Rev. Biol. Trop., 35(2): 287-297, 1987.
Troop movement and food habits of white-faced monkeys in a tropical-dry foresto
Daniel Moscow* Associated Colleges of the Midwest, Apartado 1026 5, San José, Costa Rica. Christoper Vaughan Posgrado en Manejo de Vida Silvestre, Universidad Nacional, Heredia.
(Received December 12, 1986)
Abstract: Home-range, movement and activity patterns and diet of a group of 16 white-faced monkeys (Cebus capucinus) were studied in a Costa Rican tropical dry forest at the end of the dry season (March and April) cf 1982. The troop used an area of .6 7 km2 and moved an average of 4.5 km ± 0.6 daily. Three basic daily activities were identified: resting, moving and feeding. Resting was greatest between 1000-1500 hours. Movement, which was strongly correlated with feeding, occurred most between 0500-0900 hrs and 1300-1800 hrs. The primates were omnivores, feeding on parts of 27 species of plants, four species of insects and an Anolis lizard.
The white-faced monkey or capuchin, Cebus of Nicoya, Guanacaste Province, Costa Rica. capucinus imitatar Thomas (Primates: Cebi The refuge is found at 10030' N and 85030' W. dae), is found in a wide variety of forest types Palo Verde is about 7,542 ha in size and situat from Belize to Panama (Hall 1981). This ed in the tropical-dry forest life zone (Tosi monkey is omnivorous and feeds mainly on 1969). Within the refuge, nine vegetation types fruits and seeds but also on flowers, woody tis were described: mangrove swamp, seasonal sue, young shoots, insects, and small vertebrates marsh, fresh-water flooded forest, lowland de (Defler 1979, Freese 1976, Freese and ciduous forest, deciduous forest on limestone Oppenheimer 1981, Hladik et al. 1971, Oppen hills, evergreen forest, overgrown pasture, Aca· heimer 1960). However, troop movement has cia scrubland and pasture (Vaughan et al. received little attention. The objectives of this 1982). study were to describe dai1y activity patterns Annual rainfall at nearby Puerto Humo for and food habits of a C. capucinus troop in a the 1961 to 1970 period averaged 2296 mm, tropical-dry forest during the latter part of the 91% of which fell between May and October. dry season. Mean annual temperatures of 370C are com mono Elevation varies from 1 2 to 232m. The late-dry season months of March and April have STUDY AREA AND STUDY TROOP little rain, relatively abundant fruit production, and extreme temperature differences between The Dr. Rafael Lucas Rodriguez Caballero areas of different vegetation types. National Wildlife Refuge (hereafter referred to The study area was about 100 ha in size and by its popular name, Palo Verde) is located on located on the southern shope of Cerro Guaya the northern bank of the Tempisque River, 20 cán, one of two prominent limestone hills with kilometers from the river's outlet into the Gulf in the wildlife refuge. Three forest types were described in this study area. These included: deciduous, semi-deciduous and evergreen, defi (*) Present Address: Department of Plant Pathology, University of California, Berkeley, California. ned as individual trees retaining respectively,
287 288 REVISTA DE BIOLOGIA TROPICAL
0-25%, 25-75% and greater than 75% of their large-scale directional travel was considered, leaves during the latter part of the dry season. while small-scale movements were exc1uded The study troop consisted of three old and two from the map. Daily distance traveled by the young adult males, three adult fe males, seven troop was calculated by summing all measured juveniles and one infant for a total of 16 indivi distances between consecutive estimated troop duals. positions. Home range was estimated by using METHODS the minimum convex polygon of all troop posi tons calculated (Mohr 1947). The study was conducted from 6 to 25 Statistical notation fo llowed that of Sokal March and from 13 to 29 April 1982. Our ob and Rolhf (1981). servations began about 15 min before sunrise (05 15 hrs) when capuchins were leaving their RESULTS AND DISCUSSION sleeping sites. The troop was followed until about 20 min after sunset (1800 hrs) when it F or each hour of the