A Survey of Ranging Patterns and Micro-Habitat Preference of Saguinus Midas In

Home , Callitrichidae, Tamarin

A Survey of Ranging Patterns and Micro-Habitat Preference of Saguinus midas in

Berg en Dal, Suriname

A thesis submitted

to Kent State University in partial

fulfillment of the requirements for the

degree of Master of Arts

Gina Marie Cherundolo

August, 2018

Thesis written by

Gina Marie Cherundolo

B.A., The Pennsylvania State University, 2009

M.A., Kent State University, 2018

Approved by

Mary Ann Raghanti , Advisor

Mary Ann Raghanti , Chair, Department of Anthropology

James L. Blank , Dean, College of Arts and Sciences

TABLE OF CONTENTS

TABLE OF CONTENTS ...... iii

ACKNOWLEDGMENTS ...... vii

Chapter

I. INTRODUCTION ...... 1

Suriname: Ecology and Population ...... 1

Biodiversity of Suriname ...... 3

Phylogenetic Context ...... 4

Primates of Suriname ...... 4

Family Callitrichidae ...... 5

Saguinus midas ...... 8

Tree Selection and Forest Structure ...... 10

Study Questions ...... 11

II. METHODS...... 12

Research Overview ...... 12

Survey Area ...... 13

Sampling Methods: Primates ...... 14

Sampling Methods: Habitat ...... 15

Analysis...... 16

III. RESULTS ...... 18

Habitat Assessment ...... 18

Observations of Saguinus midas and using the GPS for tracking ...... 23

Home range size and tamarin group counts at Bergendal ...... 25

iii

IV. DISCUSSION & CONCLUSION ...... 28

REFERENCES ...... 31

APPENDIX A ...... 35

APPENDIX B ...... 38

APPENDIX C ...... 42

LIST OF FIGURES

Figure 1.1: The Guiana Shield Region ...... 2

Figure 2.1 – Bergendal Resort and surrounding forest ...... 13

Figure 3.1: RSC Trail system with selected trail markers ...... 19

Figure 3.2: RSC Trail System with initial tamarin encounters with 50 x 50 m grid overlay ...... 22

Figure 3.3: RSC Trail System with white dots indicating trail marks ...... 25

Figure 3.4: Home range assessment for tamarins in survey area ...... 27

LIST OF TABLES

Table 1.1 –Home Range Comparison for Selected Saguinus (Leontecebus included as former members of Saguinus)...... 9

Table 1.2 – Genus Saguinus Group Metrics (Adapted from Porter et al. 2017) ...... 10

Table 3.1: Summary of Tree Measurements ...... 20

Table 3.2: Tamarin Observations with GPS Tracking Information ...... 24

Table 3.3: Home range and group size calculations for all four groups in survey area ...... 26

Table 4.1: Saguinus midas group size comparisons ...... 28

ACKNOWLEDGMENTS

First and foremost, I wish to extend my sincere thanks to my advisor, Dr. Marilyn

Norconk, for her assistance, encouragement and patience during this process. In addition, I would like to thank the other members of my committee, Drs. Richard Meindl and Mary Ann

Raghanti, for their much-appreciated input towards the completion of this thesis. I also want to thank all of my instructors at Kent State for their instruction and guidance, both in and out of the classroom.

Additionally, I must thank the Graduate Student Senate for assisting me in my travels to

Suriname, as well as the Sylva Koemar and the wonderful employees at Bergendal for making sure we were all taken care of. I need to also thank my assistants in the jungle, Katrina (Dietz)

Eschweiler, Jennifer Humphreys and Alex Mason for all their tireless help finding monkeys and measuring trees.

A personal thanks to Dexter Zirkle and Dr. Eric Seemiller for their great support and advice throughout this process. I could not have done this without them. I also could not have done this without the support of Caroline Tannert and Barbara Davis.

Finally, I need to thank my parents, John and Cheryl Cherundolo, for their extreme patience, support and encouragement.

vii

Chapter I

Introduction

Suriname: Ecology and Population

Suriname, formerly Dutch Guiana, is the smallest independent nation in South America.

It is situated in the northeastern part of the continent, with French Guiana to the east and Guyana to the west. These three nations are informally known as the “Guianas” and together encompass about 500,0002 km (Boggan et al. 1997). Suriname encompasses 163,8202 km of mostly forested habitat and has a population of more than 591,919 people as of June 2018 (Central Intelligence

Agency 2018). As of 2004, approximately 67% of the population lives in the Paramaribo metropolitan area, with 20% residing along the coastal regions and the remainder living in small tribal communities in the interior (National Institute for Environment and Development in

Suriname 2016). Suriname’s population density is low compared to the rest of the world, averaging three people/km2 (National Institute for Environment and Development in Suriname

2005).

Initially an English colony, Suriname fell under Dutch control in 1667 (Davis 2009).

Suriname was colonized and became a plantation economy, using slave labor to produce and export goods including cotton, coffee and sugar (Hendrison and de Graaf 2011). Timber plantations were also common, especially between 1700-1900, and were mostly located along major waterways, including the Suriname River (Hendrison and de Graaf 2011). Land closest to

the rivers were depleted quickly, due to lack of machines (all sawing was done manually) and animal power to source timber farther inland (Hendrison and de Graaf 2011). Plantation numbers decreased quickly after slavery was abolished in 1863 (Hendrison and de Graaf 2011).

Suriname is part of a unique geological and ecological region known as the Guiana

Shield (Fig. 2.1), a 4.6 billion-year-old landmass between the Atlantic Ocean and the Amazon and Orinoco Rivers and encompasses all of modern-day Suriname, French Guiana and Guyana, as well as parts of Brazil, Bolivia, Colombia and Venezuela (Boggan et al. 1997). With 30,000 endemic plants, 75% of primary vegetation retained, and fewer than five people/km2 the Guiana

Shield is considered a “good news area” for conservation of biodiversity (Myers et al. 2000).

Figure 1.1 – The Guiana Shield Region (NMHS 2015, adapted from

http:// http://botany.si.edu/bdg/)

Average annual air temperature ranges from 26˚C in January to 31˚C in October

(National Institute for Environment and Development in Suriname 2016). Average annual

rainfall ranges from approximately 2,445 mm in central east Suriname to 1,561 mm on the north central coast (National Institute for Environment and Development in Suriname 2016). The entire Guiana Shield region sees at least one, and most of the time, two dry seasons each year

(Boggan et al. 1997). This is due to the yearly oscillation of the intertropical convergence zone

(ITCZ) (Nurmohamed and Naipal 2006), a band of low pressure that encircles the earth near the equator and continually moves, or oscillates, throughout the year to correspond to each hemisphere’s summer. It is positioned north of the equator in accordance with the northern hemisphere’s summer and below the equator during the southern hemisphere’s summer.

Consistent with this pattern shift, Suriname has one long wet season spanning from April to mid-

August, followed by a long dry season from mid-August to November (Nurmohamed and Naipal

2006). After these long seasons, a short wet season runs from December to February, followed by a short dry season from February to March (Nurmohamed and Naipal 2006). These seasons can vary from year to year; sometimes the seasons last longer or shorter than expected, and sometimes they skip altogether (Boggan et al. 1997). Above or below normal precipitation can occur depending on the presence of El Niño or La Niña oscillations, as they affect sea surface temperatures (Nurmohamed and Naipal 2006).

Biodiversity of Suriname

Suriname is one of the few tropical areas on Earth to have most of its forests untouched

(Boggan et al. 1997), with more than 90% of the country covered with forest (National Institute for Environment and Development in Suriname 2016). The entire Guiana region boasts a variety of geographic regions, ranging from mountains to coastal plains, and numerous different vegetation types, including swamp forests, marsh forests and mangrove (Boggan et al. 1997),

and it is this range of topography that is largely responsible for the vast number and diversity of plant and animal species (Funk et al. 2007). Suriname has 4,984 species of plants, 730 species of birds, 175 species of reptiles, 102 species of amphibians, 450 species of freshwater fish and 192 species of mammals (including eight primates) (General Bureau of Statistics of Suriname 2016).

