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Academic Journal of 2016, 7, 643-660 Technology Full-length paper

Three years of continuous monitoring of the large terrestrial mammals of Brownsberg Nature Park, Suriname

Paul E. Ouboter*, Vanessa S. Kadosoe* National Zoological Collection of Suriname/Environmental Research Center, Anton de Kom University of Suriname*

Abstract Brownsberg Nature Park in Suriname has a rich biodiversity. The area is also threatened by mining, illegal hunting and possibly also tourism. To assess the impact of threats on the terrestrial mammal community, a long-term continuous monitoring survey was started in 2012. To assess mammal presence, abundance and activity, 27 camera traps were placed at 16 locations. In the three-year period the camera stations were triggered 55,949 times by an identifiable animal (27%), human (41%) or vehicle (32%). Twenty-nine species of large terrestrial mammals were recorded, of which six were before this study only known to occur in the area through interviews with local residents. The crab-eating raccoon, was recorded from Brownsberg and the interior of Suriname for the first time. Twenty-eight percent of the species was recorded only after the first year and 10% only after the second year of camera trapping. Most species were relatively rare. The red-rumped agouti, puma, jaguar and red brocket deer caused most triggers. The relative abundance index (RAI) of agoutis and pumas was higher than reported anywhere else in the . A seasonal pattern of activity could only be established for the agouti: highest activity in the period with most fruits on the ground, much less activity in the flowering period of most trees. None of the species seemed to decline over the three-year period. Twelve species can be considered of international conservation interest; eight of these were shown to reproduce in the park. Differences in the number of triggers per species between areas could partly be explained by differences in tourism pressure..

Keywords: large terrestrial mammals, monitoring, camera trapping, Brownsberg, Suriname

Introduction Our knowledge of the life history of most However, the occurrence of several species is Neotropical rainforest mammals is still limited. only known from interviews with local people, However, since the 1990s the improvements in without any proof by specimens or photographs. the method of camera trapping and its increased Brownsberg Nature Park faces many threats. application in wildlife studies, has enhanced this The forests and creeks of the northwestern, knowledge enormously. northern and southeastern lowlands at the foot of In Suriname one of the best studied sites is the mountain have been destroyed by small- Brownsberg Nature Park (BNP). Its attraction is scale mining. Occasionally some gold its rich biodiversity, accessibility and facilities. miners try their luck on the plateau as well. In Its diverse flora and fauna are related to the addition, illegal hunting, plant collecting and geomorphology of the area and the 500 m eco-tourism may pose threats to the biodiversity altitudinal gradient of the Brownsberg Mountain as well. In Suriname BNP is one of the prime (P-M. Forget, pers. comm.). The mammals of destinations for many eco-tourists. According to the Brownsberg area have been studied on Fitzgerald et al. (2002) 17,000 tourists annually several occasions (Genoways & Williams, 1979; visited the park. Most of these visited the park Williams & Genoways, 1980; Genoways et al., for a one-day trip to hike on one of the many 1981; Genoways & McLaren, 2003). Lim et al. trails leading to waterfalls and creeks, but some (2005) presented an extensive overview of both stayed overnight in houses that can be rented. small and large mammals of the Brownsberg Mt.

Correspondence to: Paul Ouboter, Anton de Kom University of Suriname, Leysweg 86, Paramaribo, Suriname, Tel. 597-494756, E-mail: [email protected] Available on-line December 7th, 2016

