Baird’s tapir density in high elevation forests of the Talamanca region of Costa Rica

© 2012 Wiley Publishing Asia Pty Ltd, ISZS and IOZ/CAS 381

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Integrative Zoology 2012;; 7: 381–388 doi: 10.1111/j.1749-4877.2012.00324.x

ORIGINAL ARTICLE

Baird’s tapir density in high elevation forests of the Talamanca region of Costa Rica

José F. GONZÁLEZ-MAYA,1,2,3 Jan SCHIPPER,1,4 Beth POLIDORO,1,5 Annelie HOEPKER,1 Diego ZÁRRATE-CHARRY1 and Jerrold L. BELANT31Sierra to Sea Institute, ProCAT Internacional/Colombia, Las Alturas, Coto Brus, Puntarenas, Costa Rica, 2Instituto de Ecología, Universidad Nacional Autónoma de México (IE-UNAM), Ciudad Universitaria, DF, Mexico, 3Carnivore Ecology Laboratory, Forest and Wildlife Research Center, Mississippi State University, Mississippi, USA, 4Big Island Invasive Species Committee, University of Hawaii, Hilo, Hawaii, USA and 5Old Dominion University, Norfolk, Virginia, USA

AbstractBaird’s tapir (Tapirus bairdii) is currently endangered throughout its neotropical range with an expected popula-

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lished in Valle del Silencio within Parque Internacional La Amistad (PILA). Seventy-seven tapir pictures of 15 individuals comprising 25 capture–recapture events were analyzed using mark-recapture techniques. The 100% minimum convex polygon of the sampled area was 5.7 km2 and the effective sampled area using half mean max-imum distances moved by tapirs was 7.16 km2. We estimated a tapir density of 2.93 individuals/km2 which rep-resents the highest density reported for this species. Intermountain valleys can represent unique and important habitats for large mammal species. However, the extent of isolation of this population, potentially constrained by steep slopes of the cordillera, remains unknown. Further genetic and movement studies are required to un-derstand meta-population dynamics and connectivity between lowland and highland areas for Baird’s tapir con-servation in Costa Rica.

Key words: camera trap, capture, Central America, populations, tapir

Correspondence: José F. González-Maya, Sierra to Sea Institute, ProCAT Internacional/Colombia, Las Alturas de Cotón, Coto Brus, Puntarenas, Costa Rica.Email: [email protected]

INTRODUCTIONBaird’s tapir [Tapirus bairdii (Gill, 1865)] is the larg-

est Neotropical mammal species and was listed in 2008

as ‘endangered’ on the IUCN Red List of Threatened Species (Castellanos et al. 2008). The species’ dis-tribution is disjunct and ranges from southern Mexi-co to Colombia (Reid 1997) from sea level to 3600 m (González-Maya et al. 2009). Little is known of its pop-ulation status and ecology, with most studies concen-trated in lowland areas (Naranjo 1995a,b;; Piñera 1995;; Foerster & Vaughan 2002;; Naranjo & Bodmer 2002). We are aware of only 1 study conducted in mountainous areas (Tobler 2002). However, isolated habitats within these montane areas increasingly represent the majori-

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J. F. González-Maya et al.

© 2012 Wiley Publishing Asia Pty Ltd, ISZS and IOZ/CAS

ty of the remaining suitable habitat. Thus, conservation efforts must ensure the persistence of these habitats and area-sensitive species. For Costa Rica and most of Central America, follow-

ing severe agricultural expansion in lowlands, most re-maining habitats are restricted to isolated high eleva-tion areas of mountain ranges, creating virtual islands of habitat (Schipper 2010). Several countries have started to declare mountain habitats as protected areas during implementation of land-use capacity studies. In Costa Rica, most high elevation mountainous areas are cur-rently protected, with the Talamanca mountain range representing the largest protected area in the country (González-Maya et al. 2008). This area may represent the only available forest continuum capable of maintain-ing viable populations of large mammal species in Cos-ta Rica.We present the first absolute density estimate for

species, using camera traps and mark–recapture tech-

to document high elevation forest use by Baird’s tapir which will provide baseline data for further assessments of mountain ecosystems, tapir ecology and conservation across its range.

