food adaptations of a transplanted japanese macaque troop (arashiyama west)

PRIMATES, 20(3): 399-410, July 1979 399

Food Adaptations of a Transplanted Japanese Macaque Troop (Arashiyama West)

TIM W. CLARK University of Wisconsin, Madison

ABSTRACT. In 1972 Arashiyama West troop of Japanese macaques was transplanted to southcentral United States and kept in semi-free ranging conditions. The new environment provided an opportunity to assess aspects of the species' adaptive potential. About 1,500 feeding observations were made monthly over 6.5 months. Unlimited provisioned food was available, but monkeys utilized native plants immediately and use increased until it included 50 4- ~o of the diet by weight. Shrubs provided 7570 of foods in first month and 32~ thereafter. Sorghum comprised 25 ~ in May-July. Soil, arthropods, fungi, bulbs, and roots each comprised less 5 70. Between 21 and 37 foods were utilized monthly. Monthly food uses corresponded to availability. Many food plants required unique handling by monkeys. Four general adaptive responses to potential foods are described. Evolution has clearly shaped the Japanese macaque into a highly omnivorous and behaviorally flexible animal.

I N T R O D U C T I O N

Seldom does an opportunity arise to document adaptations of an intact, integrated primate society to potential food resources of an entirely new environment outside the species' evolutionary experience. In 1972 the Arashiyama A troop of Japanese macaques (150 members renamed Arashiyama West) was transplanted to the southcentral United States and maintained under semi-free ranging conditions (CLARK MANO, 1975). The te rm"adapta t ion" here means"accommodat ion" and"adjustment ," denoting any immediate reaction in response to a suddenly changed environment and with modification of overt characteristics or elicitation of latent ones in maintain- ing individual or collective integrity (MoRRISON & MENZEL, 1972). This paper reports results of a 6.5 month study which: (1) quantitatively described patterns of food acquisition and utilization and (2) documented new behavior patterns involved in utilizing several food plants.

STUDY AREA

Transplanted to a site 48 km north of Laredo, Texas, the troop was confined to a 0.6 ha corral from February 23 to March 24, 1972 and then released into a 42 ha enclosure where they free-roamed for the study's remainder. Containment facilities and some features of the transplant site were described by CASEY and CLARK (1976). Major vegetational features of southern Texas and the macaques evolutionary home at Arashiyama near Kyoto, Japan are compared in Table 1.

400 T.W. CLARK

Table 1. Comparison of some biotic characteristics of Arashiyama, Kyoto, Japan and Laredo, Texas, U.S.A.

Evolutionary home Transplant site Biotic characteristics Arashiyama, Kyoto, Japan Laredo, Texas, U.S.A. Flora

No. of species No. of species eaten No. of families Five largest families

Plant communities No. of communities Name of communities and dominants

No. of vegetation strata

height of overstory vegetation Life-form of overstory

841 est. 250+ 192 46+ 155 27+ Compositae Gramineae Gramineae Compositae Rosaceae Leguminosae Leguminosae Verbeneae Aspidiaceae Euphorbiaceae

3 Narrow leaf (Pinus sp.) Broad leaf (Quercus & llex spp.) Sugi (Cryptomeria japonica) (3) Trees ( 2 ) -

Shrubs Shrubs Herbs Herbs

11 m 1.5m Trees Shrubs

1 Acacia-Opuntia savannah

METHODS

Feeding behavior was determined by observing feeding monkeys. Methods varied somewhat depending on the feeding t roop 's location. During the first 30 days hourly sampling of monkey activities included recording all food items being used. The troop was followed on its nearly daily foraging trips after its release into the outer enclosure. The t roop tended to feed three times daily; all three feeding times are equally represented in the quantitative sample. Every three days, one complete daily sample was made- - tha t is, the morning, noon, and evening foraging times were each sampled. While foraging, the t roop frequently spread itself over a 754- m z area. By walking through the foraging troop, I recorded food uses of all monkeys encountered, usually 50 or more animals, yielding about 150 observations every three days and nearly 1,500 per month. Data is expressed in frequency of use; frequency is one monkey's use of one plant.

