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1. Foraging efficiency and ecological risk aversion in juvenile Bornean orangutans

   Knott, Cheryl; DeLong, K (January 2017, American Association of Physical Anthropologists Annual Meeting)

   For species with low mortality and high reproductive costs, like primates, rather than maximize reproduction, natural selection should favor slow growth and slow reproductive rates. Orangutans, because of their slow life history, and the extreme fluctuations in their food supply, are hypothesized to have been selected for slow juvenile development to avoid ecological risk. Juveniles are predicted to be particularly vulnerable during periods of low food availability because of lower foraging success. Thus, we tested the hypothesis that juvenile orangutans are less efficient foragers than adults and that they are less able to both access and digest important fall-back foods. Data were collected on wild orangutans in Gunung Palung National Park, Borneo, Indonesia between 1994-2016. Analyses are drawn from 468 matched follows of mother-offspring pairs in which more than 75% of the diet has been analyzed. We found that juveniles ate fruit when their mother’s ate fruit during 98.3% of bouts. However, for other food items, juveniles were much less likely to eat the same food items (insects = 65.2%, leaves 76.5, bark 70.0%, flowers 75.0%, pith 65.8%). As expected, we found that juveniles ate significantly fewer calories than did adults overall, but this difference was particularly pronounced during periods of low food availability (p < 0.001). We show that these results were due to constraints on both the processing and digesting of fall-back foods. Thus, the long period of orangutan growth and dependency may reflect a risk-averse growth strategy in this forest characterized by dramatic fluctuations in preferred fruits. 

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2. Oral Processing Behavior in Bornean Orangutans of Different Age and Sex Classes

   Wyszynski, Sofia I; Kane, Erin E; Bakley, Tori D; Rinaldi, R; Susanto, Wahyu; Knott, Cheryl D. (November 2021, Northeastern Evolutionary Primatologists)

   Orangutan diets vary dramatically with food availability. Orangutans preferentially eat fruit when available, but due to dramatic and unpredictable fluctuations in fruit availability, orangutans often consume unripe fruit, bark, seeds, and leaves. Their robust craniodental structure suggests that they are well adapted to consume mechanically challenging foods. Since differences in jaw anatomy and body size pose physiological differences in terms of gape, exerted force, and resistance to wear and breakdown, growth and allometry likely affect an orangutan’s ability to process a mechanically challenging diet. Thus, we predict that orangutans of different ages and sexes process foods differently. Given juveniles' smaller and less powerful craniodental structure, and the time required to develop ecological competence, we hypothesized that juveniles may have more difficulty in processing foods than adults. We recorded the frequency that foods were introduced to the mouth, and chewed with different teeth (incisors, canines, and molars) in 561 feeding videos collected in Gunung Palung National Park in West Kalimantan, Borneo on wild orangutans (Pongo pygmaeus wurmbii). Videos were stratified by age and sex class and foods were categorized by type. Infants and juveniles use their canines significantly more frequently than adult females (p< 0.05) and flanged males (p< 0.05). Molar use also differed by age and sex class (F(3)=2.551, p=0.05), with juveniles chewing with their molars significantly more frequently than adult females (p=0.05). Differences in adult and juvenile oral processing profiles suggest juveniles may process some foods less efficiently than adults. 

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3. Ecological competence in wild Bornean orangutans: food sharing, processing, and nutrition

   Kane, Erin E; Susanto, Tri Wahyu; Conklin-Brittain, Nancy L; Knott, Cheryl D (October 2019, Third Lembersky Conference in Human Evolutionary Studies)

   Wild orangutans cope with dramatic, unpredictable fluctuations in food availability throughout development which have significant implications for energy balance for growth, development, and maintenance of body function. Foods that are especially important when preferred fruits are not available (e.g., bark/pith, termites, seeds) can require force, manual manipulation, and/or skill to access. Orangutans’ extended life history may mitigate ecological risk associated with the challenging environment of juvenile growth and development. Given the complexity of important foods, orangutans’ semi-solitary lifestyle, and the relatively brief period between weaning and independence, offspring transition to ecological independence during their extended nursing period. Here, we examine mother-offspring food transfer in the context of nutritional quality and complexity, with handling time as a proxy. Data were collected between July 2009-July 2019 in Gunung Palung National Park, Indonesia. We predict complex food items are transferred most frequently. In 245 observations of food sharing, fruit was most frequently transferred, followed by invertebrates and bark/pith. Handling time and transfer frequency were negatively correlated (Rho=-0.75, p<0.001): slower-eaten fruits were shared more frequently than rapidly-eaten fruits. Fruit size and sharing frequency were also negatively correlated (Rho=-0.73, p<0.001). There was no correlation between sharing frequency and free simple sugar concentration (Rho = 0.36, p=0.13), though sharing frequency and total nonstructural carbohydrate concentration were correlated (Rho=0.46, p=0.04). Food sharing was most common when mothers ate large, slowly-eaten fruits rich in nonstructural carbohydrates. Food sharing allows mothers to transfer more complex foods to their offspring, and may facilitate knowledge transfer as offspring become ecologically competent. 