Phylogenetic Context

The primates of Central and South America comprise the parvorder Platyrrhini, or New

World monkeys, the fossil record of which dates back to approximately 26 million years ago

(Wildman et al. 2009). Currently, 152 species are recognized (or 204 species and subspecies together) of New World monkeys (Rylands et al. 2012). There is debate as to how many families are classified within Platyrrhini, with Hershkovitz (1977) placing Platyrrhines into just two families, Callitrichidae and Cebidae. Groves (2001) expanded this taxonomy, using four families

(Cebidae, Aotidae, Pitheciidae and Atelidae). Rylands and Mittermeier (2009) maintain much of

Grove’s model, with the major exception of placing marmosets and tamarins in their own family,

Callitrichidae, rather than as a subfamily of Cebidae.

Primates of Suriname

Suriname is home to eight species of primates (Baal et al. 1988) that represent four families and eight genera, following the revised New World monkey taxonomy as described by

Rylands and Mittermeier (2009) and Garbino and Martins-Junior (2018). Family Atelidae has two species in Suriname: Alouatta seniculus and Ateles paniscus. The Pitheciidae also has two species: Pithecia pithecia and Chiropotes satanas. Members of the Cebidae include Sapajus apella, Cebus olivaceus and Samiri sciureus (Baal et al. 1988). The lone callitrichid in Suriname

is Saguinus midas (Baal et al. 1988). Formerly, marmosets and tamarins were considered to be a subfamily in Cebidae (capuchins and squirrel monkeys), but they have since been reclassified as their own family, Callitrichidae, composed of five families (Rylands and Mittermeier 2012).

Family Callitrichidae

Platyrrhines evolved isolated from their Old World counterparts, and thus exhibit several divergent characteristics. Prehensile tails that can grasp, serve as a fifth limb and that are strong enough to hold the animal’s entire weight, are only found in the families Atelidae and Cebidae

(Wildman et al. 2009). Members of the family Aotidae (genus Aotus) are the only nocturnal anthropoid primates (Wildman et al. 2009). It is also notable that the largest adult member of the platyrrhines (the 11.4 kg Mexican black howler monkey, Alouatta pigra) is 100 times heavier than the smallest adult member (the 110 g pygmy marmoset, Callitrhrix pymaea) (Wildman et al.

2009).

Members of the Callitrichidae also exhibit unusual and unique behavioral and reproductive characteristics among all primates, including a large maternal-to-infant mass ratio, post-partum estrus, alloparental care of infants, short interbirth intervals, rapid male and female sexual maturity (Digby et al. 2007) and reproductive suppression in females, which will be discussed later (Diaz-Munoz 2016). Members of this family are the only primates to practice polyandry, in particular, “cooperative polyandry,” or mating between a single female and at least two males with stable partnerships (Baker et al. 2002). A female’s status and seniority have an impact on her reproductive success in the group, and she will increase her reproductive potential by suppressing the potential of the subordinate females, likely in order to secure the aid of adult males and adult offspring with raising (carrying) her offspring (Diaz-Munoz 2016). Groups will

consist of typically one reproductively active female, and this female will grant reproductive access to more than one male at a given time (Baker et al. 2002). Unlike many primate species, callitrichids do not exhibit signs of physiological stress as part of their reproductive suppression

(Beehner and Lu 2013). While the exact mechanisms are unknown, adult females residing in their natal groups present non-cycling levels of gonadotropins and ovarian hormones, and females that become subordinate also show a significant drop in these hormone levels (Beehner and Lu 2013). The reproductively active female will suppress ovulation of subordinate females via neuroendocrine mechanisms and behavioral, visual and olfactory cues, as observed in the closely related common marmoset, Callithrix jacchus (Beehner and Lu 2013). However, Beehner and Lu (2013) argue that odor, visual and chemical signals were temporary and did not maintain suppression, indicating that social cues were the dominant stimulus for suppressing ovulation in subordinate females.

Further compounding callitrichid reproductive potential is rapid sexual maturation in both females and males. Females reach sexual maturity beginning at 12 months of age, with gestational length ranging from 125 days in Leontopithecus species to 184 days observed in

Saguinus oedipus (Digby et al. 2007). Males begin to produce sperm beginning at 13 months of age and can father young by 15 months of age (Digby et al. 2007). The majority of offspring are fraternal twins, with triplets and singletons (most likely the result of a pregnancy in which the other dizygotic twin was resorbed or aborted early) less common (Hershkovitz 1977).

Males and females co-parent offspring (known as alloparenting) no matter the paternity

(Digby et al. 2007). Multiple males, all with significant likelihood of paternity, will collectively rear the offspring of a single female (Baker et al. 2002), and there is strong selection pressure for this strategy due to the high reproductive demands on the female (Diaz-Munoz 2016).

Callitrichids have a short interbirth interval, with many species being able to ovulate and conceive within a month of giving birth, and lactation does not obstruct postpartum ovulation

(Digby et al. 2007). Thus, many species of callitrichids can give birth twice in one year, the highest reproductive potential of any anthropoid primate; however, total lifetime output is severely limited by overall short lifespans and high infant mortality rates, compounded with reproductive suppression in subordinate group members (Digby et al. 2007).

All Neotropical primates are strictly arboreal. As part of the Platyrrhini, callitrichids inhabit Central and South America in a variety of habitats that vary in tree density, rainfall and climate, including evergreen, secondary, coastal, lowland and riverine forested regions (Digby et al. 2007).

Several species of tamarins have been observed to form mixed-species, or polyspecific, associations. These groups travel, forage and rest together, as well as defend a common home range (Bicca-Marques and Garber 2003). S. imperator, S. mystax, S. labiatus, S. fuscicollis have all been observed in long-term, stable polyspecific associations with another member of

Saguinus (Bicca-Marques and Garber 2003, Norconk 1990).

While the majority of New World Monkeys have the dental formula 2.1.3.3, marmosets and tamarins differ from the rest of the platyrrhines in that they lack the third molar in both the upper and lower jaws, resulting in a dental formula of 2.1.3.2 (Moynihan 1976). This reduction is likely because callitrichid jaws are too short to contain a full set of teeth, or possibly because the insectivorous diet can better be masticated with fewer teeth (Moynihan 1976).

A key morphological and geographical difference between tamarins and marmosets is the dental adaptation to extract tree gums and saps as a substitute for fruit (Digby et al. 2007).

Species of marmosets use their lower incisors to gouge holes in tree trunks and branches using

derived characteristics in the lower jaw and dentition, including elongated incisors, thickened buccal enamel, jaw muscles and a V-shaped mandible (Garber 1992, Thompson et al. 2014).

They use these adaptations to repeatedly take advantage of nutrient-rich tree sap, or exudates.

This ability allows Cebuella, Callitrix and Mico to inhabit regions that are highly seasonal, poor in resources and/or disturbed where tamarins would otherwise not be able to live (Digby et al.

2007). Tree exudates are crucial to the marmoset’s diet as they are consumed year-round and account for more than 70% of overall diet during certain times of the year (Thompson et al.

2014).

Saguinus midas

Seven genera are currently recognized as part of family Callitrichidae, which is the most species-rich family of all platyrrhines. Following Rylands et al.’s (2016) model, they are

Cebuella, Callibella, Mico, Callithrix, Callimico, Leontopithecus, Leontocebus and Saguinus, the latter three genera comprising all tamarins.

Saguinus midas, also known as the golden-handed or red-handed tamarin, is found in

Guyana, French Guiana, Suriname and south into northern Brazil. Hershkovitz (1977) describes the species has having blackish ears, face, external genitalia, forequarters, thighs and tail. The face and ears are nearly hairless, with the middle and lower back hair marbled in color and discernable from the black mantle (Hershkovitz 1977). The most distinct feature of S. midas are the hands and feet, ranging in color from shades of yellow to orange and red (Hershkovitz 1977).

Hershkovitz also described the species having black feet, but this variation is now recognized as a separate species, Saguinus niger (Rylands et al. 2016).