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In 2012 we started a long-term camera trapping kilometers further south, but this part of the road project at Brownsberg with the main aim to (Pedreku Pasi; abbr. PP) was overgrown during monitor the terrestrial mammal populations and a period of social unrest in the interior in the get insight into the impacts that different threats 1990s. At the start of the 21th century part of have on these populations. By 2015 several this road was opened again for research, but not species could be verified or added to the area, for motorized traffic; this was also the case for a and insight in the status of species in the area is second road going down the mountain to Witi gained. This is the subject of the present paper. Creek (Jeeptrail). These roads were still A detailed analysis of the impact of eco-tourism, accessible by foot when we started this project, interspecific interactions or population dynamics and we included 3.2 km of these in our study will be treated separately in the future. area. After two years we had to clear these road sections from re-growing vegetation to keep Materials and Methods these accessible. Most of the main roads and the Study area hiking trails going down the mountain are The Brownsberg Nature Park (11,800 ha) is included in the study areas as well (Fig. 1). located at the northwest corner of the , just 100 km south of the Field methods capital Paramaribo between 04° 45’ 46” and 05° Our trapping cameras were located at the road 59’ 44” N, and 055° 07’ 58” and 055° 15' 23” W entering and crossing the plateau, including the (De Dijn et al., 2006). It can be reached by closed roads in the south, and in the first tarmac road. The Brownsberg Mt occupies most kilometer of trails going down from the plateau of its area. It is an approx. 500 m high mountain (Fig. 1). These sites were selected based on the with an almost flat ferro-bauxite cap on top. The high probability of the cameras being triggered plateau is approximately 34 km long and 13.5 by large cats. Data from December/January 2013 km wide at its widest point. At Brownsberg four till November/December 2015 were used for seasons can be distinguished: the short dry analysis. A total of 16 locations were fitted with season from February till April, the long rainy cameras (Figure 1). The cameras were spaced season from May till halfway August, the long 0.8 to 1.4 km apart. Usually two cameras were dry season from half August till November and placed almost opposite to each other. This is the short rainy season from December till important to maximize the chance to capture January. During the last decennium the timing of every animal that is passing, to have a spare these seasons seems to be less predictable (pers. camera in case one breaks down, and to obs.). The mountain receives a lot of rain, and in photograph identifiable animals on both flanks. addition in the night and early morning the On some of the narrower hiking trails only one upper slopes of the mountain and the plateau are camera was installed. The survey was started often wetted by mist from the clouds, especially with the use of Reconyx® HC600 cameras, but at the eastern rim. The mountain is almost Reconyx® PC900 cameras gradually replaced completely covered by mesophytic and xero- these. Both types of cameras have an infrared mesophytic rainforest (De Dijn et al., 2006). The flash that is not visible (to humans and most nature park is in long-term lease by the animals). The cameras were powered with Foundation for Nature Conservation in Suriname rechargeable Energizer® NiMH batteries. (STINASU), that exploits it as an eco-tourist Memory cards used were SanDisk® 4 Gb facility. The main access road to reach the park Extreme SD cards. Cameras were covered with a facilities on top of the plateau is on the northern steel housing, especially to limit the impact of side of the mountain (Agwago Kununu Pasi) direct raindrops on the camera (these cameras (Figure 1). This road continues to the park are not fully waterproof under tropical headquarters while a sidetrack (Mazaroni Weg; conditions). In the cameras two small cylinders abbr. MW) continues 4 km to the south on the of desiccant were placed to lower inside plateau. Formerly this road continued many humidity.

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Fig. 1: Map of study area. Triangle – camera station. Continuous thick black line – open and closed roads. Closed roads are the ones going to Jeeptrail and PP locations. Thick spotted line – trails.