MATERIALS AND METHODS

Study area

The Talamanca mountains form the continental di-vide between the Caribbean Sea and Pacific Ocean in southeastern Costa Rica and western Panama (Fig. 1). The Talamanca montane forest ecoregion represents the largest undisturbed forest patch in Costa Rica, rang-ing from 500 to 3820 m asl, and whose central mas-sif generates unique and varied climatic conditions on both slopes (González-Maya et al. 2008;; González-Maya & Cardenal 2011). This ecoregion is currently protected under various categories and comprises 4 protected ar-eas, including Parque Internacional La Amistad (PILA), the largest in the country (González-Maya et al. 2009). The region represents an important biodiversity area and consequently falls under several protection catego-ries including biosphere reserve (6126 km2), UNESCO Human Natural Heritage Area (Kapelle 1996) and En-demic Bird Area (Harcourt et al. et al. 1998), Center for Plant Diversity (Davis et al. 1997) and Global 200 Ecoregion (Olson & Dinerstein 2002). In addition, this region is also considered an important component of the Mesoamerican Hot Spot (Mittermeier et al. 2005;; González-Maya et al. 2009).

Figure 1 Location of study site for Baird’s tapir density estimation, PILA, Costa Rica, Jan–Mar 2006.

83°W

83°W

9°N

9°N

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Tapir density in Costa Rica


PILA is also one of the largest protected areas in Mesoamerica (199 147 ha) and represents the most pristine habitat in Costa Rica, due mostly to its rug-ged terrain which has kept it isolated and therefore protected from agricultural expansion. Valle del Silen-cio is an inter-montane valley located in the southern portion of La Amistad (9°6' N, 82°57' W;; Fig. 1), with elevations from 2400 to 2800 m asl, corresponding to a montane humid forest (Holdridge 1979). It is com-posed mostly of mixed forests dominated by oak (Quer-cus spp.) and other high-elevation species (e.g. Prunus spp., Drymis spp.) with dense bromeliad and Chusquea spp. understories. Valle del Silencio also contains some open areas known as ‘jardines’ or ‘turberas’, containing ferns, cycads and mosses. Our sampling area included the entire valley, surrounded by steep slopes and rug-ged terrain. Valle del Silencio is characterized by low human use;; only one trail occurs in the valley and is used for tourism.

MATERIALS AND METHODSWe estimated tapir density using camera traps and

mark–recapture techniques during Jan–Mar 2006. Cam-era trapping has been used previously for estimating presence, abundance and densities of cryptic species (Karanth et al. 2004;; Silver et al. 2004;; González-Maya et al. 2008), including tapirs (Noss et al. 2003;; Trolle et al.flash cameras (20 cameras in total;; Photoscout, PTC Technologies, Huntsville, AL, USA) were established in the area and separated by 1–3 km. At each station, cam-eras were attached to trees and positioned facing each other about 50 cm above ground along established an-imal trails. Each site was prepared by removing veg-etation between cameras and placing additional veg-etation along the trail to further encourage animals to walk between cameras to ensure we obtained pictures of both sides of each animal (González-Maya et al. 2008). We established our camera array throughout the valley along existing tapir trails to encompass all available ta-pir habitat and avoid potentially biased detection rates from cameras placed along human trails (Harmsen et al. 2010). Stations were baited with a mixture of Wildcat Lure # 2 (a mixture of cat glands, natural food odors and catnip oil;; Hawbaker & Sons Lures & Baits Inc, Fort Loudon, TN, USA) and Obsession fragrance (Calvin Klein Inc., New York, NY, USA). This study was part of a larger project to evaluate mammal distribution and fe-lid density along an elevational gradient in the Talaman-ca mountains;; therefore our objective was to account for

maximum detection rates of all medium- and large-sized mammals in the study area.Cameras operated continuously for 54 days on aver-

age at each site, representing the dry season in the pied-

camera was configured with a 10 min delay between pictures. Cameras were set to record date and time of each picture, allowing estimates of activity and the abil-ity to create capture histories for individuals. Each sta-tion was characterized by percentage forest canopy cov-er using a forest densitometer (Terra Tech Inc., Forest Spherical Densiometer Concave–Model C, Salem, OR, USA) and slope using a handheld clinometer (PM-5/360 PC, Suunto Inc., Vantaa, Finland). Although previous studies have used camera trapping

to estimate tapir density (e.g. Noss et al. 2003;; Trolle et al.

