1
Feeding Ecology, Digestive Strategies, and Implications for Feeding Programs in Captivity

The welfare of nonhuman primates in captivity depends heavily on meeting their nutrient needs in a manner that considers normal foraging and feeding behavior, structure and functions of the digestive system, and the options and constraints of captive dietary husbandry.

FEEDING ECOLOGY

In developing a system for the nourishment of captive nonhuman primates, it is helpful to examine the literature on the feeding ecology of primates in the wild. Several observational methods have been used to record foraging and feeding behavior in natural ecologic systems (Altmann, 1974; Lehner, 1996), and data derived with these methods are summarized in Tables 1-1 through 1-6. To interpret the findings properly, the reader should have a background in the methods used, and a brief discussion of them follows.

Feeding-Ecology Methods Involving Visual Observations of Behavior

Data collected during visual observations of behavior typically include length of feeding bout, plant species eaten, plant parts eaten (for example, fruit and leaf), percentage of part eaten, feeding rate (for example, number of fruits consumed per minute), diameter and height of food plant, and food-plant location.

OBSERVATION OPTIONS

Choosing a data-collection method requires, as a first step, selection of one of two animal-observation options.

Focal-Animal Observation One individual is observed during a given session of data collection (it can also be a pair or a small subgroup). Sessions can vary from 5 min to a whole day. This method is used to identify multiple behaviors in selected individuals. When sessions are only 5-10 min long, it is common to switch observations to another animal in the group for the next session.

All-Animal Observation Primates that are naturally grouped are observed simultaneously. This method is feasible only when observing a few easy-to-see behaviors. It is not recommended for detailed feeding behaviors.

SAMPLING METHODS

After selection of an animal-observation option, the second step is to select a method of sampling foraging and feeding behavior.

Ad Libitum (or Periodic) Sampling This is the classic, pre-1970s field method, used before modern statistical techniques and advanced technologies were commonly applied. Today, it is recommended only for preliminary reconnaissance or the study of rare behaviors. This method is biased toward spectacular behaviors, like hunting, thus overestimating faunivory compared with herbivory.

Continuous-Recording Sampling Method These sampling methods result in the most complete and accurate data. They are recommended for studying feeding ecology but are difficult to use with arboreal animals, such as primates.

  • All-Occurrences Sampling. All occurrences of one or a few behaviors are recorded over an extended period,



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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 1 Feeding Ecology, Digestive Strategies, and Implications for Feeding Programs in Captivity The welfare of nonhuman primates in captivity depends heavily on meeting their nutrient needs in a manner that considers normal foraging and feeding behavior, structure and functions of the digestive system, and the options and constraints of captive dietary husbandry. FEEDING ECOLOGY In developing a system for the nourishment of captive nonhuman primates, it is helpful to examine the literature on the feeding ecology of primates in the wild. Several observational methods have been used to record foraging and feeding behavior in natural ecologic systems (Altmann, 1974; Lehner, 1996), and data derived with these methods are summarized in Tables 1-1 through 1-6. To interpret the findings properly, the reader should have a background in the methods used, and a brief discussion of them follows. Feeding-Ecology Methods Involving Visual Observations of Behavior Data collected during visual observations of behavior typically include length of feeding bout, plant species eaten, plant parts eaten (for example, fruit and leaf), percentage of part eaten, feeding rate (for example, number of fruits consumed per minute), diameter and height of food plant, and food-plant location. OBSERVATION OPTIONS Choosing a data-collection method requires, as a first step, selection of one of two animal-observation options. Focal-Animal Observation One individual is observed during a given session of data collection (it can also be a pair or a small subgroup). Sessions can vary from 5 min to a whole day. This method is used to identify multiple behaviors in selected individuals. When sessions are only 5-10 min long, it is common to switch observations to another animal in the group for the next session. All-Animal Observation Primates that are naturally grouped are observed simultaneously. This method is feasible only when observing a few easy-to-see behaviors. It is not recommended for detailed feeding behaviors. SAMPLING METHODS After selection of an animal-observation option, the second step is to select a method of sampling foraging and feeding behavior. Ad Libitum (or Periodic) Sampling This is the classic, pre-1970s field method, used before modern statistical techniques and advanced technologies were commonly applied. Today, it is recommended only for preliminary reconnaissance or the study of rare behaviors. This method is biased toward spectacular behaviors, like hunting, thus overestimating faunivory compared with herbivory. Continuous-Recording Sampling Method These sampling methods result in the most complete and accurate data. They are recommended for studying feeding ecology but are difficult to use with arboreal animals, such as primates. All-Occurrences Sampling. All occurrences of one or a few behaviors are recorded over an extended period,

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 1-1 Prosimian Feeding Ecology Scientific Name Common Name Dieta Behavior Body Weightb References 100% insectivorous Tarsius T. bancanus T. dianae T. pumilus T. spectrum T. syrichta Western tarsier Dian’s tarsier Pygmy tarsier Spectral tarsier Philippine tarsier Animal prey 100%; T. bancanus example: beetles 35%, ants 21%, locusts 16%, cicadas 10%, cockroaches 8%, vertebrates 11% of feeding time (not seen eaten by all Tarsius); also eaten: crickets, mantids, moths Nocturnal, arboreal, solitary or pairs or multimale/ multifemale, group size 2-6 individuals 77.6-117 g females, 27.5-134 g males Crompton & Andah, 1986; Fogden, 1974; Gursky, 1996; Kappeler, 1991; MacKinnon & Mackinnon, 1980a; Niemitz, 1984; Nietsch & Niemitz, 1991; Tremble et al., 1993 Mostly insectivorous Allocebus A. trichotis Hairy-eared dwarf lemur In wild, unknown; in captivity, insects 70%, sweetened rice broth, fruit Nocturnal, arboreal, forage solitary or male/female pair, sleep 2-6 78-90 g females, 75-98 g males Albignac et al., 1991; Kappeler, 1991; Meier & Albignac, 1991; Mittermeier et al., 1994 Arctocebus A. aureus A. calabarensis Golden angwantibo Angwantibo Animal prey 79% (73-85%), fruit 13% (12-18%), other vegetation 8%; prey: caterpillars 77% (65-90%) crickets, beetles, ants Nocturnal, arboreal, forage solitary, sleep 1-2 A. aureus 150-270 g; A. calabarensis 200-465 g Bearder, 1987; Charles-Dominique, 1974; Charles-Dominique & Bearder, 1979; Gonzalez-Kirchner, 1995; Silva & Downing, 1995; Wolfheim, 1983 Galagoides G. demidoff G. thomasic G. zanzibaricus Demidoff’s bush baby Thomas’s bush baby Zanzibar bush baby Animal prey 75% (70-81%), fruit 17% (4-30%), gums/ resins 5% (0-18%), leaves, buds; prey:moths, beetles, grasshoppers, ants, some birds Nocturnal, arboreal (mostly), forage solitary, sleep (females) 1-10 G.demidoff 46-69 g females, 78-85 g males; G. thomasi 55-149 g; G. zanzibaricus 118-155 g females, 130-183 g males Charles-Dominique, 1974; Gonzalez-Kirchner, 1995; Harcourt & Bearder, 1989; Harcourt & Nash, 1986; Hladik, 1979; Kappeler, 1991; Nash et al., 1989; Nash & Harcourt, 1986; Silva & Downing, 1995 Loris L. tardigradus Slender loris Almost exclusively insects, small amount of young leaves, shoots, hard-rind fruits, flowers, eggs, small vertebrates; often insects strong smelling Nocturnal, arboreal, forage solitary, sleep 2-4 102-322 g Butynski, 1982; Petter & Hladik, 1970; Silva & Downing, 1995; Wolfheim, 1983 Omnivorous, gums dominate Euoticus E. elegantulus (Galago elegantulus) E. pallidusc Southern needle-clawed bush baby Northern needle-clawed bush baby Gums 55% (35-75%), animal prey 32% (20-44%), fruit 12% (5-20%), birds Nocturnal, arboreal, forage solitary, sleep 1-7 271 g female, 270-360 g males Butynski, 1982; Charles-Dominique, 1974, 1977; Charles-Dominique & Bearder, 1979; Gonzalez-Kirchner, 1995; Hladik, 1979; Kappeler 1991 Galago G. senegalensis G. moholi Northern lesser bush baby Southern lesser bush baby Gums (Acacia) 48%, animal prey 52% (butterflies, moths, beetles), gums from 2 tree species, no vertebrate prey Nocturnal, arboreal, forage solitary, sleep 1-3 G. senegalensis 126-193 g females, 125-212 g males; G. moholi 140-229 g females, 160-255 g males Bearder, 1987; Bearder & Doyle, 1974; Bearder & Martin, 1979; Doyle, 1979; Doyle & Bearder, 1977; Harcourt & Bearder, 1989; Nash & Whitten, 1989; Silva & Downing, 1995 Otolemur O. crassicaudatus (Galago crassicaudatus) Thick-tailed greater bush baby Gums 44% (18-62%), fruit 27% (21-33%), animal prey 14% (1-27%) (invertbrates and vertebrates), nectar 4% (0-8%), seeds 3% (0-7%), misc. vegetable matter 8% (0-16%) Nocturnal, arboreal, male solitary, female and offspring forage together, sleep 1-6 1122-1497 g females, 1126-1750 g males Bearder & Doyle, 1974; Butynski, 1982; Doyle & Bearder, 1977; Kappeler, 1991; Masters et al., 1988 Phaner P. furcifer Fork-marked lemur Tree gum (resins) bulk of diet, some fruit, sap, animal matter 10%, flowers, buds, nectar, secretions of Homoptera insects Nocturnal, arboreal, solitary or male/ female pairs, sleep 1-4 350-600 g Charles-Dominique & Petter, 1980; Hladik, 1979; Hladik et al., 1980; Kappeler, 1991; Pariente, 1979; Petter et al., 1971, 1975

