Terrestrial African Monkeys 1) Cercopithicini a. Chlorocebus (vervet monkey) i. Small (3-4.

5 kg), very closely related to the guenons in size and shape, it is an edge and riverine species, rarely goes into deepest forest or driest areas ii. Locomotion is intermediate, use both the ground and the trees, somewhere in between Papio and guenons, about 20% of their time spent on the ground, 30% at lower levels, 35% in the crown of the trees iii. Opportunistic omnivores, fruits make up 25-65% depending on location and season, leaves represent 5-40%, flowers are 10-35%, grasses are 1-15%, and animal prey makes up 5-40% of their diet, utilize some forest resources 1. Senegal at 17 species of plants, in another study, they fed on 66 species of plants, though a great proportion is from Acacia spp. And grasses iv. Go into the trees for most of their fruit and plant food, quick and agile for searching for insects b. Erythrocebus (patas monkey) i. Limited to the northern savannah regions, dry areas where they seem to replace the vervet monkeys ii. Males weigh about 12 kg, females are approximately 6 kg iii. Most terrestrially adapted, almost all of their locomotion and 85% of their feeding on group, slender with long limbs, not well-adapted climbers, use trees to rest and survey landscape, can run up to 35 mph iv. Depend on three species of Acacia for gums, leaves, and swollen thorns to get ants within, 80-85% of diet during dry season is made up of one species of Acacia, will sometimes eat fleshy fruits, seeds, flowers, and mushrooms when available v. Use the ground and sometimes widely-dispersed trees throughout savannah vi. Cat-like, sneak up on insect prey 2) Papionini a. Papio (savannah baboons) i. Five species, Pan-African Sub-Saharan species, four of the species are along this range, fifth is the Papio hamadryas which is found in Ethiopia and Southwest Arabia ii. Forest fringe animals, also present in gallery forest, edges of primary or secondary forest, sometimes in rocky or hilly areas, usually found near standing water iii. Males weight about 20 kg, females are approximately 10 kg iv. Stocky build, arms and legs are about the same length, spend a great deal of time on the ground, though will feed and rest in trees, P. hamadryas uses the ground slightly more often

v. Opportunistic omnivore, focuses on a few specific species that are abundant in the area (primarily Acacia spp.), eat seeds, pods, blossoms, saps, cambian layer of bark, leaves, and feed on insects, also feeds on tamarindus indica (kili trees) with small nutritious leaves, savannah resources vi. During wet season, eat blades of grasses, during the dry season, eat seeds, rhizomes, and corms (underground stems), may make up 90% of dry season diet, most other animals do not eat this resource, very adaptable vii. May go into the forest in search of fruit, but will not go very deep into the forest viii. Hunts a number of relateively large vertebrates, not enough protein in insects ix. P. hamadryas lives in even drier areas that other four species, depend heavily on Acacia (30-90%) and grasses, during dry season, focus on dry-adapted plants and underground corms, will not exploit forest resources like other baboons, will only go into forest to reach water sources, do not eat many insects or vertebrates x. Tend to feed on the ground throughout the savannah, look under rocks for insects, slow and deliberate xi. Social Organization 1. 8-200 animal groups with a mean between 30-50 animals, multi-male multi-female groups 2. When groups are smaller, spend much more time surveying the environment, larger groups are beneficial on the open plains when predation is common (Stacey) 3. Sub-adult males tend to be on the periphery of the group, some are slowly integrating from other groups, others migrating out, some alpha males in the core of the group with females, juveniles, and infants 4. Zuckerman (1932) sex and the drive of sex is what maintained group structure, probably not the case because of the seasonality of sex and breakdown of structure during these times 5. Washburn and DeVore (1965-70) first to spend long periods of time in the field, center of social organization was the dominant male, kept group stable, not really true, dominance hierarchy and dominant males are actually least stable aspect of baboon groups a. Altmann and Havsfater - changes occur by demographic factors (older leaves, younger rises, every 13.3 days) or through fights (occur every 21 days) 6. Packer (mid-1980s) all males migrated from natal group, 50 animals changed groups, 48 were males, of