Amounts eaten were estimated by direct observation. Amounts of food eaten and time spent eating, moving, or sitting were recorded. Immediately following observations, like amounts of the items eaten by the monkey were collected from adjacent plants. These were weighed, oven-dried, and reweighed. Field notes were taken on phenological events of major food plants, particularly flowering time, seed fall, and drying.

RESULTS

PATTERNS OF FOOD ACQUISITION AND UTILIZATION

From February 23 to August 31, 1972, over 8,000 observations on food intake were

recorded.

Food Adaptation of Transplanted Japanese Macaques 401

Plant Life-forms Used

Shrubs were the most used life form. Shrubs provided over three-fourths (75.2~) of the natural foods (non-provisioned) consumed during the first month and about one-third (31.9 ~) from late March through August (Fig. 1). The importance of shrubs varied monthly. The second most important food was cultivated sorghum (Sorghum vulgare), a grass, comprised over one-fourth total intake of non-provisioned foods from May through July. Use of trees as food remained relatively uniform and consistently low throughout the study. Opuntia sp., despite their numerous spines, were readily consumed; comprising 20.3 ~ of native foods consumed from February 23 to March 24. This figure dropped to a low in late spring and began to slowly increase as the year progressed. Grasses (Graminae) other than sorghum comprised a large food class. Monkeys used grasses in greatest amounts early in the year and again after other species had been exhausted or dried. Forbs made up only a small part of the total diet.

TROOP HELD IN CORRAL TROOP RELEASED INTO ENCLOSURE

...... ,'..~.:::~s: ,~:::::~< ======================= I L ::::~.~':::~:::: ~:~:~:-: .$.:-5::~, :':': .':':

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i OTHER =; ~

........ ,.., ...... ............. :~ ~ A S g'~.~..,,. " " , ~- : ~ - X - ~ , ~

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2 3 F E R , - I 2 5 M A R . -~ 1 - 3 1 1 - 3 0 i I - ~ 1 i | - ~ 1 ~4 MAR. ~0 APR. MAY JUNE dULY AUGUST

Fig. 1. Relative use of natural foods used by Japanese macaques of Arashiyama West.

Within two weeks after the troop arrived in Texas, many monkeys were ingesting soil. In March, soil made up 2.9~o of the total intake, in April 5.3~, in May 9.1 o//o, June 10.5 ~, July 9.6 ~, and in August 5.8 ~. Comparison of the chemical composi- tion of Texas to Arashiyama soils is given in Table 2.

The remainder of the diet was made up of arthropods (mostly grasshoppers and some leaf hoppers), fungi, bulbs, and roots; collectively, these comprised tess than 5 ,% of the naturat food diet.

402 T.W. CLARK

Table 2. Comparison of constituents of soil in ppm ingested by Japanese monkeys from Arashiyama, Japan, 1971, and the same troop after transport to Laredo, Texas, 1972

Amounts in ppm Location sample No. pH Ca Mg p K Zn Japan, Arashiyama 1 4.1 245 294 9.8 112 4.8 2 4.5 194 353 8.3 135 6.1 3 4.6 126 252 8.2 63 3.2 4 4.4 115 86 8.1 78 2.6 Mean 4.4 170 246 8.6 97 4.2 United States, Laredo 1 7.5 3000 250 20.5 345 "high" 2 7.8 3000 250 21.0 470 "medium" 3 7.3 2910 250 21.0 470 "high" 4 6.5 2450 250 7.0 520 "high" 5 8.1 3000 250 34.5 445 "medium" 6 7.8 3000 245 14.0 385 "high" Mean 7.5 2983 249 19.7 439 - -

Number of Food Items and Major Food Plants

At the end of the first month, the Arashiyama West monkeys consistently utilized 21 i tems--19 plant species, insects, and soil. This represented about 10 ~o of the estimated 250 species on the transplant site. By the second month, 36 items were taken; all the previously taken food items plus 14 new plants and some bulbs. In both May and June, 37 items were consumed, although not the same 37 for both months. In July only 31 items were used, in August, 27.

