COSTA RICA FOREST RESEARCH PROGRAMME (CRF) Osa … · Corcovado National Park (CNP) where it is estimated 2.5 % of all species living on Earth can be found. Species that are not only

COSTA RICA FOREST RESEARCH

PROGRAMME (CRF)

Osa Peninsula, Costa Rica

CRF Phase 141 Science Report

01 January – 23 March 2014

Nathan J. Roberts, Delyth Williams, Anna Morris, Jessica Dangerfield,

John Scott, Hal Starkie, Sophie Burns, Kirsty Hunter, Alex Caldwell

CRF141 Roberts N. J., Williams D., Morris A.,

Dangerfield J., Scott J., Starkie H., Burns S.,

Hunter K. and Caldwell A.

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Staff Members

Nathan Roberts (NJR) Principal Investigator (PI)

Chris Martin (CM) Project Coordinator (PC)

Anna Douglas-Morris (AM) Assistant Research Officer (ARO)

Delyth Williams (DW) Assistant Research Officer (ARO)

Jessica Dangerfield (JD) Assistant Research Officer (ARO)

John Scott (JS) Assistant Research Officer (ARO)

Hal Starkie (HS) Conservation Apprentice (CA)

Sophie Burns (SB) Conservation Apprentice (CA)

Jenny Collins (JC) Field Communications Officer (FCO)




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Table of Contents

1. Introduction ..................................................................................... 5

1.1 Natural History of Costa Rica and Wildlife Conservation ............................................ 5

1.2 Osa Peninsula ................................................................................................................. 5

1.3 Aims and Objectives of Frontier CRF ........................................................................... 6

2. Training ............................................................................................ 7

2.1 Briefing Sessions ........................................................................................................... 7

2.2 Science Lectures ............................................................................................................ 7

2.3 Field Training ................................................................................................................ 8

2.4 BTECs............................................................................................................................ 9

3. Research Work Programme ........................................................ 10

3.1 Survey Areas ................................................................................................................ 10

3.2 Local Population Density of The Four Primate Species Coexisting in Costa Rica ..... 11

3.2.1 Introduction .......................................................................................................... 11

3.2.2 Methodology ........................................................................................................ 11

3.2.3 Results .................................................................................................................. 13

3.2.4 Discussion ............................................................................................................ 14

3.3 Latrine Site Selection in the Neotropical Otter and Characterising the Occurence of

Mucous in Spraints ...................................................................................................... 14

3.3.1 Introduction .......................................................................................................... 14

3.3.2 Methodology ........................................................................................................ 15

3.3.3 Results .................................................................................................................. 16

3.3.4 Discussion ............................................................................................................ 17

3.4 Using Camera Traps to Estimate Species Abundance of Otters .................................. 19

3.4.1 Introduction .......................................................................................................... 19

3.4.2 Methodology ........................................................................................................ 21

3.4.3 Results .................................................................................................................. 21

3.4.4 Discussion ............................................................................................................ 21

3.5 Avian Diversity within Primary, Secondary and Riparian Forest ............................... 22

3.5.1 Introduction .......................................................................................................... 22

3.5.2 Methodology ........................................................................................................ 22

3.5.3 Results .................................................................................................................. 23

3.5.4 Discussion ............................................................................................................ 23

3.6 CRF and the Osa Conservation Sea Turtle Conservation Programme ........................ 23




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3.6.1 Introduction .......................................................................................................... 23

3.6.2 Methodology ........................................................................................................ 24

3.6.3 Results .................................................................................................................. 25

3.6.4 Discussion ............................................................................................................ 25

3.7 Diversity and Vertical Stratification of Butterflies within Nymphalidae ................... 26

3.7.1 Introduction .......................................................................................................... 26

3.7.2 Methodology ........................................................................................................ 27

3.7.3 Results .................................................................................................................. 28

3.7.4 Discussion ............................................................................................................ 29

3.8 The Three-Dimensional Space Use of Mantled Howler Monkeys (Alouatta palliata) 31

3.8.1 Introduction .......................................................................................................... 31

3.8.2 Methodology ........................................................................................................ 32

3.8.3 Results .................................................................................................................. 33

3.8.4 Discussion ............................................................................................................ 34

4. Additional Projects ....................................................................... 35

4.1 Testing the Influence of Hiking Trails on Bird Diversity and Abundance within

Neotropical Primary and Secondary Lowland Forest. ................................................. 35

4.2 Measuring Disturbance Through Novel Techniques: Soundscape Ecology within a

Heterogeneous Landscape on the Osa Peninsula, Costa Rica. .................................... 35

4.3 Systematically Comparing Methods of Studying Butterfly Diversity: Canopy Trapping

Versus Netting. ............................................................................................................ 36

5. Proposed Work Programme for Next Phase .............................. 38

6. References ...................................................................................... 39

7. Appendices ..................................................................................... 49

Appendix 1. Capture Frequency of Butterfly Species Within the Two Sample Areas ......... 49




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1. Introduction

1.1 Natural History of Costa Rica and Wildlife Conservation

Despite occupying only around 50,000 km2 of the Earth’s surface, the Central American country

of Costa Rica is home to around 5 % of all species on Earth (Sanchez-Azofeifa et al., 2002).

The extraordinary biodiversity observed here has in part been attributed to the climatic

conditions present in this area. Temperature fluctuates little throughout the year, with an

average annual temperature of 27 °C. Rainfall however, is much more variable with distinct wet

and dry seasons that are particularly prominent in the southern Pacific lowlands which receive

an average annual rainfall of 7,300 mm (Baker, 2012). The dry season begins around

November/December and continues through to April/May after which the rainy season

commences.

Costa Rica has become a world leader in conservation and ecotourism (Fagan et al., 2013).

With over 190 reserves and national parks, 11 % of the country’s total area is occupied with

these protected natural areas, all created since the 1960s. Prior to this, human activities such as

logging and hunting seriously threatened biodiversity in this region with over half of the

country’s forests having been cut pre-1960s (Henderson, 2002). Habitat fragmentation and loss

due to expanding human populations which encroach into natural areas also threaten wildlife

populations by reducing the size of available habitat and bringing them into closer contact with

humans. Though some of these issues have been controlled through the implementation of

several reforestation programmes and legislation (Sánchez-Azofeifa et al., 2001; Henderson,

2002) others continue to threaten Costa Rican ecosystems today.

Costa Rican law currently protects 166 species from hunting, capture and sale; but illegal

hunting still occurs, including in protected areas (Baker, 2012). Poaching of turtles for their

fatty calipee and the collection of turtle eggs has severely depleted populations of endangered

Black turtles (Chelonia mydas) and vulnerable Olive ridley turtles (Lepidochelys olivacea) that

use Costa Rica’s coastlines as nesting sites. Similarly, the hunting of Costa Rica’s wild cat

species, peccaries and tapirs for their meat, skins and other animal parts has significantly

reduced wild populations and ongoing human-cat conflict may present an additional threat to

Costa Rican felid species. Furthermore the projected impacts of climate change in combination

with these factors are likely to have significant adverse effects on Costa Rican biodiversity.

1.2 Osa Peninsula

The Osa Peninsula is located within the Southern Pacific lowlands, one of three endemic zones

in the country (Henderson, 2002). On the peninsula lies the world-renowned wildlife refuge,

Corcovado National Park (CNP) where it is estimated 2.5 % of all species living on Earth can

be found. Species that are not only endemic to the country of Costa Rica but also species and

subspecies endemic to the region such as the Cherrie’s Tanager (Ramphocelus costaricensis)

and the Red-backed squirrel monkey (Saimiri oerstedii) are found on the Osa Peninsula. These

endemic species are of conservation significance because local extirpation within their limited

range would result also in their global extinction.




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The Osa Peninsula has a population of around 12,000 people, mainly in small scattered

settlements. The major sources of income in the region are agriculture (rice, bananas, beans and

corn), livestock (cattle), gold mining, logging and more recently the expanding eco-tourism

industry (Carrillo et al., 2000).

Frontier’s Costa Rica Forest Research Programme (CRF) began in July 2009 in collaboration

with the local non-governmental organisation Osa Conservation at the Piro site (N 08°23.826,

W 083°20.564) in the south-east of the Osa Peninsula. There are six core study areas within

CRF in terms of focal species and groups: primates, birds, butterflies, Neotropical otters, sea

turtles and wild cats. The Neoptropical otter is one of the Frontier Costa Rica Forest Research

Programme’s key study species owing to its status by the IUCN as a Data Deficient mammal.

The shorelines adjacent to the property are used by Black turtles and Olive ridley turtles as

nesting beaches throughout the year. Beach patrols are conducted in collaboration with Osa

Conservation as a part of a long-term monitoring and protection programme. Additionally all

four Costa Rican primate species coexist on the Osa property study area, as do numerous bird,

butterfly and amphibian species.

1.3 Aims and Objectives of Frontier CRF

Under the umbrella of the research programme, the specific aims and objectives are:

i) To calculate the density and abundance of the four primate species within the Osa

Conservation property at Piro and its boundaries.

ii) To record the Neotropical otter’s local distribution, assess the environmental

characteristics around latrine sites and investigate the role of scat deposition in

intra-specific communication.

iii) To systematically test the reliability of using camera trap images to estimate the

abundance of Neotropical otters by individual identification and capture-recapture

analysis.

iv) To investigate the occurrence and scale of human-felid conflict within the Osa

Conservation Area (ACOSA) and local attitudes towards conservation.

v) To measure the avian diversity within primary and secondary forest and riparian

habitats within the Osa Conservation property.

vi) To monitor the activity and health of nesting marine turtles on Piro and Pejeperro

beaches and to support the management of the hatchery under the Osa Conservation

Sea Turtle Conservation Programme.

vii) To assess the diversity of butterflies of the Nymphalidae family within an area of

the Osa Conservation property and investigate inter-species vertical stratification.

viii) To investigate the behaviour of mantled howler monkeys and their three-

dimensional space use within the Osa Conservation property at Piro and its

boundaries.




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2. Training

Volunteer Research Assistants (RAs) and newly appointed staff receive a number of briefing

sessions on arrival (Table 1), followed by regular science lectures (Table 2) and field training

(Table 3) throughout their deployments and contracts. The Costa Rica Forest Research

Programme also supported candidates completing the BTEC Advanced Certificates and

Advanced Diplomas in Tropical Habitat Conservation (Table 4).

2.1 Briefing Sessions

All arrivals to CRF are introduced to the aims of the research programme, the methodologies

followed and the conservation significance of the individual studies and are also provided with

an update as to the achievements of CRF. This information is delivered via the Introduction to

the Frontier Costa Rica Forest Research Programme presentation. Additionally all volunteers

and staff are given a full health and safety and medical briefing and are tested before

participating in field activities, those volunteers undertaking a BTEC and/or CoPE qualification

are given an introductory brief before they begin the assessments.

