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Structure of a Lowland Neotropical Galliform Bird Guild

Permanent Link: http://ufdc.ufl.edu/UFE0021564/00001

Material Information

Title: Structure of a Lowland Neotropical Galliform Bird Guild
Physical Description: 1 online resource (64 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: galliform, guild, neotropical
Wildlife Ecology and Conservation -- Dissertations, Academic -- UF
Genre: Wildlife Ecology and Conservation thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: This study (1) examined guild structure, (2) compared guild characteristics to predictions from potentially relevant hypotheses of guild structure regulation, and (3) assessed the impacts of hunting on a guild comprised of the Great Curassow (Crax rubra), Crested Guan (Penelope purpurascens), Plain Chachalaca (Ortalis vetula), Ocellated Turkey (Meleagris ocellata), and the Great Tinamou (Tinamus major) in 2000-2002. The study area, located in the Maya Biosphere Reserve in northern Guatemala, included a national park unit with protected populations of the guild and an adjacent community concession unit subject to a range of subsistence hunting pressure. Diet niches and overlap were described from analyses of upper digestive tract contents from 267 C. rubra, 181 M. ocellata, 205 O. vetula, 142 P. purpurascens, and 55 T. major collected over a 24-month period. Descriptions of habitat niches and overlap, species abundance, and selected reproductive parameters were derived from line-transect sampling during a 30-month period in the park unit (1,770 km total) and over a 12-month period in the concession unit (995 km total). Descriptions of nest-sites, clutch-sizes, and other reproductive characteristics were based on observations made at 24 C. rubra nests, 39 M. ocellata nests, 77 O. vetula nests, 19 P. purpurascens nests, and 66 T. major nests. Dry-mass proportions of grit were highest and similar in the diets of M. ocellata (22%) and C. rubra (20%), lowest and similar in O. vetula (2.7%) and T. major (2.2%), and intermediate in P. purpurascens (9%). Seeds and pulp represented the greatest and second-greatest proportions respectively, of the diet of all species when grit and snail shells were excluded from analyses. Overlap in the seed component of the diet ranged from 50-86% for eight of the ten species-pairs, and the average combined overlap exceeded 50% for all species except O. vetula. Habitat preferences were exhibited by C. rubra and P. purpurascens for tall-forest, and by O. vetula for low-forest. T. major occurred at a greater frequency in low-forest in the two areas with the least hunting pressure, and in high-forest in the two areas with the greatest hunting pressure. The frequency of occurrence of M. ocellata did not differ among habitat types. All species exhibited consistent vertical patterns of strata occupancy. Guild characteristics were semi-consistent with structural regulation through competitive interactions with respect to habitat niches, body-size assortment, and evidence of competitive release, but were inconsistent with respect to diet niches and null-pool comparisons. The variability of habitat occupancy and abundance within the guild were consistent with expectations for non-equilibrium processes. Vertical stratification of nest-placement and other reproductive attributes of the guild were consistent with expectations for a guild subject to intense predation pressure. The guild appears to be a terrestrial vertebrate example of the non-equilibrium competition model of community structure proposed by Connell (1980). All species were most abundant in the fully-protected area; however, the abundance of C. rubra, M. ocellata, and P. purpurascens declined consistently along the increasing gradient of hunting pressure. Guild biomass in the most heavily hunted area represented 30% of the guild biomass in the protected area.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Labisky, Ronald F.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0021564:00001

Permanent Link: http://ufdc.ufl.edu/UFE0021564/00001

Material Information

Title: Structure of a Lowland Neotropical Galliform Bird Guild
Physical Description: 1 online resource (64 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: galliform, guild, neotropical
Wildlife Ecology and Conservation -- Dissertations, Academic -- UF
Genre: Wildlife Ecology and Conservation thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: This study (1) examined guild structure, (2) compared guild characteristics to predictions from potentially relevant hypotheses of guild structure regulation, and (3) assessed the impacts of hunting on a guild comprised of the Great Curassow (Crax rubra), Crested Guan (Penelope purpurascens), Plain Chachalaca (Ortalis vetula), Ocellated Turkey (Meleagris ocellata), and the Great Tinamou (Tinamus major) in 2000-2002. The study area, located in the Maya Biosphere Reserve in northern Guatemala, included a national park unit with protected populations of the guild and an adjacent community concession unit subject to a range of subsistence hunting pressure. Diet niches and overlap were described from analyses of upper digestive tract contents from 267 C. rubra, 181 M. ocellata, 205 O. vetula, 142 P. purpurascens, and 55 T. major collected over a 24-month period. Descriptions of habitat niches and overlap, species abundance, and selected reproductive parameters were derived from line-transect sampling during a 30-month period in the park unit (1,770 km total) and over a 12-month period in the concession unit (995 km total). Descriptions of nest-sites, clutch-sizes, and other reproductive characteristics were based on observations made at 24 C. rubra nests, 39 M. ocellata nests, 77 O. vetula nests, 19 P. purpurascens nests, and 66 T. major nests. Dry-mass proportions of grit were highest and similar in the diets of M. ocellata (22%) and C. rubra (20%), lowest and similar in O. vetula (2.7%) and T. major (2.2%), and intermediate in P. purpurascens (9%). Seeds and pulp represented the greatest and second-greatest proportions respectively, of the diet of all species when grit and snail shells were excluded from analyses. Overlap in the seed component of the diet ranged from 50-86% for eight of the ten species-pairs, and the average combined overlap exceeded 50% for all species except O. vetula. Habitat preferences were exhibited by C. rubra and P. purpurascens for tall-forest, and by O. vetula for low-forest. T. major occurred at a greater frequency in low-forest in the two areas with the least hunting pressure, and in high-forest in the two areas with the greatest hunting pressure. The frequency of occurrence of M. ocellata did not differ among habitat types. All species exhibited consistent vertical patterns of strata occupancy. Guild characteristics were semi-consistent with structural regulation through competitive interactions with respect to habitat niches, body-size assortment, and evidence of competitive release, but were inconsistent with respect to diet niches and null-pool comparisons. The variability of habitat occupancy and abundance within the guild were consistent with expectations for non-equilibrium processes. Vertical stratification of nest-placement and other reproductive attributes of the guild were consistent with expectations for a guild subject to intense predation pressure. The guild appears to be a terrestrial vertebrate example of the non-equilibrium competition model of community structure proposed by Connell (1980). All species were most abundant in the fully-protected area; however, the abundance of C. rubra, M. ocellata, and P. purpurascens declined consistently along the increasing gradient of hunting pressure. Guild biomass in the most heavily hunted area represented 30% of the guild biomass in the protected area.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Labisky, Ronald F.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0021564:00001


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STRUCTURE OF A LOWLAND NEOTROPICAL GALLIFORM BIRD GUILD


By

ERICK HOGAN BAUR

















A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2008

































2008 Erick Hogan Baur




























To my family, Michaelyn, Milano, and Chloe Lilyana









ACKNOWLEDGMENTS

I thank my committee chair Ronald F. Labisky, who set the gold-standard for patience, and

also committee members David Steadman, George Tanner, and especially Daniel Brooks for his

unfailing enthusiasm. The Department of Wildlife Ecology and Conservation of the University

of Florida, the Wildlife Conservation Society, and the African Safari Club provided financial

support for this research. I thank Roan McNab, America Rodriguez, and the Guatemalan staff of

the Wildlife Conservation Society for their support, and also John Polisar and Robin Bjork for

counsel and kindness early in this process. I thank Anthony Novack, whose collaboration and

friendship along the way helped make the adversities more sufferable and the successes more

significant. In Uaxactun I am grateful to Pablo Nufiez, Antonio Ramos, Victor Mendez, Manuel

Mendez, Amilcar Fajardo, Aurora Soza Duran, Guillermo Calanclan, and the many participants

in data collection efforts there. The seemingly endless processing of dietary specimens was

bearable thanks to the assistance of Millicent Butterworth, Miguel Si, Lucas Cuc, and Enrique

Barahona. I owe a debt of gratitude to Nery Solis and Victor Hugo Ramos of the Monitoring and

Evaluation Center of the National Protected Area Council, and to Pedro Pineda of the Institute of

Agriculture, Natural Resources, and the Environment at Rafael Landivar University. Thanks to

Oscar Lara, Julio Morales, and Javier Rivas of the School of Biology at the University of San

Carlos for support with permitting and specimen identification. Salvador and Libertad Brizuela

provided sanctuary during the writing stage of this process. Thanks to Kitty Emery, Delores

Tillman, and Susana de Radachowsky for overseas assistance and to Isora Labisky, Greta

McNab, Milan Hooper, and Erik and Claire Lewis for their generous hospitality. I am most

grateful to my wife Michaelyn Bachhuber and to my children Milano and Chloe Lilyana for all

of their support, patience, and encouragement.









TABLE OF CONTENTS

page

A CK N O W LED G M EN T S ................................................................. ........... ............. .....

LIST O F TA B LE S ......... ..... .............. ................................................................. 7

LIST OF FIGURES .................................. .. ..... ..... ................. .8

A B S T R A C T ......... ....................... .................. .......................... ................ .. 9

CHAPTER

1 IN TRODU CTION ............... .............................. ................. ............... 11

Regional-Level Diversity Hypotheses........................................................ ............... 14
L ocal-L evel D diversity H ypotheses .............................................................. .....................15
O b j e c tiv e s ................... .............................................................. ................1 7
Conservation Threats ......................................................... .......... ........ ....... 19
Study Area ..........................................20

2 METHODS .........................................23

Diet and Harvest Data ..................................................... ............ ............... 23
Line-Transect Data ......................................................... .......... 24
H ab itat U se .........................................................................2 5
V ertical Strata O occupancy ............................................................................ ....................26
Species Abundance and Guild Composition ........................................ ....... ............... 26
N e st S ite D ata .........................................................................................2 7

3 R E SU L T S .............. ... ................................................................29

D iet C om position and O overlap ...................................................................... ...................29
H harvest D description .............. ..... ............................. ............. 29
Size A ssortm ent ...................................... ......................................................30
H ab itat U se ............................................................................... 3 0
V vertical Strata O occupancy ........................................................................... .......... ........... 1
Species Abundance and Guild Composition ........................................ ....... ............... 32
N est Site and C lutch C haracteristics............................................................ .....................34

4 D ISC U S SIO N ..............................................................................................36

Guild D iet Characteristics............... ...... .................... .. ............ ....... 36
G uild H habitat C haracteristics................................................. .. ..... 38
Relevance of Com petition to Guild Structure ........................................ ..........................39
Relevance of Non-Equilibrium Factors to Guild Structure ................................. ...............43
Relevance of Predation to Guild Structure ............. ....................... ..................... 45









Alternative Influences on Guild Structure.. ... ............................... ................. .......... ...............48
Guild Response to Hunting Pressure .......... ........ .................................... 48
C onclu sions.......... ..........................................................50

L IST O F R E F E R E N C E S .......... .................... .......................................... .....................................57

B IO G R A PH IC A L SK E T C H .............................................................................. .....................64
















































6









LIST OF TABLES


Table page

3-1 General diet compositions of guild members based on dry-mass proportions of
contents from upper-digestive tract specimens collected in Uaxactun, Flores, El
Peten, from January 2000 through December 2001............... ................. ............... 52

3-2 Overlap of dietary seed components between species-pairs and average seed
component overlap of individual species with all other guild members. ..........................52

3-3 Overlap of habitat occupancy between species-pairs and average combined overlap
of individual species with all other guild members. ....................... ... .................53

3-4 Overlap of vertical strata occupancy between species-pairs and average combined
vertical overlap of individual species with all other guild members...............................53

3-5 Density estimates for Crax rubra, Meleagris ocellata, Ortalis vetula, Penelope
purpurascens, and Tinamus major by forest-class in the Maya Biosphere Reserve,
G u atem ala. ............................................................. ................ 54

3-6 Guild compositions based on proportional population size and relative biomass at
each transect group. ....................... ........ ..... .... ...... ............ 54

