Jaguar (Panthera onca) Food Habits in Atlantic Rain Forest of ...

BIOTROPICA 33(4): 691–696

2001

Jaguar (Panthera onca) Food Habits in Atlantic Rain Forest of Southeastern Brazil1 Ricardo C. Garla2 Universidade Estadual Paulista, Rio Claro, Saõ Paulo 13506-900, Brazil Eleonore Z. F. Setz Universidade Estadual de Campinas, Departamento Zoologia, C.P. 6109, Campinas, Saõ Paulo, 13083-970, Brazil and Nivar Gobbi Universidade Estadual Paulista, Rio Claro, Saõ Paulo 13506-900, Brazil

ABSTRACT Between January and December 1996, the food habits of a relict population of jaguars were studied in 220 km2 Linhares Forest Preserve, which comprises much of the remaining old-growth Atlantic Forest of Espı´rito Santo, Brazil. Fecal analysis indicated opportunistic feeding on 24 prey species (N 5 101 scats). Mammals represented 87 percent of the total items, followed by reptiles (9.8%) and birds (2.8%). Considering prey weight, 23.4 percent of the items weighed 1–3 kg, 40.5 percent were 3–10 kg, and 27.7 percent weighed more than 10 kg. Analysis of relative prey frequency and biomass indicated that the diet was concentrated in two prey types: long-nosed armadillo and whitelipped peccary. Literature data suggest that forest jaguars rely on the same mammal prey over their entire geographic range.

RESUMO Entre janeiro e dezembro de 1996, os ha´bitos alimentares de uma populaçaõ da onça pintada foram estudados na Reserva Florestal de Linhares (220 km2), que compreende 25 porcento da floresta Atlaˆntica prima´ria remanescente do Estado do Espı´rito Santo, Brasil. A ana´lise de fezes indicou o consumo de 24 espećies de presas (N 5 101 amostras). Mamı´feros representaram 87 porcento do total de itens, seguidos por re´pteis (9.8%) e aves (2.8%). Considerando o peso das presas 23.4 porcento dos itens pesavam entre 1 e 3 kg, 40.5 porcento de 3 a 10 kg, e 27.7 porcento acima de 10 kg. De acordo com a ana´lise de frequëˆncia relativa de ocorreˆncia e a estimativa de biomassa relativa, a dieta concentra-se em dois itens: o tatu galinha e queixada. A onça pintada parece basear sua dieta nas mesmas espećies de mamı´feros ao longo de sua distribuiçaõ geogra´fica. Key words:

Atlantic rain forest; Brazil; food habits; jaguars; Panthera onca; scat analysis.

THE JAGUAR (PANTHERA ONCA) is the largest extant Neotropical felid (Seymour 1989). Until the late 1970s, most of the information available on the species were anecdotal accounts of explorer–naturalists (Perry 1970) and hunters (Almeida 1976). This situation has changed with the recently published studies on jaguar biology in the wetlands of the Brazilian Pantanal (Schaller & Vasconcelos 1978, Schaller & Crawshaw 1980, Crawshaw & Quigley 1991) and Venezuelan Llanos (Hoogesteijn et al. 1992); forests of Belize (Rabinowitz & 1 Received 19 November 1999; revision accepted 18 December 2000. 2 Corresponding author: Rua Amador Bueno, 1342/120, Centro, Ribeiraõ Preto, Saõ Paulo, 14010-070, Brazil; Email: [email protected]

Nottingham 1986, Watt 1987), Peru (Emmons 1987), Mexico (Aranda 1994) and Costa Rica (Chinchilla 1997); and in the Paraguayan Chaco (Taber et al. 1997). The status of the jaguar in eastern South America, however, was totally unknown until recently, when Crawshaw (1995) concluded a study on its ecology in a subtropical forest of southern Brazil and Facure and Giaretta (1996) presented their data on carnivore food habits. Eastern Brazil represents the most critical region for jaguar survival (Oliveira 1992). This region encompasses the most densely populated and highly developed part of South America, and most of the natural vegetation that survives is isolated in forest fragments (Oliver & Santos 1991). Jaguars are still found in several regions, but it is uncertain

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if populations will persist. This situation clearly indicates the urgency of studies on jaguar biology in this region. An assessment of the food base is essential to predict the viability of jaguar persistence in its present habitat and to plan for its future. This investigation was developed to provide a description of jaguar food habits in a preserved forest in the state of Espı´rito Santo, which due to its large size protects one of the largest jaguar populations in Eastern Brazil. To the best of our knowledge, our study is second only to that of Rabinowitz and Nottingham (1986) in Belize and is the first comprehensive study of jaguar diet in the Atlantic rain forest of Brazil.