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Omnivorous, plants (especially fruits) dominate Cheirogaleus C. major C. medius Greater dwarf lemur Fat-tailed dwarf lemur Fruit, young leaves, flowers, nectar, leaf buds, gums, animal prey (mostly insects, some chameleons); C. major lethargic in cool, dry season; C. medius increase in body weight 120-250 g during rainy season (6 months), hibernate 7-9 months Nocturnal, arboreal, forage solitary, sleep 1-5 C. major 235-470 g; C. medius when feeding (rainy season) 142-217 g, especially tail, hibernate loses 100 g Hladik, 1979; Hladik et al., 1980; Mittermeier et al., 1994; Petter et al., 1977; Wright & Martin, 1995 Eulemur (Petterus) E. coronatus E. fulvus E. macaco E. mongoz E. rubriventer Crowned lemur Brown lemur Black lemur Mongoose lemur Red-bellied lemur Fruit 45% (7-79%), leaves 45% (20-89%), flowers 10% (1-52%), few insects; dry season diet of E. mongoz, E. rubriventer, E.fulvus in some habitats nectar 82% (81-84), fruit 17%, leaf 1%; E. fulvus tolerates high levels of toxic plant compounds Cathemeral, mostly arboreal, multimale/ multifemale groups, group size 5-18 or just family groups 1.4-2.4 kg Andriatsarafara, 1988; Colquhoum, 1993; Dague & Petter, 1988; Hladik, 1979; Kappeler, 1991; Overdorff, 1993; Richard & Dewar, 1991; Silva & Downing, 1995; Sussman, 1974, 1977; Sussman & Tattersall, 1976; Tattersall, 1977, 1979; Wilson et al., 1989; Yamashita 1996 Galago G. alleni G. gallarum G. matschiei Allen’s bush baby Somali bush baby Matschie’s bush baby (was Euoticus inustus) Fruit 74% (73-76%), animal prey 24% (23-25%) (invertebrates and frogs), 2% other vegetation (fallen fruit, seeds, gums) Nocturnal, arboreal, forage solitary 86%, sleep (females) 1-4 G. alleni 200-445 g; G. gallarum 196-225 g Butynski, 1982; Charles-Dominique, 1977; Gonzalez-Kirchner, 1995; Nash et al., 1989 Lemur L. catta Ring-tailed lemur Fruit 54% (34-70%), leaves 33% (24-50%), flowers 3% (0-8%), herbs 8% (6-15%), bark, sap, cactus, misc 2% (0-7%); Tamarindus indicus is 25% of diet: 12% leaf, 12% pods Diurnal, arboreal, terrestrial, multimale/ multifemale, with 1 alpha female, group size 5-30 1.96-2.705 kg Jolly, 1966; Kappeler, 1991; Rasamimanana & Rafidinarivo, 1993; Sauther & Sussman, 1993; Silva & Downing, 1995; Sussman, 1974; Yamashita, 1996 Microcebus M. (Mirza) coquereli M. murinus M. myoxinusc M. rufus Coquerel’s dwarf lemur Gray mouse lemur Pygmy mouse lemur Brown mouse lemur Fruit, animal matter (insects, frogs, bird eggs, chameleons), young leaves, flowers, gums, sap/resins, nectar, buds, seeds; spends up to 50%of time in dry season licking larval secretions of Homoptera off branches Nocturnal, arboreal mostly, forage solitary, some pairs, sleep 1-4; M. murinus store fat in tail and less active in dry season, do not hibernate, sleep 1-15 M. coquereli, M. myoxinus 302 g female, 308 g male; M. rufus 41-63 g females, 35-70 g males; M. murinus 40-109 g varies 50-60 g when “hibernates” Corbin & Schmid, 1995; Hladik, 1979; Kappeler, 1991; Pages, 1980; Petter et al., 1971, 1977; Wright & Martin, 1995 Nycticebus N. coucang N. pygmaeusc Slow loris Pygmy loris Fruit 50%, animal prey 30%, gums 15% (10-19%), shoots, bird eggs, insects that have repugnant taste and smell Nocturnal, arboreal, forage solitary N. coucang 375-900 g female, 850-1207 g male: N. pygmaeus 372 g female, 462 g male Bearder, 1987; Duckworth, 1994; Kappeler, 1991; Silva & Downing, 1995; Tan, 1994; Van Horn & Eaton, 1979 Otolemur O.garnettii Garnett’s greater bush baby Fruit 27% (4-50%), animal prey 61% (44-78%) (beetles, ants, termites, snails, birds), seeds 3% (0-7%), other vegetation 9% (0-18%) (resins, bark, pollen) Nocturnal, arboreal, male solitary, related females overlap 740-1460 g female, 822-1640 g male Bearder, 1987; Harcourt & Nash, 1986; Masters et al., 1988; Nash & Harcourt, 1986; Nash et al., 1989; Silva & Downing, 1995 Perodicticus P. potto Potto Fruit 74% (65-82%), gums 40% (21-60%), animal prey 20% (10-30%) (ants make up 65%of insect prey), some leaf and fungus; when fruit is scarce (dry season) Nocturnal, arboreal, forage solitary 96%, pairs 4%, sleep 1-2 850-1600 g Charles-Dominique, 1974; Gonzalez-Kirchner, 1995; Hladik, 1979; Oates, 1984 Varecia V. variegata Ruffed lemur Ripe fruit 74%, 21% leaves (2% shoots, 1% young leaves, 18% mature leaf), flowers 5% (1-40%), seeds, nectar; 74% nectarivorous in 1 month of year Diurnal, arboreal mostly, family or larger groups, 5-32 individuals 3.512 kg female, 3.471 kg male Dew & Wright, 1994; Kappeler & Ganzhorn, 1993; Morland, 1993; Richard & Dewar, 1991; Rigamonti, 1993; White, 1989