Table 1. Briefing sessions conducted during Phase CRF141

Briefing Session Presenter

Introduction to the Frontier Costa Rica Forest Research Programme NJR

Health and Safety Brief and Test CM

Medical Brief and Test CM

Introduction to the BTEC and CoPE Qualifications NJR

Introduction to Media and Communications JC

2.2 Science Lectures

A broad programme of science lectures is offered at CRF providing information and training in

various aspects of research. Lectures are typically presented utilizing PowerPoint giving an

opportunity for greater understanding of the ecology and identification of focal species, methods

of data analysis used by CRF and the considerations when planning research projects.

Lectures are scheduled with the following objectives:

Allow every volunteer and staff person to attend each presentation at least once during

deployment, regardless of length stay.

Meet the time requirements for BTEC assessments,

Avoid conflict with other activities, maximizing attendance.

Workshops provide detailed training on specific software and applications utilized in

conservation purposes.

Attendance of lectures is non-compulsory.




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Table 2. Science lectures delivered during Phase CRF141

Science Lecture Presenter Date(s) Delivered

Mariposas de Costa Rica JD 10 January 2014;

08 February 2014;

12 February 2014;

21 February 2014;

13 March 2014

Data Analysis: Studying Biodiversity with

EstimateS

NJR 20 January 2014;

29 January 2014

Common Bird Species of the Osa Peninsula AM 22 January 2014

Data Analysis: Studying Biodiversity with

DIVERSITY

NJR 29 January 2014

Data Analysis: Primate Density and

DISTANCE

DW 05 February 2014;

28 February 2014

Survey Planning: BTEC E07 Preparation NJR 07 February 2014;

22 March 2014

Introduction to Primate Behaviour DW 17 February 2014

Surveying and Monitoring: BTEC E09

Preparation

NJR 20 February 2914

Studying Abundance of the Neotropical Otter:

Ex situ Research with CRF

NJR 21 February 2014;

06 March 2014;

21 March 2014

Spatial Analyses: The Use of GIS at CRF NJR 24 February 2014;

04 March 2014

Anurans of the Osa Peninsula JS 25 February 2014

Soundscape Ecology HS 08 March 2014

Bird Identification JS 10 March 2014;

19 March 2014

2.3 Field Training

Volunteers and newly appointed staff receive field training supporting research objectives.

Training is hands-on and provides an opportunity for participants to become familiar with and

use the field equipment.

Specific training sessions to prepare volunteers and staff for accurate data collection

Identification of flora and fauna

Survey method training

Skills and tools used for research

o Turtle Tagging Training

o GPS functionality

o Camera trap

o Trapping (butterfly nets and traps)

o Proper maintenance of equipment




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Table 3. Field training provided during Phase CRF141

Field Training Provider

Reptile and Amphibian Identification JD, HS, JC, AM

Camera Trap Workshop NJR, DW

Tree Identification CM

Turtle Tagging Training NJR

Turtle Patrol Data Collection NJR, AM, DW, JD, SB

Primate Surveys by Distance-sampling NJR, AM, DW, JD, JS, SB, HS

Bird Surveys by Point Surveys AM, JS

Butterfly Canopy Trapping NJR, JD, HS, SB

Butterfly Netting JD, HS, SB

Field Equipment: GPS, Laser Rangefinder and Clinometer NJR, AM, DW, JD, JS, HS, SB

Identification of Neotropical Otters and Their Signs, and

Recording River Attributes

NJR, DW

Studying Primate Behaviour DW

2.4 BTECs

Frontier offers volunteer Research Assistants an opportunity to gain internationally recognised

qualifications based around teamwork, survey techniques, environmental conservation and

effective communication of results. The BTEC in Tropical Habitat Conservation may be studied

as an Advanced Certificate (four week program) or Advanced Diploma (ten week program).

Table 4. BTECs Undertaken in Tropical Habitat Conservation During Phase CRF141

Student Name Project Title (and qualification) Mentor

Jordan Farrant Effect of Canopy Height and Canopy Cover on Scat

Locations of the Neotropical Otter (Advanced Certificate)

DW

Alex Gordon The Relationship Between Turtle Track Width and

Nesting Success (Advanced Certificate)

NJR

Jamie Thomas Inca Project: Understanding Public Attitudes Regarding

Wild Cats of the Osa Peninsula and Finding Solutions to

Human-Felid Conflict (Advanced Diploma)

NJR

Laurence Beaumont Understanding the Important Characteristics of Scat

Locations of Neotropical Otter That Have Mucous

(Advanced Certificate)

DW

Danielle Cerrone Measuring the Effects of Disturbance in Primary and

Secondary Forest on the Abundance of Selected Bird

Species (Advanced Diploma)

JS

Jennifer Sumners Do Sea Turtles (Family: Cheloniidae) Come Ashore To

Nest At A Particular Moon Phase? (Advanced Diploma)

NJR

Nathan Norris [Title TBC] (Advanced Diploma) DW

Isabella Eddington Movement Paths and Space Use of Mantled Howler

Monkeys (Advanced Certificate)

DW

Laura Sheffield [Title TBC] (Advanced Diploma) [TBC]




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3. Research Work Programme

3.1 Survey Areas

Field research is conducted within an 11.83 km2 privately-owned property in Piro, Osa

Peninsula, Costa Rica and its boundaries (8°23’-8°26’N, 83°18’-83°25’W) (Figure 1). Largely

within the Golfo Dulce Forest Reserve and Osa Conservation Area (ACOSA) Wildlife Refuge,

this property is owned by NGO Osa Conservation (OC) and is a heterogeneous landscape

composed of lowland moist primary, secondary and coastal forest. Dominant tree species

include; Ficus insipida, Ceiba pentandra, Attalea butracea, Carapa guianensis, Castilla tunu,

Spondias mombin, Hyeronima alchorneoides, Chimarrhis latifolia, Fruita dorada, Caryocar

costaricense, Ocotea insularis, Pouteria torta, and Inga allenii. The property also borders dense

forest and pastoral properties to the southeast, encompassing pochote (Bombacopsis quinata)

and teak (Tectona grandis) plantations in the north and north east, each occupying 0.4 km2.

There are two biological research stations within the study area: Cerro Osa station in the north-

west and Piro station in the southern region of the property. Mean annual rainfall and

temperature for the area is 5,000-6,000 mm and 26-28 °C; the dry season extends from the end

of December until March.

Figure 1. The Frontier Costa Rica Forest Research Programme (CRF) study area with

surveyed beaches, rivers and trails marked. LS: Laguna Silvestre; NT: New Trail;

Rd: Road; TP: Terciopelo; PT: Piro; BT: Beach Trail; LH: Los Higuerones; AT:

Ajo; CA: Chiricano Alegre; OT: Ocelote; CO: Cerro Osa; NB: Northern Border; CT:

Chicle.




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Within the property, there is a forest trail network covering most of the southern and north-

eastern portions. Besides Cerro Osa which is an old logging road and the north-south segment

of Northern Border and Laguna Silvestre, which are both 5 m-wide cleared tracks, the trails are

narrow and machete-cut. New Trail is the most recently cut trail and is approximately one year

old, while all other trails are more than five years old.

The property is bordered to the south by the Pacific Ocean and the immediate coastline is

separated into two beaches – Playa Piro (2 km) and Playa Pejeperro (4.5 km). The primary

rivers intersecting the property are the Rio Piro and the adjoining Quebrada Coyunda; the mouth

of the former is at approximately the centre of Playa Piro.

3.2 Local Population Density of The Four Primate Species Coexisting in Costa

Rica

3.2.1 Introduction

There are four species of primate found in Costa Rica: the Central American squirrel monkey

(Saimiri oerstedii), Mantled howler monkey (Alouatta palliata), Geoffroy’s spider monkey

(Ateles geoffroyi) and White-faced capuchin (Cebus capuchinus). All four species are present

within the study site in Piro. A. geoffroyi is listed as endangered and S. oerstedii as vulnerable

(IUCN, 2013). Population declines exceeding 50% over the past 45 years have been reported for

A. geoffroyi, principally driven by stresses induced from habitat loss (Cuarón et al., 2008a). S.

oerstedii has a limited distribution, restricted to Panama and Costa Rica meaning populations

exploited for the pet trade and experiencing habitat loss and degradation are already vulnerable

to stochastic events. Land conversion for agriculture and development, clear cutting, selective

logging and the pet trade have implications for all four Costa Rican primate species (Cropp and

Boinski, 2000).

The primate guild in Costa Rica is vital for forest structure integrity. All four species perform

important roles as seed dispersers (Julliot, 1997; Garber et al., 2006) and the Osa Peninsula is

the only part of Costa Rica where all four species occur in concert (Carrillo, 2000).

Frontier CRF has been surveying the presence and absence of all four primate species in

collaboration with Osa Conservation since March 2010. The overall aim of this research is to

provide an insight into the habitat use by primates in the area to better aid management and

policy decisions at a local level. The principal and foremost objective of this project is to

produce estimates of the population density of each of the four species.

3.2.2 Methodology

Data is collected following a standardised line-transect sampling protocol (Buckland et al.,

2010; Thomas et al., 2010). The trails are sampled equally across times of day. Surveys are

taken at 0600, 0800, and 1400 hrs to cover peak primate activity, increasing the detection

probability. The transects for the most part, combinations of forest trails. Transects sampled

(Table 5) are walked by two observers at a constant speed of 1.5 – 2 kmhr-1

as recommended by

Karanth and Nichols (2002). No transect is surveyed more than once on the same day and




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sampling is only conducted in fair weather due to the reduction in detection probability in

adverse weather conditions.

Table 5. Transects Sampled and Descriptions

Transect Name (code) Transect

Length (km)

Habitat Description and

Trail Type

Ajo – Ocelote (AT – OT) 3.1 Trail through secondary and mature,

primary forest

Terciopelo – Piro (TP – PT) 2.3 Trail through riparian, primary and

secondary forest

Cerro Osa (CO) 1.9 Old logging road and forest trail through

secondary and mature, primary forest,

abandoned pochote (Bombacopsis

quinata) plantation and across a branch

of the Quebrada Coyunda River

Laguna Silvestre (LS) 2.6 Access road through secondary and

primary forest

Beach Trail – Road (BT – RD) 1.6 Trail through secondary forest and main

access road dissecting secondary forest

Chiricano Alegre – Cerro Osa –

Los Higuerones (CA – CO – LH)

1.9 Trail through secondary and mature,

primary forest

Chicle (CT) 1.8 Forest ridge trail through primary forest

and across Quebrada Coyunda River

Northern Border (NB) 2.5 Old logging road through a pochote

(Bombacopsis quinata) plantation and

trail through mature, primary forest

New Trail-Road (NT – RD) 3.8 Trail through secondary forest and main

access road dissecting secondary forest

Mean ±SD 2.4 ± 0.7

Total 21.6

All visual encounters of primates are recorded, logging the spatial location with GPS, noting the

number of individuals within the observed group and perpendicular distance from the transect

line to the geometric centre of the group at first sighting. All individuals visible at the same time

and exhibiting the same general behaviour (e.g., resting, moving or foraging) were considered to

be of the same group (Chapman et al., 1995). Agreement on group size was made between the

observers where the group could be counted together; otherwise the observers split their search

effort into non-overlapping areas (e.g., left and right of the trail) and made independent counts

which were then summed. Where possible group composition was also recorded as secondary

data. Vocal detections are not included.