3-7 Temporal reproductive patterns of Crax rubra, Meleagris ocellata, Ortalis vetula,
Penelope purpurascens, and Tinamus major based on the number of monthly
observations of reproductive activities. .................... ................ ....... ........ 55









LIST OF FIGURES


Figure page

1-1 Map of study area. Lightly shaded areas indicate the two management units in which
study data were collected. Circles depict the 50km2 areas corresponding to each
transect group .................................... ........................... .... ...... ........ 56









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

STRUCTURE OF A LOWLAND NEOTROPICAL GALLIFORM BIRD GUILD

By

Erick Hogan Baur

May 2008

Chair: Ronald F. Labisky
Major: Wildlife Ecology and Conservation

This study (1) examined guild structure, (2) compared guild characteristics to predictions

from potentially relevant hypotheses of guild structure regulation, and (3) assessed the impacts of

hunting on a guild comprised of the Great Curassow (Crax rubra), Crested Guan (Penelope

purpurascens), Plain Chachalaca (Ortalis vetula), Ocellated Turkey (Meleagris ocellata), and the

Great Tinamou (Tinamus major) in 2000-2002. The study area, located in the Maya Biosphere

Reserve in northern Guatemala, included a national park unit with protected populations of the

guild and an adjacent community concession unit subject to a range of subsistence hunting

pressure. Diet niches and overlap were described from analyses of upper digestive tract contents

from 267 C. rubra, 181 M. ocellata, 205 0. vetula, 142 P. purpurascens, and 55 T. major

collected over a 24-month period. Descriptions of habitat niches and overlap, species

abundance, and selected reproductive parameters were derived from line-transect sampling

during a 30-month period in the park unit (1,770 km total) and over a 12-month period in the

concession unit (995 km total). Descriptions of nest-sites, clutch-sizes, and other reproductive

characteristics were based on observations made at 24 C. rubra nests, 39 M ocellata nests, 77 0.

vetula nests, 19 P. purpurascens nests, and 66 T. major nests.









Dry-mass proportions of grit were highest and similar in the diets ofM. ocellata (22%) and

C. rubra (20%), lowest and similar in 0. vetula (2.7%) and T. major (2.2%), and intermediate in

P. purpurascens (9%). Seeds and pulp represented the greatest and second-greatest proportions

respectively, of the diet of all species when grit and snail shells were excluded from analyses.

Overlap in the seed component of the diet ranged from 50-86% for eight of the ten species-pairs,

and the average combined overlap exceeded 50% for all species except 0. vetula. Habitat

preferences were exhibited by C. rubra and P. purpurascens for tall-forest, and by 0. vetula for

low-forest. T. major occurred at a greater frequency in low-forest in the two areas with the least

hunting pressure, and in high-forest in the two areas with the greatest hunting pressure. The

frequency of occurrence ofM. ocellata did not differ among habitat types. All species exhibited

consistent vertical patterns of strata occupancy.

Guild characteristics were semi-consistent with structural regulation through competitive

interactions with respect to habitat niches, body-size assortment, and evidence of competitive

release, but were inconsistent with respect to diet niches and null-pool comparisons. The

variability of habitat occupancy and abundance within the guild were consistent with

expectations for non-equilibrium processes. Vertical stratification of nest-placement and other

reproductive attributes of the guild were consistent with expectations for a guild subject to

intense predation pressure. The guild appears to be a terrestrial vertebrate example of the non-

equilibrium competition model of community structure proposed by Connell (1980). All species

were most abundant in the fully-protected area; however, the abundance of C. rubra, M. ocellata,

and P. purpurascens declined consistently along the increasing gradient of hunting pressure.

Guild biomass in the most heavily hunted area represented 30% of the guild biomass in the

protected area.









CHAPTER 1
INTRODUCTION

This study examines the structure of a lowland tropical bird guild composed of the Great

Curassow (Crax rubra), Crested Guan (Penelope purpurascens), Plain Chachalaca (Ortalis

vetula), Ocellated Turkey (Meleagris ocellata), and Great Tinamou (Tinamus major) in the El

Peten department of Guatemala. These species share ecological and taxonomical or

morphological similarities, and a limited size range that make them likely competitors (Root

1967; MacNally 1983). Three of the species, C. rubra, P. purpurascens, and 0. vetula, are

members of the family Cracidae, a Neotropical galliform family represented by 11 genera and 50

species that reaches maximum diversity in South America (Brooks & Strahl 2000). These three

species are representative of the general size range of the family; Ortalis includes small cracid

species, Crax large species, and Penelope intermediate-sized species. The largest species in the

guild, M. ocellata, is also a galliform and the only tropical representative of the new world

subfamily Meleagridinae. T. major belongs to the Neotropical family, Tinamidae which is most

closely related to ratites. Tinamidae is represented by 9 genera and 47 species and also reaches

greatest diversity in South America (Bertelli & Porzecanski 2004). Although not a galliform, T.

major was included in this study because tinamids are ecologically similar to galliforms, and its

size is consistent with the range of the guild (Thompson 2004; Brooks et al. 2004).

These species exist sympatrically in the forest-dominated landscape of the study area and

share similar feeding habits. Although cracids have been described as frugivores (Karr 1971;

Thiollay 1994) or granivores (Terborgh et al. 1990), diet studies have recognized the significance

of seed, leaf, and animal materials (Santamaria & Franco 2000; Jimenez et al. 2001; Mamani-F

2001; Rivas & Morales 2003). Diet descriptions ofM. ocellata indicate an omnivorous diet









dominated by seeds (Steadman et al. 1979). T. major diet has been described as granivorous

(Thiollay 1994) or omnivorous (Howell & Webb 1995).

C. rubra is considered to function primarily as a seed predator on large-seeds and to a

lesser degree as a disperser of small seeds (Santamaria & Franco 2000; Rivas & Morales 2003).

Guans and chachalacas are considered to have lesser roles as seed predators and greater roles as

dispersers than curassows (Brooks & Strahl 2000; Rivas & Morales 2003). M. ocellata has a

muscular gizzard and consumes grit (Steadman et al. 1979) and likely functions more as a seed

predator than a disperser. An assessment of the impacts of the diet of T. major was not available;

however, due to its intermediate size relative to guans and chachalacas, the species is probably a

lesser seed predator and greater disperser than the larger guild members.

Guilds of large galliform species are not well represented in literature regarding bird

community structure and diversity. This under-representation may reflect less interest in

generalist species, which may be less suitable for niche dimension research which has dominated

community ecology, or may reflect vulnerability to hunting (Schoener 1983; Ricklefs 1990;

Terborgh et al. 1990). Large galliform species are often reduced or extirpated where they are

accessible, including sites where some of the most comprehensive field research of bird

communities have been conducted (Karr 1971; Karr 1976b; Thiollay 1989: Terborgh et al.

1990). The Cracid Specialist Group of the International Union for the Conservation of Nature

and Natural Resources (IUCN) describes Cracidae as the most threatened avian family in the

Neotropics, ranking 24 of its 50 species in need of priority conservation efforts (Brooks & Strahl

2000). Although generally less abundant than smaller bird species, large galliforms may become

common where protected (Thiollay 1989; Silva & Strahl 1991). Under natural conditions these

species may represent significant proportions of the biomass of bird communities (Terborgh et









al. 1990; Redford 1992), and may qualify as "keystone trophic" components of tropical

communities (Terborgh 1992). The combination of poor scientific representation, vulnerability,

threatened status, and ecological significance of large galliform species emphasizes the need for

research on such guilds where they persist under unexploited conditions (Terborgh et al. 1990;

Thiollay 1994; Brooks & Strahl 2000).

Rahbeck and Graves (2001) cited 120 existing causal hypotheses of bird species-richness

within the context of presenting yet a new model for predicting bird species-richness patterns.

The tendency of tropical ecosystems to be more diverse than temperate systems, or latitudinall

diversity gradient", may be the longest recognized pattern of species-richness (Hutchinson 1959;

Pianka 1966; Gill 1990; Ricklefs 1990). Many hypotheses proposed to account for the structure

of ecological communities, bird species-richness patterns, and the latitudinal diversity gradient

are potentially relevant to the diversity and structure of this guild. Assessment of the relevance

of any particular hypotheses is complicated by a variety of factors. Different hypotheses may

predict similar local effects, may identify causal factors that are not mutually exclusive, or that

may counteract one another. The interpretation of relationships among hypotheses varies among

authors, and the same research findings are cited in support of distinct hypotheses among authors

(Pianka 1966; Begon et al. 1990; Ricklefs 1990; Kricher 1997).

Regional-level hypotheses of tropical diversity generally emphasize the roles of abiotic or

historical factors operating on temporal or spatial scales that are not amenable to testing based on

the results of a single-site field study. Local-level tropical diversity hypotheses generally

emphasize the causal roles of biotic factors or environmental variability. Biotic regulation of

community structure may manifest as interspecific competition, external population controls, or

resource heterogeneity (Begon et al. 1990; Ricklefs 1990). Local-level hypotheses generally









diverge with respect to variability, either regarding communities as equilibrium systems

regulated by deterministic processes, or as non-equilibrium systems in which environmental

factors counteract deterministic processes (Begon et al. 1990; Ricklefs 1990).

Regional-Level Diversity Hypotheses

The time hypothesis attributes greater tropical diversity to the historic stability of tropical

biomes relative to temperate biomes subjected to glaciation during the Pleistocene (Begon et al.

1990; Ricklefs 1990). Evidence contrary to this hypothesis include: findings that tropical biomes

were also subject to historic periods of expansion and contraction (Haffer 1969), that temperate

biomes may be as old as tropical biomes (Ricklefs 1990) and that speciation can occur on

temporal scales that are insignificant relative to biome age (Remsen 1990). Support for this

hypothesis includes findings of comparisons of African and Neotropical bird communities in

similar forest and savannah biomes in which bird community diversity was positively correlated

to the relative age and historic extension of biomes (Karr 1976b).

The climatic stability and the productivity hypotheses attribute tropical diversity to

reduced variability of abiotic factors such as temperature, precipitation, and solar radiation

(Pianka 1966; Sanders 1968; Begon et al. 1990). Both hypotheses credit abiotic factors

operating on regional levels with increased local diversity. Stable tropical climates are favorable

to many plant and poikilothermic faunal taxa that in turn provide diverse and consistently

available resources to consumers (Pianka 1966; Kricher 1997). The productivity hypothesis

emphasizes the role of tropical stability in reducing physical stress on producers, permitting

greater investment in non-maintenance activities such as reproduction, which results in greater

abundance of producers and available resources to consumers (Connell & Orias 1964).









Local-Level Diversity Hypotheses

The regulation of community structure through competition has been a dominant

paradigm in community ecology (Schoener 1974; Schoener 1983; Gill 1990; Ricklefs 1990).

The competition hypothesis contends that when faunal communities are in a state of equilibrium,

species will occupy distinct ecological niches (Hutchinson 1959). Greater local diversity in

tropical systems increases interspecific competition, which becomes the most significant

determinant of community structure. When community structure is regulated by competitive

interactions, niche specialization is predicted to be prevalent among member species. Increased

niche specialization permits ecological separation of competitors, which are expected to exhibit

regular spacing along niche dimensions (Ricklefs 1990). According to this theory, modem

tropical assemblages are diverse because increased niche specialization has resulted in

"narrower" niches, thus accommodating greater numbers of species along resource gradients

(Ricklefs 1990; Begon et al. 1990). Because modern assemblages represent species that have

resolved historic competitive conflicts via co-adaptation, competition for resources may not

occur except under conditions of extreme resource scarcity (Wiens 1977; Fleming 1979). The

significance of competition may vary among trophic levels, with greater relevance to producers,

decomposers, and predators than to herbivores, frugivores, and some omnivores (Hairston et al.

1960; Fleming 1979).

An alternative role of competition in the regulation of community structure predicts that

competitive interactions in non-equilibrium systems will lead to greater niche generalization and

overlap (Connell 1980). Competitive interactions are inconsistent when competitor abundance is

highly variable or under conditions of diffuse-competition. Individuals of a given species may

interact with distinct competitor species, or may not face consistent competition from any

particular competitor species (Buckley 1983). This hypothesis contends that under such









conditions, species are more likely to develop evolutionary responses to food acquisition

efficiency or mortality factors than to the niches of competing species.