these, 2 were juveniles, 41 were sub-adult or mature males, 5 were old males (on average every 4.5 years), all older males had transferred in and all sub-adults had transferred out a. Natal males would migrate out when they reached sub-adulthood, usually leave in pairs move into a group with fewer males their age b. Multiple-transfer males would move into a group with more estrous females c. Dominance hierarchy was not the glue holding these groups together 7. Donald Stone-Sade studied primarily macaques, familial interactions of kin focusing on matrilines, 79% of nearest neighbors were between kin, only 15% would be expected by chance a. Studied allo-grooming, 62% were between kin, other grooming partners were male-female couples b. Males, upon reaching sub-adulthood, would not mate with mothers or sisters, but would during dominance stage, non-copulative mating, mate with mothers and sisters to prove dominance 8. Imo was a young Japanese macaque, researchers would leave sweet potatoes on the beach, she began washing them in the ocean, traced adoption of this trait throughout the group, first age-mates through her mother, then other age-mates, up to their mothers, 5 years later, 80% of 2-7 year-olds had picked up the trait, 18% over 7 (all female) picked it up a. Males were more conservative than females, except before 3 years old b. At four year old, Imo also sorted oatmeal and rice by separating it from sand in water, 2-4 year olds were the first to pick it up 9. Females have a much more stable hierarchy, only change because of demographic reasons, females inherit dominance through their mother, alphas daughters will move into hierarchy just below her, youngest adult daughter will be immediately below mother, maintains stability over time 10. Dominance is not a genetic trait, no reason for their to be relationship with reproductive success a. Hausfater found that only about 50% of mating could be explained by dominance hierarchy

b. Packer found that there was no correlation between number of days spent consorting and dominance i. Male was more successful if consort relationship developed ii. Males may have been dominant because of their size, may have lost out because iii. If two males align to fight off another male, both will mate no matter what their dominance position is iv. Older males had more success than would be expected from their position in the hierarchy, age, tenure in the group v. Dominant males may get in too many fights, may have been injured or be too aggressive for females vi. General canine condition related to mating, but not dominance vii. Fatigue of dominant males allows others to move in viii. Male selectivity and female choice were not related to dominance c. Shirley Strum focused on females first, found that males were not the center of group attention i. Better predictor of mating success was tenure in the group rather than dominance, social finesse allowed them to stay in the group for extended periods ii. 7 to 8 adults females throughout study, formed sleeping groups with 2-3 adults males, same males would have close relationships during the day, involved with only a small subset of the group d. Barbara Smutz found that every one of the 34 females in the group had a special relationship with at least one of the 18 males, friends i. Females had more friendly interactions with friends than other males ii. Tense appeasement gestures around other males iii. Males were responsible for intervening in disputes for female and her offspring iv. 97% of friendly interactions with infants were with friendly males e. Rasmussen found that special relationship were maintained in consortship

i. Closer proximity and grooming between friends translated into reproductive presentations by females more to their friends than others, males mounted more often, clearly not related to hierarchy f. Bernstein tried to determine relationship between aggression, grooming, and mounting i. Could not correlate with dominance for any of these traits in any of five species g. Fighting, mating, grooming, inventiveness, priority of access to resources, leader of group movement all used as measures of dominance i. These are not correlated to one another, so dominance is incarnated differently for different species h. Silk, Alberts, and Altmann found that social contact and social integration were positively associated with reproductive success of females, these were independent of rank b. Theropithicus (gelada baboon) i. Wet grasslands in the cool highlands of Ethiopia, gramnivore (the cow of the primates), grass-eating adaptations found in ancient baboons, 90% of diet consists of grasses, some bushes, focus on green grasses during the wet season, seeds at the beginning of the dry season, during the peak of the dry, dig for corms ii. Males weight about 20 kg, females are approximately 10 kg iii. Longer arms than legs, spends almost all of its time on the ground, sits on its haunches feeding on grasses iv. Only species that is not an omnivore, almost no animal protein in its diet v. Differences in what they eat are vary by location, certain species are more or less dense in a given area, focus on Acacia and grasses, also shoots and tamarind pods, as well as some more dry-adapted plant species, very infrequently eat vertebrates of invertebrates vi. Longer arms than legs, sit on haunches and shuffle while the pick grasses at a very high rate 3) Crook and Aldrich-Blake (1968) first study to utilize the scan-sampling method that everyone uses now, Jeane Altmann later reviewed these methods and was attributed credit a. Studied three terrestrial primates (Anubis baboon, gelada monkey, and vervet monkey) in sympatry in Ethiopia i. Vervet monkey are found mainly in gallery forest and at the edge of cliffs where there are thickets and some grasses, sometimes went into the cultivated areas