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Leavesdominate other plant parts Avahi A. laniger Woolly lemur Seasonally exclusively folivorous: 91% leaves (40% mature leaves, 51% mixed mature and young), 9% flowers, rarely fruit and bark Nocturnal, arboreal, monogamous pairs, groups 2-5 individuals 1.3 kg female, 1.0 kg male Albignac, 1981; Ganzhorn, 1988; Ganzhorn et al., 1985; Harcourt, 1991; Kappeler, 1991; Richard & Dewar, 1991 Indri I. indri Indri (babakoto) Young leaves, buds, and petioles 45% (1-75%), fruit 38% (5-75%), unripe seeds 12% (10-15%), flowers and buds 3% (1-6%), mature leaves 2% (0-3%), occasionally soil Diurnal, arboreal, monogamous family groups 2-6 individuals 7.1 kg female, 5.8 kg male Hladik, 1979; Mittermeier et al., 1994; Pollock, 1975, 1977 Lepilemur L. dorsalis L. edwardsi L. leucopus L. microdonc L. mustelinus L. ruficaudatus L. septentrionalis Gray-backed sportive lemur Milne-Edwards’ sportive lemur White-footed sportive lemur Small-toothed sportive lemur Weasel sportive lemur Red-tailed sportive lemur Northern sportive lemur Leaves primarily, some fruit, bark, seeds, flowers; L. mustelinus can tolerate high alkaloid levels; L. ruficaudatus may practice caecotrophy and have high tolerance for toxins; L. leucopus 100% leaves Nocturnal, arboreal, solitary or male/ female pairs, sleep 1-3; do not hibernate 544-915 g, L. edwardsi 1000g Albignac, 1981; Charles-Dominique & Hladik, 1971; Ganzhorn, 1988; Hladik, 1979; Hladik & Charles-Dominique, 1974; Hladik et al., 1980; Kappeler, 1990, 1991; Kappeler & Ganzhorn, 1993; Nash, 1994; Silva & Downing, 1995 Propithecus P. diadema P. tattersalli P. verreauxi Diademed sifaka Golden-crowned sifaka Verreaux’s sifaka P. diadema and P. tattersalli: young leaves 25% (5-44%), mature leaves 25% (0-46%), fruit, ripe or unripe 43% (0-72%), flowers 7% (0-23%); P. verreauxi: mature leaves 38% (2-70%), young leaves 40% (0-70%), fruit 7% (5-8%), flowers 10% (0-40%), bark 5% (4-9%) Diurnal, mostly arboreal, pairs to multimale/ multifemale groups, 2-12 individuals P. diadema: 5.6-7.2 kg, P. tattersalli: 2.1-3.8 kg, P. verreauxi: 3.5-3.6 kg Hemingway, 1998; Hladik, 1979; Jolly, 1966; Kappeler, 1991; Meyers & Wright, 1993; Richard, 1974, 1977, 1978; Yamashita, 1996 Mostly bamboo Hapalemur H. aureus H. griseus H. simus Golden bamboo lemur Lesser bamboo lemur Greater bamboo lemur Bamboo 95% (85-98%) (shoots 89%, mature leaves 6%, young leaves 1%, petioles 1%), flowers 1%, fruit 2%, fungus 2%; H. griseus also eats phragmites leaves and shoots, Papyrus pith; H. aureus eats a bamboo containing 12 × lethal dose (for humans) of cyanide Diurnal or cathemeral, arboreal, family 2-6 individuals; H. simus 1 male + multifemale or multimale/ multifemale groups 4-30 individuals H. aureus 1.5 kg female, 1.7 kg male; H. griseus 800-939 g; H. simus 1.3-2.4 kg Glander et al., 1989; Kappeler, 1990; Meier & Rumpler, 1987; Overdorff et al., 1997; Petter & Peyrieras, 1970a; Petter et al., 1975, 1977; Silva & Downing, 1995; Wright, 1986; Wright & Randrimanantena, 1989; Wright et al., 1987 Wasthought of asinsectivorousbut isomnivorous-frugivorous Daubentonia D. madagascariensis Aye-aye Seeds/nuts 47% (12-84%), nectar 8% (1-20%), larvae 20% (2-45%), canker 20% (5-42%), other (soft fruit, fungus, galls, bamboo) 5% (0-12%); larvae extracted with long thin finger; eat coconuts (0-58% where available) same way. Nocturnal, arboreal, forage solitary, sleep 1-2 2.6 kg female, 2.8 kg male Ancrenaz et al., 1994; Andriamasimanana, 1994; Erickson, 1995; Iwano & Iwakawa, 1985; Kappeler, 1991; Petter & Peyrieras, 1970b; Pollock et al.,1985; Sterling, 1994; Sterling et al., 1993 aDiet format: mean (range). bBody weights in ranges whenever possible; single numbers are not averages but indicate that only one individual of the species has been weighed in the wild. cNo data available from the wild but assumed to be similar to congenerics. often 1 day. Usually combined with focal-animal sampling, this is an excellent but difficult method for recording foraging and feeding behavior. Start-and-stop rules, independent of the behavior being studied, are required. Sequence Sampling. A sampling period starts with the beginning of a sequence of a chain of behaviors, such as foraging for insects and feeding. The sampling period ends when the observed sequence ends. This method is of lim-