Biotic environmental factors that maintain populations below carrying capacity create

non-equilibrium conditions that prevent competitive interactions from reaching the point of

competitive exclusion (Pianka 1966; Ricklefs 1990). The tropical diversity gradient applies to

predators, parasites, and pathogens just as it does to other species. Predators often exhibit

"frequency-dependent e'le. tion", defined as the tendency to select the most abundant forms of

prey. Parasites and pathogens are likely to increase in correlation to increased host populations.

Because competitive exclusion does not occur under such conditions, communities may become

more diverse (Ricklefs 1990; Kricher 1997). In assemblages of populations that are subject to

external biotic controls it is predicted that species will exhibit lesser niche specialization and

greater resource use overlap (Pianka 1966).

Abiotic environmental factors may create non-equilibrium conditions by reversing

successional vegetation trends or by increasing resource variability. Forest-gap dynamics

increase the heterogeneity of tropical forest diversity and structure (Connell 1978; Schemske &

Brokaw 1981). Large rivers that flood seasonally maintain vegetation along the banks in early

successional stages, increasing habitat heterogeneity and favoring greater bird diversity (Remsen

& Parker 1983; Brooks et al. 2004). Highly variable precipitation patterns may reduce the

availability or consistency of resources, limiting community diversity and precluding the

development of niche specializations for particular resources (Karr 1976a).

Greater local diversity in the tropics may create resource gradients that are sufficiently

heterogeneous to support a greater diversity of consumers (Hutchinson 1959; Kricher 1997). If

resource heterogeneity is significant to community structure it is predicted that niches will









exhibit greater dimensionality (Ricklefs 1990). Habitat heterogeneity has been positively

associated with bird species diversity in both tropical and temperate climates (MacArthur &

MacArthur 1961; Karr & Roth 1971). The diversity of some tropical bird communities has been

attributed to greater diversity of frugivorous, insectivorous, and nectivorous species, reflecting

utilization of resources that are more diverse and abundant in the tropics (Pianka 1966; Karr

1971; Terborgh 1990).

Objectives

The primary objective of this study was to examine guild structure in a manner that

would permit the assessment of relevant regulatory processes to the guild. Specific objectives

were (1) to determine the degree of specialization of principal niche dimensions within the guild,

(2) to determine the degree of ecological separation or overlap along corresponding resource

gradients, and (3) to compare guild characteristics to predictions derived from potentially

relevant hypotheses of structural regulation. The secondary objective of this study was to assess

the impacts of hunting pressure on guild structure and composition.

Niches were described along certain dietary and habitat dimensions. Diet is considered

the most essential niche dimension among terrestrial vertebrates (Schoener 1974; Schoener

1983) and dietary specialization has been documented in many tropical bird guilds (Ashmole

1968; Sherry 1984). Spatial separation along habitat resource gradients has been associated with

the diversity of many bird communities (Cody 1968; Gill 1990). Habitat resources may be

partitioned through preferential use, temporal variability of use, and vertical stratification (Gill

1990; Pearson 1971; Karr 1971).

Other guild characteristics that were compared to predictions based on potentially

relevant hypotheses included size assortment, responsiveness of guild members to abundance

variations of other guild species, guild membership, variability of species abundance and habitat









occupancy, and reproductive strategies. Size assortment is a mechanism of mitigating

interspecific competition that occurs when combinations of similar species with distinct sizes

persist better than similar-sized species (Case & Sidell 1983). Size assortment has been

associated with the greater diversity of some tropical bird guilds (Ashmole 1968; Brooks 1998).

If guild member populations are constrained by competitive interactions, then significant

reductions of some species may offer an opportunity for competitive release to other guild

members. The use of a greater range of habitat resources in the absence of competitors has been

documented in some tropical bird communities (MacArthur et al. 1966). Bird populations are

not static and significant temporal variations of abundance or habitat use may represent non-

equilibrium factors that reduce the predictability of competitive interactions and counteract local

processes of competitive exclusion (Karr & Freemark 1983; Gill 1990; Loiselle & Blake 1992).

Vertical stratification of nest-placement is a mechanism of reducing nest-predation pressure

through increased spatial distribution of nests (Martin 1988a; Martin 1993; Schmidt & Whelan

1998). Temporal separation of reproductive activities may mitigate interspecific competition for

nesting or brooding resources in bird communities (Ricklefs 1966). Comparisons of guild

composition to null pools of potential members that are regionally available have been used to

assess the significance of competitive interactions to guild structure in tropical bird communities

(Pearson 1977; Cornell & Lawton 1992).

It was predicted that if interspecific competition in the classic, equilibrium-system

context is significant to the regulation of guild structure that: (1) species would have specialized

niches, (2) species would exhibit regular spacing along niche dimensions, (3) species would be

sufficiently different in size to mitigate competitive interactions, (4) species would exhibit some

form of competitive release in response to significant variations in the abundance of other guild









members, (5) guild members might stagger reproductive activities to reduce competition for

nesting and brooding resources, and (6) guild membership would include fewer similar species

than are available from the regional null-pool of potential guild members.

Further it was predicted that if non-equilibrium processes were significant to guild

structure, species would exhibit limited niche specialization and ecological separation, and

populations would exhibit non-equilibrium properties such as significant temporal variations in

abundance or habitat occupancy. Also it was predicted that if predation pressure was significant

to guild structure then guild members would exhibit differential nesting strategies or

reproductive traits that are adaptive to conditions of high mortality.

Conservation Threats

The "Maya Forest" refers to multinational complex of protected areas corresponding to

the southern Yucatan peninsula regions of Mexico, Belize, and Guatemala. The Maya Forest is

the largest Neotropical forest remnant in Mesoamerica encompassing approximately three-

million hectares of contiguous forest cover (Radachowsky & Ramos 2004; CEMEC-CONAP

2006). The principal Guatemalan component of the forest is the Maya Biosphere Reserve

(MBR) created in 1990 that includes over two-million hectares (CONAP 2001). Despite legal

restrictions on the harvest of wildlife, law enforcement in Guatemala is minimal and hunting is

largely uncontrolled. Throughout most of the MBR, these species are subject to persistent

hunting pressure that is positively associated with permanent settlements and human access

(Radachowsky & Ramos 2004). Some of the most isolated parts of the reserve continue to

support unexploited populations of the guild species.

In 1960, the human population of the Peten was approximately 60,000, but has since

doubled at approximately 8-year intervals (Meyerson 1997). A census of the Peten in 2001

recorded over 700,000 inhabitants, at which time there were approximately 90,000 residents









within the MBR (CONAP 2001). Since the declaration of its protected status, the MBR has lost

over 2,000 km2 of forest cover (Ramos et al. 2007) and been fragmented into distinct eastern and

western forest blocks. Forest fires provoked by swidden (slash-and-burn) agricultural practices

occur annually. The three worst fire seasons in the last 10 years were 1998, 2003, and 2005,

during which 4,335 km2, 3,985 km2 and 3,752 km2 respectively were affected by forest fires

(Ramos et al. 2007).

The eastern forest block of the MBR retains approximately 8,000 km2 of forest cover that

links the Mexican and Belizean components of the Maya Forest. Current infrastructure

proposals include three major road-building projects within this part of the reserve (Ramos et al.

2007). One proposed project is a highway that would connect Guatemala and Mexico through

the best-conserved areas remaining in the eastern forest block. Environmental impact

assessments of that project based on models from data on existing roads in the MBR (Ramos et

al. 2007) predicted severe forest-cover loss, fragmentation into as many as three separate forests,

and the loss of continuity between the Mexican and Belizean components of the Maya Forest by

2025 (Ramos et al. 2007).

Study Area

This study was conducted in two adjacent management units of the MBR; the community

forestry concession of Uaxactun and the Mirador-Rio Azul National Park (Figure 1-1). The

Uaxactun unit includes approximately 836 km2 of the multiple-use zone and contains a single

permanent settlement, the village of Uaxactun with approximately 700 residents (NPV-OMYC

1999). The Mirador-Rio Azul National Park encompasses approximately 1117 km2 and is

uninhabited (CONAP-ONCA-CECON 2002). The Mirador-Rio Azul National Park is divided

into two sub-units, Mirador in the west and Rio Azul in the east. Data for this study were

collected in the Rio Azul unit in the northeastern part of Guatemala bordering Mexico on the









northern perimeter, Belize on the eastern perimeter, and the Uaxactun unit on the southeastern

perimeter.

Data from the nearest meteorological stations indicated mean monthly temperatures

ranging from 23 C to 300 C (INSIVUMEH), being lowest November-February (mean median

temperature = 26 C) and highest April-July (mean median temperature = 32 C). Local average

annual precipitation is approximately 1250 mm. The annual precipitation pattern exhibits peaks

in May or June and in September, and a four-month dry season from January-April.

The Uaxactun and Rio Azul units straddle the drainage basin of the Rio Tikal-Rio Hondo

watershed. Elevation ranges from 100-400 m, the lowest areas corresponding to the drainage

basin and the highest to a steep karst escarpment that borders the basin to the west and north.

The eastern edge of the basin is characterized by isolated upland areas with gentle slopes.

Seasonal flooding can be severe in low and poorly drained areas during the rainy season.

Throughout the forest, small ponds that form in depressions during the rainy season often dry-up

during the dry season, leading to seasonal scarcity of available surface water. The Rio Tikal

River is an intermittent "arroyo" that only occasionally forms a current but retains small ponds

year-round in depressions along the channel.

The landscape of the study area is dominated by a forest continuum that is classified as

Subtropical Moist (Holdridge et al. 1971). A classification of the forest community in Tikal

National Park, which is adjacent to the Uaxactun unit, found strong correlations of forest

composition and structure to the edaphic conditions along the topographical gradient (Schulze &

Whitacre 1999). Proceeding down the slope of this gradient, canopy height and basal area

decrease and canopy opening size and stem-density increase (Schulze & Whitacre 1999).









Although 11 predictable forest-classes were identified along the topographical gradient, species

richness and diversity were consistent among forest-classes (Schulze & Whitacre 1999).

The village of Uaxactun originated in the early 1900's as a collection center for "chicle",

the resin of the Manilkara zapota tree that was the original base ingredient of chewing-gum

(NPV-OMYC 1999). Although the economic importance of gum-resin extraction has since

diminished greatly, the local economy continues to be dominated by the extraction of forest

resources including timber, palm fronds, allspice, construction materials, and wildlife. Although

small-scale agriculture is locally important, deforested agricultural areas represent only 4% of

the concession area (NPV-OMYC 1999). To accommodate extraction activities primitive camps

interconnected by an extensive trail network are distributed throughout the concession.

Wildlife is harvested opportunistically during the course of other extractive and

agricultural activities on a continuous basis. Community-based hunting pressure is inversely

correlated to distance from the village (Polisar et al. 1998); however, camp-based extractive

activities have temporary but intense impacts far from the village (McNab 1998; Morales &

Morales 1998). Although subsistence hunting pressure is biased towards larger vertebrate

species, individuals are generally harvested without regard to sex or age-class. Unpublished data

provided by the Wildlife Conservation Society (WCS) in Guatemala indicated that the wildlife

harvest in Uaxactun in 1997 produced approximately 10,400 kg of game meat, which would cost

approximately $30,000 US to replace with meat produced from domestic livestock. Given that

the average annual household income is less than $2,000 US (personal observation) among

approximately 140 households in the village, the annual wildlife harvests represent a subsidy

equivalent to more than 10% of the local economy.









CHAPTER 2
METHODS

Diet and Harvest Data

Data were collected from birds harvested for subsistence purposes by residents of the

village of Uaxactun for the purpose of describing diet, body mass, and harvest pressure of guild

species. The participating hunters provided upper-digestive tracts (gizzards and also crops when

available) from harvested specimens along with relevant observations, for a reward equivalent to

the value of a pound of game-meat (<1$ US). Diet samples were separated from the tracts, dried

in a solar dryer, and stored in paper envelopes in the community. Samples were dried again in

electric dryers in the laboratory prior to processing. The contents of each sample were separated

into distinct components for which dry mass values were recorded and identified on general and

specific levels.

