Talamanca Large Mammal Study – Progress Report 2015 Prepared August 2015

Dr. Mike Mooring, Ph.D.

Dylan Poorboy, Jessica Fowler, Ellen Asselin, Tanner Mathews Department of Biology, Point Loma Nazarene University, San Diego, CA 92106 USA

Email: mikemooring@pointloma.edu

Costa Rica Research Permits Nº 098-15 – ACLAP; Nº 016-1515 – ACOPAC

Summary

Our research group is conducting a long term survey of the large mammal fauna of the high

elevation Talamanca Cordillera, the major cloud forest region of Costa Rica. The aim is to fill the

gap in our understanding of jaguar and other cloud forest predators (6 species of felids plus coyote

and tayra), and to determine the status of major prey species that these predators depend upon

(e.g., peccary, deer, tapir). Although habitat loss from deforestation has largely ended in Costa

Rica, illegal hunting from adjacent agricultural areas is producing an ‘empty forest syndrome’ in

many of the region’s premier protected areas. Since 2010, we have investigated keystone

predators and prey using camera traps, genetic scat analysis, interview projects, and community

outreach to support conservation action and promote community-based conservation among local

residents. These activities involve the participation of local partners in national parks, private

reserves, and biological corridors to maintain our camea trap network. We also collaborate with

governmental agencies, conservation organizations, universities, and ecotourism operators.

Highlights from 2015 inculde the following points, which are detailed in this report:

We expanded our survey area north into the ACOPAC and ACT conservation areas

Dylan Poorboy completed his Honors project, occupancy modeling of our camera trap data

Carol Williams and Jessica Fowler completed an analysis of activity budgets of detected species

Ellen Asselin analyzed seasonal and annual fluctuations in detections of predators and prey

Tanner Mathews reports on the successfully use of our detection dog ‘Berry’ to find felid scat

Stephanie Butera conducted an interview project in the Alexander Skutch Biological Corridor

We monitored and deployed new cameras in La Cangreja and Carara National Parks

We travelled to Cabo Blanco National Park to deploy a new camera trap network

The team climbed to the summit of Mount Chirripó to deploy new cameras in the paramo

We deployed a new and larger camera trap network at Rio Macho Forest Reserve, Villa Mills

A record number of 8 charlas were held to promote community participation in the project

Photos from a biological corridor and a national park show the presence of illegal hunters

Amazing photos continue to be taken, e.g., a baby tapir, group of 3 pumas, and tamanduas

We were invited to share our camera trap data with 3 nationwide inventory projects

Table 3 on page 33 lists the 26 mammalian species found by our camera traps since the project began.

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Analysis of Occupancy for Mammalian Species in Tropical Montane Ecosystems Honors Project by Dylan Poorboy

Abstract The Talamanca Cordillera of Costa Rica is

important habitat for many species, possessing high levels of

species richness and some of the largest populations in the

country of several wide-ranging species such as jaguar

(Panthera onca) or endangered species such as Baird's Tapir

(Tapirus bairdii). Unfortunately, our understanding of

mammalian populations within this region is limited.

Occupancy modeling can help to illuminate the ecological dynamics of a site using collections of

detection and non-detection data to estimate the proportion of habitat occupied by a species.

Data collected from camera trap surveys throughout the Talamanca Cordillera over four years was

used for occupancy modeling. With the program PRESENCE, I obtained measures of species

occupancy for the region as a whole and for specific surveys within the region using single species-

single season models, with model likelihood determined by the Akaike Information Criterion (AIC).

Region-specific occupancy estimates were obtained for the 24 large mammal species detected in

the camera trap survey, including those with rare detection data. Site-specific occupancy

estimates were also obtained for many of the detected species. Tapantí National Park, Los

Quetzales National Park, and the Savegre Valley showed a significant effect of elevation on

occupancy. Estimation of occupancy for species in the Talamanca Cordillera provides new

information on the ecological dynamics of large mammal species in this region. These findings can

be useful in informing future studies and guiding conservation action.

Introduction Occupancy modeling is an important tool for assessment of detection/non-

detection data such as produced by camera trap surveys. This form of modeling uses detection

histories to yield important variables: ‘occupancy’ (ψ), defined as the proportion of an area

occupied by a species, and ‘probability of detection’ (p), which can be defined as the probability of

detecting an individual of a species within a sampling occasion. ‘Naïve occupancy' is simply the

proportion of sites with detections out of the total pool of sites, but this measure does not

account for imperfect detection. Realistically, detection is never expected to be perfect and the

naïve occupancy value would be an underestimation of the species true occupancy. Occupancy

modeling seeks to solve this issue through use of the whole detection history for a species in a

linear regression mathematical model. This form of population modeling also allows for

incorporating variables referred to as covariates (e.g., elevation, distance to roads, human

population), which enable the determination of environmental factors that may influence

occupancy and can increase model precision. The aim of this study was to determine occupancy

and detection probability for the mammals of the Talamanca Cordillera.