The perpendicular distance to the centre of the group was calculated using the radial distance r

and the sighting angle θ, which are related to the perpendicular distance x by the formula x =

rsinθ. The sighting angle was measured with a compass and the radial distance calculated using

the formula r = dcosθ, where r is radial distance, d is the direct distance measured with a laser

rangefinder between the observer and the geometric centre of the group in the tree, and θ is the

slope angle between the observer and the geometric centre of the group in the tree measured




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with a clinometer. Alternatively, where field conditions permitted, a tape measure was used to

measure the perpendicular distance. After every recorded observation, the observers

immediately continued along the transect noting any direction of animal movement to eliminate

the chance of recording the same individuals more than once per survey.

Data were analysed using DISTANCE ver. 6.0 (Thomas et al., 2010) under the following

assumptions: i) all groups at zero perpendicular distance to the line were detected with certainty;

ii) groups were detected at their initial locations and were not counted twice, and; iii) accurate

measurements of distances were made from the line to the geometric centre of the group

(Thomas et al., 2010). As trails were concentrated around the southern edge of the property, our

survey effort was not evenly distributed within the study area. The study site was therefore

divided into two substrata to avoid bias: an area of 1.67 km2 in the southern region of the

property (encompassing trails Terciopelo-Piro, Beach Trail-Road, Ajo-Ocelote, Chiricano

Alegre-Cerro Osa-Los Higuerones and the lower 414 m of Cerro Osa) and an area of 10.16 km2

in the northern and western sections of the property (encompassing trails Chicle, Northern

Border, Laguna Silvestre, New Trail-Road and the remainder 1052 meters of Cerro Osa). These

two sections were considered as two substrata layers in DISTANCE (Buckland et al., 2010).

The conventional distance sampling (CDS) function was employed to permit stratification of the

two sections of the property. The data was truncated by 10% (Buckland et al., 2001). Cluster

size regression analysis was selected for each model except in circumstances where warnings

arose. In these cases, the mean of observed cluster sizes was chosen to estimate cluster size

(Link et al. 2010). As there were several replicates of each trail, the number of samples was

added as the multiplier to the CDS model to account for the survey effort (Leca et al., 2013).

Several candidate models were tested and the most parsimonious model was selected based on

AIC values. The suitability of each model was also determined using chi-square (χ2) goodness

of fit (GOF) tests (cf. Leca et al., 2013): significant results of χ2

analysis suggest that the model

does not fit the data (Buckland et al., 1993).

The data were stratified at the sample level and density estimates were pooled using the mean

density estimates of the two substrata, weighted by the strata area to compensate for the large

difference in substrata area. Detection function and cluster size were estimated at the global

level.

The research was non-invasive and adhered to the legal requirements of Costa Rica. Any and all

aggressive behaviour towards the observers by individual primates was responded to by moving

on as quickly as possible.

3.2.3 Results

The density of Black-handed spider monkeys on the property was calculated from 15 replicates

of each trail and 107 spider monkey encounters over a period of four months. After truncating

the data by 10%, 96 recordings were used in the analysis. The most parsimonious model was the

uniform key with a cosine polynomial expansion which estimated there to be 27 individuals on

the 11.83 km2 property and its boundaries; 2.31 individuals/km

2. At the strata level, a density of

2.34 and 2.12 individuals/km2 were estimated for the northern and southern region, respectively.




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New Trail-Road had the highest number of encounters with 29 spider monkey records, whereas

Northern Border had the lowest with only one spider monkey encounter being recorded.

3.2.4 Discussion

The recommended number of observations per species (60-80) has now been exceeded for the

howler monkey; density will be calculated and reported in Phase CRF142. However,

insufficient encounters have been made with White-faced capuchins and Central American

squirrel monkey to produce robust density estimates of these species. Baseline data of these

primate species is vital throughout global biodiversity hotspots (Mittermeier, 1987; Sechrest et

al., 2002) owing to their ecological role as seed dispersal agents (Mittermeier, 1987; Stevenson,

2001; Estrada et al., 2005) and CRF will continue with the objective of providing this for all

species witin the primate guild.

The estimated density of spider monkeys is drastically lower than anticipated and is notably

smaller than the number of spider monkeys estimated in CNP; 68.45 individuals/km2

(Weghorst, 2007). Even when considering the range of population estimates across

Mesoamerica (5.4 to 89.5 individuals/km2; (Piñeros, 1994; Ramos-Fernández and Ayala-

Orozco, 2002), our estimation falls below this range. There is an abundance of fruiting tree

species on the property that are known to provide resources to spider monkeys and promote

their occupancy; Spondias mombin, Ceiba pendandra, Inga spp., Ocotea spp. and Pouteria

campechiana (Coelho et al., 1976; Estrada et al., 2004; Méndez-Carvajal, 2013; Weghorst,

2007; White, 1986). Spider monkeys are noted to be very sensitive to disturbance, usually being

one of the first Neotropical primates to become extinct following fragmentation (Link et al.,

2010; Ramos-Fernández and Wallace 2008).The property has been subjected to extensive

anthropogenic modification and despite the surrounding landscape being part of a wildlife

corridor, it has been altered by agricultural practices potentially reducing the efficacy of the

corridor by limiting species’ natural dispersal.

Spatial data collected from each primate encounter may be used in future studies within CRF to

relate the distribution of primates with the level of disturbance, as quantified by novel

techniques of soundscaping. Primates are differentially sensitive to disturbance and the density

estimates of the remaining three species will potentially, provide more of an indication of the

impact anthropogenic activity within the property is having on the primate community.

3.3 Latrine Site Selection in the Neotropical Otter and Characterising the

Occurence of Mucous in Spraints

3.3.1 Introduction

The neotropical river otter (Lontra longicaudis) has a widespread distribution inhabiting rivers,

lagoons and streams from Mexico through to Uruguay (Emmons and Feer, 1997). Despite this

wide distribution, little is known about the species ecology, local distribution and population

status; thus hindering the development of appropriate conservation strategies. The IUCN defines

the priorities for this species as field surveying of populations and identifying key habitats in

order to establish proper conservation status of what is presently listed on the Red List as Data

Deficient (Waldemarin and Alvarez, 2008). Urgently updating the status of this species is




15

critical as its low reproductive potential and dependence on healthy aquatic environments mean

that the species cannot respond quickly to population declines, demonstrated by the local

extinction experienced between 1950 and 1970 in response to extensive hunting in parts of the

species’ former range (Cezare et al., 2002; Waldemarin and Alvarez, 2008).

Spatial use by otters can be cost-effectively determined by recording the location of their

spraints and other signs within and around aquatic environments (Davison et al., 2002). Where

spraints, urine and anal secretions are repeatedly deposited by an otter, a latrine is established

(Bowyer et al., 1995). The exact purpose of latrines is not conclusive, but is believed to play a

role in intraspecific communication (Melquist and Hornocker, 1983; Rostain et al., 2004),

signalling reproductive state to the opposite sex (Rostain et al., 2004), marking areas of key

resource use (Kruuk, 1992) and conveying the social status of males (Rostain et al., 2004). It is

further suggested that despite otters being carnivorous and generally solitary, they are not

territorial in the classical sense as it would cost too much time and energy to patrol to the polar

extents of their home range to mark territorial boundaries (Melquist and Hornocker, 1983;

Remonti et al., 2011). Latrines are, therefore located within the home range of otters (Melquist

and Hornocker, 1983) and as these sites are indicative of habitat quality and suitability,

determining which environmental characteristics otters will preferentially mark reveals insights

into their ecology.

While some aspects of the environment are relatively consistently reported to influence latrine

site selection, such as conspicuous substrata (Quadros and Montiero-Filho, 2002; Depue and

Ben-David, 2010), prey availability (Melquist and Hornocker, 1983) and vegetation structure

(Prenda and Granada-Lorencio, 1996); numerous other interacting factors are likely also

promote latrine site selection. There are however discrepancies as to the importance of each

factor. These other factors include water depth (Anoop and Hussain, 2004; Depue and Ben-

David, 2010; Remonti et al., 2011; Carrillo-Rubio and Lafón, 2004) and water edge topography

(Swimley et al., 1998; Bowyer et al., 1995; Anoop and Hussain, 2004).

Furthermore, the study of seasonal variation in detection rates of spraints and other signs has

been advocated as well as monitoring frequency of use of latrine site (Rheingantz et al., 2004;

Rheingantz et al., 2008; Santos and Reiss, 2012). Seasonal changes in sprainting behaviours are

also expected to be synchronous with the breeding season (Stevens and Serfass, 2008) and

temporal peaks may be explained by prey availability (Remonti et al., 2011; Almeida et al.,

2012) and changes in distribution (Gori et al., 2003).

3.3.2 Methodology

Two linear transects were established along the course of the Rio Piro and Quebrada Coyunda,

measuring 5.2 km and 2.8 km respectively. Each river was sampled once each week, typically

as two distinct transects. One began at CRF camp and surveyed 3.5 km north on the Rio Piro

and the other began at CRF camp, surveyed south on the Rio Piro to the river mouth (1.7 km)

and then surveyed 2.8 km of the Quebrada Coyunda east from the intersection with Rio Piro.

Surveyors walked within the river course and typically began sampling early morning.

During the course of the survey all rocks, logs, fallen trees, roots and river margins were

actively searched for otter spraint. Fresh spraint was distinguished from old spraint based on




16

moisture content and how contained the spraint was. Old spraint which was dry and broken

apart would be recorded as such for consideration in analyses as environmental characteristics

may have changed significantly since the time of spraint deposition. The presence/absence of

mucous in spraint was also recorded in addition to the substrate on which the spraint was

deposited.

Each site was determined to be a new site or a reutilised site where new spraints were found in

previously used sites; this was verified ex-situ from GPS data recorded at the time of each

spraint encounter mapped onto GIS. At each site, river width was measured as well as the river

depth at 1 m intervals of the river cross-section, escape cover distance; defined as the distance

from the water’s edge to the point where the undergrowth starts was measured for both river

margins.

3.3.3 Results

Between 1st October 2013 and 21

st March 2014, latrine utilisation by Lontra longicaudis was

recorded on the Rio Piro and Quebrada Coyunda (Figure 2). Spraint presence along with its

relative occurrence of mucous was surveyed over a length of 170.8 km. During the study

period, 497 spraints were recorded. The majority of spraints encountered were deposited on

conspicuous rocks, rock platforms, (n = 152; 39.1%) fallen tree trunks and roots (n = 215;

55.2%); very few were on finer loose substrate on river margins (n = 22; 5.7%). Further, within

months over 80% of spraints were found on previously visited latrines and excluding the

southernmost extent of the Rio Piro, the full study area was utilised for scat deposition during

the sampling period at least once.

On average, 7.6 % of spraints encountered per month contained mucous. There was an observed

peak in the frequency of spraint encountered per kilometre and the relative occurrence of

mucous in January and February, respectively.

