At least half of the world’s primate species are currently threatened with extinction. Slow life histories combined with rapid habitat loss and hunting in recent years has heightened the extinction risk for many species, including the two largest extant lemurs, the diademed sifaka (Propithecus diadema) and the indri (Indri indri). Both species belong to the taxonomic family Indriidae, have similar adult weights, and occur in sympatry in certain areas of the montane rainforests of eastern Madagascar. Both species are adapted for folivory however I. indri spend considerably more time feeding on leaves than do P. diadema resulting in several energy-saving adaptations in I.indri. In this dissertation, I explore infant development and maternal strategies of these critically endangered primates with the goal of increasing our knowledge of reproduction and ontogeny in both species. Although previous studies have elucidated key differences in adult behaviour, there is a dearth of information on infants and lactating females in either of these two species. Between June and December of 2011 and 2012, I collected continuous time focal animal data, in Maromizaha forest, to examine behavioural patterns of 12 infants and their mothers from 0 – 33 weeks. In addition, I developed a framework to define and quantify the weaning process and facilitate comparisons across different species and studies. P. diadema infants developed feeding competency and independent locomotion faster than did I. indri infants however both species were consistently feeding independently more than they were suckling by week 20. The process of feeding ontogeny in I. indri was likely accelerated by coprophagy, as all infants of this species consumed their mother’s feces regularly from 10 – 15 weeks old. Lactating females of both species spent more time feeding in mid-lactation when maternal investment was the highest. The prolonged inter-birth interval in I. indri is suggested as another adaptation that reduces energetic expenditures. In addition, the protracted period of close contact with their mother may offer infant I. indri more time for social learning of the mother’s diet and the group song and for developing competency in vertical clinging and leaping without a tail for balance and support. / Graduate

I studied the influence of microhabitat on the predator-sensitive foraging (PSF) of Verreaux’s sifaka (Propithecus verreauxi verreauxi) at Berenty Reserve, Madagascar from June 11, 2007 to August 10, 2007. Three groups of sifaka from a gallery, closed-canopy forest, a riverine forest, and the spiny/scrub forest were observed. PSF was assessed by measuring the spatial location, spatial cohesiveness, and rate of vigilance, vocal, and non-vocal alarm behaviour of foraging sifaka. While there were statistically significant between-group differences in the spatial location (i.e., terrestrial, low-, mid- and high-canopy) used while foraging, there were no statistically significant inter-group differences in spatial cohesiveness, terrestrial or aerial vigilance, or in the frequency of vocal (i.e., terrestrial and aerial calls) or non-vocal (i.e., gestural) alarms. Thus, it appears that the PSF of free-living Verreaux’s sifaka is largely uninfluenced by variation in microhabitat.