1. 135 animals/km2, studied three groups, home ranges were about 30 hectares 2. Day range was about 700 meters (600-800 m range) 3. Use of trees was about 57% of the time, on the ground about 30% 4. Ate significantly more insects than other species, ate a lot of fruits, some flowers, some leaves ii. Papio anubis went through all of the zones, range throughout the entire area, gallery forest, and thicket predominated, though sometimes went into open areas, were chased 1. Mean group size of 20, home range of about 95 hectares, only 26 individuals/km2 2. Day range was about 1220 meters (700-2000 m range) 3. Spent about 28% of their time in trees, 62% of their time on grasses and herbs (ground) 4. Ate primarily fruits and seeds, but also a significant amount of leaves (mostly of grasses) iii. Theropithecus gelada was only found in open grassland and thicket areas, slept on the cliffs, and were never far from the grasses, never went into forest 1. One group of 17 animals, small one-male units, 82 animals/km2 and home range was 84 hectares 2. Day range was about 630 meters (500-1000 m range) 3. Spent less than 1% of their time in the trees, spent almost 98% of their time on the ground 4. Over 90% of their diet is made up of leaves (mostly grasses) 4) Differences in Social Organization a. Savannah Baboons large group with females in the center (matrilineal organization) with dominant males, migrating and subadult males on the periphery i. Believed that males were expendable, so they would move on the outside of the group, however this is not the case ii. Seems that females always have large males (friends) with them when they move iii. Rarely restricted by food and water, tend to be fairly well protected by predation, though they are still vulnerable to terrestrial predators, sleeping sites are in the forest where predation is not common b. Hamadryas Baboons 12-750 animals in a troop (sleeping group) often found at these rocky outcrops, look a bit like savannah baboon groups, but in the morning, troop with split into a number of smaller groups and disperse as bands for travel to feeding sites, then split into uni-male or all-male groups which act as stable sexual units (1 male with 1-9 females), band and troop may not be the same the next night,

certain males seem to stick together in larger organizations in clans (form of male patri-locality) i. Males are the center of attention, lead the group, herd the females and keep them in line ii. Males are the same from year to year, but females migrate, male from an all-male group with adopt a daughter from a unimale group, in uni-male groups with very old male, son may stick around and later usurp power iii. Live in much drier areas, with rocky outcrops, very few trees, and no riverine forests iv. Food in highly dispersed, more predation in these open areas without forest to provide cover, protective sleeping sites are very rare, must come together to reduce vulnerability c. Gelada Baboons look similar to savannah baboon organization, near cliffs with grasslands nearby, will rarely move 300 meters from the sleeping sites at the cliffs, only permanent unit is the uni-male group that spreads out each day to feed, sometimes with a number of unimale groups in bands, and returns to the herd at night i. Food depends on seasonality, high predation risk on open plains, sleeping sites are only available at the cliffs edge ii. Females are the focus of the group (not herded), young female will adopt a male, female will sometimes form groups to choose males, if a male dies, female will attract another male rather than a male actively usurping power d. Patas Monkey (Cercopithecinii) live out in the driest, most resourcepoor areas, one-male units with very large home ranges (not an exclusive reproductive unit, non-resident males may join during reproductive season), females are resident, these are essentially their home ranges, group average 22 members with 1:17 sex ratio, males are there primarily for predator avoidance, males gain knowledge of resources from females i. Competition occurs between males when in the same group ii. High predation, so most visually obvious animals in the area without many trees iii. Rowell studied two groups, most of the time, there were no males in the group, 18 periods of males residence in these group, during 14 of these, males were there for less than 2 months, in 4 of these stayed for more than 2 months, more than one male was in the group several times iv. Males are much more vulnerable when they are not in a group

Savannah Baboon Hamadryas Gelada Baboon Patas Monkey Arboreal

Food/Water + + - + + - + +

Predation (+ high) + + + + + + + + -

Sleeping Sites + + - + - + +

Asian Monkeys 1) Colobinae a. Presbytis (langurs) has three species groups i. Semnopithecus found in India, Nepal, Sri Lanka 1. Presbytis entellus (Hanaman langur) very widespread throughout the region, sympatric both Kasi species (P. senex in Sri Lanka and P. johnii in India) a. P. entellus is very adaptable, generalized folivore, edge species, large home ranges, and variable groups structures and sizes b. P. senex is restricted to the Sri Lankan rainforest, specialized arboreal folivore, focusing on leaves and some fruits when available, small one-male groups c. P. johnii are similarly specialized folivores in western Indian rainforest ii. Presbytis Southeast Asia 1. Melalophus is a generalized frugivore and folivore, low canopy, small one-male groups iii. Trachypithecus Southeast Asia 1. Specialized folivore, focuses on mature leaves, lives primarily in the high canopy, quadrupedal locomotion, large multi-male and one-male groups b. Odd-nosed Monkeys Eastern Asia i. Nasalis (proboscis monkeys) 1. N. larvartus is basically a mangrove species where forests have flooded ii. Simias (simakobu) iii. Pygathrix (Douc langur) 1. One of the most restricted ranges and most endangered populations in Vietnam, Cambodia, Laos iv. Rhinopithecus (Golden langur) only genus with more than one species belonging to it (4) 1. Monsoon or montane forest, well-adapted to cold weather 2) Cercopithicinae