et al. (2003) provided a number of unique features for positive individual iden-

-perience, including scars, white spots and stripes, black markings on the face or sides, ears with distinctive fea-tures, tail length, coat color and body structure. Based on these features and high picture quality (i.e. pictures including the complete individual or portion of individ-ual with distinct features visible, picture not blurred and

-tures for each individual. Following Oliveira-Santos et al. (2010), we counted only new individuals as those

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4 independent classifications of individual tapirs from all tapir pictures were made by the authors and collabo-

images and was requested to sort images by individu-al tapirs and record the attributes and main features used to identify each individual. We then compared the inde-

potential bias and error.After accuracy tests and identification, capture his-

tories were constructed for each individual by combin-ing respective photograph events into 5-day sampling intervals. This interval was selected after testing multi-ple sampling intervals to obtain overall maximum cap-ture probabilities following previous recommendations to estimate population size (Otis et al. 1978;; Trolle & Kery 2003;; González-Maya & Cardenal 2011). Cap-ture–recapture for closed populations (Otis et al. 1978) was used to estimate tapir abundance (program Cap-ture;; United States Geological Survey, http://www.

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Table 1 Habitat characteristics of camera sites for detection of Baird’s tapir, Valle del Silencio, Talamanca region, Costa Rica, Jan–Mar 2006

Site Captures (# of photographs) Slope (%) Canopy cover (%) Elevation (m asl)1 2 10 70 23792 2 32 45 23743 0 40 85 24774 3 28 65 25365 12 12 95 25066 0 34 55 25447 4 13 45 25168 9 33 30 25369 45 5 68 256310 0 11 65 2438

mbrpwrc.usgs.gov/software/capture.html). To estimate density, we calculated the mean maximum distance moved (MMDM) derived from maximum distances be-tween capture sites where individual tapirs were photo-graphed. The MMDM was divided by 2 then converted into a radius and used to buffer each camera to estimate the effective sampling area (ESA;; González-Maya et al. 2008;; González-Maya & Cardenal 2011). The ESA was then used to calculate tapir density (individuals/km2) us-ing the abundance estimate obtained from mark–recap-ture analyses divided by the effective sampling area.

RESULTSWe obtained 540 trap nights total for the 10 camera

stations. Mean distance between cameras was 1.95 km (SD 0.50). We obtained 96 high-quality usable photo-graphs of 9 mammal and bird species;; 77 photographs were of tapirs including females, males and juveniles (including pictures of paired individuals). Other mam-mal species included Leopardus pardalis (Linnaeus, 1758), Mazama americana (Erxleben, 1777), Tayassu tajacu (Linnaeus, 1758), Sylvilagus dicei Harris, 1932, Nasua narica (Linnaeus, 1766) and Didelphis marsupi-alis Linnaeus, 1758.

with a maximum dissimilarity of 6.4 % recorded be-tween pairs of observers. Thus, 5 images could not be assigned to an individual tapir and were excluded from further analyses. The most commonly used features for

individual identification were ear markings and body scars (93%;; Fig. 2). We identified 15 individual tapirs during the study, including 7 females and 8 males. Us-

Figure 2 Examples of characteristics used to identify individu-al Baird’s tapirs, (a,b) natural ear-marks and (c,d) body-marks and (e,f) photographs that were not possible to identify, Valle del Silencio, Talamanca region, Costa Rica, Jan–Mar 2006.

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ing 5-day sampling intervals, we obtained 25 captures of 15 individual tapirs at 7 of the 10 stations. Mean num-ber of photographs per individual was 1.60 ± 0.82, with a maximum capture of 4 photographs for 1 individual. There was no association between number of tapir im-ages and habitat characteristics (r9 < 0.26, P > 0.57;; Ta-ble 1).Mean distance between recapture stations was ob-

tained for 6 individuals, resulting in a mean maximum distance moved of 1.04 ± 0.27 (SD) km. Using half of the mean maximum distance moved (0.5 × MMDM) to create buffers around cameras resulted in an effec-tive sampling area of 7.16 km2 (Fig. 3). We select-ed this measure since no other information regarding home range for the species in this habitat was avail-able. Additionally, our study site was clearly defined by geographic boundaries (steep slopes on all sides) and preliminary surveys in the surrounding slopes failed to record evidence of tapir signs (e.g. tracks) or trails. Thus, the ESA constructed from 0.5 × MMDM con-tained all suitable tapir habitat as recommended by Fos-ter & Harmsen (2012);; consequently, we did not use a larger ESA to avoid underestimating tapir density.Capture analysis indicated a closed population