Table 1.1 –Home Range Comparison for Selected Saguinus (Leontecebus included as

former members of Saguinus) (Adapted from Digby et al. 2007:88)

Current Species Home Range (ha) Daily Path Length (m/day) Leontocebus fuscicollis 40 1,849 L. weddelli 44+ 1,312 Saguinus imperator 30-100 1,420 S. midas 31.1-42.5 N/A S. mystax 40 1,946 S. niger 35 N/A L. tripartitus 16-21 500-2,300

S. midas has an average body mass of 515 g (±34 g) in females and 575 g (±15 g) in males, making the females slightly above-average for genus Saguinus and males the heaviest

(Smith and Jungers 1997). Body length has an average of 237 mm (±18 mm) (Hershkovitz

1977). Their home range averages between 31.1-42.5 ha (Day and Elwood 1999) in groups ranging from 2-12 individuals (Porter et al. 2017). S. midas is able to exploit a variety of food sources, such as ripe fruits, insects and small invertebrates (Digby et al. 2007). They take advantage of small fruit patches found lower in the forest canopy and tend to prefer edge habitats, which may reduce competition from larger-bodied competitors (Digby et al. 2007). S. midas is like most other callitrichids, is a pioneer species that favors secondary and disturbed habitats (Oliveira and Ferrari 2000).

Table 1.2 – Genus Saguinus Group Metrics (Adapted from Porter et al. 2017)

Species Min Max Mean Group Size SD n Saguinus bicolor 2 11 4.9 N/A 4 Leontocebus fuscicollis 1 17 6.1 1.81 8 Saguinus geoffroyi 19 6.2 0.36 2 Saguinus imperator 3 7 4.8 N/A 2 Saguinus inustus 6.0 2.70 1 Saguinus labiatus 2 13 7.3 N/A 2 Saguinus leucopus 2 12 5.2 3.20 3 Saguinus martinsi 3 8 5.0 N/A 1 Saguinus midas 2 12 5.8 2 4 Saguinus mystax 1 16 7.5 0.99 3 Saguinus niger 3 7 5.0 N/A 2 Saguinus oedipus 2 15 5.8 2.65 4 Leontocebus tripartitus 2 10 4.9 N/A 3

Tree Selection and Forest Structure

The combination of unusual behavioral and morphological characteristics make the

Callitrichidae exceptional among primates (Garber 1980). Garber even compares the family to tree squirrels in their locomotor behavior, adding that their possession of claw-like nails enables to them to routinely use vertical clinging during travel by embedding the claws into the bark, rather than grasping with the entire hand as other monkeys would (1980, 1992). While some researchers have argued that this adaptation would enable tamarins to choose large vertical trunks and supports, Garber instead noted that Saguinus oedipus geoffroyi from Panama avoids sharp and vertical supports and restricts the majority of movement to small and medium-sized branches, as well as vines and overlapping foliage. The use of large trunks enables them to gain access to food resources, such as gums and small insects (Garber 1980, 1992).

Vidal and Cintra (2006) studied the forest structure and its effect on the occurrence of S. bicolor in Brazil By recording the presence of tamarins in a gridded study area and recording habitat data for each plot, such as dead logs, snags, lianas, tree size, canopy opening, etc., they hypothesized that since tamarins prefer to cling to vertical trunks to forage for food sources, areas in the study area with a higher abundance of trees would be preferred over areas with fewer trees. Vidal and Cintra (2006) concluded that the occurrence of S. bicolor was related to a lower abundance of logs, higher frequency of snags and smaller canopy opening, suggesting the species does not utilize forest resources randomly. Since there are few studies of S. midas in the wild and S. midas is closely related to S. bicolor (Rylands et al. 2016), I will compare habitat use with S. bicolor.

Study Questions

Given phylogenetic similarity, I hypothesize that S. bicolor and S. midas have very similar habitat preferences. The aim of this study is to characterize a riverine/secondary forest habitat in eastern Suriname and to assess habitat use by following S. midas social groups. I predict that S. midas will express a preference for microhabitats with high liana density, small tree size and low basal area. I will also collect data on S. midas group size and ranging patterns at this site, since very little information is available for this species and has seldom been studied in

Suriname (but see Mittermeier 1977, Colavita 2005 and Veres 2012).

Chapter II

Methods

Research Overview

Research was conducted from May to June 2012 in Berg en Dal (5° 8.731ʹ to 5° 7.810ʹ N,

55° 4.360ʹ to 55° 3.592ʹ W), in eastern Suriname. Also known as the Bergendal Ecotourism

Resort, it is located on the right bank of the Suriname River, approximately 90 km south of

Paramaribo, the capital city (Fig. 2.1). The Bergendal Resort is on the grounds of a former plantation dating to as early as 1677 (http://www.bergendalresort.com/en/history), producing first sugarcane and then was converted to a timber plantation in 1835 (Bergendal Resort, 2015).

The latter resulted in the majority of the study area being logged (clear-cut).

Presently, the forest on the grounds of Bergendal is secondary growth and at most 100 years old. The exception is some large trees (e.g., Ceiba pentandra, Malvaceae), which are considered “sacred trees” and were not cut (Norconk, personal communication 2012, Veres

2012).

The land is relatively low-lying, with the lowest land regularly flooding after rainfall. The topography does vary, with uphill areas that are considerably drier and less saturated than lowland areas that are consistently swampy. Relative elevation on the trail systems varied between 7 m to 30 m above sea level. The forest contains a variety of trees and plant life,

including Bellucia grossularioides (Melastomataceae), Attalea maripa (Arecaceae, palm) and many types of lianas.

Figure 2.1 – Bergendal Resort and surrounding forest, as seen from atop Armadillo Hill, looking south. The study area is beyond the buildings. The Suriname River is also visible in the photo. (Photo taken by Gina Cherundolo, 2012)

Survey Area

A trail system was constructed in riverine forest flanked by the Suriname River. Trail tags were placed at 25 m intervals and each of the nine trails varied in length (Fig. 2.2) for a total of approximately 2.5 km. The main trail was the R trail, which begins from the base camp and spans approximately 950 m through relatively high, dry ground that increases in elevation and terminates at a creek that branches off the Suriname River. A secondary trail, named C trail, also

begins at the base camp site (labeled “backhouse” on Fig 2.2) and flanks the creek for approximately 300 m before terminating at a creek crossing. A third major trail, named S trail, is in an swampy and oft-flooded part of the trail system that runs parallel to R and connects two R trail tributaries. The majority of the trail tributaries branch off of the R trail, eventually rejoining it farther up or down the trail. These trails have names, such as RA, RF and RP, to denote that these trails have their origin from the R trail. This main trail system will be denoted as the RSC trail system (total area approximately 38 ha) and individual trails will be identified accordingly.

Upon arrival at the site, each trail was navigated, cleared of growth and each trail marker was entered as a waypoint on a handheld GPSmap CSx unit.

Sampling Methods: Primates

On 15 days from May 21 to June 8, 2012, trails were walked continuously in search of tamarins beginning at 7 a.m., with approximately 8 hours (estimated 67.5 hours total observation time) trail walking continuing through the day, concluding with tamarins entering final sleeping sites around 4 p.m. Five individuals were usually participating in the observations [(G.

Cherundolo, M. Norconk, K. Dietz and A. Mason (from Kent State University) and J.

Humphreys (from Central Washington University)]. Thus at any given time there could be three primate groups tracked simultaneously (two groups of two people and a single person).

When a primate group was encountered, the time of the encounter and nearest trail marker were noted, a group count was made, as well as any behaviors that were observed including the direction they were traveling. If the observers had a GPS device, the “tracking function” was enabled and the group was followed as long as possible (i.e., contact was lost or could no longer be followed due to terrain or habituation issues as tamarins were not used to

being around people). Heavy rain also interrupted the ability to follow the monkeys. Tracks that began at the beginning of the initial encounter and ended when the tamarins were lost are indicated as “complete,” while tracks that were interrupted during the observation period are indicated as “incomplete.” Significant events and places, such as feeding trees, inter-troop group encounters, etc., were marked on the GPS using “waypoints.” Information was pooled among all observers to map ranging patterns behavior of the different groups of tamarins on a daily basis and to begin to formulate daily routines and patterns. A preliminary map was constructed by hand.

When observers had different counts of the same group, the average number was used

(e.g., if counts ranged from four to six, a group size of five was used for this calculation), and the minimum was used if observers had incomplete counts (e.g., if at least five were observed, I recorded “five” as group size). When separate groups merged or came into close proximity, the initial group size estimate was noted prior to the merger (e.g., if four individuals were observed and then joined by an additional six, the original observation number of four was calculated).