Four food plants comprised over one-half the total intake of native forage: Sorghum, guajillo (Acacia berlandieri), black brush (Acacia amentacea), and prickly pear cactus. All species were available throughout the study except sorghum, which was not present during the first 30 days. Upon release into the outer enclosure, monkeys began using sorghum in quantities greater than any other single plant. This food featured prominently in their diet until the field was totally exhausted by their use in late summer. The second most important food plant was guajillo with buds, flowers, fruits, and leaves used in season. Black brush and prickly pear cactus were consumed in relatively equal amounts. Even though abundant uneaten provisioned foods were present at all times during the first 30 days, monkeys began using native Texas plants for food.

Consumption Patterns and Amounts of Food Consumed

Consumption rates were determined by following nine monkeys to observe the sequence of plants visited, amounts consumed, distances covered, and time spent in picking, ingesting, masticating, and moving (Table 3). These individuals moved from plant to plant eating a variety of foods in succession. They spent 76 ~ of their time moving between food plants and only 24 ~ of the time picking and ingesting foods. Although their activity may vary with the monkey's hunger and condition, consumption rates of the eight food plants recorded in Table 3 were utilized in a remarkably similar way. Even prickly pear cactus with its spines was ingested at a rate similar to grasses and other non-spiny plants.


Table 3. Foraging patterns exhibited by Japanese monkeys (N = 9), Arashiyama West, in the brushlands of south central Texas, April-July, 1972

Total time Picking & Masticating Distance No. of observed Age ingesting & moving moved plants Plant species

Date (min) & sex (min) (min) (m) visited utilized 4/22 5.5 Ad. female 2.0 3.5 18 2 Black brush,

guajillo 4/22 3.5 Juv. male 2.0 1.5 4 3 Black brush,

prickly pear 4/23 3.0 Juv. female 0.5 2.5 9 3 Black brush,

guajillo, grasses 4/23 2.5 Ad. female 2.0 0.5 0 1 Guajillo 4/28 1.5 Ad. female 0.5 1.0 12 1 Grasses 4/28 1.0 Ad. female 0.5 0.5 4 2 Buffel grass,

guajillo 5/I1 23.0 Ad. female 4.0 19.0 120 16 Buffel grass, black

brush, sorghum 5/22 7.0 Ad. female 1.5 5.5 29 6 Guajillo, Panicum

sp. 6/4 16.0 Ad. female 2.5 13.5 35 13 Buffel grass,

guajillo, sorghum Means 7.0 1.7 (24 %) 5.3 (76 %) 20.6 5.2

Seasonal trends in monkey food uses generally corresponded to periods of maxi- mum availability of the food items (Table 4). Shortly after the t roop's release in late winter, few insects and seeds were available. As flowers developed and seeds ripened and insects became more common, consumption of these items increased. The most heavily used plant species showed consumption at several phenological stages (Table 4).

Use patterns shifted markedly in response to availability of various parts. Bark and leaves of guajillo and black brush, of course, were present at all times, but were not used significantly except when other parts, such as buds, flowers, or seeds, were absent. Early in the year (February 23-March 24) flowers were used predominantly; these were replaced in use sequence by beans (March 25-April 30), and later by bark (June 15-July 31) and in late summer by leaves (August 1-31).

Even though abundant uneaten provisioned foods were present at all times during the first 30 days, monkeys began using native Texas plants for food more and more, until estimates showed that over 50 ~ of the total daily intake (by weight) was native foods (Fig. 2).

Estimates were made of food amounts monkeys consumed (Table 5). Total food intake was a combination of provisioned and natural foods.

From May 22 to June 11, 11 adult females were followed during their entire morning foraging sorties; their feeding periods ranged between 1 and 2 hr (7-10 A.M.). Immediately after a monkey ate a food plant and moved on, I picked a like amount from an adjacent plant and labeled it. These food items were weighed, dried in an oven for 4 hr at 121~ and reweighed. Foods consumed during the mid-day and evening foraging were also observed and weights estimated. This was the basis for estimating the amounts by weight of native foods eaten. The provisioned grain and fruits were also weighed. From these measurements, the "average" adult female ate about 60 g in

404 T.W. CLARK

Table 4. Seasonal use and phenology of guajillo and black brush by plant part by Japanese monkeys of Arashiyama West in brushlands of southcentral Texas (Figures are frequency of use and percent frequencies of use.)