day, between 220 and = = settled into trees to sleep. A group of more 360 min (x 314.4 min; S.D. 45 .2) of obser than six monkeys was arbitrarily defmed as a vatio n time was collected. The ratio of observa troop and a group of six or less, as a sub-group. tions collected for adults and juveniles was The focal animal technique (Altmann 1974) 60:40, respectiveIy. This approxirnated the age was used for collecting activity data, where a class composition of 50: 50 of adults to juveni single capuchin was initially chosen at random les for the study troop. and fo llowed. Duration of each activity was re corded to the nearest minute which was the Troop home range. During March and April, 2 smallest time unit . Activity was grouped into the troop had ahorne range of 0.67 km . A hourIy periods and for one statistical test, into significantly greater proportion of diurnal activ two periods of fo ur and one of five hours. ity occurred in the evergreen fo rest compared When the fo cal animal separated from the troop to its total area in the study area (x2 = 8.4, dJ. or was lost by us, our observations was directed = 1, P
TABLE 1
Diurnal use of fo rest types by a white·fa ced monkey (Cebus capucinus) troop. Palo Verde. (March·April, 1982) Forest Type Home Range Time Spent in Forest Type ha % hrs % Evergreen 3.3 ( 5) 19.9 (32) Semideciduous 6.0 ( 9) 9.3 (15) Deciduous 22.1 (33) 15.5 (25) Deciduous with evergreen pitches 35.6 (53) 17.4 (24)
Total 67.0 (lOO) 62.1 (lOO)
routes were highly defined, and capuchins On many occasions, especia11y in deciduous moved through specific trees. Others were gen· f o rests, several adult capuchins walked at eralized, and the monkeys trayeled in particular ground level while the remainder of the troop directions via many nearby paths. continued arborea11y. These terrestrial move Troop movements of C. capucinus were ments covered up to 50 m, lasted up to six min observed by Baldwin and Baldwin (1977) on and inc1uded foraging. Baldwin and Baldwin the Burica Peninsula in Panama, with daily path (1977) reported that C. capucinus foraged for lengths ranging from 0.6 to 1.5 km. Clutton insects for distances of more than 20 meters Brock and Harvey (1977) have listed sornedaily along the ground. Moreover, Defler (1979) path lengths for 56 primate species belonging to observed that C. albifrons spent up to 50% of eight families. Our value of 4.5 km.for C. capu their active period foraging at ground level dur cinus exceeded the daily path lengths of all 56 ing the dry season (February) in Colombia. species with the exception of three species of Pap io. In their analysis of fe eding and move Activity patterns. Feeding was significantly ment data for the 56 species, Clutton-Brock greater between 0500-0900 hrs' than in the and Harvey (1977) found a negative association other four and five hour periods of the day (H between daily path length and the proportion = 9.17, dJ. = 2, p 80 - 1 - Elevotion (m.) ------Troil Rood o 100 200 -- - Home ronge , , , METERS O Water hole Sleeping site 0 Resting site -0 Structure Fig. 1. Home range of a white-faced monkey (Cebus capucinus ) troop. Palo Verde. March-April, 1982. this was due to variation in daily temperatures another folIowed changes in the fruiting phe and food sources. nology in sorne cases. For example , ripe Bauhi nia pods were abundant in March throughout Herbivory. Capuchins fe d on 27 plant species the home range, but were absent in Aprll. Brosi (16 families), several insect species, and one ver mum alicastrum fruits were more plentiful in tebrate (Table 2). Plant parts eaten inc1uded; March than Aprll, and Muntingia calabura was seeds, fruits, flowers, flower buds,leaves, young an abundant and much used food source in shoots, sap, and woody tissue. Aprll . Fruit was the primary fo od source in both The percentage of whole food units discar months. Shifts from one plant food so urce to ded or dropped andfor the portion of a unit MOSCO W & VA UGHAN: White-faced monkeys in a tropical dry forest 291 '·0 � ______lOO ------_ ... - ,/ ,. ,- /""--...._-'- .... , / I ,.