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In 2015, Lambert and Rothman urged primate nutritional ecologists to revise the view of fallback and optimal foods from stable traits inherent in the food to variable qualities determined by the state of the consumer. Here we provide behavioral evidence to support this revision. In primates, fruit is often the preferred food category because it is typically high-energy, high-carbohydrate, and low in fiber. Orangutans in particular, are said to consume fruit preferentially and when it is available, whereas leaves, bark, and pith are often considered fallback foods. Using movement ecology, we ask if wild Bornean orangutans (Pongo pygmaeus wurmbii) navigate only to fruit, or whether navigation to non-fruit foods is supported by our data. We find that orangutans do deviate from a direct fruit-to-fruit path to consume non-fruit foods (n = 54, range 8% - 84%; p = 6.819e-07.) Next, we ask if orangutans consume non-fruit foods when in the proximity of fruit resources. We find that 25.5% of the time that orangutans eat a non-fruit food, there is an available fruit within 50m (n=308). Building on previous research finding that orangutans maintain a 10.1:1 NPE:P balance, we use this geospatial data showing that orangutans navigate to and choose non-fruit foods, even when fruit is available, to suggest that orangutans are seeking foods based on their current nutritional state and not only to maximize energy. This supports the claim that 'fallback' is not an inherent characteristic of a food, but rather is in the state-dependent eye of the consumer. Funders: National Science Foundation (BCS-1638823; BCS-1613393), NSF GRFP (DGE-1247312); Boston University; National Geographic Society, US Fish and Wildlife (F15AP00812), Leakey Foundation, Disney Wildlife Conservation Fund; Wenner-Gren Foundation; Nacey-Maggioncalda Foundation, Princeton University 

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 

Seed dispersal is important for forest growth, maintenance, and regeneration. Orangutans are large-bodied frugivores with ecological roles as seed predators and seed dispersers. However, little is known about orangutans’ ecological roles and how they relate to orangutans’ patterns of frugivory. We investigated Bornean orangutans’ (Pongo pygmaeus wurmbii) ecological roles at the Cabang Panti Research Station in Gunung Palung National Park, Indonesian Borneo. We collected orangutan feces (n=401) and analyzed them for intact seeds (August 2018 to March 2020). We observed orangutan fruit handling behavior for 306 feeding bouts for 53 fruit genera to measure how often orangutans swallow, spit, or predate seeds. We used Ivlev’s Electivity Index to analyze fruit preference using long-term feeding data and phenology data (2014-2019). Lastly, we combined fruit preference with fruit handling behavior using the seed dispersal effectiveness framework to identify which fruit taxa were most effectively dispersed. Orangutans dispersed seeds in 71.8% of fecal samples with a mean of 27.9 ±4.5 (SD=0.95) seeds (>2mm) per fecal sample. Orangutans predated seeds more often than spitting or swallowing seeds (predating= 42.1% of fruit feeding time; spitting= 21.8%; swallowing= 12.5%; mixed behaviors= 10.6%, not observed=12.0%). Additionally, the top five preferred fruit genera, (Dialium, Sindora, Scaphium, Magnifera, and Spatholobus) were highly predated (0 to 5% of seeds dispersed). We identified Alangium and Tetramerista as the most effectively dispersed genera, orangutans frequently dispersed and preferred these fruits. We found orangutans are frequent seed predators, but this overlaps with their seed dispersal role, and we describe orangutans’ seed dispersal contribution. 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; Boston University GRAF 

Due to extreme incongruence in parental investment, the potential for sexual conflict in orangutan reproductive strategies is high. Female orangutans prefer flanged males, but with intense male-male competition and sexual coercion, it is unclear how females are able to exert choice. We hypothesized that female orangutans use initiation and maintenance of associations with preferred males as a mechanism of female choice. We used encounter rates and behavioral measures of proximity maintenance to distinguish between the role of female choice and male coercion in male-female associations in Gunung Palung National Park, West Kalimantan, Indonesia. From May 2018-April 2019, we collected data on initiation, termination, and proximity maintenance during associations between males and cycling females (females without dependent offspring or with offspring over 6 years old). Encounters between cycling females and flanged males were more likely to be female-initiated (83.3%), while encounters with unflanged males were more likely to be male-initiated (80%) (N=16, p=0.035, Fisher’s exact test). Additionally, the Brown Index indicates significantly more female proximity maintenance when cycling females associated with flanged males than unflanged males (N=33, W = 198.5, p= 0.012). However, using long-term data on male-female associations, we found that dyads of flanged males and females were less likely to encounter other males compared to dyads of unflanged males and females (N=542, χ2= 3.3051, df=1, p=0.035). This indicates that flanged males may also use associations to mate guard females. These data indicate that there are behavioral manifestations of both female choice and male coercion in orangutan associations. Funders: NSF (DGE-1247312, BCS-1638823, BCS-0936199), Boston University, Leakey Foundation, Disney Conservation Fund, and US Fish and Wildlife Service (F18AP00898) 