Unknown or ambiguous counts were not included in the calculation.

Sampling Methods: Habitat

The “micro” or “local” habitat in the vicinity of the trail mark was characterized using a focal-tree method whereby focal (largest) tree and the five closest trees formed a cluster that was assessed to determine local habitat characteristics. “Focal” trees were identified at 25 m intervals on all trails for a total of 690 trees measured in 97 “local” habitats. Each focal tree and five closest trees were measured at diameter at breast height (DBH). Elevation was recorded at the trail marker as well as the number of liana (vines) covering each of the six total trees, and the

radius from the marker tree to the farthest tree was also measured. Other information was noted, such as if the site was flooded, if the area was at a steep incline, if there were many known fruiting trees in the area, etc.

Analysis

GPS information, including tracks, trail marker waypoints and encounter locations, were entered into the computer using the programs Garmin BaseCamp and Google Earth to render the waypoints and tracking in a 3D space. Several trail markers that were deemed inaccurate due to satellite error (e.g., markers were too close together, marker was too close to the river) were manually moved to a more accurate estimated location. Encounters logged visually were placed manually on the map next to the tree marker where the encounter took place. Basal area for each tree was then calculated by using a modified formula for the area of a circle, where the basal area

(BA) equals

퐷퐵퐻 2 퐵퐴 = 휋 ( ) 2

BA units are expressed in cm2. The BA of all six trees in each cluster were then added together total to calculate total basal area (TBA) in cm2. Foresters and ecologists commonly use a set amount of acreage in a forest to use for the area, but with this method, each M1 tree was approximately 25 m apart thus providing relatively fine-grained measurements of forest density throughout the trail system.

Group size metrics were compared to other data for species within the genus Saguinus.

Observations will be compared to tree measurement data using a grid analysis and a Mann-

Whitney test similar to analysis performed by Vidal and Cintra (2006). A Kruskal-Wallis test

was used to compare average DBH, TBA, lianas and radius among all ten trails, with post-hoc analysis.

A grid system was overlaid on the trail system map to divide the study area into equal sections. This was done using the image manipulation program GIMP. The survey area size and location was locked using Google Earth so that the both entire trail system was in view and the scale would remain 200 m; this ensured the grid dimensions would remain uniform. A 50 x 50 m grid was then overlaid on the survey area as the program rendered it automatically. This was to ensure that placement of the cells was an objective process. The trail markings and encounters that ended up in each square were noted and squares were listed as either containing any encounters (Yes) or lacking an encounter (No). Any encounter overlapping into multiple cells was placed in the cell where the center of the marker or bulls-eye was located. Cells in which tamarin observations overlapped with trail markers were analyzed by comparing BA, DBH and number of lianas. When multiple trail markers occurred in the same cell, these values were averaged to provide a single value for the cell.

All statistics, including Mann-Whitney, Kruskal-Wallis and descriptive statistics were performed using SPSS v.22 and hand calculations by calculator. Maps and figures were constructed using GIMP 2.8.14 for MAC OS, Google Earth, Garmin BaseCamp, and ArcGIS

10.4. Level of significance for all statistical tests was set at p < 0.05 and all tests were two-tailed.

Chapter III

Results

Habitat Assessment

A total of 690 trees were measured on the RSC trail system (Figure 3.1): each tree bearing the trail marker (at 25 m intervals) and the five trees closest to it (a “tree cluster”). Some marker trees acted as the marker for two different trails and were not measured twice (e.g., RQ

0.0 is the same tree as R 0.5). Due to a measuring error with one marker tree, basal area (BA) for that tree group could not be calculated accurately, so that a group of six trees was omitted for all basal area calculations (N = 684). However, for calculations involving DBH and averages, the five trees in this group were included (N = 689) (Table 3.1).

Figure 3.1: Google Earth image with superimposed RSC Trail system with selected trail markers. Creek location (blue line) is approximate. The Suriname River is visible in the upper right quadrant.

Each tree’s DBH, the radius from marker tree (M1) to farthest tree (M6) and total number of lianas on all six trees in the group were measured in the field (Appendix A). Calculations were also made to determine each tree’s BA, the sum of each tree group’s BA and the total basal area

(TBA) for each group (Appendix B) and summarized in Table 3.1.

Table 3.1: Summary of Tree Measurements of “tree clusters” = “marker” trees and

nearest neighbors on all Bergendal trails. Average DBH = total diameter at breast height

values for all trees in each tree cluster, was averaged by trail; BA = average basal area

of tree clusters (in cm2); radius = distance from marker tree to farthest neighbor in

cluster (in meters); and average number of lianas per tree cluster for each trail.

Complete data is found in Appendixes A and B.

Trail N Avg. DBH (cm) Average BA (cm2) Avg. Radius (m) Avg. # Lianas C 13 17.49 ± 6.79 488.13 ± 365.76 5.6 ± 2.05 5.8 ± 2.4 R 37 12.06 ± 4.58 204.19 ± 172.51 3.6 ± 1.1 3.9 ± 2.1

RA 6 9.91 ± 3.66 109.69 ± 81.10 4.2 ± 1.1 3.5 ± 2.1 RB 4 9.70 ± 5.83 200.64 ± 194.16 2.5 ± 0.8 2.0 ± 0.8 RC 9 11.12 ± 4.12 175.64 ± 114.36 3.8 ± 1.2 6.9 ± 2.9 S 11 14.38 ± 13.24 788.59 ± 1796.18 3.1 ± 1.0 7.7 ± 4 RF 8 7.60 ± 4.4 133.23 ± 18.19 3.0 ± 0.9 6.4 ± 3.6 RE 9 10.31 ± 3.92 17.46 ± 145.89 3.0 ± 0.4 4.1 ± 1.9 RP 10 13.14 ± 7.41 342.95 ± 436.96 3.2 ± 1.1 5 ± 3.2 RQ 7 8.42 ± 1.98 87.69 ± 36.79 2.5 ± 0.7 3.9 ± 2.8

A Spearman’s rank-order correlation was run to determine the relationship among the four measurements. All correlations were positive and statistically significant. The cluster radius was correlated with number of lianas (rs(114) = 0.191, p = 0.042), with the average BA for all five trees in the cluster (rs(114) = 0.410, p < 0.0005) and with average DBH (rs(114) = 0.463, p <

0.0005). The number of lianas was correlated with the average DBH (rs(114) = 0.203, p = 0.030) and with the average BA (rs(114) = 0.228, p = 0.015).

A Kruskal-Wallis H test showed that there was a statistically significant difference in the

2 cluster radii among the ten trails (χ (9) = 28.63, p = 0.001), in the number of lianas among the ten

2 2 trails (χ (9) = 26.02, p = 0.002) and in the average DBH of the tree cluster (χ (9) = 20.40, p =

0.016). Total basal area (sum of BA for all trees in the cluster) was also statistically different

2 among trails (χ (9)= 18.50, p = 0.03).

Pairwise post-hoc analysis with Bonferroni correction revealed that there was a significant difference in tree cluster radii between the C trail and the RQ trail (p = 0.002). There was also a significant difference in the liana count of clusters of S trail and RB trail (p = 0.05).

And, there was a significant difference in total basal area (p = 0.023) and average DBH of tree clusters (p = 0.006) between C trail and RF trail.

Figure 3.2: RSC Trail System with initial tamarin encounters with 50 x 50 m grid overlay.

Orange targets indicate initial encounters that correspond with tracking data. Green targets indicate initial encounters observed visually.

Mann-Whitney tests were conducted to assess statistical significance between cells with tamarin initial encounters (N = 17) and those without (N = 34) (Figure 3.2). Cells with encounters had average basal area sums that were significantly lower than cells with no encounters (N = 51, U = 167.0, p = 0.015). Cells with encounters tended to have smaller average

DBHs than cells with no encounters, but the difference was not significant (N = 51, U = 202.0, p

= 0.082). Cells with encounters had significantly more lianas than cells with no encounters (N =

51, U = 189.5, p = 0.047).