STUDY SITE The study was conducted at Linhares Forest Preserve (LFP; 198039–198169S, 398539–408059W), during 11 months between January and December 1996. The 220 km2 LFP is located on coastal lowland Atlantic Forest (30–60 m elev.) and is contiguous with the 242 km2 Sooretama Biological Reserve (SBR), the largest conservation area in Espı´rito Santo. Together they comprise 50 percent of all old-growth forest remaining in this state and are considered to be among the most important nature reserves in eastern Brazil (Oliver & Santos 1991). LFP consists of a mosaic of habitat types: primary lowland forest (137 km2) with canopy trees up to 40 m high; secondary forest (11 km2); gallery forest (9 km2); ‘‘mussununga’’ (17 km2) forest with 5 to 15 m high trees growing on sandy soils); coastal sand scrub (13 km2); brush (3 km2); and wetlands (17 km2). The remaining 11 km2 is comprised of administration and research facilities (Peixoto & Gentry 1990, Oliver & Santos 1991). SBR has a similar vegetation. The climate is classified as tropical wet (Ko¨ppen) with a mean annual temperature of 238C. The mean annual precipitation of 1383 mm occurs mostly between December and March. Mean temperature during the study was 23.88C, and rainfall in 1996 was 892.5 mm with peaks in March (148.3 mm) and November (273.4 mm). June (16.7 mm) and August (13.5 mm) were the driest months (LFP Meteorological Station). Outside the study area, land use includes coffee, sugarcane, cocoa, and tropical fruit plantations, and cattle ranching.

METHODS Jaguar diet was studied by fecal analysis. Scats were collected along the 117 km of dirt roads in LFP

monthly (except March) during 1996. We also surveyed 50 km of roads in the contiguous SBR from April to October. Survey yields between reserves were compared in terms of number of scats collected per 10 km, and the number of species consumed in SBR was compared to random samples drawn from the LFP data set. Jaguar scats were distinguished from those of puma (Puma concolor) using associated tracks, hair ingested while grooming, and circumstantial evidence. Scats were stored in plastic or paper bags and dried. In the laboratory, they were broken up and washed with water over a fine-mesh screen, dried, and examined under a stereomicroscope. Macroscopic hair characteristics were compared with a reference collection of hair from known species. Hard parts (bones, teeth, nails, and scutes) were compared to photographs or museum specimens of mammals and reptiles. Scat content was expressed in terms of percent occurrence (the number of times a particular prey species was found as a percentage of all species identified, %Oc 5 N 3 100/142). For prey weighing more than 2 kg, we estimated the number of animals killed from occurrence frequencies using the empirical relationship found by Ackerman et al. (1984) for pumas. We followed Stoen and Wegge (1996) in assuming that the digestive systems of large felids were similar to one another. This technique corrects for both the increased production of scats and the greater probability that a scat will not contain recognizable prey parts as prey size increases. The method permits estimating the relative biomass consumption and the number of individuals killed for each prey type. This analysis was based on the 50 scats that contained only a single recorded prey item. Prey identified from different scats were assumed to represent independent captures, unless there was evidence that the same individual was represented in separate scats. We recorded eight cases in which two scats containing the same prey item were found close together on the same road. We counted each of these as a single occurrence. Three scats containing jaguar hairs with unidentifiable matter were not considered in the analysis.

RESULTS At least three resident jaguars, as suggested by track surveys, produced the 101 collected scats. The mean number of jaguar scats found during monthly three-day surveys at LFP was 6.8, with a minimum of 2 and maximum of 16. The number of

Jaguar Food Habits in Southeastern Brazil

TABLE 1.