In 2015, Lambert and Rothman urged primate nutritional ecologists to revise the view of fallback and optimal foods from stable traits inherent in the food to variable qualities determined by the state of the consumer. Here we provide behavioral evidence to support this revision. In primates, fruit is often the preferred food category because it is typically high-energy, high-carbohydrate, and low in fiber. Orangutans in particular, are said to consume fruit preferentially and when it is available, whereas leaves, bark, and pith are often considered fallback foods. Using movement ecology, we ask if wild Bornean orangutans (Pongo pygmaeus wurmbii) navigate only to fruit, or whether navigation to non-fruit foods is supported by our data. We find that orangutans do deviate from a direct fruit-to-fruit path to consume non-fruit foods (n = 54, range 8% - 84%; p = 6.819e-07.) Next, we ask if orangutans consume non-fruit foods when in the proximity of fruit resources. We find that 25.5% of the time that orangutans eat a non-fruit food, there is an available fruit within 50m (n=308). Building on previous research finding that orangutans maintain a 10.1:1 NPE:P balance, we use this geospatial data showing that orangutans navigate to and choose non-fruit foods, even when fruit is available, to suggest that orangutans are seeking foods based on their current nutritional state and not only to maximize energy. This supports the claim that 'fallback' is not an inherent characteristic of a food, but rather is in the state-dependent eye of the consumer. Funders: National Science Foundation (BCS-1638823; BCS-1613393), NSF GRFP (DGE-1247312); Boston University; National Geographic Society, US Fish and Wildlife (F15AP00812), Leakey Foundation, Disney Wildlife Conservation Fund; Wenner-Gren Foundation; Nacey-Maggioncalda Foundation, Princeton University 

Due to extreme incongruence in parental investment, the potential for sexual conflict in orangutan reproductive strategies is high. Female orangutans prefer flanged males, but with intense male-male competition and sexual coercion, it is unclear how females are able to exert choice. We hypothesized that female orangutans use initiation and maintenance of associations with preferred males as a mechanism of female choice. We used encounter rates and behavioral measures of proximity maintenance to distinguish between the role of female choice and male coercion in male-female associations in Gunung Palung National Park, West Kalimantan, Indonesia. From May 2018-April 2019, we collected data on initiation, termination, and proximity maintenance during associations between males and cycling females (females without dependent offspring or with offspring over 6 years old). Encounters between cycling females and flanged males were more likely to be female-initiated (83.3%), while encounters with unflanged males were more likely to be male-initiated (80%) (N=16, p=0.035, Fisher’s exact test). Additionally, the Brown Index indicates significantly more female proximity maintenance when cycling females associated with flanged males than unflanged males (N=33, W = 198.5, p= 0.012). However, using long-term data on male-female associations, we found that dyads of flanged males and females were less likely to encounter other males compared to dyads of unflanged males and females (N=542, χ2= 3.3051, df=1, p=0.035). This indicates that flanged males may also use associations to mate guard females. These data indicate that there are behavioral manifestations of both female choice and male coercion in orangutan associations. Funders: NSF (DGE-1247312, BCS-1638823, BCS-0936199), Boston University, Leakey Foundation, Disney Conservation Fund, and US Fish and Wildlife Service (F18AP00898) 

Seed dispersal is important for forest growth, maintenance, and regeneration. Orangutans are large-bodied frugivores with ecological roles as seed predators and seed dispersers. However, little is known about orangutans’ ecological roles and how they relate to orangutans’ patterns of frugivory. We investigated Bornean orangutans’ (Pongo pygmaeus wurmbii) ecological roles at the Cabang Panti Research Station in Gunung Palung National Park, Indonesian Borneo. We collected orangutan feces (n=401) and analyzed them for intact seeds (August 2018 to March 2020). We observed orangutan fruit handling behavior for 306 feeding bouts for 53 fruit genera to measure how often orangutans swallow, spit, or predate seeds. We used Ivlev’s Electivity Index to analyze fruit preference using long-term feeding data and phenology data (2014-2019). Lastly, we combined fruit preference with fruit handling behavior using the seed dispersal effectiveness framework to identify which fruit taxa were most effectively dispersed. Orangutans dispersed seeds in 71.8% of fecal samples with a mean of 27.9 ±4.5 (SD=0.95) seeds (>2mm) per fecal sample. Orangutans predated seeds more often than spitting or swallowing seeds (predating= 42.1% of fruit feeding time; spitting= 21.8%; swallowing= 12.5%; mixed behaviors= 10.6%, not observed=12.0%). Additionally, the top five preferred fruit genera, (Dialium, Sindora, Scaphium, Magnifera, and Spatholobus) were highly predated (0 to 5% of seeds dispersed). We identified Alangium and Tetramerista as the most effectively dispersed genera, orangutans frequently dispersed and preferred these fruits. We found orangutans are frequent seed predators, but this overlaps with their seed dispersal role, and we describe orangutans’ seed dispersal contribution. 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; Boston University GRAF 

Bornean orangutans (Pongo pygmaeus wurmbii) are large bodied great apes that live in rainforests dominated by mast-fruiting dipterocarp trees with extreme fluctuations in fruit availability. Orangutans respond to this temporal and spatial variability in fruit production by ranging over large areas and adopting a semi-solitary social structure. Females have overlapping home ranges, engage in both scramble and contest competition for food, and actively avoid each other. Overlap requires individuals to share access to resources and adjust ranging to optimize energy intake, thus habitat quality likely influences ranging patterns. Here we investigate whether habitat and food availability are significant predictors of female orangutan home range overlap using data collected at Gunung Palung National Park, Indonesia, a site with 7 distinct habitats. Researchers collected GPS waypoints of orangutan movements during all day focal follows. Fruit availability was measured through monthly monitoring of over 6000 trees, across 60 plots. We used R to calculate range overlap per habitat between pairs of adult female orangutans over three-month periods (2013-2019). Our results show a trend towards a negative relationship between overlap and fruit availability (N=15, Pearson’s R= -0.322, p=0.242). We also found habitat to be a predictor of female range overlap, with overlap most likely to occur in the alluvial bench habitat and significantly less likely in the peat swamp (p<0.05). These findings reveal the independent influences of fruit availability and habitat type on female orangutan home range overlap, highlighting the potential importance of habitat-specific food availability on ranging behavior and contest competition. 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; Boston University GRAF