They were fitted to trees with a Python lock at a month between the second and third week, height of 30 to 80 cm, dependent on the position checking the condition of the camera and of the tree relative to the road or trail. changing the batteries, memory card and Settings were arranged at rapid-fire taking 5 desiccant. images per triggering, with a delay of 3 minutes. It should be emphasized that camera triggers are The delay was important to prevent battery an indication of activity and abundance. drainage and the filling of memory carts by However, the actual number of individuals animals like gray-winged trumpeters that may cannot be determined when specimens triggering linger in front of a camera for extended periods. a camera are not individually recognizable. In addition to the 3-minute delay, specimens that Notwithstanding this, a low number of triggers remained in front of the camera or kept is likely to indicate a rare species or a species appearing, were only counted after 30 minutes. with another (micro)habitat preference, while a This happened often with agoutis, armadillos, high number of triggers is likely to indicate a peccaries and pumas. Species that normally live relatively abundant species. in groups, like peccaries and coatis, were counted as one triggering despite several Data analysis specimens being visible in the photographs. The Data analysis was focused on large mammals, photographs of opposite cameras were pooled, which are defined as all mammal species in so numbers of triggers/species were counted per which adults weigh more than 1 kg. The data location, not per camera. The wildlife cameras were analyzed with Excel and SPSS. Diversity used in this research are triggered by heat and indexes were calculated with Species Diversity movement, humans and cars are therefore and Richness from Pisces Conservation Ltd®. photographed as well. These were counted to Simpsons Diversity Index (Simpson, 1949) was assess disturbance. Cameras were serviced every used, calculated as 1-D to make diversity

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increase from 0 to 1. According to Magurran Also their identification to species was not (2004) this diversity index is “one of the most always possible, and they are therefore left out meaningful and robust diversity measures from the analysis. Also left out are two species available”, while the Shannon-Wiener Index is of squirrels (under 1 kg) and two species of often difficult to interpret, however most monkeys, the brown capuchin (Cebus apella) publications report the Shannon-Wiener Index. and the wedge-capped capuchin (Cebus Therefore, the Shannon-Wiener Index was olivaceus) that triggered the cameras only once, calculated as well. To detect differences between and are here considered to be completely various areas based on different disturbance arboreal. This means that 29 species are listed factors, Principal Component Analysis (PCA) here (Tab. 1). was carried out with Community Analysis Although the first six months already produced a Package 4 from Pisces Conservation Ltd®. For good accumulation curve, additionally ten PCA the data was transformed with Ln(x+1) to species (34%) were recorded only after the first limit the impact of very abundant species. Sites 6 months, eight species (28%) only after one were grouped based on various disturbance year and three species (10%) only after two factors (number of tourists, number of vehicles, years. type of roads). The significance of the various Humans and cars triggered the cameras more divisions in groups was checked by Analysis of often than any mammal in the area: humans Similarity (ANOSIM) (Clarke, 1988, 1993). In triggered the cameras on 22,497 occasions, cars addition, for each species the Relative on 17,515 occasions (Tab 1). Abundance Index (RAI) for the total area was From the calculated Relative Abundance Index calculated as the number of camera trap triggers (RAI) shown in Table 1 it is clear that most per 100 trap days. Species of special species are rare. In the three-year period the conservation interest are defined here as listed in southern naked-tail armadillo (Cabassous the IUCN Red List of Threatened Species unicinctus), grison (Galictis vittata) , oncilla (IUCN, 2015) as Near Threatened, Vulnerable, (Leopardus tigrinus) and Brazilian porcupine Endangered or Critically Endangered, or listed (Coendou prehensilis) triggered a camera only in CITES Appendix I or II (CITES, 2015). once, the capybara (Hydrochaeris hydrochaeris) only twice. Other rare species are the southern Results tamandua (Tamandua tetradactyla), giant During 17,520 trap days the cameras were anteater (Myrmecophaga tridactyla), crab-eating triggered 55,949 times by an identifiable animal raccoon (Procyon cancrivorus) and bush dog species (27% of triggers), human (41%) or (Speothos venaticus). The mammal that vehicle (32%). Thirtyfive species of mammals triggered the cameras the most was the red- were identified in the three-year period. In rumped agouti, with 6,024 triggers. Other addition, several birds, reptiles and unidentified species that were often photographed are the small rodents and bats were photographed. Of puma (Puma concolor), jaguar (Panthera onca), the 35 mammals the brown four-eyed opossum ocelot (Leopardus pardalis) and red brocket (Metachirus nudicaudatus) and the gray four- deer (Mazama americana). eyed opossum (Philander opossum) are usually less than 1 kg in weight and can therefore not be considered as large mammals.