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 1-2 Callitrhix Feeding Ecology Scientific Name   Dieta Behavior Body Weightb References Fruit and insect foraging dominate diet, gums seasonally important Callithrix C. argentata C. aurita C. geoffroyi C. humeralifer C. kuhlii C. mauesic C. nigriceps Bare-ear marmoset Buffy tufted-eared marmoset Geoffroy’s tufted-eared marmoset Tassel-eared marmoset Wied’s tufted-eared marmoset Maues marmoset Black-headed marmoset 27% (24-30%) of total daily activity foraging for insects; therefore, total feeding time spent on insect foraging 56% (50-63%), fruit 33% (28-37%), exudates (gums) 11% (5-16%); when fruit scarce, exudate intake increased Diurnal, arboreal mostly, multimale/ multifemale group size 3-20 individuals 190-320 g females, 357-450 g males; C. nigriceps 370 g male, 390 g female Ferrari, 1993; Ferrari & Ferrari, 1989; Ferrari & Rylands,1994; Ford & Davis, 1992; Harrison & Tardif, 1994; Koenig, 1995; Muskin, 1984; Rylands, 1993; Rylands & de Faria, 1993; Stevenson & Rylands, 1988 Fruit dominates, insects important, gums or nectar seasonal Leontopithecus L. caissarac L. chrysomelas L. chrysopygus L. rosalia Black-faced lion tamarin Golden-headed lion tamarin Black lion tamarin Golden lion tamarin Ripe fruit 53% (32-78%), insect foraging 25% (14-50%) of feeding time, unripe fruit 6-7%, exudates (gums) 9% (1-20%), nectar 7% (0-43%) Diurnal, arboreal mostly, pairs or multimale/ multifemale 2-3 adults/group, 2-16 total 361-794 g females, 437-710 g males Albernaz, 1997; Butynski, 1982; Dietz et al., 1997; Ferrari, 1993; Ferrari & Ferrari, 1989; Ford & Davis, 1992; Rylands, 1993; Tardif et al., 1993 Gumsdominate, insectsimportant, fruit can depend on location Callithrix C. jacchus C. flaviceps C. penicillata C. pygmaea (was genus Cebuella) Common marmoset Buffy-headed marmoset Black tufted-eared marmoset Pygmy marmoset Exudates (gums) 45% (24-70%), fruit 16% (14-30%), insect foraging 39% (30-70%), nectar in dry season; C.pygmaea exudates (gums) 60% (30-77%), fruit 8% (0-10%), insects 30% (20-33%) Diurnal, arboreal mostly, multimale/ multifemale, groups 1-15; C. pygmaea monogamous families, up to 4 litters living together 182-354 g females, 225-406 g males; C. pygmaea 112-140 g females, 99-160 g males Coimbra-Filho & Mittermeir, 1978; Ferrari & Ferrari, 1989; Ferrari & Rylands, 1994; Ford & Davis, 1992; Ramirez, 1985a; Rylands & de Faria, 1993; Silva & Downing, 1995; Soini, 1982, 1988, 1993 Insects and fruit dominate, gums and nectar seasonally important Callimico C. goeldii Goeldi’s monkey Preferred food insects; also soft, sweet fruit in wet season, sticky coating of gum on pods in dry season; rarely buds or young leaves; diet similar to Saguinus spp, sometimes live with mixed Saguinus troops Diurnal, arboreal mostly, monogamous pairs, some within group, 2-8 individuals 400-535 g Ford & Davis, 1992; Heltne et al., 1981; Mittermeier & Coimbra-Filho, 1977; Pook & Pook, 1981, 1982 Saguinus S. bicolor S. fuscicollis S. geoffroyi S. imperator S. inustusc S. labiatus S. leucopus S. midas S. mystax S. nigricollis S. cedipus S. tripartitusc Bare-faced tamarin Saddleback tamarin Red-crested tamarin Emperor tamarin Mottled-faced tamarin Red-bellied tamarin Silvery-brown bare-faced tamarin Golden-handed tamarin Mustached tamarin Spix’s black-mantled tamarin Golden-mantled Cotton-top tamarin saddleback tamarin Insects 45% (30-77%), fruit 35% (13-74%), exudate 10% (0-37%), nectar 7% (0-35%), young leaves 3%, seeds; 34.8% of total activities foraging for insects, 17% plant foods; insect capture rate might be only 5.4% of prey-foraging time Diurnal, arboreal, multimale/ multifemale groups, 2-16 individuals; S. imperator, S. labiatus, and S. midas multimale/ multifemale, but only 1 reproducing female 272-600 g females, 242-633 g males Crandlemire-Sacco, 1988; Egler, 1992; Ferrari & Ferrari, 1989; Ford & Davis, 1992; Garber, 1984, 1988, 1993a,b; Harrison & Tardif, 1994; Lopes & Ferrari, 1994; Pack et al., 1999; Peres, 1993a; Ramirez, 1985a,b; Skinner, 1985; Silva & Downing, 1995; Soini, 1987; Terborgh, 1983 aDiet format: mean (range). bBody weights in ranges whenever possible; single numbers are not averages but indicate that only one individual of the species has been weighed in the wild. cNo data available from the wild but assumed to be similar to congenerics.

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 1-3 Cebid Feeding Ecology Scientific Name Common Name Dieta Behavior Body Weightb References More insectivorous than frugivorous Saimiri S. boliviensis S. oerstedii S. sciureus S. ustusc S. vanzoliniic Bolivian squirrel monkey Red-backed squirrel monkey Common squirrel monkey Golden-backed squirrel monkey Black squirrel monkey Animal prey, particularly insects 60% (47-100%), vertebrates 1%, fruit 25% (15-39%), flowers 5% (2-13%), leaves 13% (11-18%), seeds/ nuts, successful in 61% of insect foraging; during dry season, rely on figs S. boliviensis, S. oerstedii: diurnal, arboreal, multimale/ multifemale, groups, up to 23 individuals; S. sciureus, S. vanzolinii: groups, 22-50 individuals 0.54-1.25 kg females, 0.48-1.2 kg males Costello et al., 1993; Ford & Davis, 1992; Janson & Boinski, 1992; Mittermeier & vanRoosmalen, 1981; Rosenberger, 1992; Silva & Downing, 1995; Souza et al., 1997; Terborgh, 1983 Primarily frugivorous Ateles A. belzebuth A. chamek A. fuscicepsc A. geoffroyi A. marginatusc A paniscus White-bellied spider monkey Black-faced black spider monkey Brown-headed spider monkey Black-handed spider monkey White-whiskered spider monkey Black spider monkey Total fruit 78% (18-100%), including unripe fruit 6%; seed 5% (0-19%); total leaves 16% (0-38%), including mature leaves 3%; flowers 3% (1-10%); epiphytes 2%; dead wood, buds, insects 1% Diurnal, arboreal (salt licks on ground), fission-fusion, groups 3-35 individuals 5.0-11.0 kg females, 5.8-9.8 kg males Chapman, 1987, 1988; Ford & Davis, 1992; Hladik, 1975; Klein & Klein, 1975, 1977; MendesPontes, 1997; Milton, 1981; Mittermeier & vanRoosmalen, 1981; Nunes, 1998; Robbins et al., 1991; Silva & Downing, 1995; Simmen & Sabatier, 1996; Symington, 1988; VanRoosmalen, 1985; VanRoosmalen & Klein, 1988; White, 1986 Primarily frugivorous, seasonally seeds or leaves important Aotus A. nigriceps Southern red-necked night monkey   A. trivirgatus Northern gray-necked owl monkey Fruit (soft) 44% (16-75%), leaves 32% (5-46%), insects 13% ( 5-15%), other (especially flowers) 11%; Aotus diet similar to Callicebus but ate less vegetation, more insects in abundant season Nocturnal, arboreal, monogamous family groups, 2-5 individuals; feed in groups 25-55 at low elevations 0.78-1.1 kg females, 0.825-1.05 kg males Durham, 1975; Engqvist & Richard, 1991; Kinzey, 1992; Wright, 1981,1989, 1994 Callicebus C. brunneus C. caligatusc C. cinerascensc C. cupreusc C. donacophilusc C. dubiusc C. hoffmannsic C. modestusc C. moloch C. oenanthec C. olallaec C. personatus C. torguatus Brown titi monkey Chestnut-bellied titi monkey Ashy gray titi monkey Red titi monkey Bolivian gray titi monkey Hershkovitz’s titi monkey Hoffmann’s titi monkey Bolivian titi monkey Dusky titi monkey Andean titi monkey Beni titi monkey Masked titi monkey Collared titi or widow monkey Fruit 61% (30-87%) (of which seeds may be as much as 28%), leaves (mostly young) 21% (2-66%), insects 12% (0-28%), flowers 2% (0-18%); when food scarce, ate 25% bamboo and vine leaves Diurnal, arboreal, monogamous family 2-6 individuals 0.7-1.5 kg Crandlemire-Sacco, 1988; Easley, 1984; Ford & Davis, 1992; Heiduck, 1997; Kinzey, 1977, 1981, 1992; Kinzey & Gentry, 1979; Muller, 1996; Palacios, 1997; Robinson et al., 1987; Silva & Downing, 1995; Terborgh, 1983, Wright, 1994 Cebus C. albifrons C. apella C. capucinus C. olivaceus White-fronted capuchin Tufted or brown capuchin White-throated capuchin Weeper or wedge-capped capuchin Fruit 55% (10-95%), of which seeds are 8% (0-39%); leaves (mostly young) 8% (0-39%); insects 33% (2-100%); flowers 2% (0-14%); C. apella in Argentina ate bromeliad leaves 72%, fruit 3%, insects 25% Diurnal, arboreal mostly, multimale/ multifemale groups of 2-40 individuals; C. apella, C. olivaceus: with alpha male 1.4-3.8 kg females, 1.3-4.8 kg males Brown & Zunino, 1990; Chapman, 1987; Chapman & Fedigan, 1990; Ford & Davis, 1992; Hladik et al., 1971; Janson, 1985; Janson & Boinski, 1992; Mittermeier & vanRoosmalen, 1981; Peres, 1994a; Robinson, 1984; Simmen & Sabatier, 1996; Teaford & Robinson, 1989; Terborgh, 1983