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Methods

This study was conducted as six distinct surveys along the Talamanca Cordillera. They were (1) the

Savegre Valley, including Los Quetzales National Park and Los Santos Forest Reserve, and featuring

Cerro Buenavista (second highest mountain in Costa Rica at 3491 meters) and the Cerro de la

Muerte massif; (2) the Alexander Skutch Biological Corridor (ASBC), which connects Chirripó

National Park to the forest fragments of Las Nubes Biological Reserve and Los Cusingos

Neotropical Bird Sanctuary; (3) Chirripó National Park, featuring Cerro Chirripó (highest peak in

Central America at 3820 meters) and high elevation páramo habitat; (4) Tapantí National Park,

which receives the highest rainfall in Costa Rica (7000mm), is an enormous area much of which is

inaccessible due to lack of trails; (5) La Marta Wildlife Refuge; and (6) El Copal Biological Reserve;

both La Marta and El Copal are located north of Tapantí National Park near Pejibaye. Occupancy

analyses were conducted using detection histories for each mammal species from individual

camera trap units as occupancy sites and 12 sampling occasions of one month each over the

course of a year. Total camera trap days were recorded to measure sampling effort. Analysis of

species occupancy for the entirety of the survey was conducted by dividing the study region into

100 km2 zones over a 3600 km2 grid (Figure 1). Detection histories for each mammal detected in

any of the surveys for a year were assembled using the 36 grid zones as occupancy sites. Sampling

years were divided in the full year format, treating each month as an individual sampling occasion.

Figure 1. Occupancy sites for the full survey analysis. All numbered zones are 100km2. Camera symbols

represent locations of camera stations. Zones without camera stations were treated as “unsampled".

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Results

The complete study region of the Talamanca Cordillera was assessed for the 2012-2013 and the

2013-2014 sampling periods. Total sampling effort for this survey consisted of the combined effort

of the Tapantí, Savegre, Chirripó and ASBC surveys with 24,396 combined camera trap days during

the 2012-2013 sampling period, and 26,237 camera trap days during 2013-2014 sampling period.

Twenty-three species were detected across the region during both years, of which 19-20 species

yielded estimates of species occupancy (Figure 2, Table 1). The highest calculated occupancy

estimate for a species during 2012-2013 was 0.8661 for Dice’s cottontail, while the lowest was

0.1254 for Nine-banded armadillo. For 2013-2014, the highest occupancy estimate of 1.000 came

from coati while the lowest estimate was 0.1134 for white-tailed deer.

Within the region, the Tapantí and Savegre surveys showed the greatest support of elevational-

based models, meaning that elevation appears to influence the presence or detection of species.

Specifically, elevation gradients were associated with occupancy of Baird's tapir, red brocket deer,

collared peccary, raccoon, coati, and Dice's cottontail among prey species, and influenced

occupancy for oncilla, margay, ocelot, puma, tayra, and coyote among predators. It further

appears that puma may be affected by the elevation gradient within ASBC and Chirripo surveys.

Figure 2. Occupancy (Ψ) results for both years of the Talamanca Cordillera region assessment.

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Table 1. Occupancy (Ψ) and probability of detection (p) results for Talamanca Cordillera region.

Common Name Species Name Naïve Occupancy Occupancy(ψ) Detection Prob. (p) Model Type AIC Weight

Paca Agouti paca 0.4286 0.5351 (± 0.2024) 0.2403 (± 0.0905) Royle/Nichols 0.7766

Coyote Canis latrans 0.7500 0.8069 (± 0.1171) 0.2705 (± 0.0741) Royle/Nichols 0.9993

Mexican Hairy Porcupine Coendou mexicanus 0.3750 0.5298 (± 0.3339) 0.0946 (± 0.0773) 1 Group, Constant P 0.5198

Striped Hognosed Skunk Conepatus semistriatus 0.6250 0.7846 (± 0.1741) 0.1499 (± 0.0677) Royle/Nichols 0.7578

Agouti Dasyprocta punctata 0.1250 0.1257 (± 0.1176) 0.6658 (± 0.1366) 1 Group, Constant P 0.5117

Nine-Banded Armadillo Dasypus novemcitus 0.1250 0.1254 (± 0.1192) 0.4136 (± 0.1439) 1 Group, Constant P 0.5083

Common Opossum Didelphis marsupalis 0.2500 0.2842 (± 0.1764) 0.2648 (± 0.1128) Royle/Nichols 0.5236

Tayra Eira barbara 0.6250 0.7767 (±0.2002) 0.1316 (± 0.0703) Royle/Nichols 0.6949

Ocelot Felis pardalis 0.3750 0.4457 (± 0.1771) 0.3326 (± 0.0947) Royle/Nichols 0.7611

Oncilla Leopardus tigrina 0.7500 0.8097 (± 0.1443) 0.1524 (± 0.0627) Royle/Nichols 0.9673

Red Brocket Deer Mazama americana 0.6250 0.7190 (± 0.1443) 0.2712 (± 0.0727) Royle/Nichols 0.9989

White-nosed Coati Nasua narica 0.5000 0.6592 (± 0.1553) 0.3091 (± 0.0783) Royle/Nichols 0.9929

Gray Four-Eyed Opossum Philander opossum 0.2500 0.2746 (± 0.1684) 0.2488 (± 0.0950) 1 Group, Constant P 0.4813

Northern Raccoon Procyon lotor 0.3750 0.4748 (± 0.1853) 0.2922 (± 0.0946) Royle/Nichols 0.8343

Puma Puma concolor 0.7500 0.7474 (± 0.1436) 0.3652 (± 0.0960) Royle/Nichols 0.8129

Dice's Cottontail Sylvilagus dicei 0.7500 0.8661 (± 0.0963) 0.3634 (± 0.0852) Royle/Nichols 0.9974

Northern Tamandua Tamandua mexicana 0.1250 0.1314 (± 0.1230) 0.3273 (± 0.1384) 1 Group, Constant P 0.5093

Baird's Tapir Tapirus bardii 0.7500 0.8241 (± 0.1098) 0.3822 (± 0.0833) Royle/Nichols 0.9982

Collared Peccary Tayassu tajuca 0.6250 0.6293 (± 0.1724) 0.3373 (± 0.0683) 1 Group, Constant P 0.6431

Common Name Species Name Naïve Occupancy Occupancy (ψ) Detection Prob. (p) Model Type AIC Weight