Acad J Sur 2016 (7), 643-660 Academic Journal of Suriname 2016, 7, 643-660 Technology Full-length paper

Table 1. Camera triggerings by large terrestrial mammals, tourists and vehicles at Brownsberg Nature Park in 2013, 2014 and 2015. Also shown are the total number of triggers in 3-years’ time and the RIA (relative abundance index).

Species Common name 2013 2014 2015 Total 3 years RAI Dasyprocta leporina Red-rumped agouti 1529 2180 2315 6024 34.38 Puma concolor Puma 419 377 405 1201 6.85 Panthera onca Jaguar 287 331 281 899 5.13 Leopardus pardalis Ocelot 216 279 335 830 4.74 Mazama americana Red brocket deer 127 140 90 357 2.04 Cuniculus paca Paca 81 135 119 335 1.91 Myoprocta acouchi Red acouchy 83 60 125 268 1.53 Mazama gouazoubira Grey brocket deer 93 74 92 259 1.48 Didelphis marsupialis Common opossum 45 87 113 245 1.39 Dasypus kappleri Great long-nosed armadillo 13 81 118 212 1.21 Tapirus terrestris Brazilian tapir 31 101 25 157 0.90 Dasypus novemcinctus Nine-banded long-nosed armadillo 59 21 51 131 0.75 Leopardus wiedii Margay 18 23 30 71 0.40 Priodontes maximus Giant armadillo 10 21 38 69 0.39 Eira barbara Tayra 14 6 18 38 0.22 Puma yagouaroundi Jaguarundi 5 8 23 36 0.21 Pecari tajacu Collared peccary 10 20 3 33 0.19 Nasua nasua South American coati 11 16 4 31 0.18 Tayassu pecari White-lipped peccary 16 4 20 0.11 Sciurus aestuans Guianan squirrel 4 13 1 18 0.10 Tamandua tetradactyla Southern tamandua 4 3 7 0.04 Myrmecophaga tridactyla Giant anteater 1 3 3 7 0.04 Speothos venaticus Bush dog 5 1 6 0.03 Procyon cancrivorus Raccoon 4 4 0.02 Hydrochaeris hydrochaeris Capybara 2 2 0.01 Galictis vittata Grison 1 1 0.006 Leopardus tigrinus Oncilla 1 1 0.006 Coendou prehensilis Brazilian porcupine 1 1 0.006 Cabassous unicinctus Southern naked-tailed armadillo 1 1 0.006 Tourists 7533 7669 7295 22497 128.41 Vehicles 4182 7456 5877 17515 99.97

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Figure 2. Activity of red-rumped agouti and paca. Tree phenology is indicated as: FD – fruit development; RF – ripe fruits; F – flowering

Temporal patterns triggers over the three years: red-rumped agouti, Figure 2 shows the wide variation in the number ocelot, common opossum (Didelphis of monthly triggers, and therefore activity over marsupialis), great long-nosed armadillo time, of the red-rumped agouti with the lowest (Dasypus kappleri), margay (Leopardus wiedii), numbers during the long dry season. Red- giant armadillo (Priodontes maximus) and rumped agoutis have a limited home range and jaguarundi (Pumayagouaroundi. therefore triggers per camera is correlated to activity for this species. Other common species Reproduction have some variation in triggers as well (Figure Juveniles were seen for 12 species and pairs of 3). The puma had an exceptionally high number adult individuals for six solitary species (Tab. 2). of triggers in January-February 2013. Very Very remarkable is the high number of puma surprising is the general conformity in triggers pairs (15). Jaguar juveniles (often with their between jaguars and pumas. mother) had the highest number of triggers of all juveniles (13). Table 1 and Figures 2 and 3 do not indicate that any of the species is declining. Jaguar triggers Spatial pattern dropped almost to zero in November-December There is a slight difference between sites in the 2015, but numbers increased again the next number of species found. Only 15 species were months (unpubl. data). Brazilian tapir (Tapirus recorded for the trail to Mazaronival and the terrestris) triggers were relatively high in 2014 road junction to this fall, followed by 16 species and low in 2015 (Tab. 1). The same trend at the Leoval trail (Figure 4). Leoval is the trail applies to both species of peccaries. Several visited by most tourists (average 263 species show an increase in their number of triggers/month).