Observations of Saguinus midas and using the GPS for tracking

We encountered monkeys 45 times on the RSC trail system between May 28 and June 8

(Appendix C); of these, we obtained group counts in 36 encounters. The average number of monkeys per encounter was 4.4 ± 2.2. Groups ranged from 2-11 individuals, including juveniles and infants, if seen, although single tamarins, (i.e., not part of a group) were also spotted five times and were included in group average data.

Partial or “complete” GPS tracks were logged during 12 encounters; 18 encounters had a precise start and end time logged either with the GPS or manually (Table 3.2). “Complete” refers to a complete overlap between tracking and duration of the follow. The remaining encounters have only a start time logged with an unclear end time.

The average observation time for the 18 encounters with a clear start and end time was

99.33 ± 67.92 minutes. This includes start and end times logged electronically on the GPS and logged manually in a notebook. The shortest logged time was 15 minutes and the longest logged time was 230 minutes.

Table 3.2: Tamarin Observations with GPS Tracking Information. Table indicates the

date of observation, the approximate location on the trail where the observation began,

the time the observation began, the time in minutes the track lasted, the average walking

speed of the observer during the track, the distance traveled during the track, and the

number of monkeys observed at the beginning of the track.

Date Approximate Start End Total Time Avg. Distance # Monkeys Location (minutes) Speed (m) 5/22 R 0.600 12:00 14:50 170 0.7 km/h 1600* 5 5/23 R 0.525 8:30 12:10 220 0.5 km/h 747 * >7 5/27 Backhouse 7:30 11:20 230 1 km/h 1000 6 5/28 R 0.375 7:54 11:08 194 N/A 1200 8 5/28 Backhouse 14:00 16:15 135 0.2 km/h 220 * 5 5/29 R 0.525 8:30 10:30 120 N/A 895 11 5/29 R 0.550 10:48 11:03 15 0.4 km/h 124 1 5/31 R 0.100 7:45 8:30 45 N/A N/A 5 5/31 R 0.600 7:50 8:50 60 N/A N/A >4 6/1 R 0.175 7:30 8:21 51 N/A N/A >5 6/1 RQ 9:08 10:30 82 N/A N/A 9 6/2 C 0.0 7:45 8:30 45 N/A N/A 5 6/2 R 0.500 8:30 10:40 130 N/A N/A 5 6/3 RF 0.050 8:39 9:30 49 N/A N/A 4 6/4 R 0.600 8:34 N/A N/A N/A 785 2 6/5 Backhouse 6:53 7:53 60 N/A 299 5 6/7 RP 0.150 7:49 8:22 33 N/A 996 ? 6/8 R 0.500 8:36 9:10 34 0.3 km/h 184 ?

*Distance not true total, track not complete.

Figure 3.3: RSC Trail System with white dots indicating trail marks. Creek location (blue line) is approximate. Each GPS Track collected during a monkey follows is identified by a separate color.

Home range size and tamarin group counts at Bergendal

It is likely that at least four groups of tamarins had home ranges that intersected with the

RSC trail system. This estimation of four groups was determined through observation of group counts, track information (Figure 3.3), distance traveled, direction traveled and encounter locations. Average group size for all four groups ranged from 4.2 to 4.8 individuals per encounter (Table 3.3).

GPS trackings for each group were collated and minimum convex polygons (MCP) were used to assess home range size for each group of tamarins in the survey area (Figure 3.4,). Home range sizes ranged from 0.53 to 10.0 ha, with an average of 4.38 ha. It is notable that Group A’s and Group B’s home ranges are much smaller than the other two groups. Group A’s home range extended into a swampy area that was difficult to access off trail on foot, thus limiting GPS track information for these tamarins; thus their home range is likely to have been underestimated.

Likewise, Group B was observed to cross the creek to the northeast daily and became inaccessible after that point. Therefore, home range data for these two groups is skewed. It is also notable that the overlap point for Group C and Group D was a significant feeding tree patch and the site of several inter-troop aggressive encounters.

Table 3.3: Home range and group size calculations for all four groups in survey area.

For each group (A, B, C, D), estimate home range is calculated in ha, and estimated

average group size is calculated for all encounters associated with each group.

Group Estimated Estimated Home Range (ha) Avg. Group Size A 1.40 4.8 B 0.53 4.2 C 5.60 4.3 D 10.0 4.3

Figure 3.4: Home range assessment for tamarins in survey area (c. 38 ha). Polygons connecting tracks of each encounter were overlaid in Google Earth and identified by color, with group labels A, B, C and D. Creek location (blue line) is approximate.

In the RSC trail system, where several of the trails were statistically significant from one another in local characteristics (i.e., average tree cluster radius, DBH, basal area and lianas), the tamarins were more likely to be encountered for the first time in areas with lower average basal areas and more than an average number of lianas. I hypothesize, although more data collection is needed, that tamarins are also more likely to be encountered for the first time in areas with more space between trees and larger canopy opening.

Chapter IV

Discussion and Summary

Group number and size

Despite the short duration of the study, the average number of monkeys per encounter of

4.4 ± 2.2 corresponds to S. midas group data for the species (Porter et al. 2017) and for the genus

Saguinus. The assessment of four groups at Bergendal is also consistent with Veres’ (2012) survey of the same site. The average number of tamarins for this study could have skewed lower due to solitary encounters.

Min. Avg. Max. SD Source 6 6 2 Day and Elwood 1999 3 7 Kessler 1995 3 5.7 8 Norconk et al. 2003 3 12 Rylands and Mittermeier 2 5.8 12 Wolfheim 1983* 2 5.74 13 2.8 Veres2013 2012 1 4.4 11 2.2 This Study *secondary source

Table 4.1: Saguinus midas group size comparisons (minimum, average, maximum, standard deviation.)

Based on encounter data, GPS tracking, and daytime trail walking observations, it is likely there were four groups of tamarins utilizing the RSC trail system area. One group, “Group

A” mainly congregated near the backhouse and was found on the RA, RB and into the C trails.

This group often ranged into a swampy area that was difficult to access when they left the trail.

A second group (“Group B”) was also seen at the backhouse, but GPS tracks for these tamarins could not be logged because this group crossed a creek behind the backhouse daily, although this group was observed crossing this creek and returning. This resulted in much smaller home ranges than expected for both groups. Given the duration of the study, these should be considered minimum home range sizes.

The remaining two groups were found frequently at the same time of day at a patch of

Bellucia trees near the intersections of the R, RE and RQ trails. I observed these groups fusing into larger groups near these trees, as well as several inter-troop encounters, likely instigated over territory at the feeding site. One of these groups primarily flanked the river along R trail in both directions (“Group C”), while the other group (“Group D”) primarily stayed down trail.

However, a larger amount of group fission/fusion must have occurred in the study area, due to the number of singleton encounters and duos observed. Due to GPS tracker device failure, more

GPS tracks could not be measured after June 8, so it is possible further accurate GPS data would reflect more accurate home range and group data.

Tree Preference

The encounter and tree measurement comparison results suggest that S. midas prefers microhabitats with higher liana density, small tree size and low total basal area during daily, non- sleeping site activities. This preference corresponds with Vidal and Cintra’s (2006) assessment of S. bicolor. This behavior contrasts with sleeping site selection (Day and Elwood 1999), where trees have a larger DBH, stature and crown diameter than non-sleeping trees. The use of taller

trees with larger tree crowns for sleeping may reduce the chances of detection overhead from predators such as raptors (Day and Elwood 1999). During the daytime, however, the preference for patches of smaller trees is more common for feeding (Vidal and Cintra 2006) and traveling.

Their small body size and low weight allow both S. bicolor and S. midas to reach smaller, lower branches for food sources than a larger primate would be able to reach; larger primates did overlap with S. midas in the study area as Alouatta seniculus was observed on one occasion and heard on many mornings. However, prolonged exposure in open areas puts both species at greater risk of predation in the daytime (Vidal and Cintra 2006), particularly from hawks and eagles. Additional tree measurement data, including crown diameter and tree height, could supplement these findings. More data on tamarin encounters, including more GPS tracks, or even tagging, could give a more complete picture of the home range and group patterns at Bergendal.

A further assessment on feeding site locations in the study area could also highlight the tamarins’ preference for prolonged exposure in more open areas of the RSC trail system. Further study of predation rates and time spent in these more open microhabitats could shed more light on the risks associated with these habitats.