Paca Agouti paca 0.5000 0.5939 (± 0.1708) 0.2164 (± 0.0759) Royle/Nichols 0.8893

Coyote Canis latrans 0.6000 0.6914 (± 0.1369) 0.3140 (± 0.0787) Royle/Nichols 0.9973

Striped Hognosed Skunk Conepatus semistriatus 0.4000 0.4468 (± 0.1721) 0.3132 (± 0.0711) 1 Group, Constant P 0.5324

Agouti Dasyprocta punctata 0.3000 0.3531 (± 0.1747) 0.4575 (± 0.1390) Royle/Nichols 0.5531

Nine-Banded Armadillo Dasypus novemcitus 0.4000 0.6346 (± 0.2893) 0.1149 (± 0.0570) 1 Group, Constant P 0.5164

Common Opossum Didelphis marsupalis 0.3000 0.3865 (± 0.1696) 0.3177 (± 0.1024) Royle/Nichols 0.7169

Tayra Eira barbara 0.6000 0.7949 (± 0.2113) 0.1608 (± 0.0530) 1 Group, Constant P 0.6079

Ocelot Felis pardalis 0.5000 0.5398 (± 0.1683) 0.3711 (± 0.0698) 1 Group, Constant P 0.5454

Jaguarundi Herpailurus yaguarondi 0.3000 0.8352 (± 0.7247) 0.0504 (± 0.0473) 1 Group, Constant P 0.5014

Oncilla Leopardus tigrina 0.4000 0.5909 (± 0.2413) 0.1277 (± 0.0727) Royle/Nichols 0.6181

Margay Leopardus wiedii 0.3000 0.3964 (± 0.2000) 0.1704 (± 0.0709) 1 Group, Constant P 0.5061

Red Brocket Deer Mazama americana 0.7000 0.7182(± 0.1383) 0.3295 (± 0.0858) Royle/Nichols 0.8145

White-nosed Coati Nasua narica 1.0000 1.0000 (± 0.0000 0.4362 (± 0.0511) Royle/Nichols 0.4239

White Tail Deer Odocoileus virginianus 0.1000 0.1134 (± 0.1083) 0.2569 (± 0.1398) 1 Group, Constant P 0.5063

Gray Four-Eyed Opossum Philander opossum 0.2000 0.2763 (± 0.1180) 0.1466 (± 0.0868) 1 Group, Constant P 0.5081

Northern Raccoon Procyon lotor 0.3000 0.3050 (± 0.1474) 0.3946 (± 0.1004) 1 Group, Constant P 0.5089

Puma Puma concolor 0.9000 0.9804 (± 0.0395) 0.1670 (± 0.0779) Royle/Nichols 0.9683

Dice's Cottontail Sylvilagus dicei 0.8000 0.8872 (± 0.0799) 0.2533 (± 0.0632) Royle/Nichols 1.0000

Baird's Tapir Tapirus bardii 0.8000 0.8411 (± 0.0974) 0.3645 (± 0.0830) Royle/Nichols 0.9992

Collared Peccary Tayassu tajuca 0.8000 0.8411 (± 0.0974) 0.3645 (± 0.0830) Royle/Nichols 0.9992

Talamanca Cordillera 2013-2014

Talamanca Cordillera 2012-2013

Occupancy was not calculable for many species in each of the four major surveys, an unfortunate product of the models failing when occupancy and/or detection probability was too low. The naïve occupancy values can represent a lower bound for occupancy of these rare species (Figure 3, Table 2). In this way they are still an underestimation of the species occupancy but they are not wildly out of the potential range of occupancy.

Table 2. Occupancy (Ψ) results for Talamanca Cordillera rare species.

Common Name Species Name Naïve Occupancy

Cacomistle Bassariscus sumichrasti 0.2500

Jaguarundi Herpailurus yaguarondi 0.2500

Margay Leopardus wiedii 0.2500

Jaguar Panthera onca 0.1250

Common Name Species Name Naïve Occupancy

Mexican Hairy Porcupine Coendou mexicanus 0.1000

Jaguar Panthera onca 0.1000

Northern Tamandua Tamandua mexicana 0.2000

Talamanca Cordillera 2012-2013 Rare Species

Talamanca Cordillera 2013-2014 Rare Species

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0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1

Cacomistle

MexicanHairyPorcupine

Jaguarundi

Margay

Jaguar

NorthernTamandua

Occupan

cy(ψ)

Species

TalamancaRegionRareSpecies

TalamancaCordilleraRareSpecies2010-2014

Figure 3. Occupancy (Ψ) results of rare species analysis for the Talamanca Cordillera region.

Conclusions

This study supports the assumption that species present at a site are not always detected.

Assuming that full detection is achieved, the naïve occupancy would be representative of the true

occupancy. However, when camera trap detection histories were incorporated in calculation of

species occupancy, naïve estimates were identified to consistently underestimate occupancy. The

results presented provide improved estimates of occupancy plus or minus the associated standard

error of the modeling calculation. For the species that occupancy estimates were calculated for,

new baselines have been established. Because information on the ecological dynamics of the

Talamanca Cordillera is highly inadequate, these findings will be useful in better describing this

habitat. This study is an important step in improving our understanding of the crucial montane

cloud forest habitat and may provide direction for future study and informing improvement of

current surveys.