Diet compositions were based on the dry-mass proportions of general diet components

including: animal, leaf, pulp, seed, flower/stem, and grit. For composition analyses

unidentifiable pulverized material in each sample was assigned to the identifiable items

according to their relative proportions. Alternative diet composition descriptions were

determined without grit and snail shells, due to their bias relative to the dry mass of plant and

animal tissues. Dietary-overlap indices were calculated for all species-pairs using the equation

from Pianka (1973) shown below, based on the seed component of each species diets, where Ojk

is the overlap between species and species k, and Pij and Pik are the proportions of the ith

resource used by thejth and kth species, respectively. Unidentifiable pulverized material was

excluded from the overlap analyses.

n n

Ojk Z Pij Pik / j2 Pik2
i i









The harvest data reported by local hunters were used to describe local harvest pressure on

guild member populations. Adult body-mass values from the data were applied to describe

harvest biomass. Harvest pressure was described by distance per 4 km intervals from the village.

Birds collected by local hunters are usually cleaned in the field to avoid spoilage and reduce the

burden, so access to entire specimens was limited. Balance scales were provided to field

personnel and certain local hunters in order to record body mass values from entire specimens

opportunistically. Adult body-mass records were compared between species and sexes using an

analysis of variance (ANOVA) and a Duncan's Multiple Range Test (alpha levels < 0.05 were

considered statistically significant). Size-ratios between closest-size species-pairs were

compared to predictions from competition theory.

Line-Transect Data

A system of 12 line-transects (Buckland et al. 2001) provided data used to describe

species abundance and certain niche parameters. Transects were organized into 4 groups of

three, located along a hunting pressure gradient identified aprior (Polisar et al. 1998). The

Ixcan transect group was located in the core of the Rio Azul unit where no hunting pressure was

expected. The Cedro group was located on the park perimeter adjacent to the Uaxactun unit,

approximately 25-37 km from the village where hunting pressure was expected to be light. The

Uaxactun North group was located approximately 3-15 km north of the village where hunting

pressure was expected to be heavy. The Uaxactun South group included the area surrounding the

village where hunting pressure was expected to be heaviest.

Each transect within a group was located in a stratified-random manner in order to

sample all forest-classes. In the Uaxactun unit, transects were oriented to avoid existing trails to

reduce the probability of locals hunting on them. In the Rio Azul unit all but one of the transects

were established on modified, existing trails. Eleven transects measured 2500 m, whereas one









transect measured 2750 m to improve representation of a forest-class in that group. Transects

were flagged and labeled at 50 m intervals and mapped by forest-class.

Transects were sampled by trained observers between 0600 and 0930 hours at intervals of

5-10 days depending on personnel capacity, weather, and logistics. Observers recorded data

from detections of guild members and other species of interest to associated researchers (Novack

2003; Garcia & Radachowsky 2003). The transects in the Rio Azul unit were established in

February 2000 and sampled over a 30-month period. Each year data collection in Rio Azul was

interrupted when seasonal flooding prevented access to the site. The transects in the Uaxactun

unit were sampled from August 2000 through July 2001. The total sampling effort on all

transects was approximately 2770 km.

Habitat Use

Habitat use was analyzed based on species-encounter rates by forest-class derived from

the transect data. Two forest classifications based on alternative interpretations of the results of

Schulze and Whitacre (1999) were evaluated. One classification consisted of three forest-

classes: "Upland", representing tall forest types on the upper slopes of the topographical

gradient; "Scrub", representing low forest types at the bottom of the gradient; and "Transitional",

representing intermediate forest types. The second classification consolidated the Upland and

Transitional classes of the former classification into a single "Upland" forest-class, and used the

same "Scrub" forest-class. Monthly species-encounter rates in the Upland and Transitional

forest-classes of the three-class system were compared using a student t-test to determine if

species responded to the increase resolution of that classification. To determine if species

exhibited preferential habitat occupancy, species-encounter rates by forest-class were calculated

for each transect group and annual period, converted into relative frequencies and compared

using a student t-test. Alpha levels < 0.05 were considered statistically significant for both t









tests. Habitat occupancy overlap was assessed using proportional similarity indices (Schoener

1970), calculated for each species-pair in each forest-class. For each species the combined

average overlap with all other species was also described.

Coefficients of variation (CV) calculated from species-encounter rates were used to

evaluate the temporal variability of habitat occupation. Monthly species-encounter rates by

forest-class were plotted graphically and evaluated for temporal variations of habitat occupancy.

Monthly species-encounter rates by forest-class were examined for correlation with precipitation,

temperature, and encounter rates of other guild members using a Pearson correlation analysis.

Vertical Strata Occupancy

Vertical height records from the transect data were used to assess whether guild members

exhibited consistent patterns of forest-strata occupancy. Height records were assigned to one of

three categories, ground strata (0-1m), understory strata (Im to half of the mean canopy height),

or canopy strata (half of the mean canopy height and above). The relative frequencies of

occurrence in the three strata were compared between transect groups, forest-classes, and annual

periods for each species using an analysis of variance (ANOVA) and a Duncan's Multiple Range

Test (alpha levels < 0.05 were considered statistically significant). Vertical overlap in strata

occupancy was estimated for each species-pair using proportional similarity indices (Schoener

1970). For each species the combined average overlap with all other species was also described.

Species Abundance and Guild Composition

Variables derived from the transect data that were used to compare guild member

abundance included individual-encounter rates, density estimates by forest-class, relative

population size and relative population biomass. Monthly individual-encounter rates from the

same annual period were compared among transect groups for each species using an analysis of

variance (ANOVA) and a Duncan's Multiple Range Test (alpha levels < 0.05 were considered









statistically significant). Density estimates were calculated from the transect data using the

program DISTANCE version 5 (Thomas et al. 2005). Models used to estimate densities were

selected by the program based on comparisons of all relevant combinations of detection

functions. Data were adjusted (truncated or filtered) when necessary to control for data

concentrations that were inconsistent with detection functions ("data heaping").

Spatial forest-class descriptions derived from Landsat 7 (USGS/EROS 2000) satellite

images from the study period were provided by CEMEC-CONAP (2006). Using the program

ArcView 3.2 (ESRI 2000), 4 km-radius buffers representing 50 km2 were applied to the

approximate geographic center of each transect group and were used to determine habitat

availability within the corresponding areas (Figure 1). Density estimates by forest-class were

extrapolated by forest-class representations within each 50 km2 area to estimate corresponding

species population sizes. Guild composition descriptions for each transect group were

determined based on relative population size and relative biomass representations at each site.

Guild compositions were compared between transect groups and annual periods. Temporal

variability of species abundance was assessed using coefficients of variation calculated from

individual-encounter rates during annual and 30-month periods.

Nest Site Data

Data collected from nest-sites of guild members were used to determine reproductive

characteristics of each species. Local participants were compensated a day's wage

(approximately $7 US) to accompany data collectors to nests encountered during the course of

other activities and to assist with data collection at the nest site. Clutch sizes were included in

analyses if they met the minimum clutch size reported from a literature source, and did not

appear abandoned. Search effort was not systematic with respect to habitat so forest-class

percentages are not representative of habitat distributions for the species. Nest sites could only









be visited once and some visits were made after nesting activities had concluded so nest

depredation and clutch-size records are conservative. Vertical nest placement, tree diameter, and

clutch-size were compared between species using an analysis of variance (ANOVA) and a

Duncan's Multiple Range Test (alpha levels < 0.05 were considered statistically significant).

Nest placement patterns within the guild were evaluated for spatial separation. Temporal

patterns of reproductive activities were described from field observations, the line-transect and

the nesting data and subsequently evaluated for temporal separation.









CHAPTER 3
RESULTS

Diet Composition and Overlap

Diet content samples were derived from 267 C. rubra, 181 M. ocellata, 205 0. vetula,

142 P. purpurascens, and 55 T major (Table 3-1). Grit represented 20% of the diet in C. rubra,

22% in M ocellata, 3% in 0. vetula, 9% in P. purpurascens, and 2% in T. major. Diet

compositions without grit and snails indicated seed proportions of 88% of the diet in C. rubra,

77% in M ocellata, 85% in 0. vetula, 88% in P. purpurascens, and 90% in T. major. Excluding

grit and snails, fruit pulp comprised 8% of the diet in C. rubra, 11% in M ocellata, 7% in O.

vetula, 10% in P. purpurascens, and 7% in T. major. Leaf proportions without grit and snails

contributed 10% of the diet in M. ocellata, 7% in 0. vetula, and less than 2% of the diets of the

remaining species. Proportions of flower/stem and animal materials varied among species, but

were less than 3% of the diet of any guild member.

Guild diet included 143 seed species, with similar quantities among guild members

despite sample size differences: C. rubra (74 spp.), M. ocellata (92 spp.), 0. vetula (86 spp.), P.

purpurascens (72 spp.), T major (67 spp.). Seed-species overlap values between species-pairs

ranged from 33% forM. ocellata-O. vetula to 86% for C. rubra-T major (Table 3-2). Average

seed overlap with all other species ranged from 47-68%, with T. major ranking highest, followed

in sequential order by P. purpurascens, C. rubra, M. ocellata, and 0. vetula.

Harvest Description

The recorded harvest of guild members averaged 591 individuals representing 1625 kg of

biomass annually based on average adult, body-mass of guild members. C. rubra represented an

average of 47% of the total harvest and 58% of the harvest biomass annually. M. ocellata

represented 19% of the total harvest and 28% of the biomass the first year, and declined to 13%









and 22% respectively, the second year. 0. vetula represented 16% of the harvest and 3% of the

biomass the first year, and increased to 25% and 6%, respectively, the second year. P.

purpurascens represented 7% of the harvest and 6% of the biomass the first year, and increased

to 17% and 16%, respectively, the second year. T. major represented an average of 5% of the

annual harvest and 2% of the biomass. Harvest pressure on C. rubra, M. ocellata, and P.

purpurascens was greatest in the second distance-interval (4-8 km) from the village and declined

steadily in subsequent intervals. Harvest pressure on 0. vetula and T. major was greatest in the

first distance interval (0-4 km) and declined steadily in subsequent intervals.

Size Assortment

Adult body-mass records were collected from 67 C. rubra, 29 M. ocellata, 17 0. vetula,

26 P. purpurascens, and 12 T major. Average adult body-mass was 3.4 kg for C. rubra, 4.16 kg

forM ocellata, 0.64 kg for 0. vetula, 2.31 kg for P. purpurascens, and 1.17 kg for T. major.

Adult body-mass differed between guild members based on the results of both analyses

(ANOVA, alpha = 0.0001, F = 336.91, df = 150; Duncan's, alpha = 0.05, df = 146). Adult body-

mass of sexes differed in M ocellata (male = 4.81 kg; female = 3.51 kg) (ANOVA, alpha =

0.0001, F = 26.79, df = 28; Duncan's, alpha = 0.05, df = 27) and P. purpurascens (male = 2.16

kg: female = 2.44 kg) (ANOVA, alpha = 0.0269, F = 5.56, df = 25; Duncan's, alpha = 0.05, df=

24). Size ratios (and natural logarithm equivalents) between species-pairs were 1.23:1 (0.209)

for C. rubra-M. ocellata, 1.47:1 (0.384) for C. rubra-P. purpurascens, 1.98:1 (0.681) for P.

purpurascens-T. major, and 1.83:1 (0.609) for 0. vetula-T. major. Size ratios between M

ocellata sexes and C. rubra were 1.42:1 (0.348) for males and 1.03:1 (0.032) for females.