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Both fruit and seeds rank high, sometimes seeds dominate Lagothrix L. flavicauda L. lagothricha Yellow-tailed woolly monkey Woolly monkey Fruit 67% (6-95%), seeds 10% (0-35%); insects 7% (0-34%), leaves (mostly young) 12% (2-48%); flowers 2% (0-9%), pod exudates eaten at some sites Diurnal, arboreal, multimale/ multifemale groups of 5-70 individuals; highlands groups 6-7; lowlands groups 10-20 individuals 3.5-6.5 kg females, 3.6-10.2 kg males Butynski, 1982; Defler & Defler, 1996; Durham, 1975; Ford & Davis, 1992; Kinzey, 1997; Luna, 1987; Peres, 1994b; Ramirez, 1988; Robinson & Janson, 1987; Soini, 1987; Stevenson et al., 1994 Cacajao C. calvus C. melanocephalus Bald uacari Black-headed uacari Seeds (mostly unripe) 59% (20-97%), fruit pulp 22% (1-60%), nectar 6% (0-58%), insects 5%, leaves and so on 3%; seeds of unripe fruit important as for all pithecines Diurnal, arboreal mostly, multimale/ multifemale groups 5-30 up to 100 individuals 2.4-4.0 kg Ayres, 1989; Barnett & Brandon-Jones, 1997; Fontaine, 1981; Ford, 1994; Kinzey, 1992; Mittermeier & Coimbra-Filho, 1977 Chiropotes C. albinasus C. satanas White-nosed saki Bearded saki Seeds (mostly unripe) 53% (12-96%), fruit 37% (6-84.5%); leaves 2% (0-4%), flowers 5% (1-11%), insects 3% (0-24%); seed predators on 52 species and seed dispersers of 7 species; C. satanas ingest unripe fruit with hard pericarp Diurnal, arboreal, multimale/ multifemale groups 10-30 individuals 1.9-3.3 kg females, 2.2-4.0 kg males Ayres, 1989; Ford & Davis, 1992; Kinzey, 1992; Kinzey & Norconk, 1993; Mittermeier & vanRoosmalen, 1981; Mittermeier et al., 1983; Norconk et al., 1998; Robinson et al., 1987; van Roosmalen et al., 1981, 1988 Pithecia P. aequatorialisc P. albicans P. irroratac P. monachus P. pithecia Equatorial saki Buffy saki Bald-faced saki Monk saki White-faced saki Seeds 38% (17-88%), other fruit 43% (3-51%), leaves (mostly young) 12% (0-32%), insects 1.0% (0-6%), flowers 6% (0-15%); P. monachus may eat more leaves or insects, P. pithecia more young seed ( 60%) Diurnal, arboreal, monogomous family groups, groups 2-8; P. aequatorialis, P. monachus: cryptic 0.779-2.5 kg females, 0.964-3.1 kg males Buchanan et al., 1981; Ford & Davis, 1992; Happel, 1982; Kinzey, 1992; Kinzey & Norconk, 1993; Mittermeier & vanRoosmalen, 1981; Norconk, 1996; Norconk & Kinzey, 1990; Norconk et al., 1998; Peres, 1993b Primarily folivorous, some fruit, no animal prey Alouatta A. belzabulc A. caraya A. colibensisc A. fusca A. palliata A. pigra A. sarac A. seniculus Red-handed howler Black-and-gold howler Colba Island howler Brown howler Mantled howler Black howler Bolivian red howler Red howler A. palliata, A. seniculus, A. pigra: total leaves 54% (20-100%), including 38% young, 16% mature leaf; total fruit, especially figs, 39% (0-80%), including 34% ripe, 5% unripe; flowers 9% (0-90%); A. fusca, A. caraya: 72% leaves (45-89%); fruit 20% (2-55%); flowers 8% (0-24%) Diurnal, arboreal (drink on ground, A. palliata can swim), 1,2 or multimales/ multifemales, groups 4-21 individuals; One-male groups common 2.4-7.6 kg females, 4.2-11.4 kg males Bicca & Calegaro, 1994; Chapman, 1987; Crockett & Eisenberg, 1987; de Thoisy & Richard-Hansen, 1997; Estrada, 1984; Estrada & Coates-Estrada, 1986; Ford & Davis, 1992; Galetti et al., 1987; Garcia, 1994; Gaulin & Gaulin, 1982; Glander, 1978; Hladik el al., 1971; Julliot & Sabatier, 1993; Milton, 1980; 1981; Mittermeier & van Roosmalen, 1981; Neville et al., 1988; Oftedal, 1991; Prates et al., 1987; Simmen & Sabatier, 1996; Smith, 1977; Stoner, 1996; Strier, 1992. Brachyteles B. arachnoides Woolly spider monkey or muriqui Leaves 58% (range 41-93%); fruit 28% (7-59%), within which unripe seeds were 8% (0-32%); flowers 14% (0-38%) Diurnal, arboreal, multimale/ multifemale and fission-fusion, groups 5-45 individuals 9.4 kg female, 12.1 kg male Ford, 1994; Lemos, 1988; Milton, 1984; Neville et al., 1988; Nishimura et al., 1988; Strier, 1991, 1992 aDiet format: mean (range). bBody weights in ranges whenever possible; single numbers are not averages but indicate that only one individual of the species has been weighed in the wild. cNo data available from the wild but assumed to be similar to congenerics.