// I __--� ,./ ,. ,. ,.,/ e . . ::::::::::::::: :� ::::::::::::::::::::::::::::::::::: ::: :::::::. :: ...... ...... •...... o.·.. · . . · ... o •• • •••••••••, .' •••••• '" • ••••••••• •• •••••••••• o ••••••••••• o •••• ••••••• •••••••••••• •• • ••••• •• '" •••••••••••••••••••••• ' ••••••••••••••••••• o • • ' Elevotion (in m. ) ¡ - 180- o 200 ... Forest Troil D Rood 1: : : :1 Morsh , old fields, post ure Trovel Water hole Locotion ot given time - Structure Fig. 2. White-faced monkey (Cebus capucinus) troop. movement. Palo Verde. March 12, 1982. Directed travel =4.5 km. rejected by a monkey varied between feeding average percentages of seeds fo und in picked events for a particular plant food. Discarded fruits after capuchin feeding was 70% in Pithe Bauhinia pods, for example, still contained an cellobium saman, and 33% in Brosimun alicas average 66% of their seeds, but sorne were tmm. An average of 33% of fleshy fruit tissue picked c1ean of seeds. The capuchins ate 5 to of Spondias purpurea was rejected by capu 80% of the fl eshy mesocarp of mangos. The chins. F or C. albifrons, Defler (1979) found 292 REVISTA DE BIOLOGIA TROPICAL TABLE 2 Percent of total feeding time spent on different fo ods by a white·faced monkey (Cebus eapucinus) troop. Palo Verde. (March·April, 1982) a, b/ Speeies Family plant Feeding % of Part Time total MARCH Eaten. (min.) feeding time Fruits and seeds: el Bauhiniaglabra Caesalpiniaeeae s 178 32.0 Mangifera indica '" Anaeardiaeeae f 73 13.1 Brosimumalicastrum Moraeeae 47 8.5 Mu ntingia calabura Elaeoearpaeeae s,m 40 7.2 Spondiaspurpurea Anaeardiaeeae f 40 7.2 Bursera simaruba Burseraeeae f, ex,(s?) 40 7.2 Pithecellobium soman Mimosaeeae 21 3.8 Ficus ovalis Moraeeae s, m 10 1.8 Cecropia peltata Moraeeae fr 7 1.3 Ceiba pentandra Bombaeaeeae 6 1.1 Enterolobium cyclocarpum Mimosaeeae 2 0.4 Diospyros nicaraguensis Ebenaeeae s 2 0.4 Xy losoma flexuosum Flaeourtiaeeae fr 1 0.2 Guazuma ulmifo lia Stercu liaeeae s, (fr?) (4) Mastichodendron capiri Sapotaeeae ? (2) Canavalia sp. Fabaeeae (2) Other Plant Parts: Mastichodendron capiri Sapotaeeae flb 14 2.5 Caesolpinia eriostachys Caesalpiniaeeae lf 13 2.3 Astronium graveolens Anaeardiaeeae sf 7 1.3 Calycophyllum candidissimum Rubiaeeae flb 1 0.2 Unident. wood stick wt 1 0.2 Brosimiumalicastrum Moraeeae sap (1) Enterolobium cyclocarpum Mimosaeeae flb (1) Inseets and Other Animals: d,el .unident., from ground 16 2.9 in/e:ast. 15 2.7 in/b:ast. 11 2.0 lizard (Anolis cupreus) 5 0.9 in/w, termite (Termitida.e); Trema micrantha 4 0.7 in/w: Guazuma ulmifo lia 2 0.4 in/dl.: asto (2) Total feeding time on fruitsand seeds: 467 84.2 Total feeding on other plant parts: 36 6.5 Total feeding time on animals 53 9.6 Total observed feeding time 556 100.3 APRIL Fruits and seeds: Mu ntingia calabura Elaeoearpaeeae s,m 361 54.9 Brosimum alicastrum Moraeeae s 24 3.6 Mastichodendrom capiri Sapotaeeae 20 3.0 Cassia grandis Caesalpinael"ae m, (s?) 14 2.1 Cecropia peltata Moraeeae fr 13 2.0 Ficus ovalis Moraeeae s,m 11 1.7 Xylosoma flexuosum Flaeouritaeeae fr 10 1.5 MOSCO W & VA UGHAN: White-faeed monkeys in a tropical dry forest 293 Species Family Plant Feeding % of Part Time total Eaten. (min.) . feeding time Astronium graveo/ens Anaeardiaeeae ? 10 1.5 Guazuma u/mifo /ia Stereuliaeeae s, (fr?) 10 1.5 Mangife ra indica* Anaeardiaeeae f 9 1.4 Pithecellobium saman Mimosaeeae 8 1.2 Luehea candida Tiliaeeae s 8 1.2 Acacia collinsii Mimosaeeae f,(s?) 3 0.5 Stercu/ia apetata Stereuliaeeae s 2 0.3 Spondias purpurea Anaeardiaeeae f 1 0.1 Piper tubercu/atum Piperaeeae fr 1 0.1 Enter% bium cyclocarpum Mimosaeeae 1 0.1 Other Plant Parts: Ma nilkara zapora Sapotaeeae fl 29 4.4 Enter% bium cyclocarpum Mimosaeeae flb 21 3.2 Caesa/pinia eriostachys Caesalpiniaeeae lf 16 2.4 Pithecellobium saman Mimosaeeae fl 6 0.9 Spondias mombin Anaeardiaeeae st 2 0.3 Enter% bium cyclocarpum Mimosaeeae wt 2 0.3 Trichilia sp. Meliaeeae sap 2 0.3 Mastichodendron capiri Sapotaeeae fl 1 0.2 Inseets and Other Animals: in/b: Enter% bium cyclocarpum 14 2.1 in/b: ast 13 2.0 unident., from ground 13 2.0 in/dI. katydid (Tettigonüdae): Stercu/ia apetata 10 1.5 in/di: ast 10 1.5 in/th: ast 6 0.9 in/w, andt (Formicidae): Ceiba pentandra 5 0.8 