The study of microhabitat use and forest structure preference could be invaluable for future research in conservation efforts for members of Callitrichidae and for tropical flora and fauna in general. As tropical forests are continuing to be disturbed and reduced around the world, primates are encountering fragmented, secondary forest patches. S. midas’ role as a pioneer species and hardiness in marginalized habitats make them an ideal research candidate, as they can be common in areas where larger-bodied animals can no longer survive. Researchers are studying the capability of primates to adapt to these changes in environment (e.g., Kulp and

Heymann 2015), and their role in helping these areas regenerate (Oliveira and Ferrari 2000).

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Appendix A: All Tree and Liana Measurements

Marker Elevation M1 DBH M2 DBH M3 DBH M4 DBH M5 DBH M6 DBH Radius Lianas (m) (cm) (cm) (cm) (cm) (cm) (cm) from M1 to M6 (m) C0.0 23 25.7 24.7 6.1 3.5 23 15 7.1 2 C0.025 25 11.4 43.6 4.7 3.7 49.6 28.4 8.3 4 C0.050 24 11 54.9 23 17.4 14.2 20.3 9.5 8 C0.075 27 8.8 31.1 82 3.8 11.7 36.3 4.1 8 C0.1 29 3.8 9.8 63.5 63.3 12.7 8.3 5.7 7 C0.125 30 3.8 3.1 29.2 3 32.3 6.2 3.96 2 C0.150 30 2.3 22.9 10.8 56.4 11.8 5.9 7.3 9 C0.175 30 7.2 10.6 14 5.3 11.4 16.5 5.34 5 C0.2 30 3 5.4 8.8 8.2 6.7 16.3 4.2 8 C0.225 30 4.5 8.3 8.9 63.5 3.4 28 3 5 C0.250 31 3.3 34.9 2.7 3.1 4.1 4.4 3.3 5 C0.275 31 8.4 9 8.4 4.4 59.8 13.7 4.17 8 C0.3 30 13.3 27.6 4.2 6.2 17.4 6.2 6.83 4 R0.0 19 14.7 18.5 13 3 3 2 3.37 2 R0.050 19 2.4 9 9.2 22.5 3.5 12.8 4.92 2 R0.075 19 15.8 5.6 6.9 12.9 45.3 7.1 4.67 4 R0.1 19 6.9 19.4 34.5 9 26.4 5.5 4.7 6 R0.125 19 3.4 30.2 9.5 11 2.5 10 4.44 5 R0.150 20 46.2 5.9 3.5 11.9 4 1.5 5.5 2 R0.175 21 6.7 2.2 53.1 4.6 21.1 2.2 3.35 5 R0.2 22 12.8 19.5 36.2 3.5 3.2 23 2.7 4 R0.225 23 9.8 14.7 15 7.7 45.1 3 3.7 6 R0.250 23 9.1 12.2 4 4.4 2.2 1.2 2.12 2 R0.275 23 7.8 9.6 19.8 11.7 24.7 11.7 3.6 2 R0.3 23 18 16 11.3 9.5 10.8 8.8 5.85 2 R0.325 27 20.7 24.9 49.2 40.7 19.7 11.7 3.04 3 R0.350 29 10.8 10.4 19.4 12.5 7.6 4 4.29 6 R0.375 29 13 9.3 3.1 13.5 1.4 5.4 4.33 3 R0.4 28 15 11.1 2.7 18.3 12 10 3.4 6 R0.425 28 11.4 3.3 19.7 7 5.2 9.2 4.29 5 R0.450 27 31.9 3.7 12.2 5.3 4.9 1.9 3.87 0 R0.475 26 6.2 2.2 15.4 2.4 9 3.1 2.7 0 R0.5 25 5.5 5.2 7 10.1 5.8 22.9 2.37 3 R0.525 24 5.2 4 7.8 13.9 11.5 2.8 3.77 1 R0.550 25 10.4 5 20.1 4 23.2 7.5 4.09 3 R0.575 29 16.8 5.8 3.3 4.6 18 5 3.6 4 R0.6 31 7.5 21.4 4.4 3.5 9 17.1 1.56 5

R0.625 29 13.7 8.4 2.8 7.1 3.2 14.3 2.1 4 R0.650 27 13.4 3 10.4 4.7 4.5 6.1 2.34 3 R0.675 23 26 1.9 31 19.1 5.2 5.3 4.26 1 R0.7 22 9.3 5 8 14.8 21.5 25 3.16 2 R0.725 19 19.1 17.5 20.7 2 7.3 23.8 1.77 3 R0.750 15 5.3 10.4 6.5 11 15 39.1 5 4 R0.775 4 10.5 3.8 6.4 11 18.7 7.5 4.07 7 R0.8 4 62.3 15 3 2.7 4.2 5.2 3.3 4 R0.825 5 12.7 11.8 3.4 11.8 2.7 2.5 2.84 5 R0.850 5 8 10.5 1.6 10.5 6.2 6.4 4.9 8 R0.875 6 28.5 23.1 17.2 28 5.3 6.8 2.73 7 R0.9 5 23.3 2.6 5.8 10.9 2.9 3 3.1 5 R0.925 5 21.2 62.1 4.1 9.6 2.6 26.6 5.18 9 RA0.0 19 2.4 9 9.2 22.5 3.5 12.8 4.92 2 RA0.025 20 14.9 28 23 10 5.9 18.3 3.7 1 RA0.050 21 10.3 9.2 3.4 4.7 5 5 5.01 3 RA0.075 21 8.8 20.4 6.1 17.4 2.2 6.1 3.09 3 RA0.1 21 10.8 10.8 11.1 6 2.2 2.1 5.75 6 RA0.125 20 13.5 16 8 6.8 4 7.3 3 6 RB0.0 19 5.9 26.5 8.3 22.3 36.6 4.7 3.64 3 RB0.025 19 1.1 11 1.8 1 1.5 13.9 2.6 2 RB0.02 18 2 2 54.1 2.3 2.2 3.8 2.17 2 RBend 15 6.1 3.9 3.8 8 3.6 6.4 1.7 1 RC0.0 15 9.5 5.9 2.8 11 8.9 4.5 3.07 5 RC0.025 13 7 22.4 2.4 2.2 1.5 3 3.83 9 RC0.050 12 16 20 43 11 15.6 17.6 4 11 RC0.075 19 7.9 2.8 4 3.5 41 2.4 2.4 8 RC0.1 20 17.1 6.2 9.3 24.4 4.4 9.3 2.93 3 RC0.125 23 28.8 5.2 4 2.1 20 6.8 3.19 3 RC0.150 24 10.5 11.9 5.2 7.9 11.8 28.4 6.4 9 RC0.175 26 11.1 16.4 5.2 5.4 6.5 8.5 3.97 5 RC0.2 26 5.4 7.5 5.4 4.6 3.3 41.8 4.18 9 S0.3 27 23 9 8.5 4.9 5.5 1.4 3.46 6 S0.325 26 3.5 2.9 11 5.2 2.1 1.6 2.16 8 S0.35 25 10.6 4 23.5 22.5 11.9 213.5 4.37 13 S0.375 26 7.4 2.7 9.3 3.7 2.8 1 1.61 5 S0.4 16 58 10.9 17 25.9 13.3 18.4 4.25 10 S0.425 25 7.9 12.4 1.2 2.4 1.8 2 3 3 S0.45 26 53.1 4 2.8 15.9 3.4 14.3 3.58 3 S0.475 26 64.2 5.6 7.1 1.3 11.9 38.6 3.47 8 S0.5 25 26 1.5 3.9 19.2 3 47.3 4.01 10 S0.525 23 7.5 7.7 1.4 2.1 1 1.5 1.5 4