Habitat Use

Monthly species-encounter rates between Upland and Transitional forest-classes did not

differ for any guild member. All subsequent habitat comparisons were made using the two









forest-class system. Frequencies of occurrence between forest-classes were unequal (alpha =

0.0001) for 0. vetula (t = -7.9141, df =10), C. rubra (t =11.681, df = 10), and P. purpurascens (t

= 6.1134, df = 10) and nearly unequal (alpha = 0.0601) for T major (t = 2.1193, df = 10). M.

ocellata did not differ in forest-class occupancy. 0. vetula occurred in Scrub forest with an

average frequency of 0.77. C. rubra and P. purpurascens and T. major occurred in Upland

forest with average frequencies of 0.73, 0.66, and 0.58, respectively. Habitat overlap among

species-pairs ranged from 27-48% in Scrub forest and from 23-67% in Upland forest (Table 3-3).

Combined habitat overlap among species-pairs ranged from 50-94%. Average combined habitat

overlap with all other species ranged from 61-84%, with T major ranking highest, followed in

sequential order by M. ocellata, P. purpurascens, C. rubra, and 0. vetula.

Ratios of CV values between Scrub forest and Upland forest were 1.82:1 for C. rubra,

1.35:1 for M ocellata, 2.35:1 for P. purpurascens, and 1.94:1 for T. major, indicating greater

variability in the occupancy of Scrub forest. The CV ratio between Upland forest and Scrub

forest was 1.43:1 for 0. vetula indicating greater variability in the occupancy of Upland forest.

Graphical analyses of monthly species-encounter rates by forest-class did not reveal any

consistent temporal trends or shifts in forest-class occupancy by guild members. The Pearson

correlation analyses did not identify any (alpha = 0.05) correlations of monthly species-

encounter rates with respect to temperature, precipitation, or encounter rates of other species.

Vertical Strata Occupancy

All species exhibited differential use of forest strata (ANOVA df = 35): Crax rubra

(alpha = 0.0011, F = 4.68), M. ocellata (alpha = 0.0001, F = 43.07), 0. vetula (alpha = 0.0018, F

= 4.36), P. purpurascens (alpha = 0.0001, F = 43.41), T. major (alpha = 0.0001, F = 57).

Vertical patterns of forest-strata occupancy did not differ between transect groups, forest-classes,

or annual periods for any species (Duncan's alpha = 0.05, df = 27)M. ocellata and T. major had









average frequencies of 89% and 92%, respectively, in the ground stratum. C. rubra had an

average frequency of 59% in the canopy stratum and an average of 32% in the ground stratum.

P. purpurascens had an average frequency of 84% in the canopy stratum. 0. vetula had average

frequencies of 53% and 41%, respectively, in the canopy and understory strata. Vertical strata

overlap ranged from 14% between 0. vetula-T. major to 93% between M. ocellata-T major

(Table 3-4). Four species-pairs exhibited vertical overlap greater than 50%. Average combined

overlap with all other species ranged from 40-56%, with C. rubra ranking highest, followed in

sequential order by M. ocellata, P. purpurascens, 0. vetula, and T. major.

Species Abundance and Guild Composition

Monthly individual-encounter rates differed (Duncan's, alpha = 0.05, df = 42) among

transect groups for C. rubra (ANOVA, alpha = 0.0001, F = 9.13, df = 45), 0. vetula (ANOVA,

alpha = 0.0012, F = 6.36, df = 45), and P. purpurascens (ANOVA, alpha = 0.0001, F = 31.49, df

= 45). Encounter rates decreased along the increasing hunting gradient for C. rubra, M. ocellata,

and P. purpurascens. Encounter rates were highest in the Ixcan and Uaxactun South groups for

0. vetula. Encounter rates were highest in the Ixcan group and stable among the remaining

transect groups for T. major.

Density estimates for all species were highest in the protected Ixcan group (Table 3-5).

Density estimates generated by DISTANCE for some species in the less hunted areas seemed

intuitively robust, however the results of analyses were ultimately accepted if they met the model

selection criteria of the program, were consistent with the data and derived encounter-rates, and

were comparable published density estimates for these or congeneric species.

C. rubra exhibited overall density declines along the increasing hunting gradient by 83%

in both Upland and Scrub forest. M. ocellata exhibited overall declines along the increasing

gradient of approximately 66% both forest-classes, with highest density estimates in the Ixcan









and Uaxactun North groups. 0. vetula densities were similar and highest in the Ixcan and

Uaxactun South groups. P. purpurascens exhibited an overall decline along the gradient of 72%

in Upland forest and 43% in Scrub forest. T major exhibited little density variation among the

Cedro, Uaxactun North and Uaxactun South transect groups. T. major exhibited higher densities

in Scrub forest relative to Upland forest in the Ixcan and Cedro groups, and higher densities in

Upland forest relative to Scrub forest in the two Uaxactun groups. Comparison of densities

between years in the Ixcan and Cedro groups indicated increased or stable densities among all

species, except for slight decreases for T. major in the Cedro group. Between years the

estimated density ofM. ocellata increased by 59% in Upland forest and 14% in Scrub forest in

the Ixcan group, and by 159% in Upland forest and 569% in Scrub forest in the Cedro group.

C. rubra represented proportions of the guild ranging from 24-32% in the Ixcan, Cedro,

and Uaxactun North groups, but only 11% in the Uaxactun South group (Table 3-6). M. ocellata

averaged 18% of the guild in the Ixcan and Uaxactun North groups, and 10% and in the Cedro

and Uaxactun South. 0. vetula averaged 24% of the guild in the Ixcan, Cedro, and Uaxactun

North groups and represented 46% of the guild in the Uaxactun South group. P. purpurascens

averaged 23% of the guild in the Ixcan and Cedro groups, and 14% in the two Uaxactun groups.

T. major averaged 10% of the guild in the Ixcan and Cedro groups and 19% in both Uaxactun

groups.

C. rubra accounted for the highest biomass proportion of the guild in all groups (average

40%) except Uaxactun South where it represented only 22%. M. ocellata averaged 31% of the

guild biomass in all transect groups except the Cedro group where it accounted for only 18%.

The proportions of guild biomass represented by 0. vetula increased steadily along the

increasing hunting gradient from 6% in the Ixcan group to 18% in the Uaxactun South group. P.









purpurascens accounted for 25% of guild biomass in the Cedro group, 21% in the Ixcan and

Uaxactun South groups, and 14% in the Uaxactun North group. T. major accounted for 5% of

guild biomass in the Ixcan and Cedro groups and an average of 12% in the two Uaxactun groups.

Coefficients of variation calculated from individual-encounter rates during both annual

and 30-month periods indicated double-digit percentage changes for all species. Average CVs

for annual periods were 34% for C. rubra, 35% forM. ocellata, 26% for 0. vetula, 18% for P.

purpurascens, and 21% for T. major. CVs of variation for the 30-month period were 38% for C.

rubra, 62% for M ocellata, 32% for 0. vetula, 22% for P. purpurascens, and 22% for T. major.

Nest Site and Clutch Characteristics

A total of 225 were observed: C. rubra nests (n = 24), M. ocellata nests (n = 39), 0.

vetula nests (n = 77), P. purpurascens nests (n = 19), T. major nests (n = 66). Mean nest height

differed for 0. vetula (5.7 m, range 1-18 m), and species pairs C. rubra (10 m, range 5-25 m) -P.

purpurascens (9.75 m, range 4-15 m), and T. major (0 m) -M. ocellata (0 m) (alpha = 0.05, df=

206). Mean DBH of trees associated with nest sites differed between the species group T major

(35.4 cm) -M. ocellata (32.3 cm) -P. purpurascens (26.3 cm) and the individual species C. rubra

(20 cm) and 0. vetula (16.9 cm) (alpha = 0.05, df= 151).

Mean clutch sizes forM ocellata (8.73, mode = 9) and T. major (4.49, mode = 4)

differed from each other and the other guild members (alpha = 0.05, df = 203). Mean clutch

sizes for P. purpurascens (2.16, mode = 2) -0. vetula (2.12, mode = 2) -C. rubra (2.00, mode =

2). Depredation of nests, based on a single visit, were 20% (n = 13) for T. major, 15% (n = 6)

forM ocellata, 4% (n = 3) for 0. vetula nests, 4% (n = 1) for C. rubra, and 0% for P.

purpurascens. With respect to habitat, 96% (n = 23) of C. rubra nests, 95% (n = 19) of P.

purpurascens nests, 95% (n = 23) of T. major nests, 36% (n = 14) ofM. ocellata nests, and 16%

(n = 12) of 0. vetula nests were located in Upland forest. Nesting in Scrub forest was only









observed for 0. vetula (7%), T. major (5%), and M. ocellata (3%). Only 0. vetula (78%) and M

ocellata (62%) nests were observed in agricultural habitat.

Specifically, all C. rubra and P. purpurascens nests were placed in trees, and all T major

nests in the roots of trees. Seventy-three percent (n = 52) of 0. vetula nests were placed in trees,

20% (n = 14) were in vines. M. ocellata placed 49% (n = 19) and 36% (n = 14) of their ground

nests in bracken fern (Pteridium spp.) and the roots of trees, respectively.

All guild members commenced nesting near the end of the dry season (March-April) and

continued into the wet season; however, the length of the nesting period varied among guild

members (Table 3-7). Active nests of C. rubra and M ocellata were only observed April-June,

whereas nests ofP. purpurascens were observed from April-July. Nesting by T. major and 0.

vetula occurred during March-September and March-October, respectively.









CHAPTER 4
DISCUSSION

Guild Diet Characteristics

Relatively high proportions of grit in the diets of C. rubra and M. ocellata suggest that

they function as seed predators. Relatively low proportions of grit in the diets of 0. vetula and T.

major suggest that they function less as seed predators than the rest of the guild. Diet

compositions of C. rubra, 0. vetula, and P. purpurascens were similar to previous findings

(Rivas & Morales 2003). Flower and stem materials combined did not represent a significant

proportion of the diet of any guild member. Flowers have been reported in higher proportions in

the diets of P. purpurascens (Rivas & Morales 2003), and M ocellata (Steadman et al. 1979),

and may be seasonally important to curassows (Santamaria & Franco 2000). Animal matter was

proportionately higher in M. ocellata with snail shells included, but similar to other species when

they were excluded. Diet studies based on direct observations indicate frequent consumption of

a variety of small vertebrates by curassows (Santamaria & Franco 2000, Jimenez et al. 2001).

The relatively higher proportions of leaf material in the diets ofM ocellata and 0. vetula, in

contrast to the other guild members, are consistent with findings from other studies (Rivas &

Morales 2003; Steadman et al. 1979). Excluding grit and snails, the diet of individual guild

members had similar, proportions of pulp (7-11%) and of seeds (77-90%), thus the diet of this

guild is dominated by seeds, many of which are consumed as whole fruit.

Species-pairs 0. vetula-M. ocellata and C. rubra-0. vetula displayed relatively low

dietary overlap (33% and 39% respectively), two species-pairs, P. purpurascens-T. major and C.

rubra-T. major displayed relatively high overlap (73% and 86% respectively), and the remaining

six species-pairs had intermediate values (50-68%). Of the 143 seed species consumed by guild

members; 19% (27 seed species) were shared by all five guild members, 13% (19) by four









members, 18% (26) by three members, and 21% (30) were shared by two guild members. Seed

species shared by the entire guild represented significant proportions of the diets of individual

species: 82% of the seed dry mass in C. rubra, 76% in M. ocellata, 57% in 0. vetula, 79% in P.

purpurascens, and 83% in T major. The fruits Brosimum alicastrum and Psuedolmedia spp.

were among the top-five ranking seeds in the diets of four of the guild members and ranked 15th

and 8th, respectively, in 0. vetula.

Rivas & Morales (2003) analyzed diets of C. rubra, 0. vetula, and P. purpurascens

collected from the study area between January 2002 and July 2003. In that study the two most

significant seed species in the diet of C. rubra were from the fruits Pouteria amygdalina (47%)

and Protium copal (8%); the two most significant seeds in the diet of P. purpurascens were P.

amygdalina (17%) and Vitex gaumeri (12%); and the most significant seed in the diet of 0.

vetula was Psuedolmedia spp. (19%). In the current study P. amygdalina represented 9% and P.

copal 1% in C. rubra; P. amygdalina represented 1% and V gaumeri 5% in P. purpurascens;

and Psuedolmedia spp. represented 3% in 0. vetula.