in/e: ast 1 0.2 wasp nets of Po /ybia sp. (nest, honey, larvae) 1 0.2 lizard A nolis cupreus Total feeding time on fruits and seeds 506 76.8 Total feeding time on other plant parts 79 12.0 Total feeding time on animals 73 11.2 Total feeding time 658 min 100.0 Key to symbols -al For foods whieh laek feeding time data, the number of feeding events is givenin parentheses. -b/ A question mark in parentheses indieates that the plant part may or may not have been eaten. -e/ Key to plant parts injested s = seed(s), f = fleshy mesoearp and/or endoearp, ex = exoearp, m = sticky or juicy matrix surrounding seeds. fl flower, flb flower bud, leaf, st young shoot, wt woody tissue = = lf= = = sap = sap, fr = complete fruit including seed(s). -dI Key to inseet foods (classified by method of proeur�ment): in/b = Insects obtained by stripping off bark, in/w = Insects obtained by cracking or splitting woody tissue. in/di = Insects obtained through inspection of dead leaves on trees or shrubs, in/th = Insects removed by hand from tree holes, and in/c = Inseets eaught with cupped hand(s). -e/ The plants harboring the insects are listed after the method of procurement symbols. ast = assorted plant sourees. (*) Not native to Guanaeaste. 294 REVISTA DE BIOLOGIA TROPICAL ______� " 0 �__ _� _ 100 . .-'- e / ..- / I I / .. I I • • • • I.- • • • • • • • • f .�. ' ...... ••• • • Rf:FU6E HEADOUARTERS ...... �------. :::::::-':.. :::� : . . : . . : . ... • • • . . • • • • • • . • . . • . • • • • . . • • . • • • . • • . • • • • • • • • • . • • • . • • • • • • • • �'20 ::::::::::::::::�:::::::::::::::::::::::::::::::::::::::::::::.� -180- Elevotion (inm.l Fo rest o 100 200M. ------Troil D Rood . . . Morsh , old fields, posture �---� Trovel O· · o Woter hole .7'39 locotion ot given time - Structure Fig. 3. White-faced monkey (Cebus capucinus) troop. movement. Palo Verde. March 25, 1982. Directed travel =5.1 km. 80% of fo raging time dedicated by monkeys to During the dry season, in a tropical-dry for plant matter and 20% to animal matter. Hladik est in northwestern Costa Rica, Freese (1976) et al. (1971) reported an omnivorous diet for fo und e capucinus to spend 11% of observed capuchins. The diet consisted of 65% fruit, 20% activity time feeding on insects and 26% on animals and 15% shoots, leaves and other vege plants. The sorne author cited the most used tation. fruit and seed sources in March and April as: MOSCOW & VA UGHAN: White-faced monkeys in a tropical dry forest 295 100 ---_ FEEOING ----- RESTlNG ", -•. / , .. - MOVING 90 ' / " / \ / \ 80 I \ I / / \ 70 \ I \ >- / \.----. .... \ :; I \ ' � 60 I \ \ ..,oC / .... " . '" ; , .... 50 / . z / . '" Q. / / ', .. " '" / \ / '" / a 40 ,. � "1 . X· 30 ' / I : I � /\, I I 20 I I I I 10 / I I _---.1 5'00-6'00 7'00- 8 '00 11'00- 10'00 11'00- 12'00 15'00-14'00 15'00 - 16 '00 11'00-18'00 8'00- 7:00 8:00- 9:00 10'00- 11'00 12:00-13:00 14 :00- 15'00 16:00 - 17:00 HOUR 265 355 277 I 312 360 353 380 31 5 300 I 360 337 I 220 273 I I I I I I I I I I I OBSERVAT ION TIME (MINUTES) Fig. 4. Activity pro file of a white-faced monkey (Cebus capucinus) troop. Palo Verde. March-April.1982. Bursera simaruba, Ficus Sp ., Diptodendron cos found carbohydrate source and that searching tarricense, and Luehea candida. Although Bur for insects is difficultunder dim light conditions. sera simaruba and Ficus ovalis were also eaten Feeding on animals in our study involved a in Palo Verde, they were not major food sour variety of behaviors (Table 2). Adult capuchins ces during the study period (Table 2). frequent1y sought insects on the ground. Two Capuchins in Palo Verde spent on1y a small monkeys caught and ate small Anolis cupreus percentage of feeding time on foliage (4.6% in lizards on the ground, one taking fo ur min to March and 3.6% in April). Hladik et al. (1971), consume all but the tail. Baldwin (1977) repor in a comparative study of feeding habits of fo ur ted a capuchin grasping the tail of a large iguana sympatric monkey species on Barro Colorado (Iguana iguana) and after a struggle, breaking Island, Panama, found that the leaf-eating off the end segment and eating it. Capuchins