Feb. 23-Mar. 24 Mar. 25-Apr. 30 May 1-31

Black Black Black Guajillo brush Guajillo brush Guajillo brush

Buds 40 (20.3) 6 (6.7) 12 (4.1) - - Flowers 72 (36.5) 47 (52.8) 26 (8.9) - - Beans - - - - 141 (48.3) 160 (87.0) Bark 49 (24.9) 19 (23.3) 73 (25.0) 24 (13.0) Leaves 36 (18.3) 17 (19.1) 40 (14.0) - - Total 197 (100) 89 (100) 292 (100) 184 (100)

Guajillo Flowers Seeds x x

Black brush Flowers Seeds - - x - - - - x

m w

86 (56.2) 52 (76.5) 48 (13.4) 16 (23.5) 19 (12.4) - -

153 (100) 68 (100)

most drop off pods split open

x : Peak production. (continued)

the morning, 25 g at mid-day, 275 g at provisioning, and another 50 g in the evening, for a total of about 400 g wet weight each day.

During the first 30 days, provisioning was similar to that in Japan just prior to the transplant, 7.6 kg of whole corn plus 3.5 kg of whole wheat near 8 A.M. and a like amount at 5 P.M. The 5 P.M. feeding also included either an apple, orange, or potato

Z

- 2 <r I - . r ~

0 , - I - -

PROVISIONING TIMES t 2

IO

O " I I I I I I i I ! I I r

,oo

I l l l l l l l l l l l

~ IO0

" ; ' , b ' , ' 2 ' ~ ' ~ ' ~ '

HOURS OF DAY

Fig. 2. Forag ing t imes for A r a s h i y a m a West macaques in Texas. Hour ly sample sizes in all cases exceeded 1,000 observa t ions recorded over at least ten days dur ing Feb rua ry 2 3 - M a r c h 24, 1972.


Table 4. (continued) June 1-30 July 1-31 Aug. 1-31

Black Black Black Guajillo brush Guajillo brush Guajillo brush

67 (24.4) 52 (41.3) 39 (12.4) 28 (25.0) 18 (4.7) - - 119 (43.3) 24 (19.0) 236 (85.1) 63 (56.2) 82 (21.3) 86 (34.1) 89 (32.4) 50 (39.7) 39 (12.4) 21 (18,7) 284 (73.9) 154 (61.1)

275 (100) 126 (100) 314 (100) 112 (100) 384 (100) 252 (100)

most drop off seed pods split open

Table 5. Estimated amounts of food consumed by Arashiyama West monkeys in Texas during the first six months (February 23-August 31, 1972)

Percent Estimated Percent Provisioned foods Total estimate intake estimate

No. (kg) provisioning of total natural of total Total Periods days Grain Fruits (kg) intake foods (kg) intake (kg) Feb. 23-Mar. 24 30 945 540 1,485 77% 450 23% 1,935 Mar. 24-Apr. 30 37 583 666 1,249 48~ 1,350 52% 2,599 May 1-Aug. 31 135 1,093 2,430 3,523 332,o 7,200 67~ 10,723 Total 202 2,621 2,636 6,257 41~ 9,000 59% 15,257

for each monkey. This was more food than the monkeys consumed. After release into the enclosure on March 25, the morning feeding was suspended. From May to Sep- tember, provisioned grain was reduced by one-half although the volume of fruits or vegetables remained the same. Food reductions coincided with the phenological ap- pearance of many plant species.

The largest single food item was cultivated sorghum. The 3.2 ha sorghum field produced about 6,300 kg, all of which was consumed. Portions of the field used very early in the season regenerated by mid-summer so that monkeys used such areas twice. It was more difficult to estimate the weight of native Texas vegetation consumed, but I estimated about 2,700 kg was eaten by August 31.

It was estimated that each monkey consumed about 0.5 kg each day, or nearly 75 kg was consumed by the troop daily. The total weight of all troop members at about 900 kg at the time of the transport, the 75 kg daily food intake represents 8.3 ~o of the t roop 's biomass. This percentage is almost twice as large as consumption figures for rhesus monkeys at the Wisconsin Regional Primate Research Center (HOUSER, pets. comm.).

Feeding Behavior

Because of their unique morphology, many plants required equally unique handling by the monkeys.