Figure 2. Latrine utilisation by Lontra longicaudis in the Rio Piro and Quebrada Coyunda and

relative occurrence of mucous in spraint

The distribution of spraint containing mucous varied within the study period. In October these

spraints occurred exclusively on the Rio Piro (RP), in December the extent expanded into the

0

2

4

6

8

10

12

14

0

0.5

1

1.5

2

2.5

3

3.5

Sp

rain

ts c

onta

inin

g m

uco

us

(%)

Fre

quen

cy o

f sp

rain

t /

km

New latrines

Reutilised latrines

Spraint with mucous




17

Quebrada Coyunda (QC), and within February and into March these spraint were found

primarily at the RP-QC intersection and along the course of the QC (Figure 3). Ninety six per

cent of spraint containing mucous was also found on a reutilised site. Environmental conditions

at latrine locations with spraint containing mucous were comparable to those without mucous in

spraint (Table 6).

Figure 3. Spatial and temporal distribution of scat containing mucous in relation to all other

spraint where mucous was absent

Table 6. Environmental characteristics of latrine locations at the time of spraint recording

Spraint Type Average

River Width

(m ±SD)

Average

River Depth

(cm ±SD)

Average

Escape Distance

(m ±SD)

Spraint with mucous 7.23 ± 3.95 18.95 ± 8.87 3.01 ± 1.9

Spraint without mucous 7.39 ± 3.9 20.76 ± 14.95 3.09 ± 4.2

3.3.4 Discussion

The preference for spraint deposition on conspicuous substrata observed here concurs with

previous studies (Kasper et al., 2008), although here, use of these features is more exclusive. It




18

would suggest an abundance of these features as it has been reported that if these raised

platforms are not available, L. longicaudis will scratch at the ground surface to create a small

mound which they will then defecate on (Quadros and Montiero-Filho, 2002). These behaviours

enhance visual and olfactory cues for communication (Quadros and Montiero-Filho, 2002;

Depue and Ben-David, 2010). The even distribution of logs and rocks within the study area

would confirm a preference for these features; further investigation could be performed with

respect to the positioning of these substrata within the river course; either at the margin or

within the river course.

In the first month reported here, many of the sites were recorded as reutilised. This is due to the

study being an expansion of previous work which recorded latrine locations but did not

encompass the full extent of the present study. Reutilised sites are therefore, considered since

the former study began in July 2013. The present study is therefore more representative of the

study area than in previous phases and eliminates an inevitable spike in frequency of new sites

reported at the beginning of sampling, evident in other studies (e.g., Kasper et al., 2008).

Here all records were considered independently rather than explicitly creating spraint encounter

histories for unique latrines, reporting the frequency of spraints deposited at reutilised sites

rather than specifically how many sites were reutilised in any given period. Although it may be

possible to perform analysis of the spatial data to identify latrines for which the encounter

histories can be produced, defining latrines by spatial isolation is only one method of many

which have been used (e.g., Kruuk et al., 1986; Swimley et al., 1998; Quadros and Monteiro-

Filho, 2002; Depue and Ben-David, 2010; Almeida et al., 2012; Crowley et al., 2012). It

remains contested as to what defines a latrine, hindering the comparison of data between

location and otter species.

A methodological consideration when collecting data on otter sprainting behaviour is seasonal

variation in intensity of use. The seasonal changes observed for L. longicaudis in Argentina that

were linked with changes in distribution (Gori et al., 2003). A peak in latrine use in January

followed by a decline in February and March observed here concurs with findings in Brazil

(Kasper et al., 2008). In L. canadensis; peaks in October, November and February and March

are believed to correspond to the breeding season (Stevens and Serfass, 2008) and when the

pups begin to move around in family groups (Prigioni et al., 2005). Further, temporal variation

in L. lutra sprainting activity has been linked to prey availability (Remonti et al., 2011; Almeida

et al., 2012).

Owing to the dietary preference of otters (Gori et al., 2003) and expected sprainting behaviours

around resource areas (Melquist and Hornocker, 1983), it is likely that sites with environmental

characteristics that increase prey availability will promote latrine site selection. Deep pools

create a suitable lentic environment for the preferred slow-swimming fish, thus providing otters

with plentiful prey (Carrillo-Rubio and Lafón, 2004; LeBlanc et al., 2007). It has also been

reported that deeper water may provide an effective escape route from predators (Depue and

Ben-David, 2010). Conversely, for smooth-coated otters (L. perspicillata) a preference for

shallower water has been documented, though it may be explained by close proximity to larger

water bodies where prey abundance was greater (Anoop and Hussain, 2004). Here it is

advocated that utilised sites are directly compared against non-utilised sites to better determine




19

sprainting preferences and integration of prey availability into the present study would provide

further understanding of the contributing factors in otter space use.

Groundtruthing may be required to verify records of 0 m escape cover distance, where it was

recorded irrespective of bank slope. It is assumed that a vertical high bank will not provide

otters with an escape route and therefore these records collected here should be excluded from

future analyses, using only values of 0 m where the vegetation is at the water edge and the bank

slope and height are considered practical for escape. Whilst other studies have included water-

edge topography in their analyses of latrine site selection (Remonti et al., 2011; Crowley et al.,

2012), it has not consistently appeared to be a key determining factor in latrine site selection.

Despite local sites with spraint containing mucous being comparable with those with mucous

absent, the process of groundtruthing and reanalysing data may clarify the significance of river

margin topography in sprainting behaviour.

Considering otters are found on almost every continent, in a range of habitats; there is a distinct

paucity of data available across and within species. There may be severe limitations in the

ability to compare results between studies, but care needs to be taken when comparing sites of

latrine activity and extrapolating results, as scent marking behaviours can vary depending on

population density, habitat features, human disturbance, season and there may be interspecific

behavioural differences in sprainting behaviours (Quadros & Monteiro-Filho, 2002; Guter et al.,

2008). Therefore much more intense study on the habitat preference of specifically L.

longicaudis is required.

This study provides valuable insight and progression in understanding the habits of wild otters,

expanding on the small foundation of knowledge which exists on the ecology and specifically

latrine site selection by L. longicaudis. Provided there is a large sampling effort, it has been

advocated that relating latrine site selection and environmental characteristics is a favoured

method of accurately investigating otter ecology (Guter et al., 2008) and has to date not been

studied in this part of the species’ range. The study here has potentially expanded our

knowledge of reproductive behaviours by quantifying the occurrence and distribution of spraint

containing mucous (R. MacKenzie, personal communication). Finer scale analyses are

recommended to consider whether the apparent shift in distribution of latrines with mucous

present can be explained by changes in environmental conditions. The role of mucous in otter

ecology and the value it can present in research and conservation should continue to studied.

3.4 Using Camera Traps to Estimate Species Abundance of Otters

3.4.1 Introduction

Basic information on population abundance is vital in conservation research in order to assess a

species’ current status, monitor temporal trends and define conservation priorities (Mackenzie,

2005; Oliveira-Santos et al,. 2010). Without robust data on distribution and abundance,

conservation strategies and management practices may be misguided, ineffective and inefficient

(Langbein et al,. 1999; O’Brien, 2008; Hájková et al., 2009; Obbard et al., 2010).




20

The Neotropical river otter (Lontra longicaudis) is a Data Deficient species, (Waldemarin and

Alvarez, 2008) and at present no standardised methods to estimate the abundance of the species

exist. However, it is thought that the species is rare and declining throughout its range from

north-western Mexico to Peru and Uruguay, therefore field surveys of populations are one of the

IUCN conservation priorities for the species (Waldermarin and Alvarez, 2008). Furthermore,

developing standardised methods is essential for reliably comparing data at multiple spatial and

temporal scales.

The difficulties associated with assessing the abundance of otters (Family: Mustelidae) by direct

observation are associated with low encounter frequencies, thus obligating the implementation

of alternative approaches (Hájková et al., 2009), such as recording distinct otter spraint

(Davison et al., 2002). However, estimating abundance and therefore assessing population

status of elusive and rare species is often difficult (Balme et al., 2009; Hájková et al., 2009),

and using sprainting intensity as an index of otter abundance is unreliable as it may reflect the

abundance of suitable, conspicuous sprainting sites and activity rather than true population size

(Guter et al., 2008; Calzada et al., 2010).

Camera-trapping is a relatively new methodological advancement (Pettorelli et al., 2009) that

uses specialised equipment to detect and take photographs of passing animals (Rowcliffe et al.,

2008). Since its early applications in the 1920s, camera traps have become an increasingly

popular tool in non-invasive wildlife research (Rowcliffe and Carbone, 2008). The most

common application of camera-trapping is for estimation of abundance for wild cat species

(Rowcliffe and Carbone, 2008), which is most often calculated by simultaneously

photographing the left and right flank of animals with paired camera traps. This results in

capture histories of individuals identified by their unique stripe or spot patterns and an

estimation of population parameters by capture-recapture analysis (Karanth, 1995). However,

camera trap studies of non-striped and non-spotted species are less represented and are generally

limited to species inventories, rather than estimating abundance or density (Rowcliffe et al.,

2008). Applying camera-trapping methodology to these species will indefinitely increase the

value of this tool, no longer limited to only those species with unique identifying markers

(Rowcliffe and Carbone, 2008).

Trolle et al. (2008) argue that species lacking conspicuous stripes and spots may be identified to

individual level by subtle marks, coat colouration, scars, body structure and sex, this approach

has been applied in a number of studies concerning various taxa (e.g., coyotes (Canis latrans)

(Larrucea et al., 2007); foxes (Vulpes vulpes) (Sarmento et al., 2009); Maned wolf (Chrysocyon

brachyurus) (Trolle et al., 2007); tapirs (Tapirus spp.) (Holden et al., 2003; Noss et al., 2003;

Trolle et al., 2008); and pumas (Puma concolor) (Kelly et al., 2008). However, individuals may

be misidentified if shallow scratches fade (Trolle et al., 2008), photographs may be captured in

different light and humidity conditions, as well as varying approach path angles and distances

relative to the camera (Oliveira-Santos et al., 2010). Investigation into the reliability of

individual identification of non-spotted and non-striped species has begun using tapirs as a case

study, reporting very poor levels of accuracy (Oliveira-Santos et al., 2010).

It is believed that otters may be individually identified using markings on the chin (Chanin,

2003); however the reliability of this method has not been systematically investigated. It has




21

been strongly advocated that future studies validate these theories using data from captive

individuals to test whether inter-individual variation at the population level is sufficient to

reliably distinguish between individuals (Foster and Harmsen, 2012). Furthermore, for camera

trap data to be analysed by capture-recapture methods, unique identifying markers must be

visible in every photograph (Foster and Harmsen, 2012). Alternatively, where individuals of a

species cannot be identified, novel methods reveal potential to estimate abundance from

measurements of animal and camera parameters using the images themselves without the need

to identify individuals (Rowcliffe et al., 2008).