a. Macaca widest geographic distribution of any primate other than humans, 16-22 species, all Asian except for introduced in North Africa i. Omnivores, edge species, have taken up the niches of the guenons and the baboons ii. Species from the same group will not be sympatric with one another, may be sympatric with those from other groups 1. Certain species in each group are considered peripheral because of their isolated ranges, outside of India and Southeast Asia 2. Fooden constructed the distribution map that divides the macaque species by their primary habitat (broad leaf evergreen or secondary, savannah, and edge forest) as well as sub-tropical and tropical regions a. There are a number of different overlapping pairs throughout the distribution 3) Sri Lanka Case Study a. Presbytis senex (Purple-faced Langur) i. More specialized Kasi species, replaced by P. johnii in India ii. Small groups divided into one-male or all-male groups iii. Canopy-dweller, focused on central forests, difficult to observe iv. Two species make up the majority of its diet, and many other species are eaten in very small amounts, if fruit is available, will eat that over mature leaves (which make up the majority of its diet) v. Small home range of only 4 hectares b. Presbytis entellus (Sacred Hanaman Langur) i. Generalized folivore, much more adaptable, highly terrestrial, eat twice as much fruit and less leaves (primarily immature) ii. Live in large groups (up to 20-30 indiviudals), usually either one-male or all-male iii. Eats widely dispersed fruits, has larger home range (25 hectares) c. Macaca sinica i. Omnivorous, though 77 percent of its diet consists of fruits, unlike the other two species, eats a number of animal sources for protein (does not consume enough leaf material to get protein from there) 4) Malaysian Case Study a. Trachypithecus obscura i. Specialized folivore (58%), significantly more leaf material ii. 7 kilograms, lives in mature rainforest, high canopy-dweller, slow in its locomotion iii. Live in one-male or all-male, female is the core of the group, and males migrate iv. Home ranges are smaller, as are day ranges due to concentration on predicatble, concentrated mature leaves

b. Presbytis melalophus i. Generalist folivore (35%) and frugivore (48%) ii. Found in slightly disturbed, secondary, and riverine forest, can be found in all layers of the forest, jumps more often iii. Live in one-male or all-male, female is the core of the group, and males migrate iv. Concentrates on more unpredictable food sources c. Macaca nemastrina i. Broad-leaf specialist in closed-canopy deep forest, highly frugivorous (80%), large home range within this area ii. Tend to be omnivorous (do not have sacculated stomachs for leaves), eat about 10% animal matter iii. Sleep wherever they stop eating, different place each night iv. Large multi-male groups where females form matrilines and males migrate, between 35-50 animals, more cohesive, doesnt have great variability in group size d. Macaca fascicularis i. Savannah-adapted edge species, spend quite a lot of time on the ground, often found in riverine areas ii. Opportunistic feeders with high selectivity depending on what is available in each season, much more diverse diet iii. Refuging species (returns to the same sleeping site every night, central site foragers) iv. Large multi-male groups where females form matrilines and males migrate, between 20-100 depending on environment, within these groups, may divide into subgroups, come together at refuge sites e. Hylobates syndactylus i. Larger body size, includes more leaves and insects in their diets, less fruit f. Hylobates lar T. obscura P. melalophus M. nemestrina M. fascicularis H. lar H. syndactulus Fruits 32 48 80 64 64 45 Flowers 10 14 0 9 0.4 3.6 Leaves 58 35 10 24 31.4 43 Insects 0 0 10 4.5 4.5 8

5) Mauritius Case Study (Macaca fascicularis) a. Off the east coast of Madagascar, not part of the normal distribution of macaques, about 500 years ago, Dutch most likely introduced primates from Java, presumably about 100 individuals before

b. c. d. e. f.

colonization (increased population with human influence), population is now at 40-60 thousands individuals i. Environment is completely different from Java, diet has no species in common with the ancestral population ii. Human population is dense in some areas, and very few forested regions remain, the rest is scrub savannah, highly cultivated (primarily sugar cane) iii. Macaque populations are concentrated to areas where there is forest or savannah (some ideal mosaic areas) Not in normal Asian habitat, though habits are the same, diets differ M. mulatta is located further north the M. fascicularis in their native range (Southeast Asia), highly geographically distributed Extremely adaptable, one of the best models for human evolution, little information on variation in ecology and behavior Captured, collared, tracked for observation, weighed and measured i. Studied in a protected area that would be used for hunting deer during some portions of the year Long-tailed (Crab-eating) Macaques males and females have mustaches and Prussian helmet appearance i. Groups structure is multi-male multi-female, high variation in group size, from less than 10 to more than 100, sex ratio can range from 1:1 to 1:5, matrilineal/matrilocal organization, males migrate, males are also dominant, sub-group fission and fusion during the day (similar to savannah baboon feeding groups) involving females and friends rather than individuals, return to a single sleeping site every night ii. 8 males, 23 females, 5 sub-adult males, 39 juveniles, 10 infants, home range was 117 hectares (overlap with northern group about 55.5 hectares), density of 1.3 individuals/ha iii. Nearest neighbors, females with infants stayed together, females tended to have males near them, juveniles made sure they were with someone (no real preference), males tended not to be with other males, sub-adult males are less social 1. Mutual grooming occurs between females and between females and juveniles, juveniles sometimes groom other juveniles, males rarely groom one another iv. Activity Budget: Feeding and Foraging (29.0%), moving (23.3%), travelling (5.3%), resting (22.0%), auto-grooming (9.2%) v. Three feeding peaks throughout the day with resting peaks in between, similar to other tropical mammals vi. Move more during March and April when resources are scarce vii. In most of SE Asia, especially where pig-tailed macaques are sympatric, almost exclusively arboreal, when they are not sympatric, readily exploit terrestrial habitats (close to 80%) over the year, about 50% of feeding time on ground