Percent occurrence of prey species found in 101 jaguar scats in Linhares Forest Preserve, Brazil, from January to December 1996 (N 5 142 identified prey items). Prey species

Percent occurrence1

White-lipped peccary (Tayassu pecari) Armadillo (Dasypus spp.) Coati (Nasua nasua) Crab-eating racoon (Procyon cancrivorus) Collared peccary (Pecari tajacu) Tortoise (Geochelone denticulata) Other armadillos2 Juvenile peccaries3 Paca (Agouti paca) Deer (Mazama spp.) Agouti (Dasyprocta leporina) Sloth (Bradypus variegatus) Kinkajou (Potus flavus) Capybara (Hydrochaeris hydrochaeris) Cat (Leopardus spp.) Capuchin monkey (Cebus apella) Unidentified bird Tapir (Tapirus terrestris) Porcupine (Chaetomys subspinosus) Unidentified river turtle Colubrid snake Lizard (Tupinambis merianae) Currasow (Crax blumenbachii) Bird (Falconiformes) Total 1 2 3

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15.0 15.0 10.0 8.6 7.8 7.8 7.0 4.2 4.2 2.8 2.1 2.1 2.1 1.4 1.4 1.4 1.4 0.7 0.7 0.7 0.7 0.7 0.7 0.7 99.3

Months of occurrence all months except Apr., Oct. all months Jan., Apr., May, June, Sept., Dec. Feb., July, Aug., Sept., Oct., Nov., Dec. Jan., Feb., Apr., May, Sept., Nov. Jan., Feb., Mar., Apr., May, June Feb., Apr., May, Aug., Sept., Oct. May, Sept. Apr., May, Aug., Sept., Oct., Dec. Apr., May, Aug. Feb., Apr. Aug., Sept. Sept. Dec. Sept. Dec. May, July Aug. Sept. June Jan. Jan. Sept. Sept.

Number of times a species was identified as a percentage of all prey items identified. Includes Euphractus sexcintus (1.4% of total items), Cabassous unicinctus (0.7%), and unidentified armadillos (4.9%). Juvenile peccary species could not be identified by hair characteristics.

scats collected per month changed according to the survey period: two days in January (N 5 2 scats), two days in July (N 5 4), and seven days in December (N 5 9). Scat collection was also influenced by decreased preservation during rainy months (November, N 5 3 scats) and intensification of marking by residents, apparently in response to other jaguars (May, N 5 15, September, N 5 16). During the dry months (February–September), both fresh and old scats were found. As roads were washed frequently during the wet season (October–January), scats collected during this period were usually not more than a few days old. Surveys from April to October yielded 5 scats/10 km for pooled data from the two preserves (with a total of 25 scats collected in SBP vs. 60 in LFP). Ten scats (9.9%) were associated with jaguar tracks, and 2 scats (2%) with ground scrapes. Jaguar hairs ingested while grooming were found in 18 scats (17.8%). Of these, 3 scats (2.9%) contained only jaguar hairs and identifiable fecal matter. Plant material was found in 30 (29.7%) scats. The 101 jaguar scats with identifiable animal prey from LFP contained a mean of 1.6 prey items

each, representing 24 taxa (14 taxa in 60 scats vs. 10 taxa in 25 scats from SBR). A total of 142 items was identified (Table 1). Considering prey weight, 23.4 percent were 1–3 kg, 40.5 percent were 3–10 kg, and 27.7 percent weighed more than 10 kg. Mammals comprised 87.3 percent of the items, followed by reptiles (9.8%), and birds (2.8%). Among mammals, 76 percent were species from three orders: Artiodactyla (30% of all items), Xenarthra (24%), and Carnivora (22%). Six species had percentage occurrences above 5 percent: white-lipped peccary (Tayassu pecari), armadillos (Dasypus spp.), coati (Nasua nasua), crab-eating racoon (Procyon cancrivorus), collared peccary (Pecari tajacu), and tortoise (Geochelone denticulata; Table 1). Seventyfive percent of the scats with tortoise remains were found in the northern swampy region of the preserve occupied by a large male jaguar. In contrast, the jaguar’s main prey at SBP were armadillos, coati, and deer (Mazama spp.) (22.6% each). For the 50 scats containing only one item that were corrected for relative biomass consumption, we found armadillos (all species), peccaries (both species), and procyonids to be the main prey, com-

5.9 48.2 18.4 4.7 4.6 3.5 2.1 3.3 0.7 3.8 4.3 0.4 100.0 Estimated mean live weights from Fonseca et al. (1996). * Estimated consumption of three-quarters live mean weight from Emmons (1987).