(z = –1.497, P = 0.067) and a capture probability of 0.02. The null model was the most parsimonious (Mo = 1),

followed by the heterogeneity model (Mh = 0.83) and time-behavior-heterogeneity model (Mtbh = 0.54) (Table 2). The estimated population size using model Mo was 21 individuals with standard error of 4.4 and a 95% con-fidence interval of 17–36 individuals. Using the ESA, we estimated a density of 2.93 individuals per km (95% CI = 2.37–5.03/km2) using model Mo and 2.79 individ-uals per km (2.37–4.74/km2) using model Mh. Capture probabilities for Mo and Mh models were 0.096 and 0.1, respectively, meeting the minimum threshold proposed by Otis et al. (1978) and Trolle and Kéry (2003).

DISCUSSIONWe documented the greatest density of Baird’s ta-

pir reported in the literature. Our study represents one of the first efforts to estimate Baird’s tapir density in mountainous areas which are typically not considered im-portant habitat. Other studies of tapir density in Central America and Mexico reported lower densities. Foerster and Vaughan (2002) estimated a mean density of 0.8 tapirs/km2 in Costa Rica and Naranjo and Bodmer (2002) estimat-ed 0.22 tapirs/km2 in Mexico using radio telemetry tech-niques. Using similar methodologies, Noss et al. (2003) found densities from 0.29 to 0.8 tapirs/km2 in 5 study sites in the Bolivian Chaco, and Trolle et al. (2007) esti-mated a density of 0.58 ± 0.11 in the Brazilian Pantanal, both for the lowland tapir (Tapirus terrestris Linnaeus, 1758). Thus, current estimates in Central America and other regions suggest lower tapir densities in lowland habitats. Habitat suitability for tapirs likely varies mark-edly in the Talamanca Cordillera. Our study was con-ducted in an area of presumed high tapir density;; conse-quently, extrapolating tapir abundance estimates across larger spatial extents in this region would result in over-all lower tapir densities.Despite recent studies suggesting problems us-

ing camera trapping for studying similar species (e.g. lowland tapir;; Oliveira-Santos et al. 2010), our results suggest camera trapping can be suitable for estimating tapir abundance. Oliveira-Santos et al. (2010) were un-able to accurately identify individuals using non-perma-nent or conspicuous markings. Our methods followed the same process recommended by these authors, in-

individuals, characterizing criteria used for individual

that were not consistently assigned to individual tapirs. We are uncertain of the disparity in results of tapir iden-tifications between studies. However, we recommend

Figure 3 Camera trap station locations for Baird’s tapir densi-ty estimation, Valle del Silencio, Talamanca mountains, Costa Rica, Jan–Mar 2006.

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that future studies report error rates in assignment of photographs to individual animals or, as Oliveira-Santos et al. (2010) indicate, report an estimate of agreement in assignment of photographs. Our results support use of camera trapping as a meth-

od for estimating abundance of cryptic species. Howev-er, as previously indicated (Trolle & Kery 2003;; Foster & Harmsen 2012), it is necessary to design robust stud-ies with clear study objectives to best address ecological parameters of interest. This means that for species such as Baird’s tapir, a design including accurate methods for

error in data analyses (Oliveira-Santos et al. 2010). Our study presented unique conditions that allowed most is-sues related with tapir density estimations using cam-era trapping to be addressed. First, our small study area allowed us to perform a robust design with appropriate density of camera stations. Second, our sample size de-rived from the number and quality of photographs al-lowed positive and accurate identification of individuals with adequate numbers of recaptures. Third, the usual ad hoc problems related with density estimation, such as the estimation of ESAs and MMDM, was accounted for due to topographic features of our study site defining our area of inference.Because the extent of lowland forests has been mark-

edly reduced in Central America, including Costa Rica, high elevation forests may represent important habi-tat for large mammal species such as tapirs. The low-