Asian Apes i. Gibbons and Siamangs a. Only primates that are true brachiators, swing beneath branches, hook-like hands, reduced thumb, long arms, long fingers, no tail, upright in posture (orthograde), walk with hands up in the air b. Some of the only mammals that are true monogamists, true territoriality where they defend a specific range c. Dense rainforest and monsoon forest, completely arboreal in upper canopy and emergent layer, fastest flightless animals in the trees d. Social Structure i. Average group size is 4 individual with the range from 2-6, pair is the most common interaction, though there are variations from this structure

viii. Sleep every night at the base of the mountain, refuging is fairly rare in primates ix. Predicted diet would be 72.5% fruit and flowers (few species, change seasonally), 24% leaves, and 4.4% animal prey, based on dentition, believe there is a species-specific dietary pattern, physiologically adapted 1. About 39% fruit, 8% flowers, 23% leaves, insects were around 3%, about 5% of sugar cane, grasses, stems, molasses, and other (approximately as expected) 2. Very seasonal, fruits would decrease and leaves/grasses would increase depending on availability (adaptable) 3. Pod-specialists, hard pods with seeds in the middle, seem to concentrate more on these than fleshy fruit 4. 15-30 plants make up their entire diets, though fewer than five tend to make up over 60% 5. Had a fairly balanced diet between carbohydrates, lipids, and protein (from leaves and insects), highly nutritious 6. In natural forests, diets tend to be quite similar, even in completely different environments (same structure, different content) x. Human Interactions/Conservation 1. Some people treat them as sacred animals (Hindu) a. Actively feed them to maintain the population, leave offerings for monkeys 2. Some people view them as pests (Christians) a. Feature them in their church dinners (Curri de Jacot, curri no. 2) 3. Minor crop raiders (especially sugar cane) 4. Extinctions of endemic species have been blamed on macaques, though it was probably because of humans 5. Export of animals for medical research (second largest exporter to US after China)

ii. Lifespan of 20-30 years, may have extra-group mating, 9% of matings were outside of the group iii. 10% of siamang groups had an extra adult, some had additional female with infant iv. About 1 in 6-7 pairs was a new group forming v. Both males and females care for infants e. Territoriality i. Must have enough resources in the home range, tighter relationship between the number of members in the group and the home range areas, must be able to cover the entire area to maintain borders of the homer range ii. Long (loud) calls are used for spacing, but not when they are interacting (0.2 - 5 vocalizations per day), displays between groups occur between 1-5 days (average about 37 minutes), never make contact with opposing individuals, energy expensive (males move around, females call, groom males) 1. Can tell the species and sex of the animal, may also give indication of the health of the caller 2. Helps form pair bonds as well as acting as a spacing mechanism iii. Group Formation 1. Animals leave at about 6-8 years of age, nearly full size, must develop exclusive area to attract mate a. Male will establish territory by extending parents territory (with their help) and taking control of new portion b. Male or female may replace a dead partner, may even be a younger animal from that group (due to extra-group mating c. Males may coalesce to create territory, attract female together, whichever gets the female will gain exclusive use of range, other will leave to form new territory f. Hylobates (11 species) most are allopatric, replace each other, though there are some small areas of overlap where hybrids exist i. 5-7.5 kg, found throughout Asia ii. Distinct color patters, territorial calls (duets), geographical distributions between the 11 species iii. C.R. Carpenter (studied Howler monkeys first) was one of the first to study them in 1937 in Thailand iv. Robert van Gulik was a statesman (ambassador to China and Japan), essay on the gibbon in Chinese folklore, determined the historical range from folklore (throughout China, but human population has caused in to shrink to current southern range) v. Consume 60-70% fruit (20-30% of which is figs), 30-40% leaves, 2-4% insects, some other supplements