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Activity Budget Analysis

By Jessica Fowler and Carol Williams

Abstract Little is known about the behavior of large mammals in the montane cloud forests of the Talamanca Cordillera of Costa Rica. Using time stamp data from the camera trap study, we analyzed activity budgets for large mammal species across several study sites to explore the relationship between predators and their prey. Activity budgets of predators (puma, ocelot, oncilla, coyote) were compared to their prey species (peccary, brocket deer, agouti, coati, and cottontail) to determine if activity cycles coincided, which would be indicative of predator-prey interaction. Activity budgets were also compared across study sites to determine whether site-specific differences

existed, possibly indicating disturbance (e.g., hunting). Finally, the activity budgets of invasive coyotes were compared with similar-sized native predators to determine if competitive displacement might be occurring. Results indicated that each predator had comparable activity budgets with at least one of their known prey species, confirming that predator-prey interactions are taking place. There were no major discrepancies in activity budgets across study sites that we could detect. Coyotes did not appear to influence puma or ocelot activity, implying that competitive displacement is not yet occurring, although the situation needs to be monitored. These results will help guide conservation action in this cloud forest region.

Methods

We constructed activity budgets by extracting the date and time stamp data from Camera Base entries for 13 mammals detected on our camera traps in the Savegre Valley, Tapantí National Park, Alexander Skutch Biological Corridor (ASBC), and Chirripó National Park from summer 2010 through summer 2014. We subsequently compared the activity patterns of all predators and their potential prey to assess the degree of overlap in activity budgets. We compiled time stamp data for each species based on ‘events’, with an event defined as an independent detection of an animal or group of animals that was separated by a minimum of ten minutes from other such events. Rounding to the closest hour in a 24-hour period, we then categorized each event into hourly increments. For each species, we divided the events detected in each hour by the total number of events to determine the proportion of activity occurring during each hour. To compute the percent overlap between two species, we compared the hourly proportion of activity for both species to calculate the overall percent overlap of the two species. Using the same method detailed above, we compared the predator and prey species of Savegre Valley to the same species in Tapantí, ASBC, and Chirripó. We selected coyote, puma, and ocelot as representative predators, and Dice’s cottontail, coati, collared peccary, and red brocket deer as resentative prey. These species were chosen because they had the most detections.

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Results Activity budget data were converted into graphs for visual inspection. To illustrate the activity data for a single species, bar graphs were prepared with ‘Hour of Day’ (military time) as the X-axis and ‘Events’ as the Y-axis (Fig. 4). By this method, a species could be categorized by its dominant activity pattern as either diurnal (mostly active during the day), nocturnal, (mostly active at night), or cathemeral (active during both day and night); examples are given in Figure 5. Predator species of the Savegre Valley were first examined for dominant activity budget (Fig. 6). Puma were the most cathemeral of the predators, being active during all hours of the day and night except at midday. Ocelot, oncilla, and coyote were primarily nocturnal, although all were sometimes active during the day as well (Fig. 6).

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We next examined the activity budgets of Savegre Valley prey species (Fig. 7). Some prey species were primarily active during the day (peccary, red brocket deer, coati), while others were mostly nocturnal (Dice’s cottontail, paca, Baird’s tapir). However, all ‘diurnal species’ could be active at night, whereas ‘nocturnal species’ (especially tapir) were sometimes active during the day (Fig. 7). Thus, based on overlap of activity patterns, one could expect cathemeral puma to prey more on peccary (or brocket deer or coati) than the more nocturnal predators. To analyze the degree of overlap between two species, we constructed line graphs with ‘Hours of Day’ as the X-axis and ‘Percentage’ as the Y-axis. To quantify the activity during each hour of the day, we used percentage of total events (rather than events) to standardize activity per hour among species with different number of events observed, thus allowing for comparison. We compared the activity budgets of all predators of the Savegre Valley (Figs. 8 and 9). Figure 8 shows predators with a high degree of activity overlap, being active during the same hours for at least 50% of their activity budget. Puma overlapped about 60% of their activity budget with other felids (ocelot and oncilla) and nearly 80% with coyotes, with the main difference being that puma were more active during the day. Coyotes had high overlap with ocelot (73%) and oncilla (61%), with the main difference being that coyotes were more active at night. Ocelot and oncilla (65% overlap) appeared to be active at essentially the same parts of the day.

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Figure 9 compares the activity budgets of tayra with other predators in the Savegre Valley. The degree of overlap was relatively low, ranging from 26% overlap with ocelot, to 50% overlap with puma. In all cases the low degree of overlap was due to tayra being much more active during the day and less active at night than the felids and coyote. Tayra illustrate a diurnal activity pattern and thus one would expect to see tayra preying on mostly diurnal species of prey. Figures 10 and 11 compare the overlap in activity between a representative predator (puma) and the major prey species of Savegre. Species for which activity overlap with puma was relatively high (Fig. 10) include Baird’s tapir, Dice’s cottontail, raccoon, and paca. The highest overlap (72%) was between puma and tapir activity, although it is unlikely that puma regularly hunt tapir given that Baird’s tapir (up to 400 kg) is five times larger than Central American puma (up to 80 kg).

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The other prey species (raccoon, cottontail, paca) from Figure 10 had a very similar degree of overlap with puma, between 56% to 60%. Figure 11 compares the relatively lower degree of activity overlap between puma and brocket deer (54%), peccary (53%), and coati (46%). In the case of each of these prey species, overlap was lower due to the prey species being more active during the day and less active in the early morning hours. All things considered, we would expect puma to hunt these species less frequently than those in Figure 10.

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Conclusions Activity budget data suggest that all Savegre Valley predators are active to some extent during most hours of the day, with ocelot, oncilla, and coyote being the most nocturnal, tayra being the most diurnal and puma being the most cathemeral (active both day and night). Thus we would predict that tayra will specialize on the mostly diurnal prey species (peccary, brocket deer, coati), ocelot, oncilla, and coyote would tend to hunt the mostly noctural prey (cottontail, paca), while the cathemeral puma would being able to prey on all these species plus tapir. One possibility is that puma prey on a higher diversity of species than the other predators. To test these predictions will require extensive scat collection for analysis of prey species consumed, which we hope to do with more extensive use of a scent detection dog in the future.