S0.55 23 19.3 6.1 7.8 4.5 2 2.5 3.23 15 RF0.0 34 9.4 3.9 1.8 6.9 7.2 2.9 2.78 4 RF0.025 32 6.5 4.1 2 8.8 7.2 1.8 2.61 5 RF0.05 29 8.8 5.5 3.2 2 2.1 3.2 1.69 6 RF0.075 26 7 8.8 6.3 60.3 3.2 1.9 4.25 5 RF0.1 27 4.5 8.7 3.8 1.4 1.9 1.8 2.5 6 RF0.125 27 19.4 2.9 4.2 5.7 47.5 6.8 2.6 6 RF0.150 23 12.2 15.2 5.1 3.2 1.5 1.9 4.44 4 RF0.160 23 19.3 6.1 7.8 4.5 2 2.5 3.23 15 RE0.0 14 23.2 7.6 7.3 2.6 5 11.5 3.02 4 RE0.025 13 9.3 3.5 7.3 4 3.3 6 4.15 5 RE0.05 13 27 4.8 2.5 28.6 3.7 19.2 3.36 4 RE0.075 13 8 5.5 3.7 3.5 17.1 3.1 3.13 4 RE0.1 13 8.1 2 2.4 10 4.3 1.3 2.75 3 RE0.125 12 18.5 3.5 5.6 6.5 22 9.5 3.24 5 RE0.15 10 49.7 3.5 8.2 10.6 4.3 3.8 3.3 8 RE0.175 9 20.5 5.3 8.7 14.5 20.9 1.8 3.6 1 RE0.186 9 53.1 4 2.8 15.9 3.4 14.3 3.58 3 RP0.0 12 7.7 6.2 11.4 22.2 53.7 32.2 4.5 6 RP0.025 11 6.9 3.2 31.4 1.9 10.6 2.5 2.23 3 RP0.05 7 6.4 7.5 9.4 4.9 12.5 3.8 4.17 0 RP0.075 11 14.7 99.5 4 6.7 8.2 1.2 3.6 8 RP0.1 13 34.8 4.9 4.6 2.9 5.7 1.4 2.35 6 RP0.125 14 19.3 9 8.4 5.3 5.1 4.9 3.2 8 RP0.150 14 7.5 2.4 16.5 4 15.1 6.3 1.45 0 RP0.175 13 9.9 13 2.7 8.9 11.2 13.2 2.05 6 RP0.2 8 18.4 23.3 22.4 64 25.4 5.3 4.8 9 RP0.225 12 16.9 4.7 5.7 3.8 8.1 4.6 3.19 4 RQ0.0 14 7.5 2.4 16.5 4 15.1 6.3 1.45 0 RQ0.025 14 11.8 9.8 3.6 29.5 3.5 4.4 1.58 6 RQ0.05 14 4.7 7.3 2.2 15.2 3.1 2.7 3.3 4 RQ0.075 14 5.2 13 20.9 14.3 1.6 2.65 1 RQ0.1 12 10 1.2 5.4 3.5 8 12.5 3.28 8 RQ0.125 14 4.6 12.5 10.4 9.4 3.2 1 2.83 5 RQ0.150 25? 5.5 5.2 7 10.1 5.8 22.9 2.37 3

Appendix B: Basal Area Calculations

Marker M1 BA M2BA M3BA M4BA M5BA M6BA Sum of Average (cm2) (cm2) (cm2) (cm2) (cm2) (cm2) BAs (cm2) BA (cm2) C0.0 518.748 479.164 29.225 9.621 415.476 176.715 518.748 271.491 C0.025 102.070 1493.010 17.349 10.752 1932.205 633.471 102.070 698.143 C0.050 95.033 2367.198 415.476 237.787 158.368 323.655 95.033 599.586 C0.075 60.821 759.645 5281.017 11.341 107.513 1034.911 60.821 1209.208 C0.1 11.341 75.430 3166.922 3147.004 126.677 54.106 11.341 1096.913 C0.125 11.341 7.548 669.662 7.069 819.398 30.191 11.341 257.535 C0.150 4.155 411.871 91.609 2498.320 109.359 27.340 4.155 523.775 C0.175 40.715 88.247 153.938 22.062 102.070 213.825 40.715 103.476 C0.2 7.069 22.902 60.821 52.810 35.257 208.672 7.069 64.589 C0.225 15.904 54.106 62.211 3166.922 9.079 615.752 15.904 653.996 C0.250 8.553 956.623 5.726 7.548 13.203 15.205 8.553 167.809 C0.275 55.418 63.617 55.418 15.205 2808.615 147.411 55.418 524.281 C0.3 138.929 598.285 13.854 30.191 237.787 30.191 138.929 174.873 Average 2928.773 514.234

R0.0 169.717 268.803 132.732 7.069 7.069 3.142 588.530 98.088 R0.050 4.524 63.617 66.476 397.608 9.621 128.680 670.526 111.754 R0.075 196.067 24.630 37.393 130.698 1611.708 39.592 2040.087 340.015 R0.1 37.393 295.592 934.820 63.617 547.391 23.758 1902.572 317.095 R0.125 9.079 716.315 70.882 95.033 4.909 78.540 974.758 162.460 R0.150 1676.385 27.340 9.621 111.220 12.566 1.767 1838.900 306.483 R0.175 35.257 3.801 2214.517 16.619 349.667 3.801 2623.662 437.277 R0.2 128.680 298.648 1029.217 9.621 8.042 415.476 1889.684 314.947 R0.225 75.430 169.717 176.715 46.566 1597.508 7.069 2073.003 345.501 R0.250 65.039 116.899 12.566 15.205 3.801 1.131 214.641 35.774 R0.275 47.784 72.382 307.907 107.513 479.164 107.513 1122.263 187.044 R0.3 254.469 201.062 100.287 70.882 91.609 60.821 779.131 129.855 R0.325 336.535 486.955 1901.166 1301.004 304.805 107.513 4437.979 739.663 R0.350 91.609 84.949 295.592 122.718 45.365 12.566 652.799 108.800 R0.375 132.732 67.929 7.548 143.139 1.539 22.902 375.789 62.632 R0.4 176.715 96.769 5.726 263.022 113.097 78.540 733.868 122.311 R0.425 102.070 8.553 304.805 38.485 21.237 66.476 541.626 90.271 R0.450 799.229 10.752 116.899 22.062 18.857 2.835 970.634 161.772 R0.475 30.191 3.801 186.265 4.524 63.617 7.548 295.946 49.324 R0.5 23.758 21.237 38.485 80.118 26.421 411.871 601.890 100.315 R0.525 21.237 12.566 47.784 151.747 103.869 6.158 343.360 57.227 R0.550 84.949 19.635 317.309 12.566 422.733 44.179 901.370 150.228 R0.575 221.671 26.421 8.553 16.619 254.469 19.635 547.368 91.228

R0.6 44.179 359.681 15.205 9.621 63.617 229.658 721.962 120.327 R0.625 147.411 55.418 6.158 39.592 8.042 160.606 417.227 69.538 R0.650 141.026 7.069 84.949 17.349 15.904 29.225 295.522 49.254 R0.675 530.929 2.835 754.768 286.521 21.237 22.062 1618.352 269.725 R0.7 67.929 19.635 50.265 172.034 363.050 490.874 1163.787 193.965 R0.725 286.521 240.528 336.535 3.142 41.854 444.881 1353.461 225.577 R0.750 22.062 84.949 33.183 95.033 176.715 1200.725 1612.666 268.778 R0.775 86.590 11.341 32.170 95.033 274.646 44.179 543.959 90.660 R0.8 3048.358 176.715 7.069 5.726 13.854 21.237 3272.958 545.493 R0.825 126.677 109.359 9.079 109.359 5.726 4.909 365.108 60.851 R0.850 50.265 86.590 2.011 86.590 30.191 32.170 287.817 47.970 R0.875 637.940 419.096 232.352 615.752 22.062 36.317 1963.519 327.253 R0.9 426.385 5.309 26.421 93.313 6.605 7.069 565.102 94.184 R0.925 352.989 3028.817 13.203 72.382 5.309 555.716 4028.417 671.403 Average 1225.142 204.190

RA0.0 4.524 63.617 66.476 397.608 9.621 128.680 670.526 111.754 RA0.025 174.366 615.752 415.476 78.540 27.340 263.022 1574.496 262.416 RA0.050 83.323 66.476 9.079 17.349 19.635 19.635 215.498 35.916 RA0.075 60.821 326.851 29.225 237.787 3.801 29.225 687.710 114.618 RA0.1 91.609 91.609 96.769 28.274 3.801 3.464 315.526 52.588 RA0.125 143.139 201.062 50.265 36.317 12.566 41.854 485.203 80.867 Average 658.160 109.693