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 1-4 Colobine Feeding Ecology Scientific Name Common Name Dieta Behavior Body Weightb References Strongly folivorous Colobus C. guereza C. vellerosusc Abyssinian, guereza, or eastern black-and-white colobus Geoffroy’s or white-thighed black-and-white colobus C. guereza: young leaves and buds 64% (52-90%), mature leaves 13% (2-22%), whole fruit 15% (0-34%), flower and bud 6% (0-17.1%); seeds 1%, stems 0.5%; other 0.5% Diurnal, arboreal mostly; C. guereza: 1 male or multimale/ multifemale group 2-50; others: multimale/ multifemale 6.8-8.92 kg females, 9.7-13.5 kg males Clutton-Brock, 1975; McKey, 1978; Napier, 1985; Oates, 1977, 1978, 1994, Oates et al., 1994; Struhsaker, 1978a; Struhsaker & Oates, 1975 Folivorous but >30% seed-eating Colobus C. angolensis C. polykomos C. satanas Angolan black-and-white colobus King or western black-and-white colobus Black colobus Young leaves 31% (2-85%), mature leaves 18% (4-75%), fruit 8% (0-55%), seeds 35% (0-89%), flowers and buds 5% (0-31%), stems 1% (0-15%), other 1.5% Diurnal, arboreal mostly; C. angolensis 1 male or multimale/ multifemale groups 2-50; others multimale/ multifemale 4.32-9.67 kg females, 9.7-13.5 kg males Dasilva, 1992, 1994; Groves, 1973; Harrison & Hladik, 1986; Maisels et al., 1994; McKey, 1978; McKey & Waterman, 1982; McKey et al., 1981; Silva & Downing, 1995; Tutin et al., 1997 Strongly folivorous, some seed Procolobus (Piliocolobus or Colobus) P. badius P. pennantii P. preussic P. rufomitratus P. verus Western red colobus Pennant’s red colobus Preuss’s red colobus Tana river red colobus Olive colobus Young leaves and buds 52% (7-85%) mature leaves and petiole 16% (1-60%), fruit (especially unripe) 9% (0-41%), seeds 12% (0-31%), flowers and buds 9% (0-36%), stems and miscellaneous 1% (0-17%) Diurnal, arboreal, multimale/ multifemale groups 5-80; P. rufomitratus, 1 male or multimale/ multifemale P. verus: 1 or 2 males + multifemale 4.2-8.2 kg females, 4.7-11.0 kg males Brandon-Jones, 1985; Clutton-Brock, 1975; Decker, 1994; Gatinot, 1977; Maisels et al., 1994; Marsh, 1981, 1983; McKey, 1978; Mowry et al., 1996; Oates, 1988; Oates & Whitesides, 1990; Oates et al., 1994; Silva & Downing, 1995; Struhsaker, 1975, 1978a; Struhsaker & Oates, 1975; Wachter et al., 1997 Folivorous/frugivorous (>50% leaf, <50% fruit) Nasalis N. larvatus N. (Simias) concolor Proboscis monkey Pig-tailed langur Young leaf 45% (38-48%), mature leaves 4%, fruit 40% (17-50%), of which seeds are 15-20%, flowers and buds 3%; stems 3%; other 2.5%; insects <1%; fruit eaten usually unripe; frugivorous January-May, folivorous June-December Diurnal, arboreal, swimmers, 1 male + multifemale and bachelor troops, groups 2-20; N. (Simias) concolor also in pairs or multimale/ multifemale 7.1-11.8 kg females, 8.8-23.6 kg males Bennett & Davies, 1994; Bennett & Sebastian, 1988; Ross, 1992; Watanabe, 1981; Yeager, 1989 Presbytis P. comata P. femoralis P. frontatac P. hosei P. melalophos P. potenziani P. rubicunda P. thomasi Grizzled leaf monkey Banded leaf monkey White-fronted leaf monkey Hose’s leaf monkey Mitered leaf monkey Mentawai Island leaf monkey Maroon leaf monkey Thomas’s leaf monkey Young leaves 41% (15-71%); mature leaves 4% (0-11%); fruit 42% (3-80%), of which about 7% is seeds (1-30%) and unripe fruit and seeds up to 30%; flowers and buds 10% (1-30%); other 3%; very little insect eaten <1%. P. rubicunda: seed predators Diurnal, arboreal, male + multifemale, monogamous pairs, groups 2-21; P. melalophos: 1 male or multimale/ multifemale 3.0-6.7 kg females, 5.6-8.2 kg males Adiputra, 1994; Aldrich-Blake, 1980; Bennett & Davies, 1994; Brandon-Jones, 1985; Chivers, 1994; Curtin, 1980; Davies, 1991; Davies et al., 1988; Goodman, 1989; Gurmaya, 1986; Leutenegger & Cheverud, 1982; MacKinnon & MacKinnon, 1980b; Rodman, 1978; Ruhiyat, 1983; Silva & Downing, 1995; Ungar, 1995; Watanabe, 1981 Pygathrix P. nemaeus P. nigripesc P. (Rhinopithecus) avunculus P. (Rhinopithecus) bieti P. (Rhinopithecus) brelichi P. (Rhinopithecus) roxellana Red-shanked douc langur Black-shanked douc langur Tonkin snub-nosed monkey Black or Yunnan snub-nosed monkey Guizhou snub-nosed monkey Sichuan golden snub-nosed monkey Young leaves and buds 37% (7-93%), mature leaves 37% (31-88%), fruit 15% (5-47%), seeds 3% (0-15%), flowers 7% (0-28%), lichen 5% (0-50%); figs important, almost no insects Diurnal, arboreal (some also terrestrial), multimale/ multifemale or 1 male + multifemale, groups 3-200 individuals 6.5-10 kg females, 10.9-20.3 kg males Bennett & Davies, 1994; Bleisch & Xie, 1994; Bleisch et al., 1998; Ji & Bleisch, 1994; Kirkpatrick, 1994; Lippold, 1995; Long, 1994; Nhat, 1993, 1994; Silva & Downing, 1995

