Fosa spatial dynamics and activity patterns 843


in 2017) utilized these forest patches to cross grassland areas. Fosas used patches at similar times as corridors (8.00–18.00). Activity counts for F1 andM3 reached a resting point (activity count = 0) in several forest patches while traversing grasslands.


Discussion


Fine-scale movements and activity patterns of collared fosas revealed several overall patterns in home range and habitat selection: (1) dry season home range sizes in Ankarafantsika National Park weremuch larger than documented for fosas in Kirindy Forest, another dry deciduous forest, or protected rainforests in eastern Madagascar, with varying degrees of overlap between individuals; (2) individuals generally avoid- ed open habitats and villages; (3) all fosas utilized forest patches or corridors to cross deforested areas. Additionally, accelerometer data documented that fosas remained active when traversing non-forested areas.


Home ranges


The high degree of home range overlap among the three individuals collared in 2016 is similar to the pattern observed in Kirindy Forest (Lührs & Kappeler, 2013), with male home ranges overlapping those of females and other males. Pre- vious studies at Kirindy documented males with larger home ranges than females (Hawkins, 1998), whereas home range sizes for males and females were similar in Anka- rafantsika National Park. Overlap patterns for collared indi- viduals were very different across years; the reasons for this could not be determined because of the small sample size. Home range sizes estimated by kernel densities in Anka-


rafantsika National Park far exceeded those reported in Kirindy during the same season (Hawkins, 1998; Lührs & Kappeler, 2013). This could potentially be a result of the longer monitoring duration in our study (GPS-collared fosas were tracked for a maximum of 1 month by Lührs & Kappeler, 2013); however, fosas tracked via radio telemetry during both dry and wet seasons also had significantly smaller home ranges (Hawkins, 1998). Larger fosa home ranges in Ankarafantsika National Park may reflect the fact that the landscape ismore fragmented than in Kirindy. Prey availabil- ity could be lower in fragmented habitats, resulting in fosas requiring larger home ranges tomeet their nutritional require- ments (McNab, 1963;Gittleman&Harvey, 1982). Habitat frag- mentation appears to influence the distribution of at least two of the large-bodied lemurs in the Park (Propithecus coquereli and Lepilemur edwardsi), which historically comprisedmuch of the fosa’s diet (Dollar et al., 2007), andmay also affect other lemurs in the region (e.g. Eulemur fulvus and Eulemur mon- goz). Home range overlap documented for fosas in the Park suggests there may be intraspecific competition for prey (Craul et al., 2009; Kun-Rodrigues et al., 2014).


Habitat selection and avoidance of villages


All collared individuals exhibited similar habitat selection for forest environments. Because their habitat is fragmented and their home ranges overlap, fosas in Ankarafantsika National Park are probably affected by the same villages, agricultural plots and deforested areas. Habitat preferences varied slightly between individuals,


but all generally avoided highly human-modified landscapes (agricultural areas and villages). All fosas used forest frag- ments, occupying edge habitat up to 2.5 km outside the Park boundary on several occasions. A recent camera-trap study in north-eastern Madagascar also reported open habitat avoidance, with similar habitat-use patterns outside contiguous forest patches (Farris et al., 2014), and work in central-eastern Madagascar documented fosa use of forest fragments up to 15 km from contiguous forests, with higher population density in fragments closer to contiguous forest habitats (Gerber et al., 2012). Fosas in the Park tended to avoid areas of human habitation, including areas with only a few buildings. If fosas preyed upon poultry or other domestic animals, they could be expected to select positively for village areas. There may be areas of human occupation too small to be visible in satellite images, or hidden by tree canopy, thus documenting village encroachment into the Park’s forested areas at a fine scale is necessary to gain further insights into fosa interactions with livestock and humans. The individual (M2) that most frequently entered villages was killed by a vehicle on the RN4 road, confirming that motor traffic is a threat to fosas (Wyza et al., 2018). Persecution and hunting threaten fosa populations


(Farris et al., 2015b; Hawkins, 2016) and occur mostly be- cause people believe that fosas prey on poultry (Kotschwar Logan et al., 2015). In our study, , 1% of all recorded loca- tions of the three collared maleswere in villages, and females never entered villages. Furthermore,,1%of all recorded lo- cations were within a 30mbuffer around village areas, indi- cating that during our study fosas did not spend a significant amount of time in or near villages. The median distance (across all individuals) of recorded locations to the nearest village boundary was . 2 km, suggesting that fosas prefer the forest interior, rarely approach human settlements and probably do not rely on poultry as a major food source.