1

22.5* 2.21 4.65 5.50 7.75 14.25* 4.00 8.22 13.2* 2.16 3.90 30.56* 21.4 23.2 17.9 5.4 7.1 8.9 1.8 5.4 1.8 1.8 3.6 1.8 100 White-lipped peccary Armadillo (Dasypus spp.) Other armadillos Coati Crab-eating racoon Collard peccary Tortoise Paca Deer (Mazama spp.) Agouti Sloth Capybara Total

12 13 10 3 4 5 1 3 1 1 2 1 56

25.5 20.6 16.5 5.0 6.9 9.5 1.6 5.2 1.9 1.6 3.3 2.3 100

% Relative number of individuals consumed % Relative biomass consumed Body weight1 (kg) Frequency of occurrence

The study in LFP revealed more jaguar prey taxa than any previous study on jaguar diet (24 spp. vs. 7–19 spp., 13–228 scats; Rabinowitz & Nottingham 1986, Emmons 1987, Aranda 1994, Facure & Giaretta 1996, Chinchilla 1997). As in many other sites, mammals (25–98% occurrences; Rabinowitz & Nottingham 1986, Aranda 1994, Chinchilla 1997) predominated in the diet. Birds and reptiles were consumed occasionally. In Peru associated with the use of water-edge habitats, reptiles constituted a major fraction of the jaguar diet (Emmons 1987, 1989). Our data on tortoises in jaguar diet support both the association between reptile prey and water-edge habitats and Emmons’ (1989) suggestion that larger and stronger individuals are better equipped to overcome tortoise mechanical defenses. Although subsisting on a wide variety of animals, jaguars concentrate their feeding on a small number of prey. Six species comprised 64 percent of the identified prey items at LFP, but in other studies up to two-thirds of the jaguar diet consisted of two or three prey taxa. Armadillos, mainly the forest long-nosed armadillo Dasypus novemcinctus, appear among the three main prey in Mexico (Aranda 1994), Costa Rica (Chinchilla 1997), and SBR and LFP (this study), and alone comprise 54 percent of the jaguar diet in Belize (Rabinowitz & Nottingham 1986). Its absence in Peru was related to severe flooding in the study year (Emmons 1987). Either collared

N

DISCUSSION

Prey species

prising 37, 35 and 12 percent of the biomass consumed, respectively (Table 2). Considering the number of individuals consumed, armadillos remained the most important item, representing 66 percent of the animals eaten, followed by peccaries (9.4%) and procyonids (9.3%; Table 2). In eight cases the same prey individual appeared to be represented in two scats. The prey species were armadillos, crab-eating racoon, coati, sloth (Bradypus variegatus), monkey (Cebus spp.), curassow (Crax blumenbachii), collared and whitelipped peccaries, and tapir (Tapirus terrestris). Information of sequential prey consumption was gathered on 24 September from five scats along 10 km of a road associated with tracks of a resident jaguar female. Prey remains identified from these scats were, respectively, (1) armadillo, crab-eating racoon, grass; (2) armadillo, crab-eating racoon, grass, curassow; (3 and 4) curassow, sloth; and (5) sloth.

Calculation of relative biomass and relative number of prey individuals killed by jaguars in 1996: (based on 50 scats collected in Linhares Forest Preserve, southeastern Brazil; (N 5 56 identified prey items). N 5 number of scats the item was present in. Frequency occurrence 5 N 3 100/56.


TABLE 2.

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Jaguar Food Habits in Southeastern Brazil