RB0.0 27.340 551.546 54.106 390.571 1052.088 17.349 2093.000 348.833 RB0.025 0.950 95.033 2.545 0.785 1.767 151.747 252.828 42.138 RB0.02 3.142 3.142 2298.711 4.155 3.801 11.341 2324.292 387.382 RBend 29.225 11.946 11.341 50.265 10.179 32.170 145.126 24.188 Average 1203.811 200.635

RC0.0 70.882 27.340 6.158 95.033 62.211 15.904 277.528 46.255 RC0.025 38.485 394.081 4.524 3.801 1.767 7.069 449.727 74.954 RC0.050 201.062 314.159 1452.201 95.033 191.134 243.285 2496.875 416.146 RC0.075 49.017 6.158 12.566 9.621 1320.254 4.524 1402.140 233.690 RC0.1 229.658 30.191 67.929 467.595 15.205 67.929 878.507 146.418 RC0.125 651.441 21.237 12.566 3.464 314.159 36.317 1039.184 173.197 RC0.150 86.590 111.220 21.237 49.017 109.359 633.471 1010.894 168.482 RC0.175 96.769 211.241 21.237 22.902 33.183 56.745 442.077 73.680 RC0.2 22.902 44.179 22.902 16.619 8.553 1372.279 1487.434 247.906 Average 1053.8181 175.636

S0.3 415.476 63.617 56.745 18.857 23.758 1.539 579.993 96.665

S0.325 9.621 6.605 95.033 21.237 3.464 2.011 137.971 22.995 S0.35 88.247 12.566 433.736 397.608 111.220 35800.215 36843.593 6140.599 S0.375 43.008 5.726 67.929 10.752 6.158 0.785 134.358 22.393 S0.4 2642.079 93.313 226.980 526.853 138.929 265.904 3894.059 649.010 S0.425 49.017 120.763 1.131 4.524 2.545 3.142 181.121 30.187 S0.45 2214.517 12.566 6.158 198.557 9.079 160.606 2601.482 433.580 S0.475 3237.128 24.630 39.592 1.327 111.220 1170.212 4584.110 764.018 S0.5 530.929 1.767 11.946 289.529 7.069 1757.163 2598.403 433.067 S0.525 44.179 46.566 1.539 3.464 0.785 1.767 98.300 16.383 S0.55 292.553 29.225 47.784 15.904 3.142 4.909 393.516 65.586 Average 4731.537 788.589

RF0.0 69.398 11.946 2.545 37.393 40.715 6.605 168.601 28.100 RF0.025 33.183 13.203 3.142 60.821 40.715 2.545 153.608 25.601 RF0.05 60.821 23.758 8.042 3.142 3.464 8.042 107.270 17.878 RF0.075 38.485 60.821 31.172 2855.778 8.042 2.835 2997.134 499.522 RF0.1 15.904 59.447 11.341 1.539 2.835 2.545 93.612 15.602 RF0.125 295.592 6.605 13.854 25.518 1772.055 36.317 2149.941 358.324 RF0.150 116.899 181.458 20.428 8.042 1.767 2.835 331.430 55.238 RF0.160 292.553 29.225 47.784 15.904 3.142 4.909 393.516 65.586 Average 799.389 133.232

RE0.0 422.733 45.365 41.854 5.309 19.635 103.869 638.764 106.461 RE0.025 67.929 9.621 41.854 12.566 8.553 28.274 168.798 28.133 RE0.05 572.555 18.096 4.909 642.424 10.752 289.529 1538.265 256.378 RE0.075 50.265 23.758 10.752 9.621 229.658 7.548 331.603 55.267 RE0.1 51.530 3.142 4.524 78.540 14.522 1.327 153.585 25.597 RE0.125 268.803 9.621 24.630 33.183 380.133 70.882 787.252 131.209 RE0.15 1940.004 9.621 52.810 88.247 14.522 11.341 2116.546 352.758 RE0.175 330.064 22.062 59.447 165.130 343.070 2.545 922.317 153.719 RE0.186 2214.517 12.566 6.158 198.557 9.079 160.606 2601.482 433.580 Average 1028.735 171.456

RP0.0 46.566 30.191 102.070 387.076 2264.845 814.332 3645.080 607.513 RP0.025 37.393 8.042 774.371 2.835 88.247 4.909 915.798 152.633 RP0.05 32.170 44.179 69.398 18.857 122.718 11.341 298.663 49.777 RP0.075 169.717 7775.638 12.566 35.257 52.810 1.131 8047.119 1341.186 RP0.1 951.149 18.857 16.619 6.605 25.518 1.539 1020.287 170.048 RP0.125 292.553 63.617 55.418 22.062 20.428 18.857 472.935 78.823 RP0.150 44.179 4.524 213.825 12.566 179.079 31.172 485.345 80.891 RP0.175 76.977 132.732 5.726 62.211 98.520 136.848 513.014 85.502 RP0.2 265.904 426.385 394.081 3216.991 506.707 22.062 4832.131 805.355

RP0.225 224.318 17.349 25.518 11.341 51.530 16.619 346.675 57.779 Average 2057.705 342.951

RQ0.0 44.179 4.524 213.825 12.566 179.079 31.172 485.345 80.891 RQ0.025 109.359 75.430 10.179 683.493 9.621 15.205 903.286 150.548 RQ0.05 17.349 41.854 3.801 181.458 7.548 5.726 257.736 42.956 RQ0.075 21.237 132.732 343.070 0.000 160.606 2.011 659.656 109.943 RQ0.1 78.540 1.131 22.902 9.621 50.265 122.718 285.178 78.736 RQ0.125 16.619 122.718 84.949 69.398 8.042 0.785 302.512 50.419 RQ0.150 23.758 21.237 38.485 80.118 26.421 411.871 601.890 100.315 Average 499.372 87.687

Appendix C: All Tamarin Encounters

Date Approximate Start End Total Time # Monkeys Location (minutes) 5/19 backhouse 15:30 >4 5/21 R 0.675 9:44 1 5/21 R 0.225 13:40 >3 5/22 R 0.600 12:00 14:50 170 5 5/22 C 0.175 17:00 4 5/23 R 0.525 8:30 12:10 220 >7 5/23 backhouse 12:30 4 5/27 backhouse 7:30 11:20 230 6 5/27 R 0.100 8:40 1 5/27 R 0.575 13:20 1 to 3 5/28 R 0.375 7:54 11:08 194 8 to 4 5/28 R 0.550 10:45 1 5/28 backhouse 14:00 16:15 135 5 5/28 backhouse 16:28 3 5/29 R 0.525 8:30 10:30 120 4, 10 to 11 5/29 R 0.550 10:48 11:03 15 1 5/29 R 0.025 14:00 >5 5/31 R 0.100 7:45 8:30 45 5 5/31 R 0.600 7:50 8:50 60 >4 5/31 R 0.575 10:30 6 5/31 R 0.250 10:50 12:45 115 5 6/1 R 0.175 7:30 8:21 51 >5 6/1 backhouse 7:35 ? 6/1 RQ 9:08 10:30 82 9 6/1 R 0.475 11:02 ? 6/1 R 0.675 12:30 ? 6/2 C 0.0 7:45 8:30 45 5 6/2 R 0.500 8:30 10:40 130 5 6/2 R 0.475 10:30 ? 6/3 backhouse 6:45 4 to 6 6/3 R 0.175 7:30 1 6/3 R 0.675 7:50 5 6/3 R 0.675 8:18 5 plus 3 6/3 RF 0.050 8:39 9:30 49 4 6/3 R 0.500 9:51 ? 6/3 R 0.700 9:53 ? 6/3 R 0.625 14:00 7 6/4 R 0.500 8:26 3 6/4 R 0.600 8:34 2

6/5 backhouse 6:53 7:53 60 5 6/5 R 0.625 8:00 4 6/5 R 0.575 10:00 >3 6/7 RP 0.150 7:49 8:22 33 ? 6/7 RBEND 11:15 ? 6/8 R 0.500 8:36 9:10 34 ?