1. Scattered resources that are not in great quantities vi. Sleep in the canopy and emergent layer, tend to be spread out, female is often apart from the male (up to 100 meters away) 1. Female was separated from male by 10 meters or more over 50% of the time, sometimes as far apart as 30 meters during activity g. Symphalangus (S. sindactylus) siamangs, sympatric with gibbon species with which they overlap, do not hybridize i. 10-12 kg, only found in Sumatra and Malaysia ii. Eat about 30-50% (70% of fruit diet is figs), 40-65% leaves, approximately10% insects 1. Use clumped resources, massed figs and leaves iii. Sleep in the canopy and emergent layer, tend to be cohesive, male and female will sleep in the same tree 1. Will call when they wake up to separate from other groups, alternate between feeding and resting throughout the day 2. Will go to their sleeping sites fairly early, about 2-4 hours before dusk, female usually returns before males 3. Whole group will be within 10 meters of each other 50% of the time during waking hours, and 75% of the time they are doing the same activity, 90% of the time, all members of the group were visible at the same time iv. Predation is not a major threat, pythons and large raptors may eat young, felids may eat adults 1. Primary threats are humans, war v. After the first 8-10 months, when female begins weaning, male will carry the infant until it is up to three years old, longer care period than in gibbons ii. Orangutan (Pongo) a. Only found in Sumatra (only 7500 in northwest of island) and throughout Borneo, historically found throughout Indonesia, deforestation, commercial timbering are primary threats b. Distribution of body hair, color, facial structure, chromosome number, c. Males are about 180 lbs and females are only 80, most dimorphic of all primates, males also have throat sack used for loud calls i. Arm-leg length is highest among all apes, hands, fingers, and toes are slender, reduced thumb and big toe for arboreal suspensory locomotion ii. Locomotion is slow and laborious (swing branches to reach the next one, no jumping), often come to the ground to travel (primarily males) d. Primary forest dwellers, avoid humans, deepest forest fragments, some lowland, montane, and swamp forest, also diptorocarp forest (consisting of only one genus of plants, many species, but no dominant ones, four distinct layers)

e. Mainly frugivores (63%), but do include some leaves (23%), bark (11%), flowers (2%) and insects (1%) i. Social insects (mainly termites) like other large primates, eaten primarily by juveniles and females, not males ii. Most feeding is done in the high canopy, eat about 7 species per day, but focus on a few at a time (depending on season) f. Galdikas found that they depend on three sources of fruits, large trees with large crops of highly preferred fruits (seasonal, but abundant), scattered trees which were common but not very abundant fruit, or scattered stands with many fruits (only times when they come together) i. Overall, rarely occurring and not abundant resources, thus, they require a large home range ii. Able to eat fruit before it is ripe, while it is still hard because of their size (other primates cannot eat these), also consume thick, hard, and spiny fruits, can detoxify some species of fruit that other species cannot eat iii. Leaves are eaten very selectively, usually mature leaves, barks make up a substantial portion of their diet, chew on wedges to obtain carbohydrates (smaller primates cannot do this) g. Act as pruners that led to more branching and as seed dispersers for over 70% of the species they consume h. Active for most of the day (11.5 hours of the day), feeding, travelling make up the majority of this time i. Have two peaks of feeding with an extended rest period i. Build nests for sleeping, watch mothers to learn skills required to construct, also create umbrellas to protect from rain, provide shade, avoid detection (camouflage), and play behavior i. Tigers and hunting dogs are the only real predators, clouded leopards are probably too small, crocodiles are extinct in most areas of orangutan density, but traditional predators j. Social Structure only diurnal primate that lives in solitary, but social organization, much like prosimians i. 1.5-1.8 individuals (solitary or mother with infant) ii. Regular pattern of interactions, usually move alone iii. Female home ranges are stable over time (5-8 km), there is extensive overlap between females, with very few areas of exclusive use, male ranges are much larger and overlap with other males very little, but with females a great deal 1. Some males have stable home ranges (resident males) 2. Some males home ranges are much more flexible, usually larger than resident, roam more often, sometimes even resident males that will use this method 3. Level of overlap between males is dependent on resource availability