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Semnopithecus (Presbytis) S. entellus Hanuman langur In remote, wild areas: young leaves 26% (0-69%), mature leaves 26% (0-79%) fruit 34% (0-72%), of which about 3% is seeds (0-45%), flowers and buds 9% (0-43%), stem 1.6%; other 3%; more insects than other colobines; near farms: 90% of diet is cultivated crops; Himalayan subspp eat pinecones, bark, twigs during snowy winter months Troops near temples eat offerings, raid crops; diurnal, terrestrial, and arboreal, 1 male or multimale/ multifemale, groups 11-262 individuals 6.7-15.6 kg females, 10.6-20.9 kg males Bennett & Davies, 1994; Hladik, 1975, 1988; KarGupta & Kumar, 1994; Newton, 1992; Oppenheimer, 1977; Silva & Downing, 1995; Srivastava, 1991; Starin, 1978 Trachypithecus (Presbytis) T. auratus T. cristatus T. delacouric T. francoisic T. geei T. (Kasi) johnii T. obscurus T. phayrei T. pileatus T. (Kasi) vetulus Ebony langur Silvered langur Delacour’s langur Francois’s langur Golden langur Nilgiri langur Dusky or spectacled leaf monkey Phayre’s leaf monkey Capped leaf monkey Purple-faced leaf monkey Young leaves and shoots 32% (9-52%), mature leaves and petioles 26% (1-61%), fruit 32% (1-55%), of which 7% is seeds (0-40%); flowers and buds 10% (0-43%), insects <1%, other 0.5%; more mature leaves than Presbytis and much of fruit eaten unripe; raids crop; T. pileatus - animal prey 1.6%, gum and termite soil Diurnal, arboreal (some also terrestrial), 1 (some 2) male and multifemale, groups 2-40 individuals 3.0-10.9 kg females, 6.0-13.6 kg males Aldrich-Blake, 1980; Bennett & Davies, 1994; Brandon-Jones, 1985; Brotoisworo & Dirgayusa, 1991; Chivers, 1994; Curtin, 1980; Curtin & Chivers, 1978; Fleagle, 1978; Hladik, 1975, 1977, 1988; Hladik & Hladik 1972; Islam & Husain, 1982; Kool, 1992, 1993; Kumar-Gupta & Kumar, 1994; Li, 1993; MacKinnon & MacKinnon, 1980b; Mukhergee, 1978; Oates et al., 1980; Silva & Downing, 1995; Stanford, 1988; 1991a, 1991b; Whitten, 1987, Wrangham et al., 1993 aDiet format: mean (range). bBody weights in ranges whenever possible; single numbers are not averages but indicate that only one individual of the species has been weighed in the wild. cNo data available from the wild but assumed to be similar to congenerics. ited use for quantifying a diet, because the time required for food acquisition varies. Time Sampling Methods Less complete but more manageable methods for recording feeding behavior, these are probably the most commonly used today. These methods also require independent start-and-stop rules, and dawn and dusk are often used. One-Zero Sampling. A behavior is scored only once per observation period, regardless of the number of times it occurs. This method is adequate for preliminary reconnaissance. It is not recommended for detailed feeding-ecology studies, because it generally yields poor ‘‘time-spent’’ estimates. Instantaneous Sampling. The observer records a focal-animal’s behavior at predetermined times. This method works well with ongoing behavior that can be timed with a stopwatch, such as feeding behavior. For example, during a feeding bout, what the animal is eating every 30 or 60 seconds is recorded. Another approach is to observe the focal animal every 15 min and record all behaviors for 5 min. A limitation of this approach is that rare events often are not recorded. However, when continuous observations prove impossible, this generally is considered the next-best method. Scan Sampling. Instantaneous observations are made of several animals simultaneously. This is useful for studying less-detailed behavior. Alternative Feeding-Ecology Methods In some circumstances, particularly if terrestrial primates are being studied in dense rainforest, visual observations of feeding behavior are impractical. It can be impossible to see the animals well enough to determine what they are eating or how much time they spend eating it, and alternative methods might be needed. Some researchers studying nocturnal animals use both visual observations and alternative methods (Nash 1983). Alternative methods for studying feeding ecology are outlined below. ANALYSIS OF STOMACH CONTENTS Measurement of stomach contents, now rare, can be used to estimate the mass of different food categories consumed (for example, fruit, leaves, or insects); with care and skill, one can identify the species eaten (Booth, 1956; Fooden, 1964; Charles-Dominique, 1974; Gautier-Hion et al., 1980). However, because the animal must be killed, only a single measure per animal is obtained. An additional

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 1-5 Non-colobine Cercopithecine Feeding Ecology Scientific Name Common Name Dieta Behavior Body Weightb References Omnivorousbut predominantly frugivorous(depending on habitat) Macaca M. arctoides M. assamensisc M. cyclopis M. maurac M. ochreatac M. radiata M. silenus M. tonkeanac Stump-tailed macaque Assamese macaque Formosan rock macaque Celebes moor macaque Booted macaque Bonnet macaque Lion-tailed macaque Tonkean macaque Fruit and seeds, young leaves, flowers and buds, other plant parts, gums, grass, clover, sprouts, roots, bark, resin, animal prey (insects and vertebrates), fungus, raid crops/dumps; most species, little field work; M. radiata eat temple offerings Diurnal, arboreal and terrestrial, multimale/ multifemale, some 1 male + multifemale, group size 5-50; M. silenus: rarely on ground, but swim, as do many macaques 3.69-8.5 kg females, 4.86-12 kg males Bynum, 1994; Krishnamani, 1994; Kurup & Kumar, 1993; Kuruville, 1980; Leutenegger & Cheverud, 1982; Richard et al., 1989; Silva & Downing, 1995; Wolfheim, 1983 M. fascicularis M. nemestrina M. nigra M. sinica Long-tailed or crab-eating macaque, or cynomolgus monkey Pig-tailed macaque Celebes or crested black macaque Toque macaque Fruit 67% (2-100%); flowers and buds 3% (0-68%); leaves 12% (1-62%); bark, roots, pith and other 6% (0-73%); grass 1%; fungi, resins and other 2%; prey items 11% (0-46%); M. fascicularis’ diet at one site was 51% temple offerings Diurnal, arboreal and terrestrial, multimale/ multifemale, group size 10-90 individuals M. fascicularis, M. sinica: 1.5-5.7 kg females, 3.9-8.39 kg males; M. nemestrina, M. nigra: 3.5-10.9 kg females, 6.2-14.5 kg males Aldrich-Blake, 1980; Butynski, 1982; Caldecott, 1986a,b; Davies et al., 1983; Dittus, 1977; Hladik, 1975; Lucas & Corlett, 1991; MacKinnon & MacKinnon, 1980b; O’Brien & Kinnaird, 1997; Richard et al., 1989; Rodman, 1978; Silva & Downing, 1995; Sussman & Tattersall, 1981; Temerin et al., 1984; Ungar, 1995; Wheatley, 1982; 1987; Whitten & Whitten, 1982; Wolfheim, 1983; Yeager, 1996 M. fuscata Japanese macaque Fruit 47% (0-100%); flowers 5% (0-40%); leaves 22% (0-94%); herb/grass 6% (0-65%); roots, bark, twigs, and other 13% (0-95%); fungi, resins, and other 1% (0-18%); prey 9% (0-50%); winter diets high in seeds in cedar forest, high in winter buds in other habitats Diurnal, arboreal and terrestrial, multimale/ multifemale group size 40-194 individuals 8.3-18.0 kg females, 11.0-18.0 kg males Agetsuma, 1995a,b; Agetsuma & Nakagawa, 1998; Agetsuma & Noma, 1995; Hill, 1997; Iwamoto, 1982; Maruhashi, 1980; Nakagawa, 1997, 1989a; Suzuki, 1965 M. mulatta M. sylvanus Rhesus macaque Barbary macaque Fruit 24% (0-70%); flowers 5% (0-40%); leaves 47% (2-99%); bark, pith, roots, and other 11% (0-34%); herbs or grass 9%, (0-56%); fungi or sap 1%; prey 6% (0-66%); in some sites, M. sylvanus feed heavily on acorns and cedar leaves, cones, and cambium; M. mulatta eat temple offerings Diurnal, multimale/ multifemale; M. sylvanus: group size 12-59 individuals; M. mulatta: group size 10-200 individuals M. sylvanus: 10.2-11.2 kg females, 15.3-17.0 kg males; M. mulatta: 3.0-10.9 kg females, 5.08-10.9 kg males Deag, 1983; Goldstein & Richard, 1989; Lindburg, 1977; Malik, 1986; Mehlman, 1988, 1989; Menard & Vallet, 1986; Richard, et al., 1989; Seth & Seth, 1986 M. thibetana Tibetan macaque Reproductive plant parts 35% (10-59%), ground-layer foods 22% (11-33%), leaves and other vegetative parts 43% (30-56%), prey not quantified; fed by humans near temples Diurnal, mostly terrestrial, multimale/ multifemale 7.81-14.2 kg females, 10.7-13.0 kg males Richard et al., 1989; Silva & Downing, 1995; Zhao & Deng, 1988; Zhao et al., 1991 Allenopithecus A. nigroviridis Allen’s swamp monkey Fruit 81%; pith 2%; roots, flowers, nectar, animal prey (vertebrates and invertebrates) 17%; little studied Diurnal, arboreal and terrestrial (swim), multimale/ multifemale, group size up to 40 individuals 3.7 kg female, 5.95 kg male Gautier-Hion, 1988a,b; Zeeve, 1991 Cercocebus C. agilisc C. galeritus C. torquatus C. torquatus atys Agile mangabey Tana river mangabey White-collared mangabey Sooty mangabey Fruit 76% (14-100%); leaves 12% (0-83%); flowers and buds 1% (0-5%); other plant parts 4% (0-50%); prey 8% (0-22%) Diurnal, arboreal and terrestrial, multimale/ multifemale, group size 14-95 individuals 4.7-5.47 kg females, 9.2 - 10.8 kg males Davies et al., 1983; Fleagle, 1988; Gautier-Hion, 1978, 1983; Gautier-Hion et al., 1980; Homewood, 1978; Mitani, 1989, 1991; Napier, 1981; Quris, 1975; Ross, 1991; Silva & Downing, 1995; Waser, 1984; Wolfheim, 1983