The specific objectives of this study are to: i) investigate the ability of researchers to identify

individual Neotropical river otters (Lontra longicaudis) in a set of photographs with known a

population size; ii) investigate which characteristics are being used by researchers to identify

individuals within the population; iii) quantify the proportion of camera trap photographs which

are discarded on the basis of the inability to identify individuals; iv) assess the effect of any

inter-researcher variability in individual identification on reliability of species abundance

estimates by capture-recapture analysis, and; v) provide informed recommendations towards

developing standardised methodologies for estimating the abundance of Neotropical river otters

in the wild.

3.4.2 Methodology

Camera-trapping will be conducted within the captive areas of animal collections holding at

least one individual Neotropical river otter. Cameras will be active 24 hrd-1

. The resulting

images will be grouped into three subsets: (1) right flank of animal; (2) left flank of animal; (3)

animal facing camera. Cameras will be active until more than 50 images within each subset

have been captured and it is agreed with animal collection staff that all individuals were

photographed at least once. Animal collection staff will also be asked to identify the

individual(s) in each photograph.

The photographs will be sent to members of the IUCN Otter Specialist Group and other otter

researchers who will be invited to respond to the following questions: (1) How many

individuals were identified?; (2) What parts or characteristics of the animals were considered

relevant in the identification of individuals?, and; (3) How many photos were discarded as

inappropriate for identification? Respondents will also be asked to identify the individual(s) in

each photograph, giving a unique numerical identifier to each individual.

The number of individuals identified by each participant will be expressed as a simple

calculation of error, proportion of over-estimation or under-estimation in relation to the known

population size. Characteristics used by respondents to identify individuals will be assessed to

investigate any trends in accuracy of estimate and characteristic(s) used. The proportion of

photographs discarded will be expressed as a percentage of the total number of photographs

collected. Capture-recapture methods will be applied to the capture histories created from

participants’ responses containing unique numerical identifiers for individuals in each

photograph and abundance estimates calculated in CAPTURE. These values will be compared

with the estimate derived from the capture history based on responses from animal collection

staff.




22

3.4.3 Results

Camera trap images are still being collected at the time of writing.

3.4.4 Discussion

No results to discuss this Phase.

3.5 Avian Diversity within Primary, Secondary and Riparian Forest

3.5.1 Introduction

Costa Rica has a very rich avifaunal diversity, hosting approximately 850 species, more than the

United States and Canada combined (Henderson, 2010; Sánchez-Azofeifa et al., 2001), 160 of

these bird species are endemic to Costa Rica (Henderson, 2010). The most frequently

historically encountered bird species for the Frontier Costa Rica Forest Research Programme

include: Blue-crowned Manakin (Pipra coronata), Black-hooded Antshrike (Thamnophilus

bridgesi), Chestnut-backed Antbird (Myrmeciza exsul), Black-mandibled Toucan (Ramphastos

ambiguus), Great Tinamou (Tinamus major), Green Kingfisher (Chloroceryle americana),

Long-billed Hermit (Phaethornis longirostris), Scarlet Macaw (Ara macao), Short-billed Pigeon

(Patagioenas nigrirostris) and White Ibis (Eudocimus albus).

The distribution of birds and the species richness of a given area can often be explained by the

habitat type and characteristics of the environment (Wilme and Goodman, 2003). From this

information, it has been possible to classify species as habitat generalists or specialists (based on

the extent of their occurrence in different environments), the former being more resilient to

landscape change through deforestation, fragmentation and anthropogenic activities (Pejchar et

al., 2008). Birds are good indicators of the health of the habitats they occupy due to their

sensitivity to change, therefore recognising habitat specialists and monitoring trends in

avifaunal diversity may help to identify species at greater risk of population decline as a result

of anthropogenically-induced environmental change. Furthermore, declining bird populations

will affect rates of pollination and seed dispersal, two fundamental functions of a healthy

ecosystem, facilitated by birds (Pejchar et al., 2008). Considering this and the high levels of

endemism in Costa Rica, it is important that bird communities in this region are regularly and

closely monitored.

The aim of this study is to identify and compare the bird communities found within primary and

secondary forest as well as within the course of a river and to further understand which species

are habitat generalists and specialists based on presence/absence data at each sampling point.

3.5.2 Methodology

A total of 20 survey sites were established within three different habitat types at the Piro site,

comprised of primary and secondary forest as well as a river course. Stations were set along Rio

Piro, New trail, Ocelot trail, Piro trail, Terciopelo trail, Ajo trail and Cerro Osa trail at 200 m

intervals. Each station was visited a minimum of three times, with a total of 75 points completed

between the 3rd

and 22nd

of February 2014. In order to reduce the ‘time of day’ effect, the order

in which stations were visited was reversed for each survey. Surveys were performed in the

morning between 0600 and 0800 hrs. Each point count lasted 10 minutes, with a one minute

settling period occurring before the commencement of each survey. Any species seen or heard




23

within a 50 m radius of the station point was recorded during each point count. Each survey was

led by ARO Anna Morris with assistance from ARO John Scott. Birds were identified to species

level by sight using binoculars or by song recognition. When birds could not be identified to

species level, the bird group was still recorded (e.g. as Woodcreeper spp.) but was not included

in the data analysis. An audio recording was taken during each survey and when possible,

photographs of unidentifiable birds were taken to aid with species identification. The main

references used were Garrigues and Dean (2007) and Jongsma et al., (2005).

3.5.3 Results

Now that the data collection phase has been completed; ARO Anna Morris has begun analysing

audio recordings in order to make alterations and add any species which were previously

unidentified by the survey team. The data will then be analysed to identify patterns in the

occurrence of each species and family with relation to habitat in order to achieve the aim of the

study, which is to gain an understanding of which species are more likely to be habitat

specialists and which are likely to be habitat generalists. The full results and discussion of the

study are expected to be published in Phase CRF142 or CRF143.

3.5.4 Discussion

No results to discuss this Phase.

3.6 CRF and the Osa Conservation Sea Turtle Conservation Programme

3.6.1 Introduction

Sea turtles are both a flagship species for conservation due to their iconic nature and an

excellent indicator species for climate change due to their temperature-dependant sex

determination; where increased temperatures create a sex bias skewed toward females which

could cause entire populations to collapse. Additionally, temperature-induced changes in plant

community composition together with rising sea levels may result in increased incidences of

beach erosion and inundation of nests (Janzen, 1994). Turtle late maturation in conjunction with

anthropogenic threats such as beach development, long line fishing and pollution mean that

turtle populations are highly vulnerable and often unstable (Govan, 1998). Poaching and the

illegal trade of turtle eggs cause further reductions to turtle populations, which may result in

entire clutches being destroyed. Turtle hunting was most severe from the 1950s until the 1970s,

with half of the world’s turtle catches being made in Mexico where as many as 350,000 turtles

were harvested annually (Marquez et al., 1996).

A number of conservation strategies have been established throughout Costa Rica, including

legalised limited commercial egg harvesting on a nesting beach in Ostional during the first 36

hours of wet season arribadas (Campbell, 1998) and an annual catch of 1,800 Black turtles

being granted to fishermen in Limόn (Troëng and Rankin, 2004). Though the latter may have

increased extractive use along with illegal hunting in the mid-1990s, the ban on black turtle

fishing and increased law enforcement since 1999 may have largely increased female turtle

survivorship (Troëng and Rankin, 2004). In other regions such as Tortuguero, the Costa Rican

government has made egg poaching illegal, in addition to prohibiting the trade of calipee (the

edible part of the shell) (Government of Costa Rica 1963 and 1969; Troëng and Rankin, 2004).

Meanwhile, the growing ecotourism industry in Costa Rica has provided locals with an




24

alternative source of income and has promoted conservation throughout the country. To

evaluate the effectiveness of such strategies, it is imperative that monitoring programmes are

long-term as it can take decades for species with late maturity to show a population response

(Troëng and Rankin 2004; Bjorndal et al., 1999).

On the Osa Peninsula where turtles are threatened primarily from predation by dogs, coastal

development, illegal trade of turtle eggs and to a lesser extent turtle meat (Drake, 1996), NGO

Osa Conservation has been patrolling Piro and Pejeperro beaches since 2003. Their aim is to

monitor the frequency and health of the local nesting turtle population and manage nest

relocations to the associated hatchery. The Frontier Costa Rica Forest Research Programme

works in partnership with the Osa Conservation Sea Turtle Conservation Programme

concerning the four species which nest here: Olive ridley (Lepidochelys olivacea), Pacific black

turtle (Chelonia mydas agassizii), leatherback (Derochelys coriacea) and hawksbill

(Eretmochelys imbricate). Of these, the latter two nest only rarely on these beaches. The Olive

ridley is most commonly found nesting here and is listed by the IUCN as Vulnerable, the black

turtle as globally Endangered (Troëng and Rankin 2004; Honarvar et al., 2008; IUCN, 2013).

3.6.2 Methodology

Phase 141 is in the peak black turtle season; night and morning patrols are conducted on both

Playa Piro and Playa Pejeperro managed between Frontier and Osa Conservation. Morning

patrols commenced at 0400 hrs on Playa Piro and 0300 hrs on Pejeperro (high tide permitting)

to minimise surveyor exposure to unavoidable direct sunlight and high temperatures. Night

patrols typically started at 1930 hrs on both beaches. To minimise disturbance to nesting

females, surveyors used red lights during night patrols and survey teams were a maximum of six

persons. In this phase CRF led 37 turtle patrols across Piro (n = 19) and Pejeperro beaches (n =

18).

The transect area for both beaches was divided into 100 m sectors; this constituted a two

kilometre stretch of Playa Piro and a 4.4 km stretch of Playa Pejeperro. For every turtle track

encountered details of the previous days date were recorded on morning patrols (i.e., the date of

turtle activity) or the date of the night patrol along with time, name of data recorder, sector

number; the nest type associated with the tracks either as in situ (IS), false crawl (FC; i.e., turtle

returned to sea without nesting), or NA when the nest type could not be determined; track

symmetry defined as either symmetrical (S) or asymmetrical (A), and distance to the vegetation.

The track was then crossed through with a deep heel drag in the sand to avoid the track being

recorded again in subsequent patrols. Track characteristics were used as indicators of species

where the turtle was absent (i.e., asymmetrical tracks suggest olive ridley, symmetrical tracks

suggest black). If a turtle was encountered, the species and distance to the tide was recorded, a

health assessment conducted and tagging of both the individual’s flippers completed.

In-situ nests were confirmed by inserting a stick into the sand to locate the egg chamber

(indicated by a marked change in resistance with applied pressure) followed by careful digging

to confirm the presence of eggs. A false crawl was defined by the absence of a nest or where it

was clear that the turtle returned to sea without digging a nest. In the case of predated nests

(typically evident by the presence of predator tracks, egg shells and signs that the nest had been

dug up), the predator was identified by the tracks and egg shells counted and recorded.




25

All turtles encountered pre- or mid-nesting were tagged whilst they were laying, during which

time the female enters a haze and therefore the process reduced the associated stress. To reduce

the risk of infection during tagging, the tag sites were washed firstly with water, then alcohol

and iodine before applying the alcohol-cleaned tag. National’s ear tags (National Band and Tag

Co., KY, USA) each with a unique identifying number engraved were used. Two tags were

used, one on each front flipper, both through the third nail from the body (Figure 4).

