790 C. L. Johnson et al.


FIG. 3 Predicted occupancy across the mainland native range of M. nigra. Eight potentially important landscapes, defined as spatially distinct continuous forest blocks .50 km2 with predicted occupancy .0.7, are identified by numbers (Supplementary Fig. 2, Supplementary Table 3).


thereby reducing its influence on occupancy. Detection probability, however, varied across sites, and was higher in closed canopy forests and in areas with a lower Human Footprint Index. We attribute this to a preference by M. nigra for forests and a sensitivity to human disturbance, which cause a reduced abundance of the species in non- forested and disturbed habitats, with less abundant species typically being less detectable (Royle & Nichols, 2003). However, this could also be a consequence of M. nigra being more elusive and cautious in more distur-bed areas. Although occupancy was significantly higher in closed


canopy forest than in bush/scrub habitats, we were not able to test for effects of forest type (pristine vs degraded/ logged) because of the limited resolution of available satel- lite imagery. We used NDVI as a remotely sensed proxy for vegetation structure and green biomass (Myneni et al., 1995) but, although retained in the best models, it was not signifi- cant. This is probably the result of the overriding import- ance of forest presence over the quality of that forest in determining occupancy. The dependence of M. nigra on forest is nevertheless apparent, highlighting the species’ susceptibility to forest conversion to other land uses. Along with a preference for forest cover, estimated occu-


pancy was higher within protected areas, which is probably a result of both the continuous extent of optimal habitat and lower anthropogenic disturbance. Deforestation ob- served within protected areas during 2001–2017 was much lower than outside (26.4% outside protected area vs 6.2%


loss within protected area; Hansen et al., 2013), a factor that probably implies reduced hunting and disturbance. However, protected areas currently cover only 20%ofthe area estimated to be occupied by the species. Considering that M. nigra, like many other primates (Estrada et al., 2017), is vulnerable to anthropogenic activities, the species remains at risk across 80% of its range. Our modelling approach predicted eight spatially dis-


tinct regions that are likely to support important sub- populations of M. nigra (Fig. 3, Supplementary Table 3, Supplementary Fig. 2). Five of these landscapes repre- sent the first scientific record of the species’ presence (Supplementary Table 3). In combination with evidence of range expansion (Johnson et al., 2019), this finding is of conservation relevance and contradicts speculations that Tangkoko contains the last viable population (Supriatna & Andayani, 2008; Palacios et al., 2011;Kyeset al., 2013; Engelhardt et al., 2017). Although Tangkoko does appear to hold an important and viable population (Engelhardt et al., 2017) it comprises just 3% of the total area occupied by the species (Supplementary Table 3). Therefore, although we acknowledge that population density and viability of sub- populations will vary across the species’ range, the signifi- cance of these additional areas for the species cannot be ignored. Our discovery of previously undocumented popula-


tions, in addition to theincreased rangesizeof 220 km2 recently reported (Johnson et al., 2019), results in an


Oryx, 2020, 54(6), 784–793 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000851


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