Sleep patterns, daytime predation, and the evolution of diurnal sleep site selection in lorisiforms.

OBJECTIVES: Synthesize information on sleep patterns, sleep site use, and daytime predation at sleep sites in lorisiforms of Asia and Africa (10 genera, 36 species), and infer patterns of evolution of sleep site selection. MATERIALS AND METHODS: We conducted fieldwork in 12 African and six Asian countries, collecting data on sleep sites, timing of sleep and predation during daytime. We obtained additional information from literature and through correspondence. Using a phylogenetic approach, we established ancestral states of sleep site selection in lorisiforms and traced their evolution. RESULTS: The ancestral lorisiform was a fur-clinger and used dense tangles and branches/forks as sleep sites. Use of tree holes and nests as sleep sites emerged ∼22 Mya (range 17-26 Mya) in Africa, and use of bamboo emerged ∼11 (7-14) Mya in Asia and later in Africa. Fur clinging and some sleep sites (e.g., tree holes, nests, but not bamboo or dense tangles) show strong phylogenetic signal. Nests are used by Galagoides, Paragalago, Galago and Otolemur; tree holes by Galago, Paragalago, Sciurocheirus and Perodicticus; tangles by Nycticebus, Loris, Galagoides, Galago, Euoticus, Otolemur, Perodicticus and Arctocebus; all but Sciurocheirus and Otolemur additionally sleep on branches/forks. Daytime predation may affect sleep site selection and sleep patterns in some species of Nycticebus, Galago, Galagoides, Otolemur and Perodicticus. Most lorisiforms enter their sleep sites around sunrise and leave around sunset; several are active during twilight or, briefly, during daytime. CONCLUSION: Variations in sleep behavior, sleep patterns and vulnerability to daytime predation provide a window into the variation that was present in sleep in early primates. Overall, lorisiforms use the daytime for sleeping and no species can be classified as cathemeral or polycyclic.

Slow lorises (Nycticebus spp.) display evidence of handedness in the wild and in captivity.

It has been suggested that strepsirrhines (lemurs, lorises, and galagos) retain the more primitive left-hand preference, whilst monkeys and apes more regularly display a right-hand preference at the individual-level. We looked to address questions of laterality in the slow loris (Nycticebus spp.) using spontaneous observations of 7 wild individuals, unimanual tests in 6 captive individuals, and photos of 42 individuals in a bilateral posture assessing handedness at the individual- and group-level. During the unimanual reach task, we found at the individual-level, only 4 slow lorises showed a hand use bias (R: 3, L: 1), Handedness index (HI) ranged from -0.57 to 1.00. In the wild unimanual grasp task, we found at the individual-level two individual showed a right-hand bias, the HI ranged from -0.19 to 0.70. The bilateral venom pose showed a trend toward a right-hand dominant grip in those photographed in captivity, but an ambiguous difference in wild individuals. There are many environmental constraints in captivity that wild animals do not face, thus data collected in wild settings are more representative of their natural state. The presence of right-handedness in these species suggests that there is a need to re-evaluate the evolution of handedness in primates.

Environment shapes sleep patterns in a wild nocturnal primate.

Among primates, the suborder Haplorhini is considered to have evolved a consolidated monophasic sleep pattern, with diurnal species requiring a shorter sleep duration than nocturnal species. Only a few primate species have been systematically studied in their natural habitat where environmental variables, including temperature and light, have a major influence on sleep and activity patterns. Here we report the first sleep study on a nocturnal primate performed in the wild. We fitted seven wild Javan slow lorises (Nycticebus javanicus) in West Java, Indonesia with accelerometers that collected activity data, and installed climate loggers in each individual's home range to collect ambient temperature readings (over 321 days in total). All individuals showed a strictly nocturnal pattern of activity and displayed a striking synchronisation of onset and cessation of activity in relation to sunset and sunrise. The longest consolidated rest episodes were typically clustered near the beginning and towards the end of the light period, and this pattern was inversely related to daily fluctuations of the ambient temperature. The striking relationship between daily activity patterns, light levels and temperature suggests a major role of the environment in shaping the daily architecture of waking and sleep. We concluded that well-known phenotypic variability in daily sleep amount and architecture across species may represent an adaptation to changes in the environment. Our data suggest that the consolidated monophasic sleep patterns shaped by environmental pressures observed in slow lorises represent phylogenetic inertia in the evolution of sleep patterns in humans.