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Community outreach

Every year our research team gives presentations (“charlas” in Spanish) to different communities to explain our project and promote community-based conservation (Fig. 12). This year our team presented a record number of charlas to partners and communities, giving eight (8) charlas that were attended by about 175 people. In June we spoke in 3 communities in the Alexander Skutch Biological Corridor (ASBC) to the south of Chirripó National Park. Over two days, we spoke in Santa Elena, Quizarrá, and Montecarlo to a total of about 70 people. The next day we went to Villa Mills to speak to about 20 local residents at the Rio Macho Forest Reserve station. We subsequently returned to put up new camera stations on several kilometers of trails. In mid-July we travelled to Cabo Blanco National Park to set up a new camera trap network and gave a well-received charla to about 15 staff and volunteers. This was followed by a presentation given at San Rafael de San Pedro de Cajón to speak to members of the Bosque Agua and the Rio Peñas Biological Corridors, located south of Chirripó and La Amistad National Parks. About 20 leaders attended from the community and committed to deploying and monitoring two arrays of camera traps, one in each corridor. We in turn will provide 10 cameras total, 5 in each corridor. The following day we travelled to a well-attended charla at the Chesperitos truck stop in Ojo de Agua, attended by 25 key community leaders. The presentation was followed by a lively discussion and there was great interest in putting up new cameras on private lands in the area. Our final charla two days before our departure was our annual community informational meeting at QERC in San Gerardo de Dota, with a packed room of about 25 neighbors who came to hear the latest news.

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Community Outreach in the Alexander Skutch Biological Corridor

By Stephanie Butera, York University

Stephanie Butera is beginning a Masters in Environmental Studies in the York University Faculty of Environmental Studies in Toronto, Canada. She came to Costa Rica this summer to participate in a field course during May and subsequently spent 3 weeks in the Alexander Skutch Biological Corridor (ASBC) working on a community outreach project in collaboration with our camera trap mammal survey. Stephanie is also developing a website where local residents can report wildlife sightings that would be available to others. The following is a summary of her report:

To assess the success of biological corridors, it is essential that biodiversity be measured. The focus of my project was to conduct a thorough survey of the current abundance of endemic mammal species in the Alexander Skutch Biological Corridor (ASBC). With the help of Luis Angel Rojas (field coordinator of York University’s Las Nubes Project), I conducted personal interviews with 46 members of the communities of Quizarrá, Montecarlo, and Santa Elena. Each participant was asked about their perception of the abundance of wildlife and how it may have changed over time. The interviews were conducted orally in Spanish as informal conversations, mostly within the participant’s home or place of work. The residents are rural people who mostly make a living growing coffee and other crops. I also organized public meetings (charlas) by inviting local community members to attend. Community meetings consisted of PowerPoint talks given by Dr. Mike Mooring to explain the camera trap survey, and by myself to report the results of the interview surveys. At the end of each meeting, we solicited community participation in the mammal survey project through assisting with camera trap monitoring. Farm owners were also asked to volunteer their land to install new cameras. The goal is to develop the capacity within each community for them to conduct their own community-based monitoring program so that the camera trap project can ultimately be taken over by the local people.

Interview Results All of the subjects that were interviewed had lived in the corridor for a minimum of 20 years and 60% were born in ASBC and have lived there their entire lives. The survey participants were asked to list the mammal species that they believe have increased this year compared to previous years, and those that they believe are in decline (Figs. 14-15). Squirrels were said to be very common, along with foxes, agouti, white-tailed deer and paca. Coyotes are also frequently encountered, although most people mentioned hearing them more than actually seeing them. The white-faced capuchin monkey is also quite abundant in the corridor, although it is the only monkey that is seen with frequency. People have reported seeing puma in Montecarlo (camera traps have captured the puma in Quizarrá as well). The puma in Montecarlo is said to have been travelling with a cub, and was spotted in the last six months. A few people in both Montecarlo and Quizarrá have also spotted jaguarundi. Some of the species were only spotted in the forested mountains north of the corridor, i.e., jaguar, tapir, and peccary.

The mammal that is said to have declined most noticeably is the armadillo. In each of the three towns there was consensus that armadillos were disappearing from the corridor, and they noticed a drastic drop in the number of armadillos starting 5-10 years ago. A few of the men spoke about

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hunting when they were younger and that armadillos were hunted for their meat because of its good taste. Some men also wondered whether agricultural chemicals could have contributed to the decline of armadillos. Only 12 people have seen a river otter this year and the general consensus is that their population is decreasing. The decline in river otters is believed to be caused mainly by habitat loss.

The results agree that there are three species that are experiencing population growth in the corridor: coyote, white-tailed deer, and squirrels (Fig. 14). Most people saw significant increases in their populations in the past 5-10 years. Many said that previously it was impossible to spot a deer, and now they have seen several. The population increase could be due to an increase in food availability. The coyote population has exploded recently, although they are not endemic to the area. Human activity provides suitable prey and habitat for coyotes, enabling coyote ranges to expand with the expansion of agricultural systems and urbanization. Many people see the coyote as a nuisance and some have blamed them for the disappearance of smaller prey species. Multiple individuals have seen the fur of armadillos, peccary, and other small mammals in what they identified as coyote scat.

The people in Santa Elena seem to have a smaller population of mammals than the rest of the corridor (Fig. 15). The difference could have been due to the attitudes of people living within the towns. People living in Santa Elena showed the least interest in the project and conservation, while people in Montecarlo were extremely interested and wished to gain further knowledge and/or participate. Every person interviewed had a positive outlook on the conservation of mammal species in their communities. Many expressed joy in seeing other species around them and they hope that their children and grandchildren will be able to witness the existence of the species that they saw growing up.