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Table 2 . Evidence of reproduction as indicated by pairs of species that are solitary outside the reproductive period and by juveniles (often with mothers) triggering the cameras (grey shading). * usually not solitary but in pairs or groups.

** not the whole group is seen, so more juveniles may have been present. Crosshatched – open hunting season 2015 as determined by Nature Conservation Division Suriname.

Species Pairs of solitary species Juveniles Pairs J F M A M J J A S O N D Records Number J F M A M J J A S O N D Didelphis marsupialis 4 4 to 5 Tamandua tetradactyla 1 1 Priodontes maximus 1 1 1 Leopardus pardalis 1 5 1 Puma yagouaroundi 4 1 to 3 Puma concolor 15 8 1 Panthera onca 3 13 1 to 3 Tapirus terrestris 1 4 1 Tayassu pecari 2 1 to 3* Mazama americana 5 1 Mazama gouazoubira 1 1 Cuniculus paca 1 Dasyprocta leporina 5 1 to 4

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The highest number of species was recorded for PP2.8 (Figure 4) (1-D resp. 0.87, 0.84 and 0.83). the second most visited site, Mazaroni Top, with This is as expected, but these sites are followed 22 species. Twenty-one (21) species were found by the most visited Leoval (1-D=0.82), with at the site furthest from any human activity only 16 species. Another contradiction is seen at PP2.8, but also in the middle of the Mazaroni Mazaroni Top: despite many tourists visiting the Road MW2. Diversity (Simpsons and Shannon trail the number of species is highest, but Wiener Index) is often in contradiction with diversity is the lowest (1-D=0.39). species number. Highest diversity is calculated for the least disturbed sites PP1.4, Jeeptrail and

Figure 3 . Activity of ocelot, puma, jaguar and red brocket deer. Seasons are indicated as: SDS – short dry season; LRS – Long rainy season; LDS – Long dry season; SRS – short rainy season

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Figure 4. Number of species, Simpsons Diversity Index (x10; indicated as 10 (1-D)) and Shannon-Wiener Index (x5; indicated as 5 (H’)) of all camera localities. Localities are arranged in decreasing order of the Simpsons Index

Analysis of the mammal community with more or less pointing to the trail and slope Principal Component Analysis resulted in Figure locality of Witi Creek. These are all species of 5. Localities that are closed roads, and therefore armadillos, South American coati, bush dog and with hardly any human disturbance, are plotted puma. Species not mentioned above have less to the left of the graph, open roads with traffic impact on differences in the number of triggers and tourists mostly to the right. Species that /species between sites (they may occur seems to avoid human disturbance are the anywhere), including the jaguar and common Brazilian tapir, both species of peccaries, opossum. jaguarundi, red acouchy and gray brocket deer. Differences in triggers/species are most Species that seem to prefer open roads (with significant when the sites are divided based on more human disturbance) are the red-rumped tourist numbers (ANOSIM; R=0.293, P=0.001), agouti, paca, ocelot, margay and red brocket or road/trail characteristics (trail, accessible deer. Several species are plotted in between, road, closed road; (ANOSIM; R=0.348, P=0.002).