In the current study, the most significant seed species in the diet of C. rubra were B.

alicastrum (20%) and Calophyllum brasiliensis (19%), and the most significant seed species in

the diet of P. purpurascens was Chrysophyla argentea (14%). Rivas and Morales (2003) did not

detect B. alicastrum in the diet of C. rubra and C. brasiliensis represented 6% of the diet

furthermore they found that C. argentea represented 5% of the diet in P. purpurascens. Those

authors noted only minor representation of B. alicastrum which represented (3%) in 0. vetula

and (0.05%) in P. purpurascens. Rivas (1995) analyzed diet contents of C. rubra from the same

area in 1995 and identified B. alicastrum as the most important diet species.









In summary members of this guild members consume many of the same seed species,

some of which are important to the diet of most members of the guild. Common usage of

important seed species by guild members suggests that food resources were not limited. The

contrasting results of these two studies indicate that the importance of seed species varies with

time. Temporal variability in the importance of some seeds may reflect variability in production

due to fluctuations in annual precipitation. To illustrate, Peters (1989) found that fruit

production of B. alicastrum is strongly correlated with precipitation. Local meteorological data

indicated mean annual variations in precipitation of 200mm (16% of annual mean) with extremes

of as much as 800mm (64% of annual mean).

Guild Habitat Characteristics

Overall guild occupancy was greatest in Upland forest, which may reflect a greater

volume of habitat space relative low-canopy Scrub forest. Habitat occupancy patterns of O.

vetula, C. rubra, and P. purpurascens were consistent between transect groups and annual

periods, and highly skewed towards a particular forest-class, indicating habitat selection by these

species. Preferential habitat use was exhibited by all six cracid species in a similar guild in

Bolivia (Wallace et al. 2001). Preferential occupancy of Scrub forest by 0. vetula, and of

Upland forest by C. rubra and P. purpurascens were consistent with habitat-use descriptions for

the species (Gonzalez et al. 1998; Howell & Webb 1995; Brooks et al. 2004). Open, grassy

clearings have been identified as important habitat for reproductive activities and feeding for M

ocellata, and are known to be important brooding habitat forM. gallopavo in North America

(Steadman et al. 1977; Gonzalez et al. 1998; Williams & Austin 1988). Open, grassy clearings

are almost nonexistent in the study area therefore local habitat may not be optimal forM.

ocellata.









Overlap values calculated using the proportional similarity index are expected to be

conservative (Krebs 1989). The two species that exhibited the strongest and most divergent

habitat preferences were 0. vetula and C. rubra, which subsequently had the lowest combined

habitat overlap among species-pair with 50%. The species-pairs with the most similar patterns

of habitat use, M. ocellata-T. major and C. rubra-P. purpurascens, shared the highest overlap

values in the guild at 94%. Based on that range, three other species-pairs (P. purpurascens-T

major, M. ocellata-P. purpurascens, and C. rubra-T. major) had high combined habitat overlap

(85-91%), two species-pairs (M ocellata-O. vetula, C. rubra-M. ocellata) had intermediate

overlap (71-79%), and two species-pairs (0. vetula-P. purpurascens, 0. vetula-T major) had

low overlap (56-65%).

All species that exhibited habitat preferences also exhibited greater variability of

occurrence in the alternative forest-class. M. ocellata exhibited the most variability in habitat

occupancy and the least. C. rubra, 0. vetula, and P. purpurascens exhibited similar variability

of habitat occupancy. Despite variability of habitat occupancy among all species, consistent

temporal patterns of differential habitat occupancy were not exhibited by any guild member.

The latter finding is in contrast with seasonal habitat shifts reported forM ocellata by Gonzalez

et al. (1998).

Consistent and distinct patterns of vertical strata occupation indicate vertical stratification

within the guild, which is considered to be important in tropical bird communities (Karr & Roth

1971; Pearson 1971). The vertical patterns of strata occupation exhibited by guild members

were consistent with descriptions for these species (Howell & Webb 1995; Brooks et al. 2004).

Relevance of Competition to Guild Structure

Diet compositions, high diversity of dietary resources, and significant dietary overlap

among most species-pairs indicate generalist diet niches for all guild members. The hypothesis









that guild members have specialized dietary niches is not supported by the data. The only

evidence of differential dietary resource use in the guild were the proportionally greater leaf

material in the diets ofM. ocellata and 0. vetula, and the low overlap values between species-

pairs 0. vetula-M. ocellata and C. rubra-0. vetula. The hypothesis that the guild members

exhibit regular spacing along a gradient of dietary resources is not supported by the comparisons

of guild member diets.

Differential habitat preferences between 0. vetula and the two larger cracids indicate

specialized use of habitat by these three species. The limited habitat preference exhibited by T.

major and the lack of preference exhibited by M ocellata suggest generalist habitat resource use

by these species. The hypothesis that the guild members have specialized habitat niches is only

partially supported by the data. Less selective use of habitat by M. ocellata and T. major

increased the representation of the guild in Scrub forest, and may represent deliberate spatial

separation from C. rubra and P. purpurascens populations in Upland forest. Overall habitat

occupancy patterns among all species indicate spatial separation along a gradient of habitat

resources. The hypothesis that the guild exhibits regular spacing along the gradient of habitat

resources is supported by comparisons of habitat occupancy patterns.

The hypothesis that guild members would have specialized niches with respect to the

vertical occupation of forest strata is supported by the consistent patterns exhibited by all

species. The hypothesis that the guild exhibits regular spacing along a vertical gradient of forest

strata is supported by the vertical stratification observed within the guild.

Size ratios in a community structured by competitive interactions are predicted to be

either constant, have minimum values from 1.6-2.0:1 (natural logarithm values between 0.405

and 0.683), or else to increase with species size (Hutchinson 1959; Diamond 1962; Oskanen et









al. 1979). Ratios of size differences between guild members are not constant and decrease

among the largest three species. The size differences between species-pairs P. purpurascens-T.

major (0.681) and T. major-O. vetula (0.609), were similar and also consistent with predicted

minimum differences. The size differences between species-pairs M. ocellata-C. rubra (0.209)

and C. rubra-P. purpurascens (0.384), are not similar to one another or to the size differences

between other species-pairs, and are less than predicted minimums in assemblages structured by

competitive interactions. The hypothesis that size assortment of the guild is sufficient to mitigate

competitive interactions is supported by size differences among half of the guild species.

C. rubra, M. ocellata, and P. purpurascens populations exhibited significant reductions

along the increasing hunting pressure gradient. 0. vetula and T. major abundance did not

increase significantly nor did they exhibit changes in vertical strata occupancy in the transect

group where C. rubra, M. ocellata, and P. purpurascens populations were most reduced. T

major exhibited a relative shift in habitat occupancy from Scrub forest in the two groups where

hunting pressure was light, to Upland forest in the two groups where hunting pressure heavy.

Considering that T. major had the lowest relative abundance in the transect groups where hunting

pressure was light and exhibited the highest overlap with all other species in dietary and habitat

resources, it may experience the greatest competitive pressure from the other species. T major

was the only species that exhibited an inter-annual density decrease in the Cedro group, where

M. ocellata concurrently exhibited extraordinary density increases, which may indicate

competitive inhibition of T. major by M. ocellata. These two species had the highest overlap in

the guild along diet and vertical niche dimensions as well as relatively high dietary overlap. The

hypothesis that guild members would exhibit competitive release in response to the decreased

abundance of other species is partially supported by the data.









Although guild members exhibited variable nesting period lengths, reproductive activities

peaked during the same time of year for all guild members. The hypothesis that guild members

may stagger reproductive activities to avoid competition was not supported from the temporal

nesting patterns exhibited by the guild.

Aside from the five existing guild members, the only potential guild members available

within a 200 km radius are the Horned Guan (Oreophasis derbianus), the Highland Guan

(Penelopina nigra), and the White-bellied Chachalaca (Ortalis luecogastra). The two guans are

not legitimate null pool species because they are highland species with lower distribution limits

at elevations of 1000 m for Penelopina and 2000 m for Oreophasis. Although 0. luecogastra

exists at low elevations, regional Ortalis species were previously considered con-specific, and

current taxonomic descriptions reflect distinct geographic distributions (Howell & Webb).

Because regional Ortalis species are geographically separate, they may not qualify as legitimate

null pool species capable of colonizing a distinct geographic area. 0. luecogastra is confined to

the Pacific slope of the isthmus, which is separated from the study area by mountain ranges

along the entire length of the species distribution, so the absence of the species in the study area

is not likely to be the result of competitive exclusion. A regional null pool including O.

luecogastra would consist of only six species, in which case the local absence of 0. luecogastra

could be attributable to competitive exclusion by 0. vetula. If 0. luecogastra is excluded from

the regional null pool, then all available potential guild members are represented locally.

Competitive interactions within the guild do not appear sufficient to exclude regionally available

guild members except possibly in the case of 0. luecogastra. The hypothesis that the guild

would include fewer similar species than are regionally available is not supported based on null

pool comparisons.









Guild characteristics are partially consistent with predictions based on competition theory

in an equilibrium system. Guild members exhibited niche specialization with respect to vertical

occupancy of forest strata, and to some degree with respect to habitat use, but did not exhibit

niche specialization with respect to the use of dietary resources. Guild members appeared to be

ecologically separated along habitat resource and vertical forest strata gradients, but not along

the gradient of dietary resources. Size differences appeared to be sufficient to mitigate

competitive interactions for 0. vetula, T. major, and P. purpurascens, but not for C. rubra, M.

ocellata, and P. purpurascens. Overlap between C. rubra, and P. purpurascens was relatively

high along all niche dimensions. Overlap between M. ocellata with both C. rubra and P.

purpurascens was relatively high with respect to habitat and dietary resources and low with

respect to vertical strata use. If competitive interactions were significant within the guild, C.

rubra, M. ocellata and P. purpurascens would be expected to exhibit greater specialization and

separation along the other niche dimensions. T. major exhibited inverse responses to variations

of the abundance of other guild members. Guild membership was not notably different from the

regional null pool. Although some guild characteristics are consistent with an assemblage

structured by competitive interactions, the degree is less than would be expected based on

predictions from competition theory in an equilibrium system. The limited evidence of

competitive interactions displayed by the guild may be consistent with the non-equilibrium

competition model proposed by Connell (1980).

Relevance of Non-Equilibrium Factors to Guild Structure

Niches should evolve in response to mortality factors and dietary resources as well as

competition, therefore some niche specialization and ecological separation would be expected

whether or not competitive interactions are important to the guild. Guild member niches were

generalist with respect to diet, semi-specialized with respect to habitat, and specialized with









respect to vertical occupation of forest strata. The guild did not exhibit ecological separation

with respect to dietary resources, but did appear to be distributed along habitat and vertical strata

dimensions. The hypothesis that guild species would exhibit limited niche specialization and

separation along niche dimensions is supported by the niche characteristics of the guild.

Significant temporal variability of guild member abundance would constitute

environmental non-equilibrium. CVs of individual-encounter rates during annual and 30-month

periods represented double-digit percentages for all species. Although an objective basis of

comparison of the coefficient values is not available, these percentages appear to represent

significant variation in guild member abundance. CVs were consistently highest for C. rubra

and M. ocellata, which may reflect greater movement in foraging strategies. Significant local

temporal variation of curassow abundance has been attributed to movement between patchy fruit

resources (Thiollay 1989). CVs were relatively consistent between inter-annual and the 30-

month periods for all species except M ocellata, which had a much higher coefficient for the 30-

month period. This observation suggests that inter-annual variation may be greater than intra-

annual variation in this species. Inter-annual changes of guild member densities in the two, Rio

Azul transect groups also indicated significant variation of abundance for all species. The

hypothesis that guild members may exhibit significant variation in abundance is supported by the

variability exhibited by individual-encounter rates and density estimates.

