820 J. C. Tinsman et al.


FIG. 1 Previous extent of occurrence polygons for Eulemur macaco and Eulemur flavifrons, from IUCN (used with permission). All known occurrence records for E. flavifrons, E. macaco and putative hybrids, and for Eulemur fulvus, in the area are noted.


area encompassing all known or projected occurrences of a species; IUCN, 2001)is ,2,700 km2 (Andriaholinirina et al., 2014a). Its area of occupancy (AOO; i.e. the area of suitable habitat that is actually occupied within a species’ EOO; IUCN, 2001) must be even smaller, although it had not been estimated previously. The EOO of E. flavifrons is bounded by the Mozambique


Channel to the west and the Maevarano River to the south (Koenders et al., 1985; Petter & Andriatsarafara, 1987; Randriatahina & Rabarivola, 2004; Schwitzer & Lork, 2004; Andriaholinirina et al., 2014a; Fig. 1). It extends east to the Sandrakota River and Manongarivo Special Reserve, which comprises 32,000 ha of protected Sambirano rainfor- est (MEF&MNP, 2010). Previous studies have identified the Andranomalaza River, also called the Maitsomalaza in the local Sakalava dialect, as the boundary between E. flavifrons and its parapatric sister species, the black lemur Eulemur macaco (Koenders et al., 1985; Andriaholinirina et al., 2014a). However, there is conflicting evidence regarding whether the lemurs between the Andranomalaza River and the more northern Manongarivo River are hybrids, intermediate-appearing forms on a phenotypic cline, or typical members of either species (Meyers et al., 1989; Rabarivola et al., 1991; Andrianjakarivelo, 2004;Randriatahina & Rabarivola, 2004).


The first report of phenotypic variation was by Meyers


et al. (1989), who observed two distinct groups of lemurs un- like typical E. flavifrons or E. macaco. The first group was at Beraty: individuals had light brown eyes and short ruffs of hair around their ears, characteristics that are intermediate between E. flavifrons and E. macaco. The second group was atAmbodivoahangy: individuals had darker eyes and redder coats than is typical of Sahamalaza E. flavifrons. However, when Andrianjakarivelo (2004) visited Ambodivoahangy he found animals that ‘greatly resembled’ E. flavifrons. Goodman and Schütz (2000) surveyed the eastern slopes of Manongarivo Special Reserve, north of Ambodivoahangy, and identified groups containing both E. macaco and ‘hy- brid’ individuals but did not detail their criteria for these distinctions. Their assessment was complicated by the presence of E. fulvus in that area, which may be perceived as having a reddish coat (Goodman & Schütz, 2000). Updated assessments of the lemurs in this region are ne-


cessary to establish effective conservation initiatives for these two species (Rakotonirina et al., 2011, 2014). They could also improve estimates of the species’ ranges, especially considering the area has not been assessed since 2004 (Schwitzer et al., 2014). We investigated the presence and phenotypes of Eulemur species from Sahamalaza–Iles Radama National Park to Manongarivo Special Reserve,


Oryx, 2020, 54(6), 819–827 © 2019 Fauna & Flora International doi:10.1017/S0030605318000868


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