Figure 4. Turtle flipper tag placement

While the turtle was laying (or at latest when returning to the ocean, stopping the turtle en route

by covering the eyes and kneeling in front of her), a health assessment was completed. In

addition to the date, flipper tag numbers and sector; the following health parameters were

observed: head (including the eyes, ears and nostrils), the front and rear flippers, the carapace

and skin, fibropapilomas (e.g., barnacles) and body fat were all recorded and anything unusual

noted.

3.6.3 Results

Between the 1st of January and the 23

rd of March 2014 a total of nine night and 16 morning

turtle patrols were conducted by Frontier and nine night and 28 morning patrols were led by Osa

Conservation on Piro beach. On Pejeperro beach 10 morning and 10 night patrols were carried

out by Frontier and Osa Conservation conducted 19 and 7 morning and night patrols

respectively. During these patrols two Olive ridley turtles and one black turtle were tagged on

Piro beach and four Olive ridley and five black turtles were tagged on Pejeperro. On Piro beach

a total of 29 Olive ridley tracks and 114 black turtle tracks were recorded and 13 predated nests

were excavated. On Pejeperro beach 39 Olive ridley tracks and 206 black turtle tracks were

observed and 14 predated nests were excavated.

3.6.4 Discussion

The greater frequency of encounters of black turtles than Olive ridley turtles is in line with the

peak seasons of these two species. The intensity of tagging has been very insignificant this

phase owing to management of the programme by Osa Conservation such that efforts are

concerntrated on morning patrols when turtle encounters are comparatively rare. New sampling

strategies to increase night patrol effort by means of two simultaneous patrol groups per night




26

patrol have been trialled successfully this phase. It is still under trial and any decision as to

developments or changes to patrol survey effort and protocol will be made by the programme

managers at Osa Conservation. An increased frequency of tagged turtles will progress towards

the aim of estimating population size by capture-recapture analysis and also to monitor the

health of individual turtles through recaptures.

3.7 Diversity and Vertical Stratification of Butterflies within Nymphalidae

3.7.1 Introduction

It is widely thought that we may be currently undergoing a sixth mass extinction event (Wake

and Vredenburg, 2008; Barnosky et al., 2001; Thomas et al., 2004) largely driven by

anthropogenic influences. Some estimates suggest extinction rates have increased to levels

between 50 and several thousand times higher than the background level (Ceballos et al., 2010).

Understanding and documenting biodiversity is therefore of ever increasing importance.

Despite being one of the most biodiverse taxonomic groups; insects are not well studied

compared to other groups such as mammals and birds. Lack of knowledge concerning particular

groups has prevented an accurate estimation of how much biodiversity is being lost and the rate

it is being lost at (Pimm et al., 1995). The study of insects has been impeded by the difficulties

concerned with sampling and issues involved with their identification. This has driven the study

and search for indicator species which provide researchers with a useful short term solution for

rapid biodiversity assessments of insects and potentially other taxonomic groups.

Butterflies have long been considered indicators of biodiversity (Kremen, 1992; Beccaloni,

1995; Blair, 1999; Thomas et al., 2004; Thomas, 2005), habitat disturbance (DeVries et al.

1997; Lien and Yuan, 2002) and environmental change (Thomas et al., 2004). Through studying

butterflies, much information can be learnt about the environment and potentially act as a proxy

for levels of biodiversity for other insect groups which may be more difficult to study (Elmes et

al., 1999).

Compared to other insect groups, butterflies are easily surveyed by methods such as pollard

transect walks and point counts, commonly used in temperate regions, with netting and bait

trapping used in the tropics. Species are comparatively easy to identify and have gained

popularity as a charismatic invertebrate species; combined, these characteristics have resulted in

much volunteer-based research and monitoring programmes in Europe (The UK Butterfly

Monitoring Scheme; Botham et al., 2008). However, despite being one of the more well studied

insect groups, very little is known about many species, particularly those in the tropics where

many species have yet to be evaluated by the IUCN. Currently, very little work has been

conducted on butterfly diversity on the Osa Peninsula, despite values of species richness often

being used in making conservation decisions, delineating protected areas and to monitor

temporal trends.

Many factors impact butterfly diversity, including habitat heterogeneity (Bonebrake and Sorto,

2009; Horner-Devine et al., 2003), vertical stratification and resource partitioning (Schoener,

1974; DeVries, 1997), understanding these is essential to uncovering the mechanisms which




27

support high levels of biodiversity in tropical environments. Much research on butterfly

diversity has determined a positive correlation between species diversity and habitat disturbance

(Johnson, 2011; DeVries et al., 1997; Lien and Yuan, 2002). However, it is important to note

that habitat specialists of conservation importance may not benefit, but in fact be vulnerable to

the effects of habitat disturbance (Lien and Yuan, 2002; Horner-Devine et al., 2003). For

example the Nymphalid species Zaretis ellop, a species present on the Osa Peninsula, requires

undisturbed habitat such as primary forest to thrive (Thomas, 1991).

Species assemblages also vary within vertical space and research has found a level of height

stratification between species occupying the canopy and understory. Despite species richness

being similar in both canopy and understory, species trapped in these two height classes differ,

supporting the theory of vertical stratification amongst butterflies (DeVries, 1997).

Morphological differences have also been observed between canopy and understory species,

suggesting different selective processes acting in the two microhabitats in Costa Rican

rainforests (DeVries, 1997). Vertical stratification has been explained by varying light levels

acting on the ectothermic nature of insects; however it is unlikely that this is the sole factor

maintaining vertical stratification (Walla et al., 2004). Vertical stratification may be in part

related to the height of butterflies’ food plants and therefore resource partitioning determines

height as opposed to light levels. There is also a positive correlation between height of larval

host plants and adult flight height (Beccaloni, 1997). The underlying causes responsible for

vertical stratification currently remain unresolved and require further investigation to clarify

fully.

Although extensive research has been conducted establishing the state of vertical stratification

between canopy and understory butterfly assemblages (DeVries, 1988; DeVries and Walla

2001; DeVries et al. 2010), only a small amount of research has been focused exclusively on

vertical stratification within the understory or canopy layers individually.

Here alpha diversity and vertical stratification of frugivorous nymphalid butterfly species in the

understory were assessed on the Osa Peninsula using a bait trapping methodology. The

investigation aimed to inventory butterfly species in this area to gather baseline data on species

presence for future monitoring. Members of the family Nymphalidae were targeted owing to

their ability to act as an indicator species of other lepidopteran families (Horner-Devine et al.,

2003) and their ease of sampling. It was hypothesized that butterfly species would be randomly

distributed vertically in the canopy.

3.7.2 Methodology

Data were collected by baited live capture canopy traps from 10th December 2013 to 31

st March

2014. Twelve Van Someren-Rydon style (Rydon, 1964) traps were constructed, each with a

cylinder net height of 1m sealed to a plastic tray above and a base tray secured 5cm beneath the

cylinder. This trap design maximized the capture success by exploiting butterflies’ flight take-

off pattern. To increase the completeness of the inventory and representativeness of the study,

traps were non-identical to increase probability of capturing both shade-loving and photophillic

butterfly species (Shuey, 1997); traps were either constructed with opaque or semi-transparent

trays.




28

Twelve trapping locations were systematically chosen where it was practical to erect canopy

traps. Some occupied light spots whilst others were in more shaded areas, all traps were set

within the southern portion of the property. Traps were placed either directly on, or within 5 m

of the river or trail. Six traps were set on the Rio Piro and six on a forest trail (Chiricano

Alegre). The precise trap locations were subject to minor changes during the study period owing

to fallen trees which interfered with the original trap placements.

Traps were suspended from branches at least 11 m above the ground to allow for sampling at

heights of one, two, four, six, eight, and 10 metres at each trap throughout the sampling period

to survey low- and high-flying species. In any given week within each of the two sampled areas,

no two traps were set at the same height; between weeks individual traps rotated between these

heights to ensure sampling effort was equal across traps, heights and sample areas. Therefore, in

a six week period all traps had sampled each height.

Traps were baited with approximately 20g of fermenting banana to attract fruit-feeding

nymphalid butterflies (Family: Nymphalidae); bait was placed in the centre of the lower tray.

Bait was prepared 48 hrs in advance and administered on alternate days unless all bait was

consumed within the first 24 hrs (Hughes et al., 1998). Traps were checked five consecutive

days each week for the duration of the sampling period; traps were opened and baited one day

prior to checks and closed on the fifth day each week. Baiting and checking was conducted once

daily in the afternoon.

Captured individuals were identified in the field to species level using field guides (Chacon

Gamboa and Montero, 2007). Photographs were taken whilst individuals were held in the hand

(where possible) to create an image database of both the dorsal and ventral sides to allow for

identification and three-way ex-situ independent verification following identifications made in

the field. All individuals were released alive. Ex-situ identification verification was undertaken

using additional reference guides (DeVries, 1997; Warren et al., 2013).

Data were analysed using EstimateS (Colwell, 2009) and DIVERSITY (Henderson and Seaby,

1998). Total species richness was estimated in DIVERSITY by Chao Presence/Absence.

Species accumulation curves were formed using Sobs (Mao Tau) values produced in EstimateS.

In EstimateS data were pooled by sampling date, in order to eliminate bias 100 randomisations

were run without replacement. On occasion, individuals escaped traps during the trap check

process; these individuals were excluded from final analyses as were butterfly individuals

whose identity could not be agreed upon by the independent identifiers.

3.7.3 Results

A total sampling effort of 867 trapping days was achieved using twelve traps operating five

consecutive days a week over the course of 15 weeks. Over that sampling period, a total of 266

butterfly individuals were captured across 45 species. Of those individuals; nine (3.38%)

escaped when manoeuvring traps, when handling butterflies or immediately prior to trap checks.

These individuals were excluded from data analyses, as were individuals whose wing condition

was too poor for confident identification to be made (n = 2; 0.75%).




29

The diversity of butterfly species and subfamilies within the family Nymphalidae are sampled

on the property at the riparian, Rio Piro and secondary forest, Chiricano Alegre, locations). A

total of 129 and 137 butterflies were caught on the river and forest trails respectively (Appendix

1). Thirty six species were caught on the river and 37 in the forest. Seven species were caught

exclusively on the river that were not sampled on the forest trail and eight species exclusively

on the forest trail, but not on the river. Thirty species (66.7 %) were captured on both the river

and forest trails suggesting high similarity in species assemblages between the two sites.

As expected, all individuals captured belonged to the family Nymphalidae. The butterflies

sampled comprised five subfamilies. Subfamily diversity was equal across the two sites. The

greatest diversity was sampled in the Charaxinae subfamily, for which a total of 16 species were

observed across all traps at both sites. At the species level, Archaeoprepona amphimachus

amphiktion was the most frequently trapped across the entire survey composing 10.5 % of all

individuals trapped. On the river the most frequently trapped species was Taygetis laches laches

(n = 10) and on the forest trail A. a. amphiktion (n = 19).