(Aranda 1994, Facure & Giaretta 1996) or whitelipped peccaries (Emmons 1987, Chinchilla 1997, this study) were prevalent in many sites, being taken more often than expected in Peru (Emmons 1987) and Mexico (Aranda 1994). Coatis were also prevalent in Mexico (Aranda 1994) and LFP (Facure & Giaretta 1996, this study). Results from these studies suggest that forest-inhabiting jaguars tend to rely on the same mammalian prey in different parts of their geographic ranges. The highly audible foraging habits of the white-lipped peccary and the long-nosed armadillo (Kiltie & Terborgh 1983, Montgomery 1985) and formation of herds by peccaries probably increase their likelihood of being detected by jaguars (Emmons 1987). Collared peccaries form smaller groups and are usually quiet, while white-lipped peccaries are found in large noisy herds (Fragoso 1998) and leave stronger odors (Sowls 1984). When both peccary species are in the same area, collared peccaries also avoid contact and restrain their movements (Taber et al. 1994; A. Keuroghlian, pers. comm.). Of the two species of peccaries in LFP, we recorded a higher predation on the white-lipped peccaries. We suggest that this predation probably reflected the more conspicuous habits and social organization of this species. The absence of both peccary species remains in the SBR samples may indicate the poaching pressure in public preserves (Chiarello 1999). Poachers selectively hunt for peccaries (Jorgenson & Redford 1993), and although reported for SBR, neither peccaries nor their tracks were observed during our seven-months of fieldwork (R. Garla, pers. obs.) or during another study (Chiarello 1999). LFP is managed by a private company and has an especially efficient security system (R. M. Jesus, pers. comm.). It is noteworthy that the rarity of peccaries at SBR was associated with an increased importance of deer in the jaguar diet. It is relevant that deer are also markedly less abundant at SBR than LFP (0.6 vs. 2.5 visual encounters/10 km; Chiarello 1999). Deer appear as a secondary jaguar food item in other forest sites (Emmons 1987, Aranda 1994, Chinchilla 1997), and in frequencies similar to faunal surveys in Mexico (Aranda 1994). They surpass peccaries in importance only in Belize (Rabinowitz & Nottingham 1986, Taber et al. 1997). Although there was much variation in jaguar size between populations (30–120 kg; Emmons 1990), in LFP as in other forest sites, most jaguar prey weighed between 3 and 10 kg (Emmons 1987, Aranda 1994, Facure & Giaretta 1996,

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Chinchilla 1997; cf. Rabinowitz & Nottingham 1986). The series of scats produced consecutively along a road showed that remains of smaller-bodied prey can appear in sequentially deposited scats. We also found remains of single larger prey (peccaries and a juvenile tapir; 20–60 kg) in more than one scat. Because the larger species consumed naturally by jaguars, particularly the peccaries, were relatively smaller (20–30 kg) than the big prey (50 kg) used in digestibility studies (Ackerman et al. 1984), divergences in number of scats produced may be expected when prey in this size categories are eaten. This highlights the necessity of conducting feeding trials that use native prey for each cat species. By correcting for relative biomass and relative number of prey consumed (Ackerman et al. 1984), we found striking differences in the numerical importance of smaller prey to the jaguar’s diet compared to simple occurrence frequencies in scats (Table 2). The two peccary species were estimated to comprise only 9.4 percent of the animals eaten; armadillos (all species combined) became the most important prey item, representing 66 percent of the animals consumed. About seven armadillos were eaten for every peccary consumed, rather than 1:1 as might be inferred from percent occurrence. Although a large part of LFP and SBR’s perimeter area support cattle ranching, we neither recorded nor were made aware of any evidence of cattle consumption by jaguars. Both reserves are fenced, and cattle roam in fenced pastures around them and may even penetrate forest where marshes make fence maintenance along SBR’s northern limit difficult. This was an area of high jaguar scat collection in SBR, but no cattle remains were found in scats there. Interviews conducted with people living around the reserves revealed no known cases of jaguar attacks in the last 20 years. Native prey seem sufficiently common to support jaguars in both areas; however, this isolated population, due to stochastic factors and genetic deterioration (Franklin & Frankham 1998), may have limited chances of long-term survival. There is an urgent need for efficient programs to control poachers, especially at SBR, and for accurate numerical surveys of jaguars in both areas (Karanth & Nichols 1998).

ACKNOWLEDGMENTS This paper represents partial fulfillment of a master’s thesis by the senior author. The study was supported by Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico (CNPq) and Fundaçaõ de Amparo a Pesquisa

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no Estado de Saõ Paulo (FAPESP-1995/4971-0). We thank R. M. Jesus for authorization to work in LFP, B. Brito for invaluable assistance in the field, and S. L. Mendes (Museu Mello Leitaõ) for logistic support. A. Keuroghlian kindly identified peccary species from hairs. We

are grateful to M. De Vivo, J. C. Motta Jr., E. Wang, W. W. Benson, A. Rabinowitz, and two anonymous reviewers for valuable comments on an earlier version of the manuscript. W. W. Benson kindly helped with the final English version.

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