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4. Developmental tradeoffs: Fruit availability and offspring age influence mother and offspring activity in Bornean orangutans (Pongo pygmaeus wurmbii)

   Kane, EE; Susanto, TW; Knott, CD (January 2021, American journal of physical anthropology)

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   Bornean orangutans' extended life history may be an adaptation to their forest habitat, characterized by dramatically fluctuating fruit availability. We hypothesize that juveniles rest and are carried more when fruit availability is low, spending less time in developmentally important behavior including play and independent travel to conserve energy. We test this using generalized linear mixed models and data derived from 976 follows of infant and juvenile orangutans collected during periods of varying fruit availability. Age (p<0.001), but not fruit availability (p>0.05) significantly affected offsprings’ time being carried (β=-8,1) and resting (β=-2.6, p<0.001). Younger individuals spent more time clinging and resting regardless of fruit availability. Fruit availability and offspring age interact to affect the proportion of time juveniles play (β=1.5, p<0.001) and travel (β=-0.8, p<0.001). Fruit availability impacts younger juveniles’ play behavior more dramatically than older juveniles: younger juveniles play more when fruit availability is high than when it is low, while older juveniles exhibit little variation in time spent playing depending on fruit availability. Juveniles aged 6-8 travel more when fruit availability is medium and high, juveniles aged 2-5 travel slightly less when fruit availability is high, and juveniles under 2 rarely travel independently. We also examined relationships between fruit availability, offspring age, and maternal travel distance using 2065 mother-offspring follows, demonstrating that females travel shorter distances with offspring under four, and when fruit availability is low. Thus, orangutans shift activity in response to fruit availability and throughout development, buffering young orangutans against energy depletion but suppressing developmentally important activities. Funders: NSF (9414388, BCS-1638823, BCS-0936199); National Geographic; USFish/Wildlife (F18AP00898, F15AP00812, F13AP00920, 96200-0-G249, 96200-9-G110); Leakey; Disney Conservation Fund; Wenner-Gren; Nacey-Maggioncalda; Conservation-Food-Health; Orangutan Conservancy; Woodland Park Zoo 

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5. The influence of fruit availability on energy allocation in infant and juvenile Bornean orangutans

   Kane, Erin E.; Susanto, Tri Wahyu; Knott, Cheryl D. (February 2020, American journal of physical anthropology)

   The juvenile risk aversion hypothesis suggests an extended life history is an adaptation to unpredictable environments, such as those that characterize Bornean orangutans. We examined how changes in infant and juvenile activity budgets changed over time and with varying periods of food availability. We hypothesized that during periods of low food availability juvenile orangutans would trade-off developmentally important behavior, such as play, in order to conserve energetic resources. We tested this hypothesis using 924 follows of infant (0-4 years) and juvenile (5-8 years) orangutans, collected over a 25 year period, encompassing period fo high and low food availability at Gunung Palung National Park, Indonesia. Using a GLMM to control for individual and examining both age and food availability we found that overall, infant and juveniles played less during period of low food availability compared to medium and high food availability (b= -14.3, p < 0.0001) and that play behavior varied significantly by age (b=-5.9,m p < 0.0001). Infants in the 1-2 age category showed the highest percentage of play, comprising 42% of their overall activity budget, with play dropping to 14% by age 4. We also found signficant interaction effects between age and food availability on play behavior (b=2.1, p < 0.0001). We discuss the ways that infants and juvenile trade off energetically expensive behavior through different development periods as well as the role that maternal behavior may play in buffering infants and juveniles of coping with periods of low food availability. 

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