Species richness was estimated by Chao Presence/Absence (Figure 5); therefore, our sampling

effort resulted in a survey completeness of 89 %.

Figure 5: Species accumulation curve and expected number of species computed by Chao

Presence/Absence.

3.7.4 Discussion

Obtaining a survey completeness value of 89 % suggests that the survey effort was sufficient for

encountering the majority of the species in this area within the sampling period. The data

collection is still underway, as is the verification process so these figures and species reported

here may be subject to change.

A large proportion of the species sampled on the river were also sampled on the forest trail

(66.67 %). High species assemblage similarity between the two sampling sites is expected given

the close proximity of the river and the trail as well as the biological similarity of the two sites

(both located on or close to a river and bordered with or within secondary forest habitat).

However, there are some differences between the species sampled on the river and within the

forest. Comparing the species caught in each of the two sites provides an interesting view of

some site-specific differences in species assemblages.

0

10

20

30

40

50

60

1 7 13 19 25 31 37 43 49 55 61 67 73

species number

Tra

pp

ing

eff

ort

(d

ays)

Sobs (Mao Tau)

50.556




30

The high frequency of A. a. amphiktion may be explained given that this species is known to

spend a lot of time on the forest floor (DeVries, 1988) so we would expect to capture this

butterfly when trapping in the understory. Consul fabius cerops was captured 15 times within

the sampling period (five on the river and 10 on the forest trail) and is known to fly within both

the canopy and understory layers of the forest. It has also been observed flying along forest and

river edges which may explain its presence in this study (DeVries, 1988). Another species that

was commonly encountered was the Colobura dirce, which is associated with secondary forest

habitat between 3-8 m (DeVries, 1988). This species was caught at heights ranging from 1-8 m

suggesting a greater vertical range in flight height than originally reported. The presence of the

closely related Tigridia acesta acesta suggests that the disturbance level of this area to be

moderate owing to the fact that Tigridia spp. is intolerant of high disturbance levels (DeVries,

1988).

Hamadryas amphinome mexicana is also associated with secondary forest growth. Catonephele

mexicana is also reported to be tolerant of secondary forest habitat, (DeVries, 1988) for which

further support is found in the results here. Many of the species encountered here, including

Hamadryas spp. occur in all rainforest environments (DeVries, 1988).

Moderate levels of disturbance have been found to have positive impacts on butterfly diversity

(Horner-Devine et al., 2003; DeVries et al., 1997; Johnson, 2011). So it would be expected that

high species diversity would be sampled in a secondary growth area. CNP has 220 butterfly

species (belonging to all families, not just Nymphalids). Considering that 460 of the 1250

species of butterfly in Costa Rica are Nymphalid butterflies (36.8%), if we assume that roughly

this percentage of butterfly species present in CNP belong to the Nymphalidae family,

approximately 81 species of Nymphalid butterfly occur here. This may seem unusual given that

higher species diversity should be expected on the property surveyed at Piro given the level of

disturbance compared to CNP which should be relatively undisturbed. This disparity in results

may be in part due to the small area surveyed here and the surveying of the understory species

only and therefore direct comparisons with CNP should be interpreted with caution.

Some species encountered are thought to be rare and of conservation concern such as Historis

acheronta acheronta, which was only encountered once within the sampling period. Some more

unusual findings include the presence of Memphis artacaena, known to occur mostly within

primary forest habitats and occupies the canopy layer. However this species was documented

four times during the study and the majority of these were along the river. This may indicate

that this butterfly is more of a generalist with respect to the heights it occupies and the habitat it

can inhabit. However males are known to perch at riparian edges which may account for this

species’ presence, particularly given that three out of the four were sampled along the Rio Piro.

Similarly Zaretis ellops is reported to occur in primary growth and undisturbed habitats; yet was

sampled in our study within secondary forest, although not often. This may suggest that either

the property is not significantly disturbed or that these butterflies are not as restricted with

respect to their ecology and habitats as previously alluded to.

Hamadryas arinome ariensis is also documented to occur on the Atlantic slope within the

canopy, so this is also an unexpected find on the Osa Peninsula. Nessae aglaura aglaura was




31

sampled twice over the course of this study and is usually noted for inhabiting swamp

environments within primary forests. Interestingly, this species was only sampled along the

river traps and never along the forest trail. N. a. aglaura is noted for its intolerance of habitat

disturbance (DeVries, 1988), which could account for the low encounter frequency of this

species here. The greatest species diversity was within the Charaxinae subfamily and the genus

Memphis which would be expected given the high diversity of this genus, particularly when

compared to other genera of the area (for example Tigridia acesta is the only species of its

genus).

Seasonality impacts butterflies’ abundance and although the wet season is a better time for

butterfly diversity, it is impractical to sample via bait trapping in the wet season. Over the

course of the study some trends have arisen; for example Nessae aglaura aglaura was only

captured at the beginning of the sampling period. These trends may be related to seasonality of

specific butterfly species. A year round study would make an interesting addition to the data

collected here.

In conclusion, the butterflies present on the Osa property suggest that the area has some degree

of moderate disturbance. A number of species known to inhabit primary forest have now been

documented on the site, suggesting that the habitat requirements of some species are not as

restricted as previously documented. However, the low frequencies at which these species were

sampled may suggest that they are present but rare in this environment; perhaps because they

are not well suited to the more disturbed secondary growth habitats. The study area contains

areas of primary forest and plantation. Sampling in these areas would make an interesting

comparison and perhaps supplement the inventory. The presence of some unexpected species

suggests that this area should be monitored to assess change in butterflies in this area. Mark-

release-recapture studies could also complement the information found in this study and provide

an idea of the abundance of different species.

3.8 The Three-Dimensional Space Use of Mantled Howler Monkeys (Alouatta

palliata)

3.8.1 Introduction

Mantled howler monkeys (Alouatta palliata) have an extensive range in Central and South

America from Mexico through to Peru. The IUCN classifies A. palliata as Least Concern

however their current population trend is not known (Cuarón et al., 2008b). They are primarily

folivorous but will supplement their diet with other vegetative substrate. Mature fruits of Ficus

species are an important dietary component and there is a correlation between scarcity of fruits,

dietary stress and mortality (Milton, 1982). The large range and different forest types that A.

palliata can occupy has been attributed to their relatively flexible diet (Cuarón et al., 2008b).

Unlike other species of New World primates, A. palliata is fairly resilient to anthropogenic

disturbance and can tolerate moderate levels of habitat fragmentation as observed in the A.

palliata mexicana population in Los Tuxtlas, Mexico. However, the level of fragmentation and

patch size does impact their population density, group size and behaviour, meaning they are still

under threat from habitat loss (Arroyo-Rodriguez and Dias, 2010).




32

The diversity and density of vegetative species has a positive impact on Mantled howler

monkey populations (Cristóbal-Azkarate et al., 2005; Estrada and Coates-Estrada, 1996). An

abundance of large and fruting trees (diameter at basal height >60cm) are important for howler

monkey populations and scarcity in either of these resources leads to altered foraging behaviour;

groups will feed from smaller, more widespread food resources when big, fruiting trees are

scarce (Dunn et al., 2009). An increase in distribution of food resources inevitably increases a

group’s foraging effort, thus allowing less time for other behaviours and an elevated energetic

output (Dunn et al., 2009) which may be detrimental to the health of individuals as their diet is

primarily composed of leaves - a low energy source. There are numerous large fruiting tree

species on the property, including Ficus species, so investigating their behaviour in these trees

and the time spent foraging may provide an indication about the suitability of the area for

mantled howler monkeys.

The aim of the project is to investigate the behaviour of Mantled howler monkeys and their use

of three-dimensional space within the Osa Conservation property at Piro and its boundaries.

Specifically, the objectives are to: i) record the behaviour of all visible individuals every 10

minutes using a scan sampling method; ii) map the distributions of the different groups within

the study region, and; iii) estimate the average canopy height of the group.

3.8.2 Methodology

Data were collected from 0600 until 1730 hrs, effectively dawn until dusk, six days per week

during March 2014. Attempts were made to find different groups on the property by varying the

search area to ensure that data were representative of the population. However, due to safety

concerns, northern areas of the property were not utilised due to the topography of the area.

When a group was encountered, they were followed for the full day. On the hour and every 10

minutes thereafter, scans were taken and the following data collected: canopy height; GPS

point; and behaviour of visible individuals. As the members of the group could not all be seen at

once, three minutes was used to gain as much behaviour data as possible.

The canopy height was calculated using a laser range finder, clinometer and trigonometry:

sin(angle) *radial distance. Notes were also made regarding the substrate the group was feeding

on and any identifiable characteristics of individuals in the group. In addition, any intergroup

species interactions were recorded, which ranged from the presence of another species in the

area to any physical contact between individuals (for example, we observed a fight between a

female howler monkey and a spider monkey where the howler monkey fell 20 m to the ground).

The GPS waypoints will be mapped using GIS and the behavioural data will be statistically

analysed to see whether the canopy height has a significant impact on mantled howler monkey

behaviour.

An ethogram (Table 7) was developed and used to assist in the definition of the behaviours. Any

additional behaviours were recorded and added to the ethogram.




33

Table 7. Ethogram describing the different behaviours observed

Behaviour (acronym) Definition Classification

Resting (R) Individual lying/sitting and did not move for the

duration of the scan

Resting

Hanging Tail (HT) Individual was hanging from tail Resting

Feeding Leaves (FL) Individual was feeding on leaves Feeding

Feeding Fruits (FF) Individual was feeding on fruits Feeding

Feeding Stems (Fstems) Individual was feeding on the stems of leaves Feeding

Locomotion (L) Individual was moving in the same tree for more

than 10 seconds (i.e. not engaged in another

behaviour)

Locomotion

Travelling (T) Individual was travelling from tree to tree with

obvious purpose

Locomotion

Playing (P) Individual was playing either alone or with a

conspecific

Social Interaction

Aggressive (AGG) Individual screeches or fights with another

individual (in these cases the recipient and the

aggressor was noted)

Social Interaction

Howling (H) Individual was howling Social Interaction

Grunting (G) Individual was grunting Social Interaction

3.8.3 Results

Data collection is still underway for this project so as of yet, there are no presentable results.

The spatial data from 1st March until 17

th March has been mapped onto GIS as part of a

Research Assistant’s BTEC project (Figure 6). Each coloured line denotes the daily movement

of the group followed that day. The green circles represent where each waypoint was taken; the

larger the diameter of the circle, the more time the howler monkeys spent at that location.

Resting and feeding were the main behaviours observed in these large circles.




34

Figure 6: Movements of the howler monkey groups and an indication of the duration at each

location.

3.8.4 Discussion

It can be seen that going off trail and following the groups for the whole day provides a great

deal of data on the movement of the groups (Figure 6). The clumping of some locations

suggests that the same group is being followed on different days, so it is anticipated that the

continuation of this project will provide information about group home ranges. Gaining an

insight into the movement and activity patterns of the Mantled howler monkey population on

the property expands the primate project within CRF. Hopefully, as we learn more about the

population, new research projects that will promote the conservation of these important seed

dispersal agents can be carried out.