Temporal shifts in habitat use by bird species in response to microhabitat variability may

result in dynamic assemblages representing non-equilibrium environmental factors (Karr &

Freemark 1983). The CVs of species-encounter rates in Upland and Scrub forests indicated

considerable variation of habitat occupancy. The hypothesis that guild members may exhibit

significant variation in habitat occupancy is supported by the variation observed from species-









encounter rates in each forest-class. In summary the combination of limited niche specialization

and ecological separation and the high variation of species abundance and habitat occupancy, are

consistent with predictions of non-equilibrium systems.

Relevance of Predation to Guild Structure

If guild members are subject to significant or differential predation pressure, it should be

reflected by reproductive characteristics (Martin 1988a). Predation is the primary cause of nest

failure in most bird species (Cody 1971). Nest losses to predation have been reported as high as

80% for T. major (Brennan 2004) and almost 50% forM ocellata (Gonzalez et al. 1998).

Potential mammalian nest predators observed on the line-transects included coatimundi (Nasua

narica), raccoon (Procyon lotor), collared peccary (Tayassu tajacu), white-lipped peccary

(Tayassupecari), and spider monkey (Ateles geoffroyii). Other potential mammalian nest

predators occurring locally include: two large opossums (Didelphis spp.), nine-banded armadillo

(Dasypus novemcinctus), ring-tailed cat (Bassariscus sumichrasti), and striped hog-nosed skunk

(Conepatus semistriatus) (Garcia & Radachowsky 2003; personal observation). Although avian

nest predators may be less significant for large species, corvids have been reported to be minor

nest predators onM. gallopavo (Williams & Austin 1988). Also curassow predation on tinamou

nests has been documented (Santamaria & Franco 2000). Although evidence of avian nest

predation is not available locally, the study area supports diverse and abundant populations of

corvids and rhamphastids (toucans).

Spatial separation of nest-sites through differential vertical or habitat placement may

favor bird community diversity by mitigating predation pressure (Martin 1988a). The

combination of terrestrial nest placement by M. ocellata and T. major, arboreal nest placement

by C. rubra and P. purpurascens, and shrub-arboreal placement by 0. vetula indicates vertical

nest-separation within the guild. Although nest-site data collection was not systematic with









respect to habitat, there is evidence of selective habitat use within the guild. All species had

significant but variable proportions of their nests in Upland forest. C. rubra and P. purpurascens

nests were almost exclusively in Upland forest. M. ocellata, 0. vetula, and T. major were the

only species that nested in Scrub forest. Only M ocellata and 0. vetula exhibited substantial use

of agricultural habitat.

Predation on ground nests may be greater than in other strata in tropical forests (Loiselle

& Hoppes 1983; Martin 1993; Gibbs 1991). This phenomenon is supported by the relatively

higher proportions of nests lost to predation by M. ocellata and T. major in this study. The

perception of greater predation on ground nests in the tropics may be attributable to greater

general predation pressure in the ground stratum, and therefore on terrestrial species (Martin

1988a). Gonzalez et al. (1998) reported that 40% of the M ocellata hens that had successfully

nested were killed by predators within 15 days of leaving the nest, and poult survival for the

breeding season was only 15%. Predation on poults during the first two-weeks after hatching

may represent the greatest limit to population size in M. gallopavo (Williams & Austin 1988).

Increased clutch sizes of some terrestrial birds are adaptive traits for conditions of high

natural mortality (Martin 1988b). The significantly larger clutch sizes ofM ocellata and T.

major are consistent with the prediction of greater mortality on terrestrial birds. The diversity of

mating systems within the guild reflects the variation observed in clutch sizes. Curassows, guans

and chachalacas not only share similar, small clutch sizes but also, as monogamous breeders,

exhibit biparental care of young. T. major is polygynandrous; both males and females have

multiple partners, but uniparental care is provided by males (Brennan 2004). M. ocellata is

polygamous, with uniparental care provided by females. Polygynandry and polygamy, which are









life history traits consistent with high population turnover, may increase reproductive capacity or

may improve population response to environmental variability, and are life history.

Relative to other general categories of birds, galliforms have lower annual survival rates

and a higher rate of population turnover (Gill 1990). Although local impacts of predation on

guild populations could not be directly assessed, existing evidence suggests that natural

predation pressure on the guild may be significant. Potential predator species observed on the

line-transects included jaguar (Panthera onca), puma (Puma concolor), ocelot (Leopardus

pardalis), tayra (Eira barbara), and grey fox (Urocyon cineoargenteus). Other locally occurring

predators include margay (Leopardus weidii), j aguarundi (Herpailurus yagouaroundi), ornate

hawk-eagle (Spizaetus ornatus), black hawk-eagle (Spizaetus tyrannus), crested eagle (Morphnus

guianensis), and solitary eagle (Harpyhaliaetus solitarus) (Garcia & Radachowsky 2003,

personal observation).

The only information regarding the local contribution of guild members to predator diets

is from research on jaguar and puma conducted on the same study area (Novack 2003). The

reported biomass proportions of "mostly cracids" were approximately 4% and 6% of jaguar and

puma diet, respectively. Based on the daily caloric requirements of jaguar reported in that study,

and applying a 5% correction for non-edible parts, the average jaguar or puma consumes

approximately 28 kg of guild biomass annually, or the equivalent of about 8 adult C. rubra.

Applying the density estimates for jaguar and puma from Novack (2003) to the current study

area of approximately 1000 km2, the local populations of these two predator species consume

approximately 1,400 kg of "mostly cracids" annually. Considering that the human off-take from

the guild in the study area averaged 1,625 kg of biomass annually and appeared to have a









significant impact on local populations, the natural predation pressure on the guild appears to be

significant.

Alternative Influences on Guild Structure

The influence of other potentially relevant biotic factors whose impacts on the guild

could not be directly assessed included resource heterogeneity, disease, and parasitism. If

heterogeneity is significant to the guild, resource gradients should be diverse and species niches

should have greater dimensionality. The heterogeneity of the forest habitat and the broad diet of

the guild were consistent with predictions of diverse resource gradients. One potentially relevant

niche dimension that was not addressed was specialized daily-activity patterns of guild members.

Brooks et al. (2004) reported substantive nocturnal activity for T major in a diverse tinamou

community, indicating that other niche dimensions may be important to the guild. Anecdotal and

circumstantial support for the potential relevance of disease comes from local reports of an

historic regional crash in M. ocellata populations, and regional population crashes ofM.

gallopavo in North America (Williams 1981; personal observation). Parasitism may be

especially relevant in tropical environments (Kricher 1997), especially for terrestrial species,

which due to increased contact with the soil, often have greater parasite loads than arboreal

species (Gill 1990).

Guild Response to Hunting Pressure

The recorded harvest is not representative of all subsistence hunting pressure in the study

area. Although local participation in the collection effort was significant, there is no evidence

that all local hunters participated or that regular participants did so all the time. In particular,

harvest pressure from camp-based activities far from the village is unlikely to be well

represented in the study data. During the initial stages of specimen collection, 0. vetula and T.

major were poorly represented. Through the deliberate efforts of some regularly participating









hunters, collection rates for 0. vetula were increased significantly, although rates for T major

remained low throughout the collection period. Relative to the actual impacts of local

subsistence hunting, the number of individuals harvested is expected to be slightly conservative

for C. rubra, M. ocellata, P. purpurascens, and T major, and robust for 0. vetula. Because the

proportion of sub-adult birds harvested could not be determined, the application of average adult

body-mass values to the recorded harvest for each guild species probably overestimated the

recorded harvest biomass. However, additional harvest pressure attributable to camp-based

extraction activities (McNab 1998; Morales & Morales 1998) may compensate for any

overestimation.

The harvest data indicated a disproportionately greater mean annual harvest of C. rubra

(47%) than of other guild members, which are consistent with previously findings from the area

(Morales & Morales 1998). Mean annual harvest proportions were intermediate and relatively

similar for 0. vetula (20%), M. ocellata (16%), and P. purpurascens (12%), and were

consistently low for T major (5%). These harvest levels were inconsistent with previous

findings, which indicated that the harvest of P. purpurascens exceeded that ofM. ocellata by a

factor of more than 2:1, and the harvest ofM. ocellata exceeded that of 0. vetula by a factor of

more than 5:1 (Morales & Morales 1998).

Selectivity for C. rubra, M. ocellata, and P. purpurascens by local hunters is consistent

with population trends along the hunting gradient. Based on the changes observed in abundance

estimates and guild composition between transect groups, local hunting pressure appeared to

have significant impacts on populations of these three species. Although 0. vetula and T. major

abundance was greatest in the protected Ixcan transect group, abundance among the other three

groups did not appear to be related to distance from the village of Uaxactun.









The combined population size of the guild in the heavily hunted Uaxactin South group

(1,930 individuals) represented 45% of the combined guild population size in the protected Ixcan

group (4,332 individuals). Estimated population sizes for Uaxactun South equated to 17% of the

Ixcan population for C. rubra, 28% forM ocellata, 89% for 0. vetula, 31% for P. purpurascens,

and 79% for T. major. Population biomass estimates were calculated using average adult body

mass values and were therefore robust; however, these estimates should adequately reflect the

relative differences between populations and sites. The combined biomass of the guild

populations in the Uaxactin South group (3,215 kg) represented 30% of the combined biomass

in the Ixcan group (10,587 kg). Based on the changes observed in the guild along the gradient,

local subsistence hunting pressure has significantly reduced local guild populations.

Conclusions

Guild characteristics that were consistent with predictions of structural regulation by

competitive interactions included: separation along habitat dimensions, vertical stratification,

limited size assortment, and apparent competitive release by T. major. Guild characteristics that

were inconsistent with regulation by competition included: lack of diet-niche specialization,

considerable dietary overlap, limited specialization of habitat niches, and the similarity of guild

membership to the regional null pool. Additional evidence contrary to regulation by competition

derives from the high abundance and inter-annual increase all guild members in the protected

Ixcan transect group. If the fitness of individual guild members is constrained by competitive

interactions with other guild members, then it is counterintuitive that all species would exhibit

greatest abundance in the same area at the same time, and that all species would exhibit

abundance increases in the same area during the same time period.

Guild characteristics that were consistent with non-equilibrium processes included: limited

niche specialization, limited ecological separation, and the high variability of species abundance









and habitat occupancy. Guild characteristics that were consistent with external limitations by

predation included: spatial separation of nest sites and high reproductive potential of M. ocellata

and T. major. Based on limited evidence of competition, evidence of non-equilibrium processes,

and evidence of the importance of predation, the characteristics of the guild are most consistent

with the non-equilibrium model of competition proposed by Connell (1980). The limited niche

specializations and separation along resource dimensions may be sufficient to minimize direct

interactions among species. The unpredictable local abundance of competitor species and

external limitation of populations by predation create conditions under which dietary

generalization is more adaptive than dietary specialization. Guild-member populations are

unlikely to reach the carrying capacity of the environment, therefore, competitive interactions are

held below levels that would lead to the competitive exclusion of potential guild members.









Table 3-1. General diet compositions of guild members based on dry-mass proportions of
contents from upper-digestive tract specimens collected in Uaxactun, Flores, El
Peten, from January 2000 through December 2001.
Diet compositions with all sample contents
Crax Meleagris Ortalis Penelope Tinamus
(n = 267) (n = 181) (n = 205) (n = 142) (n = 55)
Flower/Stem 0.004 0.008 0.013 0.002 0.001
Animal 0.015 0.121 0.007 0.011 0.024
Leaf 0.015 0.65 0.063 0.014 0.007
Pulp 0.066 0.07 0.068 0.087 0.065
Grit 0.204 0.222 0.027 0.093 0.022
Seed 0.696 0.514 0.822 0.793 0.881


Diet compositions with grit and snail contents excluded
Crax Meleagris Ortalis
Flower/Stem 0.005 0.012 0.013
Animal 0.016 0.011 0.006


Penelope Tinamus
0.002 0.001
0.012 0.024


Leaf 0.019 0.098 0.064 0.015 0.007
Pulp 0.083 0.105 0.07 0.096 0.066
Seed 0.877 0.774 0.847 0.875 0.902


Proportions include pulverized material detected in samples. The dry-mass of pulverized material was assigned to
the values of separable items in each sample according to their relative proportions.