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Figure 5. PCA plot of terrestrial mammals at all camera localities

Species confirmed for the Brownsberg Nature Brazilian porcupine ( Coendou prehensilis Park or species new to the area (Linnaeus, 1758)), 1 trigger. Several mammal species listed for Brownsberg Apart from these six species confirmed for the by Lim et al. (2005) were included based on Brownsberg Mt., a mammal was photographed interviews with local inhabitants only. Our at Brownsberg that is not only new for the survey confirmed the presence of the following mountain but also for the interior of Suriname in species: Southern naked-tail armadillo general, the Crab-eating raccoon ( Procyon (Cabassous unicinctus (Linnaeus, 1758)), 1 cancrivorus (Cuvier, 1798) (Fig. 6). The crab- trigger. Giant anteater (Myrmecophaga eating raccoon is common in the coastal area. tridactyla Linnaeus, 1758), 7 triggers. The most southern locality mentioned by Bush dog ( Speothos venaticus (Lund, 1842)), 6 Husson (1978) is Domburg, only 25 km from triggers, usually two individuals. Oncilla the coast (Brownsberg is approx. 100 km from (Leopardus tigrinus (Schreber, 1775)), 1 trigger. the coast). No specimens were known from the Grison ( Galictis vittata (Schreber, 1776)), 1 interior of Suriname. In 2015 this species was trigger, two individuals. photographed four times by the camera traps.

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Figure 6. Crab-eating Raccoon ( Procyon cancrivorus ) at Mazaroni Road, Brownsberg, 19 June 2015

Species of special conservation interest Of the conservation sensitive species (with Species of special conservation interest, as well sufficient numbers) none is showing an overall as the number of triggers, RAI and trend over decline over the three years, four species even the three years are shown in Table 3. In addition, seem to be increasing in numbers over the evidence of reproduction is indicated. Of the 12 period and five fluctuating in numbers. species of international conservation concern However, the jaguar shows a sudden decrease in three are quite rare at Brownsberg, the oncilla, number of triggers during the last three months bush dog and giant anteater; and three are of the study. Reproduction in the Brownsberg common, the puma, jaguar and ocelot (Fig. 7). area is proven for eight species (67%).

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Table 3. Conservation sensitive mammal species (see text) of Brownsberg Nature Park, showing IUCN Red List and CITES status, number of triggers, Relative Abundance Index (RAI), trend, and data on reproduction.  numbers increasing over three-year period; Ï numbers almost stable; Ú numbers fluctuating over three-year period.

Species Common name IUCN Red List CITES Triggers RAI Trend Reproduction App. Myrmecophaga tridactyla Giant Anteater Vulnerable II 7 0.04 no data Priodontes maximus Giant Armadillo Vulnerable I 69 0.39  pair and juvenile Near Speothos venaticus Bush Dog Threatened I 6 0.03 no data pair and Leopardus pardalis Ocelot Least concern I 830 4.74  juveniles Near Leopardus wiedii Margay Threatened I 71 0.40  no data Leopardus tigrinus Oncilla Vulnerable I 1 0.006 no data Puma yagouaroundi Jaguarundi Least concern II 36 0.21  juveniles pairs and Puma concolor Puma Least concern II 1201 6.85 Ï juveniles Near pair and Panthera onca Jaguar Threatened I 899 5.13 Ï juveniles pairs and Tapirus terrestris Brazilian Tapir Vulnerable II 157 0.90 Ú juveniles Pecari tajacu Collared Peccary Least concern II 33 0.19 Ú juveniles Tayassu pecari White-lipped Peccary Vulnerable II 20 0.11 Ú juveniles

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Figure 7. Relative abundance index (RAI) of conservation sensitive mammal species

Discussion elusive common species like puma and ocelot, This project shows the importance of long-term but it takes a long time to record very rare continuous camera trapping of a terrestrial species. mammal fauna: 34% of the species was recorded The former camera trapping survey at after the first 6 months, 28% after the first year Brownsberg Nature Park documented only 17 and 10% only after the second year. Camera species (Lim et al., 2005), this one 35 species, trapping surveys often have a duration of four to with 29 species which can be considered large six months, and may even show a good terrestrial or terrestrial/arboreal mammals. With accumulation curve for this short period. Despite this number of large terrestrial mammals, it is this, many rare species occurring in the area may one of the richest sites in the world. Ahumada et still be missing. Even with surveys of several al. (2011) compared 7 tropical rainforest sites months each year, the chance of missing rare around the globe with camera trapping as species is considerable. Camera trapping is very method. In that comparison the Central efficient for assessing common species, even Suriname Nature Reserve location was the