-ly affected during agricultural expansion in the 1970s and currently no large intact forests remain below 1500

m asl. This land-use change appears to have result-ed in severe isolation of tapir populations. Tapir pop-ulations persist in the Osa Peninsula including Corco-vado National Park (Foerster & Vaughan 2002) on the Pacific slope and Hitoy-Cerere Biological Reserve on the Caribbean slope;; however, no other evidence of tapirs occurs below 1500 m asl (González-Maya et al. 2009). Previous studies on the elevational distribu-tion of Baird’s tapir in Costa Rica (González-Maya et al. 2009) suggests variation in capture rates among el-evations, with mid-elevations (2000–2600 m asl) hav-ing the greatest capture rates. The Talamanca region is comprised of several ridges and high slopes, especially

-evations primarily in protected areas. As this rugged to-pography is typical of the region, inter-mountain valleys may represent the best available habitat for tapirs, sug-gesting suitable habitats of this cordillera are restricted to valleys and moderate slopes in the Caribbean.Because of severe fragmentation and discontinuity of

potentially suitable lowland habitats, maintaining via-ble tapir populations at higher elevations is important to conservation as it may ultimately provide the only po-tential population source for recolonizing restored low-land areas. However, there is no information on the de-gree of isolation of these populations. The Talamanca cordillera retains the largest intact forest block in Costa Rica (González-Maya et al. 2008) and, despite recent and historical records of tapirs from other parts of the coun-try, Talamanca is likely the only continuous habitat large enough to support viable populations of this species. Many questions remain;; for example, it is unclear how large mammals move across the cordillera and recent

Table 2 Capture–recapture model parameters and estimates of abundance for Baird’s tapir, Valle del Silencio, Talamanca region, Costa Rica, Jan–Mar 2006

Model Criteria Estimated abundance SE 95% CI Capture probability*Mo 1 21 4.4 17–36 0.096Mh 0.83 20 3.9 17–34 0.100Mb 0.33 24 15.48 16–103 0.074Mbh 0.5 24 15.4 16–103 n/sMt 0.02 20 3.7 17–33 0.100*Mth 0 21 4.7 17–38 0.097Mtb 0.37 16 0 16–15 0.187*Mtbh 0.54 26 11.5 18–74 n/s

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evidence suggests the Continental Divide may restrict movements of other large mammal species (González-Maya & Cardenal 2011). Understanding population con-nectivity and potential corridors across the divide is crit-ical for long-term viability of tapir populations.Talamanca represents an important region for large

mammal conservation in Costa Rica. Although poorly studied, recent efforts are improving our understand-ing of the importance of this region. Comprising al-most 25% of the country, the Talamanca region is con-sidered the ‘last frontier’ for wildlife conservation in Costa Rica (González-Maya et al. 2008). The Talaman-ca region also has comparatively high biogeographi-cal importance, species endemism and species richness (González-Maya et al. 2008). However, few studies have attempted to understand the role of this region for conservation of large species as they become increas-ingly isolated from the lowlands (González-Maya et al. 2008, 2009;; Schipper 2010;; González-Maya & Cardenal 2011). We suggest the Talamanca region is an important area for Baird’s tapir and other large mammal species and requires further studies to fully understand their role in the conservation of Costa Rica.

ACKNOWLEDGEMENTSWe thank Edward Jackson and Juan Mata for field

assistance. Special thanks to MINAET (Environmen-tal, Energy and Telecommunications Ministry of Costa Rica) staff for permits, logistics and support of the en-tire study, especially ACLA-P and PILA staff, Luis Sán-chez and Nelson Elizondo. Angélica Benítez, Elizabeth Navarro and Karla Rojas helped with data organization

-nando Casanoves and Bryan Finegan for their support through the entire process. MINAET/ACLA-P provided the permits for this research. Funding was provided by CATIE, The Nature Conservancy, ProCAT Internacio-nal, Sierra to Sea Institute and Finca Las Alturas.

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Baird’s tapir density in high elevation forests of the Talamanca region of Costa Rica

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