in howler monkey, Alouatta vil/osa, caught in our study located a katydid, and thrust a sects, while C. capucinus and Saquinus geoff cupped hand at the springing insecto During a roy i, spent 20 and 30% respectively, of their one-hour period on 24 April, a young adult feeding time on animal prey. male fed on ten paper wasp nests (Polybia sp .), four of which harbored wasps. Swatting at the Insectivory and Carnivory. Capuchins spent stinging insects, the monkey removed the nests Httle time foraging on animals between thin outer layer and ate the honey and larvae 0500-0700 hrs and 1700-1800 hrs. Freese and ftlled comb. Oppenheimer (1981) suggested that daily feed C. capucinus demonstrates considerable ingbegins with fruits because they are a readily manual dexterity in removing seeds from fruits, 296 REVISTA DE BIOLOGIA TROPICAL extracting insects from woody tissue, catching December and May-June when movement and insects and other small animals, and other feed home ranges increased and less than optimal ing behaviors. The superior manual abilities of foods were found in fe ces. this species were noted by various authors (Freese 1976, Jolly 1972, Moynihan 1976). RESUMEN Drinking. The waterhole in the study area El radio de acción, patrones de movimiento, was found within the evergreen forest (Fig. 1) actividad y dieta de una manada de 16 indivi in close proximity to major sleeping and resting duos de mono carablanca (Cebus capucinus) sites. Both evergreen forest and water availabili fueron estudiados en un bosque seco-tropical de ty influenced capuchin home range and move Costa Rica a finales de la época seca (marzo y ments during the dry season. At least during the abril) de 1982. La tropa se desplazó en un are a dry season in Palo Verde we agree with Freese de 0.67 km2 , y recorrió un promedio de 4.5km (1978) that Cebus troops are limited to areas ± 0.6 diarios. Se identificaron tres actividades where water is presento basicas diarias: descanso, movimiento y alimen tación. Descansaron más entre las 1000-1500 Drinking was observed during all five study hrs. El movimiento de la manada estaba fuerte days in Apnl, but on on1y two of five days in mente correlacionado con la alimentación y March. Our presence might have affected the ocurrió más entre las 0500.0900 hrs y monkey's behavior more in March when the 1300-1800 hrs. troop was less accustomed to us In April, adult Los primates son omnívoros y consumieron males drank from one to three times daily be partes de 27 especies de plantas, cuatro de in tween 0641 and 1736 hrs while females and sectos y una lagartija del género Anolis. juveniles drank less often, perhaps because they were move timid to our presence. Capuchins approached the waterhole cautiously, and of ACKNOWLEDGMENTS the three occasions in which the first drinker was observed, the leader was an adult maleo We thank Colin Chapman, Curtis Freese, Freese (1978) found that adult males led drink Winnifred Hallwachs, Bruce Marcot and Michael ing processions. McCoy for comments on the manuscript, Jorge Barrantes for typing severaldraftsand Miguel Ro Stress periods. This study was carried out dríguez fo r plant identifications. Friendship of during the latter part of the dry season, when workers and scientists in Palo Verde was appre water was a limiting factor and may have ciated during the study periodo The Associated caused troop concentration around waterholes Colleges of the Midwest, National University in addition to defense of these water sources by and Organization of American States provided the monkeys. We did not observe any other.C. logistical support. The project is part of Natio capucinus troops in the evergreen forest during nal University research project No. 782085 this study. F ood sources seemed sufficiently "Study and management of wildlife in Costa abundant for the monkeys, as evidenced by the Rica" . prolonged resting periods taken around midday. McCoy (1985) found that collared peccary REFERENCES (Tay assu tajacu) herds in Palo Verde showed variation in movement patterns, home range Altmann, J.S. 1974. 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