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Cercopithecus C. campbelli C. dryasc C. erythrogaster C. erythrotis C. hamlyni C. mona C. petaurista C. preussic C. sclateri C. solatusc Campbell’s guenon Dryas guenon White-throated guenon Red-eared guenon Owl-faced monkey Mona monkey Lesser spot-nosed guenon Preuss’s monkey Sclater’s guenon Sun-tailed guenon Fruit (and seeds) 54.6-90%, animal prey 5.0-25%, leaves 6.0 -18.9%, flowers 3-6%, gums 1.9-2.8%, shoots, mushrooms, nectar; eat more leaves when fruit is scarce; raid crops; many species little studied Diurnal, arboreal (C. campbelli most terrestrial of all the guenons); 1 male, multifemale; group size: C. campbelli, C. hamlyni, C. preussi., C. solatus 2-15; C. erythrogaster, C. erythrotis, C. mona, C. petaurista, C. sclateri 4-35; C. sclateri multimale/ multifemale 1.8-4.5 kg females, 2.4-7.0 kg males Bourliere et al., 1970; Caldecott, 1986a; Colyn, 1994; Napier, 1981; Oates, 1985; Silva & Downing, 1995; Wolfheim, 1983 C. ascanius C. cephus C. mitis C. neglectus C. nicitans C. pogonias C. wolfi Red-tailed guenon Mustached guenon Blue monkey DeBrazza’s monkey Putty-nosed or greater spot-nosed guenon Crowned guenon Wolf’s guenon Fruit 67% (5-100%) (seed only 8%); leaves 15% (0-96%); flowers 4% (0-51%); bark, pith, and other 2% (0-30%), fungi 2% (0-39%), invertebrates 14% (0-45%); C. mitis eat bamboo; C. pogonias eat more prey when food is scarce; C. mitis in southern Africa: fruit 21%; leaves 27%; cambium, pith, twigs 46%; fungi 6%; invertebrates less than 1% Diurnal; arboreal; 1 male-multifemale groups. (C. neglectus; some monogamous pairs); group size: C. ascanius, C. cephus, C. neglectus: 5-35; C. mitis, C. nictitans: 7-70; C. pogonias, C. wolfi: 1-19 individuals C. ascanius, C. cephus, C. pogonias, C. wolfi: 2.4-3.4 kg females, 3.2-4.8 kg males; C. mitis, C. neglectus, C. nictitans: 2.7-8 kg females, 4-9.99 kg males Beeson, 1989; Butynski, 1982, 1990; Colyn, 1994; Conklin et al., 1998; Cords, 1986, 1987; Gautier-Hion, 1978, 1980, 1983, 1988a; Gautier-Hion & Gautier, 1974, 1978, 1979; Gautier-Hion et al., 1980; Kaplin & Moermond, 1998; Kaplin et al., 1998; Lawes, 1991; Lawes et al., 1990, Moreno-Black & Maples, 1977; Napier, 1981; Rudran, 1978; Schlichte, 1978; Silva & Downing, 1995; Struhsaker, 1978b, 1980; Tutin et al. 1997; Wahome et al., 1993; Wolfheim, 1983; Wrangham et al., 1993 C. diana Diana monkey Fruit 39%; leaves 10%; flowers and buds 12%; bark, pith, and so on 1%; fungi 10%; invertebrates 31%; some reports claim more fruit or leaf Diurnal, arboreal, 1 male-multifemale, group size 5-50 individuals 4.3-7.1 kg Oates & Whitesides, 1990; Ross, 1991; Silva & Downing, 1995; Wachter et al., 1997; Wolfheim, 1983 C. lhoesti L’Hoest’s monkey Fruit 42% (22-80%), leaves 19%, herbs 35% (because are terrestrial), flowers 4%, prey 9% Diurnal, terrestrial, somewhat arboreal, 1 male-multifemale, group size 5-17 individuals 3-4 kg females, 6-7 kg males Colyn, 1994; Kaplin & Moermond, 1998; Silva & Downing, 1995; Wolfheim, 1983 Chlorocebus C. (Cercopithecus) aethiops Vervet, grivet, green, or tantalus monkey Fruit 46%; leaves 23% (more mature leaves than young); flowers and buds 10%; bark, twig, or pith 6%; fungi or gums 3%; grass 1%; prey 13%, raid crops; take handouts Diurnal, terrestrial and arboreal; multimale/ multifemale, group size 5-76 individuals 1.5-5.23 kg females, 3.1-8 kg males Butynski, 1982; Davies et al., 1983; Dunbar & Dunbar, 1974; Galat & Galat-Luong, 1977, 1978; Harrison, 1983, 1984; Kavanagh, 1978; Moreno-Black & Maple, 1977; Napier, 1981; Silva & Downing, 1995; Whitten, 1983, 1988; Wolfheim, 1983; Wrangham & Waterman, 1981 Erythrocebus E. patas Patas monkey Fruit 20% (5-34%); leaves 17% (6-27%); flowers and buds 36% (7-65%); stems, shoots and pith 3%; sap and gum 10%; prey 16% (except Kenya: fruit and seeds 6%, leaves 3%, flowers 7%, gum 39%, prey 43%). Diurnal, mostly terrestrial, 1 male-multifemale, group size 5-34 individuals 4.08-7.1 kg females, 7.48-12.6 kg males Isbell, 1998; Koster, 1985; Nakagawa, 1989b; Napier, 1981; Olson & Chism, 1984; Silva & Downing, 1995 Lophocebus L. (Cercocebus) albigena L. aterrimus Grey-cheeked mangabey Black mangabey Fruit 69% (21-91%), up to 32% of which was figs; leaves 7% (0-65%); flowers and buds 4% (0-35%); bark, pith, or stems 3% (0-22%); other plant parts 1% (0-33%); prey 17% (2-44%); raid crops Diurnal, arboreal, occasionally come to ground to drink, multimale/ multifemale, group size 6-28 individuals L. albigena: 5.4-6.4 kg females, 6.8-8.98 kg males; L. aterrimus: 13.0-18.0 kg females, 21.0 kg male Conklin-Brittain et al., 1998; Davies et al., 1983; Freeland, 1979; Gautier-Hion, 1977, 1978, 1983; Gautier-Hion et al., 1980; Horn, 1987; Mitani, 1991; Napier, 1981; Olupot et al., 1997; Olupot, 1998; Silva & Downing, 1995; Struhsaker, 1978b; Tutin et al., 1997; Waser, 1975, 1977, 1984

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