warm temperate zones. A total of 215 species (12.8%) are found worldwide, either circumtropical or circumtropical including warm temperate zones. An additional 453 species (27.0%) are Indian Ocean endemics, including 233 western Indian Ocean endemics (13.9%), 73 southwestern Indian Ocean endemics (4.4%), 16 species endemic to Madagascar and Mascarenes (1.0%), 4 species endemic to Madagascar and Comoros (0.2%), 3 species endemic to Madagascar and Madagascar Ridge (0.2%), and 37 marine species endemic to Madagascar (2.2%). Most of the autochthonous freshwater fishes are endemic to Madagascar (87 species, 96.7% of the native freshwater species). Keywords: Checklist, Pisces, Southwestern Indian Ocean, Madagascar EEZ, New records, Zoogeography. Zoobank: urn:lsid:zoobank.org:pub:D6B0C9D2-746B-4434-837F-3BEAEE4256AD Introduction Madagascar, which is inhabited by more than 22 million Malagasy people, is the fourth great island of the world with an area of 587,295 km2 (Anonymous 2014a). This large island is situated in the southwestern part of Indian Ocean, separated from the African continent by approximately 400 km large Mozambique Channel. The length of Madagascar is 1,600 km between 11°57'S and 25°39'S, its width is 570 km between 43°14'E and 50°27'E. With 5,600 km of coastline, its Exclusive Economic Zone (EEZ) is one of the largest in the Indian Ocean with a surface area of 1.14 million km² (Barnes-Mauthe et al. 2013). The continental shelf area is estimated to be 117,000 km², and extends more in the northwestern and southern part of the island (Breuil and Grima 2014). The estimated total coral reef area is 1,230 km2, the estimated diversity in species being between 200 and 300 (Spalding et al. 2001). On the other hand, the eastern coast is relatively straight and featureless, with few estuaries, capes and bays (Breuil and Grima 2014) but with extensive brackish ecosystem such as the Pangalanes Canal (Moreau 1987). Several brackish lagoons also exist in the south-east of the country (Moreau 1987) (Fig. 1). The total volume of the surface renewable freshwater resources, including lakes and rivers, is estimated about 337 km3 per year (Anonymous 2015a). Most of the Malagasy lacustrine systems are shallow and small in comparison with the African lakes, and although they are more than 3,400 lentic areas, they represent only 0.3% Accepted: 30 June 2017; Published: 14 February 2018 FISHTAXA (2018) 3(1): 1-432 2 Figure 1. Madagascar with its Exclusive Economic Zone (EEZ), dotted lines corresponding to contested limits. Shape files of rivers and coastlines are from BD500 developed by Foiben-Taosarintanin’i Madagasikara (Institut géographique et hydrographique de Madagascar, http://www.ftm.mg/framepro.htm). Rivers and streams have been simplified in order to improve readability. The shape file of coral reefs corresponds to Andréfouët et al. (2009); the shape file of mangrove cover of Madagascar coast was developed by Anonymous (2015b) using LandSat 7 ETM and Landsat 8 OLI data and a classification method. Numbers correspond to localities cited in the text, see Table 1 for their names. of the country’s area (Elouard and Gibon 2001). Lake Alaotra, situated in the east of Madagascar, is known as the largest lake of the country but does not exceed 200 km2 during the high water period (Moreau 1987) plus 350 km2 of swamps (Moreau 1979). The other main water bodies are Lake Kinkony (139 km²) and Lake Ihotry (97 km²) along with Lakes Anketraka and Tsimanampetsostsa in western Madagascar, the total area of lakes being close to 1,500 km² (Breuil and Grima 2014). Malagasy large rivers form an important and dense hydrographic network that can be separated in five sub-systems (Elouard and Gibon 2001): the Montagne d’Ambre Basin (11,200 km2), the Tsaratanana Basin (20,000 km2), the eastern, western and southern basins 2 (respectively 150,000, 365,000, and 48,750 km ). The most recent checklist of the fish fauna of Madagascar is limited to a publication by Fischer and Bianchi (1984) recording 350 fish trade species from the island. Sauvage (1887, 1891a) revised the fishes of the greater Madagascar region, but also included many species not known from Madagascar but occurring at the coast of East Africa and nearby islands including the Comoros, Seychelles and Mascarenes. Sauvage's Madagascar Region stretched between 2°S and 26°S, and between 42°E and 65°E (Maugé 1978: 95). This has caused confusion in the literature in many cases; one of the most striking cases is that of Chaetodon madagaskariensis Ahl 1923 which was based on Sauvage's (1891a) description of Choetodon (Tetragonoptrus) miliaris (non Quoy Fricke et al.