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richest with 28 species recorded by camera occasions the RAI is increased by a single or a trapping. few individuals triggering one or several The most abundant species, the red-rumped cameras multiple times. This is especially the agouti fluctuated in number of triggers case with red-rumped agoutis, armadillos and considerably over the years, with numbers in the pumas. On the other hand, all studies reporting long dry season three to four times lower than in RAI’s will be biased by this phenomenon, unless the short dry and long rainy seasons. This is individuals are identified, which is only possible probably correlated to the phenology of most for a few species. Camera triggers should be trees in the forest and consequently the amount considered principally as an indicator of activity, of fruit available on the ground. The period of secondly of abundance! Despite these high activity coincided with the period in which considerations, the number of puma and jaguar most trees develop or have ripe fruits (see De triggers are unusually high. The identification of Dijn et al., 2006). The period of low activity individuals in the near future will demonstrate if coincides with the flowering period. For the only also the density of these species at Brownsberg other frugivore, the paca, this seasonal trend is is high. less obvious. Only in 2014 pacas seemed to be In contrast to these high RAI’s, the RAI’s for the less active in the long dry season. Possibly this collared peccary and white-lipped peccary are species prefers other fruits than the agouti (also unusually low compared to other studies see Dubost & Henry, 2006). Seasonality in (Gomez et al., 2005; Srbek-Araujo & Chiarello, activity in other mammal species is even less 2005; Tobler et al., 2008; Blake et al., 2012). clear. The three cat species ocelot, jaguar and This could be caused by high predator puma fluctuate considerably in number of abundance, but other explanations are possible triggers, but no pattern related to seasonality too. All peccaries were seen at sites with low could be detected. In puma and jaguar, the tourist numbers, indicating the possibility of highest numbers of triggers are often related to a avoidance of humans. Peccaries were seen most period in which males and females are seen at sites near the edge of the plateau; possibly the together, with the pair walking back and forth in peccaries prefer the slopes of the mountain that front of several cameras. have a thicker soil layer than the hard and rocky plateau substrate. Very high RAI’s were obtained for the red- rumped agouti, puma and jaguar. The RAI for Jaguars and pumas are top predators of almost the agouti was almost ten times the number in equal size, with an overlap in prey items other studies (Maffei et al., 2002; Srbek-Araujo (Harmsen et al., 2010). Because of its stouter & Chiarello, 2005; de Souza Martis et al., 2007; body the jaguar can handle slightly larger prey Gonzalez-Maya, 2007; Ouboter et al., 2011; like tapir and more dangerous prey like white- Isasi-Catalá, 2012; Jax et al., 2015). Also for lipped peccaries (Foster et al., 2010), and pumas no study could be found with RAI’s jaguars are known to kill pumas in intraguild exceeding the 6.85 found for Brownsberg (e.g. encounters (de Oliveira & Pereira, 2014). Maffei et al., 2002; Srbek-Araujo & Chiarello, Research in other Neotropical areas has shown 2005; Kelly et al., 2008; Soria-Diaz et al., 2010; that pumas and jaguars avoided using the same Blake et al., 2012). Also the jaguar RAI of 5.13 locality at the same time (Harmsen et al. 2009). is very high compared to other studies (see At Brownsberg there is no indication of overview in Maffei et al., 2011). Of a total of 39 avoidance of jaguars by pumas. In general, the studies, only four studies produced similar or number of triggers of both species go up and higher RAI’s for the jaguar: for Mountain Pine down at the same time and also per site there is Ridge, Belize (M. Kelly, unpublished data); no negative correlation between these species Fazenda Sete, (Soisalo & Cavalcanti, (unpubl.data). 2006); Serra da Capivara, Brazil (Silveira et al., The number of jaguar triggers dropped since 2009); Tikal, Guatemala (Garcia et al., 2006). September 2015. Triggers below 10/month The very high RAI’s found at Brownsberg never happened before. It is of course possible should be considered with caution. On many that some of the jaguars migrated to parts of