4. Residence is not equivalent to dominance iv. Rijksen found that adult females were seen to interact extensively and lasted a long time 18 times over three year study (probably based on friendship rather than kinship) 1. 30 times, adolescents interacted with females (often times to interact with their infants) 2. 63 times, sub-adult males interacted with one another, more sociable than adults 3. Males and sub-adult males avoid other adult males, often have an agonistic interaction, exchange loud calls about twice a day as distance-maintaining or increasing, can be heard up to a mile apart 4. Receptive females will decrease distance upon hearing a loud call, if not receptive, will increase distance v. Mating 1. When a female is not receptive to a male, an agonistic interaction may occur (often involves sub-adult males), may be perceived as dominance interaction 2. Consortship initiated by female almost always with fully adult male, male will alter movement pattern to move with female for extended period, may consort with one male for a short period and then switch to another, or may stay with him for a long time vi. Ranging Behavior 1. Females spend most of their time in the trees, sometimes with their infants, day range is about 700 meters (rarely over 2.5 km in a single day) 2. Males will move in the trees if branches can support them, but will often come to the ground to travel, may move 4 km a day (2500 ha home range) 3. Either move slowly within a relatively small area or quickly in a straight-line manner towards a new food source 4. Availability of fruit and social interactions determine ranging behavior (females may move towards other females, adolescents spend time with other individuals, males move away from other males or towards estrous females, females with infants avoid contact) k. Pongo pygmaeus Bornean species l. Pongo abelii Sumatran species African Apes 1) Gorillas a. Gorilla g. gorilla western lowland Africa b. Gorilla g. graueri eastern lowland Africa c. Gorilla g. berengei mountainous regions d. Males are 300-400 pounds, males are about 200 pounds

e. All subspecies are endangered, difficult to study i. Robert Yerkes (psychologist who did comparative intelligence) sent out two researchers to study gorillas and chimpanzees, though they did not last long in the field ii. Schaller and Fossey studied mountain gorillas in the 1960s and 1970s, some studies continue to the present iii. First to study lowland gorillas was Fay in the 1990s f. Habitat and Locomotion i. Extremely variable habitats, though primarily in highly herbaceous areas, density is tied to terrestrial herbaceous vegetation ii. Found in primary, secondary, edge, bais (swamp forest) iii. 80% of their locomotion is terrestrial, though the often rest and keep watch from trees 1. Very agile in the trees, though they are often too heavy, females and juveniles go up more often 2. Knuckle-walkers g. Diet i. Adapted for folivorous diet with an enlarged hindgut and grinding dentition ii. Use mainly perennially available herbs and vines, actually help perpetuate this layer of the forest, limits the growth of larger trees by sitting on the ground and rolling when they rest, this destroys any nearby saplings iii. 3-4 species may make up 60% of their diet (most research has been done on mountain gorillas) iv. Darcy found that bed straw (up to 30 km a day, contains 25 liters of water), thistle, wild celery, and bamboo shoots (terrestrial vegetation) made up 95% of the diet in mountain gorilla, also may supplement with soil v. 85-90% is leaves, shoots, and stems, but may supplement with bark, pith, flowers, and fruits, though this is very different in the lowland gorilla species where they will eat 30-40% fruit vi. Little or no insect material (on purpose) h. Activity Budget i. Spend most of their time feeding, long rest in the middle of the day ii. Build nests in the evening, usually on the ground, may move up to 3 meters into the trees if proper support is available iii. Up to 3 animals in the mothers nest, though at 1.5 years of age, individuals will begin building their own nests i. Humans were throught to be the only predators of gorillas, though apparently leopards will eat j. Social Organization i. Small (up to 42 individuals, though average is only 16.9 individuals), cohesive, stable, multi-male multi-female group

1. 1.7 silverback males, 1.5 black back males, 6.2 female, 2.9 juveniles, 4.6 infants 2. Females carry infants for 3-4 years, but there is high reproductive rate ii. Core of the group is the silverback male, dominant animal, center of the groups attention, females with infants interact with each other more, males interact with females without infants more 1. Young females leave the group (and sometimes males) at about 6 years old when they reach sub-adulthood, usually do not reproduce until 9 years of age, may continue migrating until they have their first infant 2. All-male groups until males leave to form their own group with migrating female 3. Some males follow groups with older dominant males waiting to usurp power iii. Group will move about 1 km a day, home range is about 30 square kilometers 1. Females are tolerant to one another (not kin-related), but spend very little time interacting iv. In bais, different groups were found to interact with each other with differing levels of agonism, probably depending on previous interactions 1. Males are not very tolerant of single males or all-male groups (presumably trying to usurp females) 2) Chimpanzees a. Pan troglodytes (common chimpanzee) widest distribution of any age (across equatorial Africa, 4 allopatric subspecies) i. Male weigh 140 lbs and females weigh 100 lbs ii. Sympatric with gorillas throughout much of their range iii. Garner studied them in 1896, animal collector for a zoo 1. First to use recording devices in the wild iv. Habitat and Locomotion 1. Live in all types of habitat, debate as to whether they are more adapted to savannah or forest 2. Bimodal activity cycle, like most other tropical mammals, though males feed and move more than females throughout the day 3. Build nests every night, animals up to 5 or 6 years will nest with mothers, up to 5 to 20 meters up v. Diet 1. Highly omnivorous, fruit is the focus of their diet, insects and small vertebrates make up a portion of their diet as well, use manufactured tools 2. Rare fruit specialists, patches, 60-80% fruit