35

4. Additional Projects

4.1 Testing the Influence of Hiking Trails on Bird Diversity and Abundance

within Neotropical Primary and Secondary Lowland Forest.

Within the project’s study area there are 16 km of hiking trails, primarily focused around the

southern region within primary and secondary forest. There is evidence that hiking trails create

an internal edge effect within natural habitat, with the implication that this edge may lead to

changes in the species composition of bird communities (Miller et al., 1998). Human presence

on such trails is also linked to negative impacts on some species, affecting feeding strategies

(Fernández-Juricic, 2000) and nesting (Fernández-Juricic, 2002).

The methodology was designed and trialled between 23rd

February and 13th March 2014 by

ARO John Scott and PI Nathan Roberts. Geographical Information System (GIS) mapping was

used to generate 20 locations to carry out bird point counts. Fifty percent of the points occur on

hiking trails, while 50% of the points were placed 100 m away from any hiking trails. The

selected points are also split between primary and secondary forest. There is expected to be a

difference in diversity and abundance of different species between on-trail and off-trail points,

and the aim of the study is to discover which species are likely to be affected most (either

negatively or positively) by the presence of hiking trails.

This study will focus only on resident species, with data being collected over a four month

period finishing at the end of July 2014. During this period, a target has been set to visit each

point 12 times, amounting to 240 points surveyed. It is expected that this work will be reported

in Phase CRF142.

4.2 Measuring Disturbance Through Novel Techniques: Soundscape Ecology

within a Heterogeneous Landscape on the Osa Peninsula, Costa Rica.

Surveying a habitat such as a tropical forests, assessing biodiversity can be long, arduous and

impractical (Lawton et al., 1998). Traditionally these assessments have depended on species

inventories, however soundscaping offers a viable way to conduct rapid biodiversity

assessments which is cheap, less labour intensive, less invasive and can be carried out by

amateurs (Penman et al., 2005, Frommolt et al., 2008, Snaddon et al., 2013). Currently

soundscape ecology offers a great platform to conduct general biodiversity assessments.

Soundscape ecology differs from other fields of acoustic biology by focusing not on species or

individual levels, but by looking at the broader picture and studying macro or community

acoustics (Pijanowski et al, 2011). Like landscape ecology, soundscape ecology looks at the

interaction of pattern and ecological processes across broad spatial regions.

A soundscape is everything that can be heard in an environment. Soundscapes are comprised of

three components, biophony (sounds created by organisms), geophony (non-biological sounds;

rain, wind, thunder etc) and anthrophony (sounds that emanate from humans and their

inventions) (Krause, 1987, Pijanowski et al., 2011).




36

Due to sound being a core property of nature that can be radically affected by anthropogenic

disturbance, it is easy to find a relationship between biophony and environmental quality; a rich

and diverse biophony will likely reflect a healthy ecosystem (Pijanowski et al., 2011). This

assumption, along with the acoustic complexity index, is vital in the analyses of soundscapes in

determining the disturbance levels of a ecosystem.

The aim of this study is to make a health assessment of the Osa Conservation property at Piro.

This will be done be done by soundscaping nine locations to gain an understanding of past and

present disturbance levels.

Nine random locations have been chosen across the Osa conservation property at Piro.

Recording will take place twice a day, with one hour being recorded at the dawn chorus

(5:30a.m.-6:30a.m.) and one hour during the dusk chorus (5:00pm-6:00pm). From these

recordings, the first minute of every five minutes will be analysed, using the package

soundecology in the statistical program R for a rapid biodiversity assessment and acoustic

complexity index.

While waiting for the soundscape proposal to be cleared by Osa conservation, the project is set

up and ready to start as soon as permission is granted.

4.3 Systematically Comparing Methods of Studying Butterfly Diversity: Canopy

Trapping Versus Netting.

Accurately assessing species diversity is crucial to assessing levels of biodiversity in a given

area. Often conservation depends on protecting certain areas based on levels of biodiversity and

species richness (Kery and Plattner, 2007). Butterflies (specifically Nymphalidae) are often

studied to gain an understanding of biodiversity, disturbance and environmental conditions

because they act as indicator taxa for other butterflies, insects as well as vertebrate species

(Horner-Devine et al., 2003, Thomas, 2005).

Despite high species diversity of butterflies in the tropics, very little work has been conducted

on butterfly diversity and abundance. The work that has been conducted used bait traps which

exploit the feeding behaviors of certain butterfly species depending on the bait used (Austin and

Riley, 1995). A second method for assessing butterfly diversity is netting specimens in the field.

Which method is best is still debated, as there have been no direct comparisons between these

two methods to determine which is more effective and efficient for compiling species

inventories of tropical butterflies.

This study will compare the two methods, aiding future research into butterfly diversity in

tropical environments. The site for this study is the road that runs along the coast of the Osa

Peninsula connecting Puerto Jimenez with Carate. The two kilometre section of road to be

surveyed cuts through the Osa conservation Piro property and has dense tropical forest on either

side. Ten Van Someren-Rydon style bait traps will be used, each spaced 200 m apart from their

neighboring traps. All traps will be set at 1.5 m above ground and will be monitored every 24

hrs with re-baiting occurring every 48 hours. Along the same stretch of road, netting will occur

every day that the bait traps are open. The hours of netting will equal the same number of

human effort hours that occur when monitoring the traps.




37

After the data collection, comparisons between the two methods in terms of species/number of

individuals caught will take place in order to determine which, if any method is most efficient in

tropical habitats. The work is subject to permit approval.




38

5. Proposed Work Programme for Next Phase

Primate surveys will continue sampling evenly across trails and times until sufficient data is

collected for all four primate species. It is expected that in Phase CRF142 it will be possible to

report on howler monkey density and that this result will be prepared for peer-reviewed

publication as the research article of spider monkey density.

The in-situ otter research will continue to follow the same protocol in Phase CRF142. Towards

the end of Phase CRF141 there were no encounters of spraint on the Rio Piro (N) transect for

two consecutive weeks. This will continue to be explored as will any temporal changes in

distribution and environmental variables. In the next phase it is also anticipated that the ex-situ

project will make considerable progress.

The next phase will be an intermission between the two peak turtle nesting seasons experienced

on Piro and Pejeperro beaches. It is expected that turtle patrols will continue at the same

frequency and we will continue to work with Osa Conservation on the programme. Leading on

from a successful BTEC project, CRF also anticipates further investigation on the effects of

lunar phase and tide on marine turtle nesting behaviour. This will involve using historical data

and new data from night patrols in Phase CRF142.

In the next phase butterfly monitoring will continue. Further analysis is to be completed

following completion of species verification after which, the hypothesis regarding vertical

stratification will be tested. Future work will incorporate additional study sites for canopy

trapping in the primary forest and plantation areas of the property, which will provide a more

comprehensive species inventory of the property as a whole. Trapping in these locations would

also make for an interesting comparison study of the butterfly diversity present in these different

areas. This expansion of the butterfly programme will probably be best suited to the next year

owing to the approaching wet season and the associated impracticalities for canopy trapping.

Following the success of the new primate behaviour project, it is anticipated that this work can

be expanded upon, continuing with the recording and analysis of spatial data, supported by

individual group identification. This study would aim to give us a clearer understanding of the

species use of the property.

Between June and December 2013, 40 interviews were conducted with workers at livestock and

agricultural produce properties, with the objective of: i) understanding the occurrence and scale

of human-wildlife conflict; ii) understanding local attitudes towards conservation, and; iii)

providing a rapid assessment of the status of local wildlife populations. The sampled area was

between Puerto Jimenez and Carate on the Osa Peninsula. To maximise the representativeness

of this study prior to analyses, it is anticipated that more farm surveys will be conducted within

the Osa Conservation Area (ACOSA) in the northern part within Phase CRF142. This is

expected to form the first phase of a larger interdisciplinary study which will shed light on the

human dimensions and ecological constructs of conflict in this previously unreported locality

and will build a novel framework for resolving human-wildlife conflict. Ultimately, this project

will allow for the conservation of wildlife, including one of the most human-impacted wildlife

species, the jaguar (Quigley and Crawshaw, 1992).




39

6. References

Almeida, D., Barrientos, R., Merino-Aguirre, R. & Angeler, D. (2012). The role of prey

abundance and flow regulation in the marking behaviour of Eurasian otters in a Mediterranean

catchment. Animal Behaviour, 84 (6), 1475-1482.

Anoop, K. R. & Hussain, S. A. (2004). Factors affecting habitat-selection by smooth-coated otters

(Lutra perspecillata) in Kerala, India. Journal of Zoology, 263, 417-423.

Arroyo-Rodríguez, V. & Dias, P. A. D. (2010). Effects of habitat fragmentation and disturbance

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7. Appendices

Appendix 1. Capture Frequency of Butterfly Species Within the Two Sample Areas

Sub Family Species Chiricano

Alegre

Rio Piro Total

Biblidae Catonephele mexicana 3 12 15

Catonephele numilia esite 4 3 7

Hamadryas amphinome mexicana 2 3 5

Hamadryas arinome ariensis 0 2 2

Hamadryas februa ferentina 9 3 12

Hamadryas feronia farinulenta 0 2 2

Hamadryas ipthime ipthime 0 1 1

Hamadryas laodamia saurites 1 2 3

Nessaea aglaura aglaura 1 1 2

Nica flavilla 1 2 3

Charaxinae Archaeoprepona amphimachus

amphiktion

19 9 28

Archaeoprepona camilla Camilla 4 1 5

Archaeoprepona demophon centralis 3 2 5

Consul fabius cerops 10 5 15

Fountainea eurypyle confuse 3 4 7

Fountainea halice chrysophana 2 1 3

Memphis arginussa eubaena 1 2 3

Memphis artacaena 1 3 4

Memphis beatrix 1 0 1

Memphis lyceus 1 0 1

Memphis moruus boisduvali 6 9 15

Memphis oenomais 5 1 6

Memphis pithyusa pithyusa 1 0 1

Prepona laertes demodice 1 0 1

Prepona omphale Octavia 1 1 2

Zaretis ellops 2 1 3

Morphinae Caligo atreus dionysos 3 0 3

Caligo brasiliensis sulanus 5 2 7

Caligo illioneus oberon 0 1 1

Caligo telamonius memnon 1 3 4

Morpho menelaus amathonte 7 5 12

Morpho peleides marinita 3 2 5

Opsiphanes bogotanus Alajuela 3 3 6

Opsiphanes cassina chiriquensis 0 1 1

Opsiphanes tamarindii tamarindii 0 1 1

Nymphalinae Colobura annulata 6 7 13

Colobura dirce dirce 4 6 10

Historis acheronta acheronta 0 1 1

Temenis laothoe hondurensis 2 6 8

Tigridia acesta acesta 3 0 3

Satyrinae Chloreuptychia arnaca 3 0 3

Cissia confuse 0 2 2

Magneuptychia alcinoe 1 1 2

Pareuptychia metaleuca 0 1 1

Pareuptychia ocirrhoe ocirrhoe 1 4 5

Taygetis laches laches 2 10 12

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