- Annotated checklist of the fishes of Madagascar 3 & Gaimard 1825), allegedly from Madagascar, but in fact based on C. chrysurus Desjardins 1834 from Mauritius. The present paper provides an updated checklist of the fishes of the Madagascar EEZ, based on all available sources. It includes the fish species of both inland and marine waters. Material and Methods The present paper provides a list of all fishes recorded from Madagascar. The study area includes the waters of the EEZ of Madagascar, but excludes the Éparses Islands in Mozambique Channel (e.g. see Fricke et al. 2013), the Comoros and Mayotte (see Wickel et al. 2014), and Walters Shoals on the Madagascar Ridge (e.g. see Collette and Parin 1991). For each species, the first record from Madagascar is provided. Species in Sauvage (1887, 1891a), a work that deals with the greater Madagascar region including neighbouring islands, are only considered as recorded from Madagascar if the occurrence is expressedly stated. The checklist of McKenna and Allen (2005) partially uses Sauvage's records; therefore only positive records of species that the authors observed in northwestern Madagascar are used for the present checklist. Fourmanoir (1957) also deals with the greater Madagascar region including the Éparses and Comoro islands; again species are only considered to be recorded from Madagascar if a Madagascar locality is expressedly mentioned. In the checklist, an asterisk indicates families or species endemic to Madagascar. In the material lists, "C&S" stands for specimens which have been cleared and stained for osteological analysis. Higher classification follows Nelson (2006), family classification follows Laan et al. (2014), updated according to Eschmeyer et al. (2017). The more recent version of fishes of the world by Nelson et al. (2016) has not been used here, due to various errors and inconsistencies (some of those were pointed out by Britz 2017). For the distribution analysis, the following works were consulted to assess the fish fauna of islands and island groups in the Western Indian Ocean: Europa Island, Fricke et al. (2013); Seychelles, Smith and Smith (1963) and Randall and Egmont (1994), updated using various papers including Allen (1972), Dawson (1967, 1968, 1977a), Fraser and Lachner (1985), Golani (1984), Hoese and Reader (1985), Holleman (2005, 2007), Keith et al. (2004), Kim and Amaoka (2001), Kovačić and Bogorodsky (2013), Lachner and Karnella (1978, 1980), Matsunuma et al. (2017), Polunin and Lubbock (1977), Randall (1999c, 2001), Randall and Bogorodsky (2016), Randall and Clements (2001), Randall and Heemstra (1985, 2009), Randall and Lourie (2009), Randall et al. (2008, 2013, 2014, 2015), Randall and Victor (2015), Schwarzhans and Møller (2007), Springer and Williams (1994), Starck (1969), Tyler (1966), Uiblein (2011), Uiblein and Gouws (2014), Uiblein and Heemstra (2011b); Maldives, Randall and Anderson (1993) and Anderson et al. (1998), updated using various papers including Allen and Erdmann (2006), Allen and Randall (2002), Gon and Randall (2003), Fukui and Motomura (2016), Kuiter and Debelius (1999), Randall (2001, 2011b), Randall et al. (2013, 2014, 2015), Randall and Victor (2014, 2015), Smith-Vaniz (2010), Victor (2016b); Chagos Archipelago, Winterbottom and Anderson (1997), updated using various papers including Böhlke (2000), Craig (2008), Gill and Edwards (2004), Holleman (2005), Randall et al.
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