3. One of the most diverse diets of any primate, require a large home range to acquire patchy resources, probably a major limiting factor a. Supplement fruit diet with buds, flowers, seeds, and some animal protein (social insects, meat) b. Only 200 reports of kills during first 25 years of observation, 80% were at Gombe, most of these were of baboons, fighting over provisioned bananas c. Since then, about 400 kills have been reported, 250 of these were of red colobus monkeys d. At Gombe, 64% of vertebrate diet, but with increased human influence have increased to 87 e. At Mahale, a similar increase in portion of the diet made up by colobus from 14% to 56% to 83% today, not a natural phenomenon 4. Females and juveniles tend to be the ones who consume the insects, they are also the ones using the tools a. Termite Searching break sticks and twigs, lick them so that they are sticky, lick termites off i. Different ways in different populations, for different termite species b. Leaves for collecting water c. Collect rocks and anvils to open nuts (females are the only ones who break the hardest nuts), only found in some forest in Tai (cultural) i. Females will teach young how to do this, males do not learn as quickly 5. Males are the ones hunting vertebrates, males are larger, spend more time on the ground where vertebrates are more easily caught, males are more active and take more chances (not pregnant or caring for a baby), all similar to Cebus male-biased hunting 6. Food sharing between females and infants (especially in provisioning), meat is also shared because it is usually too big for one animal a. In other mammals and birds, females share with offspring, older will share with younger, and some males with females b. In chimp, 80% of sharing was between close kin c. Meat can also be used as a means of coersion vi. Predation 1. High predation rate (about 4%), primarily by leopards 2. As in all great apes, human-induced deforestation and habitat destruction vii. Social Organization

1. Communities of 50-100 individuals, interact in a fissionfusion manner within, there are no stable sub-groups other than mother and infant 2. In 498 interactions, 13% were solitary, 69% were between 2-6 animals, 9 or fewer animals seen 91% of the time, and 20 or more were seen 4 times 3. In 667 associations, average was 2.6 animals (range of 1-24), mothers with infants tend to be alone (over 50%) 4. Savannah chimps tend to be found in larger groups 5. Distribution of fruit resources (if spread, small groups) and the number of females in estrous (will be surrounded by more than average) determine the size of sub group size 6. Social networks seem to develop, particular associations with friends are seen more often 7. Males have stronger ties to one another (46% of grooming between males), females migrate out of their natal groups 8. Female choice is the major factor in who mates with whom, male dominance is not as present as perceived a. Only two real mating studies i. Promiscuous males and females have between 1 and 14 pairings ii. Possessive male exerts dominance iii. Consort male and female go off together, must chose to maintain consort iv. Most are promiscuous, dominance plays very little role viii. Warfare and Male Killing 1. In over 200 years of collective observation at 9 different sites, there have been only 10 cases of males killing other males, 10 other cases of mysterious disappearances (probably leopards) 2. Gombe studied since 1962, nothing happened there until 1974, two groups interacted in an aggressive way, competing over provisioned bananas a. Amount of aggression increased from 0.2 per day to 5 per day, wanted to close the site b. The site has been decreased from 30 square miles (connected with contiguous forest on all sides), now only 13 square miles, 150 chimps to 100, so density has increased, non-contiguous c. Some poaching and a lot of disease present due to human influence (development) 3. All sites except Goualougo have experienced a great deal of human influence, population encroachment (16

times increase at Tai), snare injuries at high rates (255 of individuals at Budongo), chimp populations are decreasing as well a. Bossou is so highly influenced that only male remains, move as one group rather that fissionfusion sub groups ix. Language and Culture 1. No chimpanzee teaches its offspring language in the same way we do, must re-label human language if we call what they do language 2. There are differences between populations throughout their range, differences do not have serious effects on their lives, very unlike human cultural differences 3. Live in the present, cannot express symbolically anything not currently present (future, past, absent individual) x. Many of the unusual characteristics that have been reported do not seem to occur at undisturbed Goualougo nor at Goodalls site during the first 20 or so years, all influenced highly by humans b. Pan paniscus (bonobo, pygmy chimpanzee) found only in a small region on the northern bank of the Zaire river (restricted range) i. Males are about 100 lbs and females are 70 lbs ii. Not sympatric with gorillas iii. Habitat and Locomotion 1. Diverse habitat, but mostly primary forest 2. Seem to be more arboreally adapted than the chimpanzee, though both knuckle-walk iv. Diet 1. 80% fruit