406 T.W. CLARK

Usually food items were picked manually, although certain foods were removed from the plant solely by oral manipulation. Often foods were sniffed before being ingested. Foods sometimes were temporarily stored in the cheek-pouches.

Food Plants

Guajillo, a leguminous shrub, is almost spineless. It reaches heights of over 2.5 m and diameters of 2.5 m, although most plants are not this large. Flowers are in glo- bose heads, and flattened bean pods are four to six times as long as wide. This was the single most important native food plant; all parts except roots were consumed. Leaves, buds, and flowers were plucked with the hands or removed by biting. Bark was broken offwith canines or incisors and peeled off. Bark strips were turned over with the hands and the inner surface placed in the mouth where the cambium layer was removed orally. Seed pods were manually removed and torn open with teeth and hands. Beans (usually 5-9 per pod) were eaten and occasionally the pods were also consumed.

Prickly pear cacti possess large numbers of needle-like spines, frequently over 5 cm long, each surrounded at its base by a small clump of tiny glochid spines. This feature did not prohibit monkey use, however. Their small dextrous hands easily manipulated the pads and seemed to avoid spines, although occasionally a monkey paused while eating to look at his palms and remove a number of glochids with teeth or the other hand. Monkeys frequently stood bipedally close to the plant and pulled at a pad which usually broke off after a few tugs. Once removed cactus pads were frequently dragged over the ground to break off spines. Through manipulations with the teeth and hands, the outer tissue layers of the pad were stripped away, exposing the favored inner tis- sues. I,n one series of observations, a monkey ate three pads in 15 rain. Buds, flowers, and "prickly pears" (i.e., fruits) were removed in a manner similar to pads, although occasionally they were bitten right off the plant. The cholla cactus tasajillo has cylin- drical stems comprised of short, brittle joints, 6-12 mm in diameter and few meters long. Whenever the plant is mechanically disturbed, these joints readily fragment. It has many branches and grows about 1 m tall and over one-half meter in diameter. Short spines covers the plant. The ripe scarlet fruits were readily consumed; small portions of new stem joints were also eaten. Picked fruits as well as stem joints were vigorously rolled between the palms a few times before ingestion, presumably to remove any small spines.

Yucca treculeana (Yucca), a unique plant in the Texas landscape, reaches 1.5 m. The upper one-half of the single stalk is covered by a dense growth of narrow, sharply pointed leaves 0.5 m long which project radially from the stem. A large, spiked flow- erhead, sometimes nearly 1 m tall, tops this plant and gives way to dense clumps of seed pods. Monkeys climbed taller trees nearby and jumped with great accuracy, sometimes over 3 m, landing directly at the base of the flower stalk. With poor aim a monkey would have impaled himself on the sharp leaves. Monkeys also climbed yuc- cas staying close to the stem and the base of the leaves, thereby avoiding the sharp leaf tips.

Animal Matter

Insects were taken in large numbers and a few spiders were also eaten. One indivi-


dual was seen eating an unidentified snail. Three juveniles captured a lizard, but it was not eaten. One monkey was seen carrying a broken egg shell, and another egg shell species was found broken; it was not known if the monkeys ate the egg's contents.

Grasshoppers and leaf hoppers were promptly placed in the mouth. On several occasions, monkeys removed a grasshopper's abdomen, ate it, and discarded the remainder.

Monkeys consumed soil daily during the hotter months. Soil was either excavated with the hands or picked up from the surface.

FORAGING PATTERNS AND MOVEMENTS

The time spent and locations of feeding sites varied somewhat with seasonal changes in distribution of foods.

In Corral (0.6 ha)

During the first 30 days, the troop was confined to the corral, hourly surveys were made of the numbers of monkeys eating and the specific food items used (Fig. 2). The three daily peaks of feeding activity were only partly regulated by provisioning, since native foods were constantly available. At least 5 ~o of the population was eating at all times during the day; this sometimes rose as high as 25 ~o except during provisioning when the entire troop was eating.

The total foraging time which exceeded the time actual spent eating, was about 1-2 hr in the morning (7-10 A.M.), 1 hr in mid-day (12-1 P.M.), and 1-2 hr in the evening (between 4:30 and 7:30 P.M., in about 45 min eating provisioned foods and remainder on native foods).