Activity patterns


Fosas in Ankarafantsika National Park did not appear to shift temporal activity patterns to avoid humans. A shift to more nocturnal activity patterns has been observed in other carnivore species in areas with frequent human activ- ity. For example, brown bears Ursus arctos restrict active periods to twilight and night-time in areas with high road densities (Ordiz et al., 2014), and avoid humans by staying


Oryx, 2020, 54(6), 837–846 © 2020 Fauna & Flora International doi:10.1017/S0030605319000498


Page 1  |  Page 2  |  Page 3  |  Page 4  |  Page 5  |  Page 6  |  Page 7  |  Page 8  |  Page 9  |  Page 10  |  Page 11  |  Page 12  |  Page 13  |  Page 14  |  Page 15  |  Page 16  |  Page 17  |  Page 18  |  Page 19  |  Page 20  |  Page 21  |  Page 22  |  Page 23  |  Page 24  |  Page 25  |  Page 26  |  Page 27  |  Page 28  |  Page 29  |  Page 30  |  Page 31  |  Page 32  |  Page 33  |  Page 34  |  Page 35  |  Page 36  |  Page 37  |  Page 38  |  Page 39  |  Page 40  |  Page 41  |  Page 42  |  Page 43  |  Page 44  |  Page 45  |  Page 46  |  Page 47  |  Page 48  |  Page 49  |  Page 50  |  Page 51  |  Page 52  |  Page 53  |  Page 54  |  Page 55  |  Page 56  |  Page 57  |  Page 58  |  Page 59  |  Page 60  |  Page 61  |  Page 62  |  Page 63  |  Page 64  |  Page 65  |  Page 66  |  Page 67  |  Page 68  |  Page 69  |  Page 70  |  Page 71  |  Page 72  |  Page 73  |  Page 74  |  Page 75  |  Page 76  |  Page 77  |  Page 78  |  Page 79  |  Page 80  |  Page 81  |  Page 82  |  Page 83  |  Page 84  |  Page 85  |  Page 86  |  Page 87  |  Page 88  |  Page 89  |  Page 90  |  Page 91  |  Page 92  |  Page 93  |  Page 94  |  Page 95  |  Page 96  |  Page 97  |  Page 98  |  Page 99  |  Page 100  |  Page 101  |  Page 102  |  Page 103  |  Page 104  |  Page 105  |  Page 106  |  Page 107  |  Page 108  |  Page 109  |  Page 110  |  Page 111  |  Page 112  |  Page 113  |  Page 114  |  Page 115  |  Page 116  |  Page 117  |  Page 118  |  Page 119  |  Page 120  |  Page 121  |  Page 122  |  Page 123  |  Page 124  |  Page 125  |  Page 126  |  Page 127  |  Page 128  |  Page 129  |  Page 130  |  Page 131  |  Page 132  |  Page 133  |  Page 134  |  Page 135  |  Page 136  |  Page 137  |  Page 138  |  Page 139  |  Page 140  |  Page 141  |  Page 142  |  Page 143  |  Page 144  |  Page 145  |  Page 146  |  Page 147  |  Page 148  |  Page 149  |  Page 150  |  Page 151  |  Page 152  |  Page 153  |  Page 154  |  Page 155  |  Page 156  |  Page 157  |  Page 158  |  Page 159  |  Page 160  |  Page 161  |  Page 162  |  Page 163  |  Page 164