Community Meeting Results The charlas attracted a total of about 83 persons over two days. The meeting in Montecarlo was the most successfully, attracting 50 people out of which 16 locals volunteered to participate, with two volunteering their land for camera trap locations. The most common local reaction was pleasure to see that such a wide array of biodiversity exists within their community, and all were grateful that the information was being shared with them. The comments indicated their astonishment at the number of living creatures we have documented in their community.

Conclusions The data produced through personal interviews compares nicely to the data acquired from the cameras, providing evidence that traditionally acquired knowledge has ecological value. The camera trap data from the corridor reveals that the perceptions of the locals interviewed generally reflects the species makeup of the area. However, the data indicates that locals are incorrect in some instances; many interviewees expressed the belief that the populations of paca and agouti had diminished, whereas the camera traps capture them frequently. These animals are nocturnal however, making it more difficult for humans to observe. The interviews are important because they (a) reveal historical trends that began before the camera trap project, (b) may provide important insights into why certain trends are occurring, and (c) help solicit community involvement and interest in the project.

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Editorial Note: The increase in populations of white-tailed deer and coyote detected in the 2015 interview study agrees with the 2014 interviews we conducted in which many interviewees reported an apparent increase in white-tailed deer and coyote sightings over the past five years. White-tailed deer may be increasing due to the increase in agricultural land and the 20% loss in forest since 1998. Coyote may also be expanding their presence with the increase in open lands. With the loss of forest lands, increase in human population, and associated expansion of hunting activity, there may have been a decline in the predator community leading to a trophic cascade in which frugivores like agouti have been able to expand their populations. As noted by Stephanie, the interview project also revealed that there is geographical variation in the mammalian species detected in different communities, with interviewees in Santa Elena reporting only 21 species compared with residents of Montecarlo that reported sighting 35 species (Fig. 15).

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Illegal Hunters and Vandalism of Camera Traps

One of the reasons we do community outreach activities such as the interviews and charlas previously described is because illegal hunters are present in many of the areas where we work. These hunters are occasionally seen on the cameras, and sometimes they vandalize the cameras in an attempt to avoid detection (Fig. 16). We have had the most cameras destroyed in the Alexander Skutch Biological Corridor, and one stolen at the Rio Macho Forest Reserve. The photos in Figure 16 indicate that illegal hunting still occurs regularly on private lands, wildlife reserves, and within national parks. Some hunters are apparently afraid that the cameras will be used to identify and capture them, and so they destroy the cameras. For this reason, we always emphasize in our charlas and community outreach activities that the purpose of the cameras is to survey wildlife and NOT to capture hunters. From our perspective, we are not the police and do not intend to use the cameras to prosecute hunters because we believe this would be counter-productive. Our hope is that through the camera trap project we can empower communities to engage in their own citizen science, community-based monitoring, and environmental education, and in this way win hunters over to the side of conservation. We feel that community-based conservation activities will increase awareness about the human benefits of healthy ecosystems and the importance of protecting wildlife and ecosystem services. Through these strategies we hope in time to convince hunters that their future is with conservation and not with hunting.

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Camera trap study in Paso de las Lapas Biological Corridor

In late 2014, our team collaborated with officials from ACOPAC (Pacific Central Conservation Area) to initiate a new camera trap survey in the Paso de las Lapas (Path of the Macaws) Biological Corridor and the two protected areas that it connects (La Cangreja and Carara National Parks). Spearheading and coordinating the project is Leonel Delgado (stationed at La Cangreja), with assistance from Adrián Arce (ACOPAC Wildlife Coordinator) and Julio Bustamante (ACOPAC Research Coordinator). The purpose of the study is to compare the abundance of large mammals in Carara, La Congreja, and Las Lapas in order to assess the effectiveness of the corridor in facilitating the movement of large mammals, with a focus on felid predators. The project is designed to link wildlife management and research monitoring, and to promote environmental education in communities. This year we travelled twice to ACOPAC to assist in field surveys. In June, the team spent three days moving, deploying, and monitoring cameras in each area: La Cangreja National Park (PNLC), Carara National Park (PNC) and Las Lapas Biological Corridor (see Fig. 17).

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Each of these conservation areas now have 5 camera trap stations, enlarged from 4 cameras each last year. We first travelled to La Cangreja National Park where we arrived at the ranger station late at night after many hours driving up the Coastal Highway. The next day, Leonel led us as we hiked the Sancho and Rio Negro Trails and monitored the camera stations (Fig. 18). We later returned to deploy a fifth camera. The following day we drove throughout the Biological Corridor in order to move cameras to more productive areas. We stayed that evening at the Punta Mala ranger station on the ocean and the next day hiked the Quebrada Bonita and Laguna Meandrica trails at Carara National Park to deploy new cameras (Fig 19).

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New Camera Trap Project at Cabo Blanco National Park

This year we had the opportunity to travel to the Nicoya Peninsula to help deploy a new camera trap survey at Cabo Blanco National Park. (Although it’s formal name is the “Cabo Blanco Absolute Nature Reserve”, it has national park status.) Cabo Blanco was Costa Rica’s very first nature reserve, formed in 1963 as the result of the efforts of Nicolas Wessberg (a Swede) and Karen Mogensen (his Danish wife) who had immigrated to Costa Rica in 1954 and had a nearby farm. Our friend and collaborator Leonel Delgado (La Cangreja National Park) organized the trip through his extensive contacts within the national parks system. To get to the Nicoya Peninsula we travelled with Leonel and his family, taking the ferry from Puntarenas to Paquera and driving to the ranger station at the end of the peninsula (Fig. 20). Over two days, with the help of Andres Jimenez (administrator of Cabo Blanco), Leonel, and a number of volunteers, we deployed six cameras along two trails that access the interior of the park (Fig. 21 and 22). Andres and his staff were generous in their hospitality, providing the team with free accommodation and meals during our stay. As mentioned earlier, we were able to give a presentation to staff and volunteers to explain our research project. We are eager to see the first photos from this new survey.