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Habitat degradation and hunting pose the most proximate threats to many primate species, while climate change is expected to exacerbate these threats (habitat and climate change combined henceforth as "global change") and present new challenges. Madagascar's lemurs are earth's most endangered primates, placing added urgency to their conservation in the face of global change. My dissertation focused on the critically endangered golden-crowned sifaka (Propithecus tattersalli; hereafter, "sifaka") which is endemic to fragmented forests across a gradient of dry, moderate, and wet forest types in northeastern Madagascar. I surveyed sifakas across their global range and investigated factors affecting their densities. I explored sifaka diets across different forest types and evaluated if nutritional factors influenced sifaka densities. Lastly, I investigated sifaka range-wide genetic diversity and conducted a connectivity analysis to prioritize corridor-restoration and other potential conservation efforts. Sifaka densities varied widely across forest fragments (6.8 (SE = 2.0-22.8) to 78.1 (SE = 53.1-114.8) sifakas/km²) and populations have declined by as much as 30-43% in 10 years, from ~18,000 to 10,222-12,631 individuals (95% CI: 8,230-15,966). Tree cutting, normalized difference vegetation index (NDVI) during the wet season, and Simpson's diversity index (1-D) predicted sifaka densities range-wide. Sifakas consumed over 101 plant species and spent 27.1% of their active time feeding on buds, flowers, fruits, seeds, and young and mature leaves. Feeding effort and plant part consumption varied by season, forest type, and sex. Minerals in sifaka food items (Mg (β = 0.62, SE = 0.19) and K (β = 0.58, SE = 0.20)) and wet season NDVI (β = 0.43, SE = 0.20) predicted sifaka densities. Genetic measures across forest fragments indicated that sifaka populations are becoming more isolated (moderate FIS values: mean = 0.27, range = 0.11-0.60; high M-ratios: mean = 0.59, range = 0.49-0.82; low overall effective population size: Ne = 139.8-144 sifakas). FST comparisons between fragments (mean = 0.12, range = 0.01-0.30) supported previous findings that sifakas still moved across the fragmented landscape. Further validation of these genetic results is needed. I identified critical corridors that conservation managers could protect and/or expand via active reforestation to ensure the continued existence of this critically-endangered lemur. / Doctor of Philosophy / Worldwide, many species of primates are threatened with extinction due to habitat degradation, hunting, and climate change (habitat and climate combined threats, henceforth, "global change"). These threats work at different time scales, with hunting being the most immediate and climate change likely to have its fullest impact experienced from the present to a longer time frame. Lemurs are a type of primate found only on Madagascar, an island experiencing rapid global change, which puts lemurs at a heightened risk of extinction. My dissertation research focused on the critically endangered golden-crowned sifaka (Propithecus tattersalli; hereafter, "sifaka"), a species of lemur found only in a few isolated forests across a dry to wet gradient in northeastern Madagascar. To better understand their extinction risk, I conducted surveys to estimate the number of sifakas remaining and investigated several factors that might determine how many sifakas can live in one place. I then explored how sifaka diets varied depending on the forest type that they inhabit and tested whether nutrients in their food might determine sifaka numbers. Lastly, I calculated sifaka genetic diversity to assess their ability to adapt to new environmental conditions and to determine whether sifakas can move across the landscape to find new mates and to potentially colonize new areas of habitat. Sifaka densities varied widely across their range (6.8-78.1 sifakas/km² ). Only 10,222-12,631 sifakas remain, which is 30-43% less than the range of estimates obtained 10 years ago (~18,000 sifakas). Tree cutting, normalized difference vegetation index (NDVI; a measure of plant health or "greenness" obtained from satellite data), and a tree species diversity index were useful measures to predict sifaka densities. Sifakas ate different plant parts (buds, flowers, fruits, seeds, and leaves) from over 101 plant species. The amount of time they spent eating each day varied by the time of year, forest type, and sex. On average, they spent a quarter of their day eating. Magnesium and potassium concentrations in sifaka food items also were useful nutrition-related measures to predict sifaka densities. Genetic analyses suggested that sifaka populations are becoming more isolated and inbred, meaning sifakas are breeding with other sifakas to which they are closely related. However, it appears that sifakas still can move between forest patches to find new mates and to potentially colonize new areas, if such areas are created. Further validation of these genetic results is needed. I also identified critical areas that will be important to protect and reforest to ensure that movements between populations can continue.

To advance conservation in the increasingly fragmented landscape of Madagascar, we must examine the persistence and interactions of species in human-disturbed habitats. I investigated lemur-predator interactions in southeastern Madagascar through a comparison of predator communities and anti-predator behaviors of Milne-Edwards' sifakas (Propithecus edwardsi) in the continuous rainforest of Ranomafana National Park, and the forest fragments of Ialatsara Forest Station. I confirmed the presence of potential aerial predators at each site, but the sifakas' confirmed native mammalian predator, fossa (Cryptoprocta ferox), was absent from the fragmented site. Playbacks of predator vocalizations did not suggest that fragment-living sifakas have weakened anti-predator responses, but that their responses may be less specific than those of conspecifics in the continuous forest. I found that fragment-living sifakas displayed less downward vigilance and more frequently used low canopy heights; these behaviors may increase their vulnerability to recolonizing ground predators. I investigated local ecological knowledge (LEK) of carnivore ecology in communities 0–20 km from continuous forest to explore the potential for such recolonization. My findings from 182 interviews in 17 communities suggest that the fossa is especially sensitive to anthropogenic disturbance; it was only observed in communities ≤ 2.5 km from the continuous forest within the last five years. In contrast, the introduced small Indian civet (Viverricula indica) and wild cat (Felis silvestris) were distributed ubiquitously and displayed an affinity to human-dominated habitats. LEK surveys can provide information on the poorly understood responses of the Malagasy carnivores to the threats they face in a changing landscape. / Master of Science

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