Natural foods were utilized more than provisioned foods during 7 of the 11 hourly intervals sampled daily, provisioned foods were used more during 3, and 1 hr showed relatively equal amounts eaten (Fig. 2). Collective use of 18 species showed three major feeding times. Guajillo was used throughout the day in large amounts with increased utilization during the three major daily feeding periods. Cactus and black brush were most heavily used in the early morning and evening. Ten species lumped together and plotted as a group were used mainly in mid-day and lightly in early evening. Grasses were used throughout the day with peak periods in late after- noon and mid-morning.

In Enclosure (42 ha)

The troop exhibited the same three feeding periods after it was released into the outer enclosure (March 25) and provisioning was greatly reduced. Variations in this pattern were influenced by weather conditions. On rainy days, feeding periods were shortened, and during hot summer months, the noon feeding period was shortened or omitted.

During foraging, individuals moved in a loosely organized group and not as a com- pact troop. Frequently, the troop spread itself over 100 m in diameter. When moving, the troop often o:cupied an area about 30 m wide and 100 m long. Seldom was a

408 T.W. CLARK

"leader" discernible; sometimes Dai (lst ranking male) was near the front and frequently W (2nd ranking male) brought up the rear of the central portion of the troop which was always followed by an all male group.

Although foraging trips took the troop over most of the enclosure, marked pref- erences were shown for certain areas. The corral served as a focus where nights were spent and provisioning occurred.

DISCUSSION

Transplanted to an environment completely different from their evolutionary his- tory, the Arashiyama macaques provided an opportunity to assess aspects of the species potential for adaptability. Adaptive responses of Arashiyama West to potential foods of the transplant site included the following:

(1) Numerous individuals began almost immediate use of several plant species as food. Use of new foods spread rapidly throughout the troop. Six months after the transplant at least 54 food items were being consumed. The acquisition and propagation of new foods, by age, sex, family, social rank, and other factors will be reported later. Utilization of new foods was not motivated by "hunger" as abundant foods were provided for the first 30 days after transplant. Possibly a nutritional deficiency was a motivating factor.

Food ecology and behavior studies of Japanese macaques under natural conditions have not been widely published. SuzuKr (1965) examined a troop's winter foods, IZAWA and NISHIDA (1963) investigated food uses of a troop at the northern limits of the species' range, and HAZAMA (1968) identified 192 food plants of the Arashiyama monkeys.

Of at least seven Japanese monkey troops transplanted, little has been published on their adaptability to new environments. The responses shown by Arashiyama West sharply contrasted with the artificially formed 82-member Ohirayama Troop (KAwAI, 1960). There an initial die-off was attributed to the relatively rigid food culture of the group's members which would not allow them to find edible plants. The monkeys were very reluctant to try new foods even when they were starving (KAWAI, 1960).

The greater adaptability of Arashiyama West monkeys may have been influenced by the long established, well integrated nature of the troop. Also, because monkeys were frequently in visual contact with one another while confined to the small corral, they had the opportunity to observe other monkeys eating new foods. This may have promoted the relatively rapid spread of new foods throughout the troop.

(2) As new foods became available phenologically, they were immediately utilized. The local distribution and abundance of foods partially accounts for the troop's movements to feeding sites away from resting and sleeping areas. Detailed aspects of resource distribution and troop movements will be reported later. Similar patterns of group movements and the relationship to resource distribution and phenology have been noted by STRUHSAKER (1974) and WASER (1975).

Gross aspects of the nutrient content of plant species eaten by the Arashiyama


West monkeys are known, but digestibility of the plants was unknown during this study so it was impossible to determine just what quantities of what nutrients were available to the monkeys. A wide variety of plants with wide ranging nutrient contents were used by the monkeys. At least in gross terms, the troop did not appear to be nutritionally deficient during the spring and early summer of 1972. Four lines of evi- dence support this conclusion. First, the animals looked and behaved as though they were in good health. Second, a number of monkeys weighed in June had all gained weight. Third, a number of peripheral males kept from any provisioned food by central troop members for over four months and forced to depend only on native Texas foods appeared to be healthy. Fourth, the birth rate was comparable to previous years in Japan.