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Figure 21. Cameras deployed at Cabo Blanco National Park: (A) Location of cameras on eastern side of park, (B) close-up of camera stations.

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Ascent of Mount Chirripó

Working with Enzo Vargas (Fig. 23A), the research and community outreach official at Chirripó National Park, the research team hiked 14 km on the Main Trail of the park to bring new camera equipment to the Base Crestones station for deployment at high elevations. The plan was to install metal posts into concrete bases so that camera stations could be established at Sabana de los Conejos (Savanna of the Rabbits) in paramo habitat. Because there are no trees in the paramo, we have been prevented until now from monitoring the mammals at these high elevations. The volunteer crew was delayed by a day, so our team climbed the peak of Mount Chirripó the following morning (Fig. 23B and 24) and returned down the mountain, a trek of 20 km. The posts and camera trap stations were later installed in Sabana de los Conejos and we will soon see what animals frequent this highest and least studied mountain habitat of Costa Rica.

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Berry the Detection Dog

For the past year we have been training our mini Australian Shepherd ‘Berry’ to locate felid scat using her finely developed sense of smell. Using scat samples of jaguar, puma, and ocelot from the San Diego Zoo and Safari Park, we spent around 180 hours in the field training Berry to find hidden samples in response to the command “find scat!” and to sit down to indicate that she has found it. Although her training sessions went very well, finding scat on the trails can be very challenging and so we were eager to test her in a field situation. To get her to Costa Rica we had to ship her as live cargo, requiring us to navigate the complicated and expensive import and export regulations and to hire an agent to get her in and out of Costa Rica. We now understand the process and have made reliable contacts in Costa Rica to help us in the future. Having overcome the travel issue, we then encountered the challenge of getting a collection permit through SINAC, the National System

of Conservation Areas. Although our project has the support of the University of Costa Rica, a delay in the process prevented us from getting the collection permit for ACLAP (La Amistad Pacific Conservation Area), where much our work is done. Thus we were unable to collect scat at Los Quetzales, Tapantí, and Chirripó National Parks this year. We were, however, able to use Berry on the Savegre Valley trails, and at La Cangreja and Carara National Parks in ACOPAC (Fig. 25). Handled by Tanner Mathews, Emma or Mike Mooring, she performed excellently on the trails and enabled us to collect 13 samples, of which many appeared to be puma. The scat samples were frozen and delivered to our collaborator Gustavo Gutierrez of the University of Costa Rica Wildlife Genetics Lab for analysis. Next year we hope to use Berry on many other trails.

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Fluctuations in Mammal Detections in the Savegre Valley

Over the years since the start of the study in 2010 we have noticed fluctuations in the number of photo detections of mammal species, especially in the Savegre Valley where we have continuously maintained a camera trap network for five years. For example, there have been years in which puma were frequently sighted on the trails, and other years in which puma detections were rare. This year, Ellen Asselin conducted a preliminary analysis of seasonal and annual fluctuations in detections to formally test our intuition. In agreement with our prediction, analysis of Savegre mammals showed regulation fluctuations over 15 sampling periods from summer 2010 through summer 2015. Of predators, coyote had the greatest year-to-year variation in detection, followed by puma (Fig. 26). Examination of prey species detections also revealed seasonal and annual oscillations, with the greatest fluctuations by Dice’s cottontail, collared peccary, and paca (Fig. 27).

Overlaid on the oscillations, we also saw a gradual increase in detections for some species (tapir, cottontail, armadillo, peccary), suggesting that these populations may be expanding (Fig. 28).

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Revised Camera Trap Array at Villa Mills

Last year a 5-camera network was established at the Rio Macho Forest Reserve in Villa Mills. One of the cameras was stolen in February 2015, and others were no longer working. The research team conducted a tour of the trails on July 7th, accompanied by Roberto Delgado. We were able to deploy a total of 7 cameras in a new camera trap array that now extends further into the reserve (Fig. 29). To avoid future vandalism, we attached to each camera a laminated card that explains that the camera is part of a mammal survey. Because the internet access at the station is limited, Adrian Ugalde (administrator) will in future bring the memory cards to Jordan Young at QERC in to share the photos. The Villa Mills site is important because it provides a point of access into the vast backcountry of Rio Macho Forest Reserve and Tapantí National Park that we have been unable to survey. We hope to obtain many more photos of elusive mammals this year (Fig. 30).

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Camera Trap Data Sharing and Collaboration

In 2014-2015 we had 3 invitations to share our camera trap data with Costa Rica projects for the purpose of preparing collaborative databases and publications. In each case, we contributed our data from the Talamanca Cordillera survey that will contribute to a nationwide database (Fig. 31). Lorena Lobos, a Masters student at the National University of Costa Rica (UNA) in the Institute for Conservation and Wildlife Management (ICOMVIS), invited us to share our data for use in her thesis on "Factors associated with the occurrence and distribution patterns of medium and large mammals of Costa Rica." Her thesis is supervised by Dr. Manuel Spinola and will be submitted for publication soon. The main objective is to use occupancy modelling to analyze the distribution of mammal species throughout Costa Rica, to determine habitat characteristics, and to reveal any conservation gaps.