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their territory outside our study area. We Brownsberg is an important area for considered the relative dryness of the 2015 long conservation. That was already indicated by a dry season as a possible cause for jaguars to priority setting workshop that resulted in migrate down the slopes of the mountain, where designating the area as a High Priority creeks still have sufficient water. However a Conservation Area (Conservation International, similar dryness occurred in 2014 (see Ouboter et 2011). Jairam (2015) studied the use of the al., 2016), without a drop in jaguar triggerings. Brownsberg Nature Park by scientists and the occurrence of endemic and threatened species In Suriname, 15 species of mammals may be and came to the same conclusion. During this hunted during part of the year or the whole year, and other surveys Brownsberg proofed to have a dependent on the species. The hunting season is very rich biodiversity; for large terrestrial supposed to be closed in the season of mammals 29 species. Of these mammals,12 reproduction. species had a high conservation status based on In the present study, no juveniles were seen in their occurrence in the threatened categories of the open hunting season for white-lipped the IUCN Red List of Threatened Species or/and peccaries, but for the tapir and both species of the Appendices of CITES. deer juveniles were seen in the open season. A Three of these species, the ocelot, puma and revision of the open- and closed hunting seasons jaguar are common at Brownsberg. All 12 is advisable. species seem to be stable. Eight of the 12 species were proven to reproduce in the area, the four The contradiction between species numbers and species without proof of reproduction are diversity is mainly caused by the enormous extremely elusive and/or rare (giant anteater, abundance of certain species, lowering the bush dog, margay and oncilla) and may evenness and with that the Simpsons and reproduce in the area as well. Shannon-Wiener Diversity Indices. This is most obvious at Mazaroni Top and MW2 with the Acknowledgements highest species numbers but the lowest diversity. This work was supported by funding from the Here it is the red-rumped agouti that triggered Belgian Directorate-General for Development the cameras more than a thousand times! Cooperation (DGDC) and the Flemish Two factors that seemed to contribute Interuniversity Council (VLIR-UOS) significantly to differences in species triggers We like to thank all persons that on occasion between areas are the number of tourists and the helped us in the field: Geeta Thakoerdien, Usha type of road/trail the cameras were positioned Satnarain, Niradj Hanoeman, Mercedes on. However, these two factors are not Hardjoprajitno, Cindyrella Kasanpawiro, independent, as hardly any tourist reaches the Richero Kasanwidjojo, Carlo Koorndijk, closed roads, and several trails were the most Gunovaino Marjanom, Dimitri Ouboter, Naomi frequented areas of the park. We therefore Reussien, Artie Sewdien and Sergio Doelasan. suggest that the number of tourists is the main The coordinator of the VLIR MSc Sustainable factor influencing and changing species relative Management of Natural Resources, Riad abundances in highly frequented areas. Nurmohamed and the VLIR Secretariat always A species that seems to have a preference for supported us fully. STINASU granted us disturbance is the red-rumped agouti, while large permission to carry out this research project at predators seem to avoid areas with many the Brownsberg Nature Park. tourists. The preference for disturbed areas of red-rumped agoutis may very well be caused by References a release of predation pressure on/near tourist Ahumada, J.A., Silva, C.E.F., Gajapersad, K., frequented roads and trails. Consequently, the Hallam, C., Hurtado, J., Martin, E., choice of locations of camera traps, near roads McWilliam, A., Mugerwa, B., O’Brien, and trails, could explain the high RAI found for T., Rovero, F., Sheil, D., Spironello, red-rumped agoutis. W.R., Winarni, N., Andelman, S.J., 2011. Community structure and diversity of

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