(3) The troop foraging strategy consisted of using a general area for several days to a few weeks, then moving to another area for a similar time, and then on to another new area. A similar pattern was noted in a stumptail macaque troop transplanted to a small island off Mexico (ESTRADA & ESTRADA, 1976).

The strategy used by individual monkeys included visiting a variety of plants in succession and taking a few food items of each plant. Often several plants were by- passed before additional food was consumed. Approximately 76 ~ of the foraging period was spent moving from plant to plant and 24 ~ spent picking food and ingesting. For every 20 m traveled approximately five plants were visited. Individuals tended to feed three times daily.

Evolution has clearly shaped the Japanese macaque into a highly omnivorous and behaviorally flexible animal. A portion of the species adaptive potential was documented here. Ultimate causes for this flexibility may have arisen as the species was, over evolutionary time, associated with complex and seasonally and annually varying environments. These varying environments may have been sufficiently unpredict- able that the species' adaptive strategy to these habitats resulted in evolution of a generalized, flexible behavioral complex.

Acknowledgements. The Arashiyama West transplant required major efforts in scientific organization and cooperation. To all Japanese and Americans who contributed, thank you. Dr. JOHN EMLEN offered invaluable encouragement and support. Mr. TETSUZO MANO'S friend- ship and extensive knowledge of the troop was greatly appreciated. Dr. GORDON STEPHEN- SON'S physical support, good cheer and humor aided in completion of this study. Dr. MIKE WELLS and Ms. DENISE CASEY provided critical advice on the manuscript. Ms. CHERYL HUGHES produced the graphic work. This project was supported by grants to Dr. EMLEN from the Grant Foundation, the National Science Foundation, the Wisconsin Regional Primate Research Center, and the Department of Zoology at the University of Wisconsin. Ms. SUSANN KOHLER, Biology Department of Idaho State University typed the manuscript.

REFERENCES

CASEY, D. E. & T. W. CLARK, 1976. Some spacing relations among the central males of a ~ransplanted troop of Japanese macaques (Arashiyama West). Primates, 17 : 443--450.

CLARK, T. M. t~ T. MANO, 1975. Transplantation and adaptation of a troop of Japanese

410 T . W . CLARK

macaques to a Texas brushland habitat. In: Contemporary Primatology, S. KONDO, M. KAWAI, & A. FHARA (eds.), S. Karger, Basel, pp. 358-361.

ESTRADA, A. & R. ESTRADA, 1976. Establishment of a free-ranging colony of stumptail macaques (Macaca arctoides) : Relations to the ecology 1. Primates, 17 : 337-355.

HAZAMA, N., 1968. Plants of Arashiyama and Plants Eaten by Wild Japanese Monkeys. Iwata- yama Monkey Park Publ., Kyoto.

IZAWA, K. & T. NISHIDA, 1968. Monkeys living in the northern limits of their distribution. Primates, 4(2): 67-88.

KAWAI, M., 1960. A field experiment on the process of group formation in the Japanese monkey (Macacafuscata), and the releasing of the group at Ohirayama. Primates, 2: 181-255.

MORRISON, J. S. & E. W. MENZEL, JR., 1972. Adaptat ion of a free-ranging rhesus monkey group to division and transplantation. Wildl. Monogr., 31 : 1-78.

STRUHSAKER, T. T., 1974. Correlates of ranging behavior in a group of red colobus monkeys ( Colobus badius tephrosceles). Amer. Zool., 14: 177-184.

SoztrK1, A., 1965. An ecological study of wild Japanese monkeys in snowy areas--Focused on their food habits. Primates, 6: 31-72.

WASER, P. M., 1975. Monthly variations in feeding and activity patterns of the mangabey, Cercocebus albigena (Lyddeker). East Aft. Wildl. J., 13:249-263.

- - R e c e i v e d June 8, 1978; Accepted December 2, 1978

Author's Present Address: TIM W. CLARK, Department of Biology, Idaho State University, Pocatello, Idaho 83209, U.S.A.; reprints to: Box 2705, Jackson, Wyoming 83001, U.S.A.

food adaptations of a transplanted japanese macaque troop (arashiyama west)

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