Adolfo Artavia invited us to contribute to the MAPCOBIO database of camera traps operating in Costa Rica with community participation. MAPCOBIO is a project funded by JICA, a Japanese aid organization, and supported by SINAC, the national system of protected areas. The goal of the project is to promote participatory management of wildlife conservation, what we call community-based conservation and monitoring. The MAPCOBIO report will soon be published. Daniela Araya and Roberto Salom of Panthera invited us to contribute our jaguar sightings from 2010 to the present to be included in a nationwide album of jaguar detections. Although we do not see jaguar on a regular basis in our surveys, we contributed 8 sightings from the Savegre Valley, Tapantí National Park, El Copal Reserve, Chirripó National Park, and La Marta Wildlife Refuge. Chirripó contributed the largest number of jaguar sightings (3 from the San Jeronimo cameras), La Marta contributed the best quality photos of jaguar, and the Savegre Valley contributed the most unique jaguar, a melanistic one with cut tail.

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Amazing Photos from the Year

During the past year our cameras have picked up some special sightings of mammals. On June 20, one of our cameras at Las Vueltas Biological Reserve caught a tapir mother and baby (Fig. 32). The newborn tapirs have a striped fur pattern that acts as camouflage in the forest light. On April 25, one of the new cameras that we deployed last year in a remote part of Tapantí National Park caught a rare sight of 3 pumas moving together down the path (Fig. 33). We believe this involved a mother puma and her two juvenile offspring. Tamandua (or lesser anteater) have always been a rare sighting on our cloud forest cameras, but with more camera traps located in lower elevations, we are starting to see a lot more of them (Fig. 34). Finally, a selection of very nice photos of predators and prey from our various study areas is presented in Figure 35.

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Salute to our Collaborators

None of our research would be possible without the participation and commitment of a large number of local partners in national parks, private reserves, and biological corridors, as well as collaborators at universities and conservation organizations. A full listing of these partners is given in the acknowledgements, but here are some of the participants in the project that we spent time with in the field this year (Fig. 36). Because our team is only in Costa Rica for two months out of the year, we depend on these dedicated individuals to maintain the camera trap survey and relay the photographic data to us via the internet. We extend out warm appreciation to them all!

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Common Name Latin Name Spanish Name Mass (kg)

Predator Species

Jaguar Panther onca Tigre 80.0

Puma Puma concolor León 50.0

Ocelot Felis pardalis Manigordo 12.0

Coyote Canis latrans Coyote 10.0

Tayra Eira barbara Tolomuco 5.0

Jaguarundi Herpailurus yaguarondi Yaguarundi 5.0

Margay Leopardus wiedii Caucel 3.5

Oncilla Leopardus tigrina Tigrillo 2.0

Prey Species

Baird’s Tapir Tapirus bairdii Danta 250.0

White-tailed deer Odocoileus virginianus Venado 30.0

Red Brocket Deer Mazama temama Cabro de monte 25.0

Collared Peccary Tayassu tajacu Saino 19.0

Paca Agouti paca Tepezcuintle 10.0

White-nosed Coati Nasua narica Pizote 5.0

Northern Tamandua Tamandua mexicana Tamandua 4.5

Northern Raccoon Procyon lotor Mapache 4.0

Nine-banded Armadillo Dasypus novemcitus Armadillo 4.0

Central American Agouti Dasyprocta punctata Guatusa 3.0

Striped Hog-nosed Skunk Conepatus semistriatus Zorro hediondo 2.5

Kinkajou Potos favus Martilla 2.5

Mexican Hairy Porcupine Coendou mexicanus Puercoespin 2.0

Common Opossum Didelphis marsupalis Zorro pelón 1.5

Central American Cacomistle Bassariscus sumichrasti Cacomistle 1.0

Dice’s cottontail Sylvilagus dicei Conejo de monte 1.0

Common Gray Four-Eyed Opossum Philander opossum Zorro de cuatro ojos 0.5

Red-tailed squirrel Sciurus granatensis Ardilla 0.4

Table 3. Large mammal species documented in the Talamanca high-elevation study site, 2010-2015

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Acknowledgments We are grateful for the partnership and assistance of the following persons. Pura vida!

Jordan and Meghan Young and Nancy Rayo of QERC

Jeff Griffitts of Southern Nazarene University, Director of QERC

Don Efrain Chacón and his family of the Savegre Mountain Lodge

Karen Soto Núñez and the staff of Los Quetzales National Park

Meryll Arias, Mahyar Shirazinia, and the staff of Tapantí National Park

Adrian Ugalde and Roberto Delgado of Rio Macho Forest Reserve, Villa Mills

Enzo Vargas and the staff of Chirripó National Park

Marisol Rodriguez of SINAC ACLA-P

Leonel Delgado and the staff of La Congreja National Park

Gerardo Varela and the staff of Carara National Park

Adrián Arce Arias and Julio Bustamonte of SINAC ACOPAC

Gustavo Induni of SINAC Central

Andres Jimenez and the staff of Cabo Blanco National Park

Luis Angel Rojas and Felipe Montoya of Las Nubes Project

Stephanie Buteria of York University, Faculty of Environmental Studies

Manuel Viquez of Refugio La Marta

Beto Chavez of Reserva El Copal

Gustavo Gutierrez of the University of Costa

Lorena Lobos of the National University

Adolfo Artavia of MAPCOBIO

Esteban Brenes of the National University

Stephanny Arroyo-Arce and Roberto Salom of Panthera

Jorge Villalobos of Air Cargo Logistic Group

José Daniel Seco of Pet Lounge

Alex Balmeceda and Tiziana Sambucci of VAMOS Rent a Car