Table 3-2. Overlap of dietary seed components between species-pairs and average seed
component overlap of individual species with all other guild members.
Species-pair Seed component Species Average
Overlap Overlapb
Crax Meleagris 57.9% Crax 62.6%
Crax -Penelope 67.9%
Crax -Ortalis 39.1% Meleagris 53%
Crax -Tinamus 85.6%
Meleagris -Penelope 57.4% Ortalis 46.8%
Meleagris -Ortalis 32.9%
Meleagris -Tinamus 63.6% Penelope 66%
Penelope -Ortalis 65.4%
Penelope Tinamus 73.3% Tinamus 68.1%
Ortalis -Tinamus 49.8%
a Calculated using equation five from Pianka (1973), pulverized material in samples was not included.
b Average seed component overlap with all other guild members.


I









Table 3-3. Overlap of habitat occupancy between species-pairs and average combined overlap of
individual species with all other guild members.
Species-pair Habitat Species Average
Overlap a Overlap b
Crax Meleagris 79.5% Crax 77.3%
Crax -Penelope 94.2%
Crax -Ortalis 50.3% Meleagris 82.6%
Crax -Tinamus 85%
Meleagris -Penelope 85.4% Ortalis 60.7%
Meleagris -Ortalis 70.8%
Meleagris -Tinamus 94.5% Penelope 81.7%
Penelope -Ortalis 56.2%
Penelope -Tinamus 90.9% Tinamus 83.9%
Ortalis -Tinamus 65.3%
a Calculated using proportional similarity index equation from Schoener (1970).
b Average combined habitat occupancy overlap with all other guild members.


Table 3-4. Overlap of vertical strata occupancy between species-pairs and average combined
vertical overlap of individual species with all other guild members.
Species-pair Vertical Species Average
Overlap a Overlapb
Crax Meleagris 43.5% Crax 56.4%
Crax -Penelope 73.8%
Crax -Ortalis 68.3% Meleagris 55.6%
Crax -Tinamus 40%
Meleagris -Penelope 17.3% Ortalis 41.8%
Meleagris -Ortalis 17.1%
Meleagris -Tinamus 93.1% Penelope 43.3%
Penelope -Ortalis 68.3%
Penelope -Tinamus 13.8% Tinamus 40.1%
Ortalis -Tinamus 13.6%
a Calculated using proportional similarity index equation from Schoener (1970).
b Average combined vertical overlap in both forest-classes with all other guild members.









Table 3-5. Density estimates for Crax rubra, Meleagris ocellata, Ortalis vetula, Penelope


purpurascens,
Guatemala.


and Tinamus major by forest-class in the Maya Biosphere Reserve,


Transect group Ixcan Ixcan Cedro Cedro Uaxactun Uaxactun
North South
Sampling 2000- 2001- 2000- 2001- 2000-2001 2000-2001
period 2001 2002 2001 2002
Upland forest densities (km2)
Crax 31.19 31.05 21.37 26.27 14.44 5.46
Meleagris 15.51 24.63 6.44 16.7 8.04 4.7
Ortalis 11.25 12.56 14.51 22.91 3.37 18.51
Penelope 24.31 28.29 16.43 15.78 7.27 6.73
Tinamus 8.44 14.64 5.46 5.16 10.96 9.13
Scrub forest densities (km2)
Crax 14.17 19.54 6.12 9.94 6.18 2.42
Meleagris 15.19 17.41 3.4 22.73 7.87 5.2
Ortalis 37.22 56.03 24.19 25.29 19.03 25.01
Penelope 11.11 15.81 6.12 6.28 4.56 6.32
Tinamus 10.37 8.97 6.8 6.54 6.39 4.51
Density estimates calculated using DISTANCE version 5, release "Beta 5".



Table 3-6. Guild compositions based on proportional population size and relative biomass at
each transect group.
Transect group Ixcan Ixcan Cedro Cedro Uaxactun Uaxactun
North South
Sampling 2000- 2001- 2000- 2001- 2000-2001 2000-2001
period 2001 2002 2001 2002
Proportional population size
Crax 0.28 0.24 0.317 0.289 0.236 0.106
Meleagris 0.172 0.167 0.098 0.197 0.181 0.109
Ortalis 0.227 0.247 0.248 0.264 0.25 0.455
Penelope 0.219 0.213 0.246 0.168 0.135 0.15
Tinamus 0.102 0.133 0.091 0.082 0.198 0.18
Proportional population biomass
Crax 0.391 0.351 0.464 0.399 0.355 0.216
Meleagris 0.294 0.302 0.177 0.336 0.334 0.274
Ortalis 0.059 0.068 0.068 0.069 0.071 0.175
Penelope 0.207 0.212 0.245 0.157 0.138 0.208
Tinamus 0.049 0.067 0.046 0.039 0.102 0.127
Population sizes were based on density estimates by forest-class extrapolated by the availability of each forest-class
within the 50km2 area corresponding to each transect group. Population biomass estimates were based on
population size and average adult body mass of each species.









Table 3-7. Temporal reproductive patterns of Crax rubra, Meleagris ocellata, Ortalis vetula,
Penelope purpurascens, and Tinamus major based on the number of monthly
observations of reproductive activities.
Category Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.
Crax rubra
Calls 8 14 38 36 46 30 8 4 1
Nests 8 1 5
Young 2 5 7 10 1 1 2
Meleagris ocellata
Calls 1 21 43 38 2 1
Nests 3 9 4
Young 1 1 4 1 2 2 2
Ortalis vetula
Nests 1 7 30 31
Young 3 5 7 5 3 3 1
Penelope purpurascens
Nests 3 3 4 3
Young 3 5 8 1
Tinamus major
Nests 1 6 9 12 1 1
Young 2 3 6 8 3
Calls include records of vocalizations specific to reproductive behavior which were recorded only for C. rubra and
M. ocellata. Nests include only observations of nests with laying or incubating adults present. Young include
observations of dependent young and hatched nests observed during the nesting period.















































Figure 1-1 .Map of study area. Lightly shaded areas indicate the two management units in which
study data were collected. Circles depict the 50km2 areas corresponding to each
transect group.













56









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BIOGRAPHICAL SKETCH

At his point I have lived roughly equal proportions of my life in Texas, New Jersey, and

Guatemala. I received a B. S. in zoology and another in wildlife management from Texas Tech

University in Lubbock, Texas. After graduating from Texas Tech I entered the U. S. Peace

Corps and was stationed in Guatemala where I worked with non-governmental organizations

working in the two largest protected areas in that nation, first in the Sierra de las Minas

Biosphere Reserve and later in the Maya Biosphere Reserve. My interests in terrestrial

vertebrate ecology and sustainable wildlife resource-use lead me to pursue graduate studies at the

Department of Wildlife Ecology and Conservation at the University of Florida. Since 1999 I

have co-developed and managed the Guatemalan operations of a community-based, sport-

hunting, conservation project for the Ocellated Turkey working in community forestry

concessions in the multiple-use zone of the Maya Biosphere Reserve.





PAGE 1

1 STRUCTURE OF A LOWLAND NEOTRO PICAL GALLIFORM BIRD GUILD By ERICK HOGAN BAUR A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2008

PAGE 2

2 2008 Erick Hogan Baur

PAGE 3

3 To my family, Michaelyn, Milano, and Chloe Lilyana

PAGE 4

4 ACKNOWLEDGMENTS I thank m y committee chair Ronald F. Labisky, who set the gold-standard for patience, and also committee members David Steadman, George Tanner, and especially Daniel Brooks for his unfailing enthusiasm. The Department of Wildlife Ecology and C onservation of the University of Florida, the Wildlife Conservation Society, and the African Safari Club provided financial support for this research. I thank Roan McNab, Am erica Rodriguez, and th e Guatemalan staff of the Wildlife Conservation Society for their suppo rt, and also John Polisar and Robin Bjork for counsel and kindness early in this process. I thank Anthony Novack, whose collaboration and friendship along the way helped make the adversities more sufferable a nd the successes more significant. In Uaxactn I am grateful to Pa blo Nuez, Antonio Ramos, Victor Mendez, Manuel Mendez, Amilcar Fajardo, Aurora Soza Duran, Gu illermo Calanclan, and the many participants in data collection efforts there. The seemi ngly endless processing of dietary specimens was bearable thanks to the assistance of Millicen t Butterworth, Miguel Si, Lucas Cuc, and Enrique Barahona. I owe a debt of gratitude to Nery Solis and Victor Hugo Ramos of the Monitoring and Evaluation Center of the National Protected Area Council, and to Pe dro Pineda of th e Institute of Agriculture, Natural Resources, a nd the Environment at Rafael La ndivar University. Thanks to Oscar Lara, Julio Morales, and Javier Rivas of the School of Biology at the University of San Carlos for support with permitting and specimen iden tification. Salvador and Libertad Brizuela provided sanctuary during the writing stage of this process. Thanks to Kitty Emery, Delores Tillman, and Susana de Radachowsky for overs eas assistance and to Isora Labisky, Greta McNab, Milan Hooper, and Erik and Claire Lewis for their generous hospitality. I am most grateful to my wife Michaelyn Bachhuber and to my children M ilano and Chloe Lilyana for all of their support, patience, and encouragement.

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........7 LIST OF FIGURES.........................................................................................................................8 ABSTRACT.....................................................................................................................................9 CHAP TER 1 INTRODUCTION..................................................................................................................11 Regional-Level Diversity Hypotheses....................................................................................14 Local-Level Diversity Hypotheses......................................................................................... 15 Objectives...............................................................................................................................17 Conservation Threats..............................................................................................................19 Study Area..............................................................................................................................20 2 METHODS.............................................................................................................................23 Diet and Harvest Data.............................................................................................................23 Line-Transect Data............................................................................................................. ....24 Habitat Use.................................................................................................................... .........25 Vertical Strata Occupancy......................................................................................................26 Species Abundance and Guild Composition.......................................................................... 26 Nest Site Data.........................................................................................................................27 3 RESULTS...............................................................................................................................29 Diet Composition and Overlap............................................................................................... 29 Harvest Description............................................................................................................ ....29 Size Assortment......................................................................................................................30 Habitat Use.................................................................................................................... .........30 Vertical Strata Occupancy......................................................................................................31 Species Abundance and Guild Composition.......................................................................... 32 Nest Site and Clutch Characteristics....................................................................................... 34 4 DISCUSSION.........................................................................................................................36 Guild Diet Characteristics..................................................................................................... ..36 Guild Habitat Characteristics.................................................................................................. 38 Relevance of Competition to Guild Structure........................................................................ 39 Relevance of Non-Equilibrium Factors to Guild Structure.................................................... 43 Relevance of Predation to Guild Structure............................................................................. 45

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6 Alternative Influences on Guild Structure.............................................................................. 48 Guild Response to Hunting Pressure...................................................................................... 48 Conclusions.............................................................................................................................50 LIST OF REFERENCES...............................................................................................................57 BIOGRAPHICAL SKETCH.........................................................................................................64

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7 LIST OF TABLES Table page 3-1 General diet com positions of guild members based on dry-mass proportions of contents from upper-digestive tract specime ns collected in Uaxactn, Flores, El Petn, from January 2000 through December 2001........................................................... 52 3-2 Overlap of dietary seed com ponents between species-pairs and average seed component overlap of individual spec ies with all other guild members........................... 52 3-3 Overlap of habitat occupancy between speci es -pairs and average combined overlap of individual species with all other guild members........................................................... 53 3-4 Overlap of vertical strata occupancy be tween species-p airs and average combined vertical overlap of individual spec ies with all other guild members................................. 53 3-5 Density es timates for Crax rubra Meleagris ocellata Ortalis vetula Penelope purpurascens and Tinamus major by forest-class in the Maya Biosphere Reserve, Guatemala...................................................................................................................... ....54 3-6 Guild com positions based on proportional population size and relative biomass at each transect group............................................................................................................ 54 3-7 Temporal reproductive patterns of Crax rubra, Meleagris ocellata Ortalis vetula Penelope purpurascens and Tinamus major based on the number of monthly observations of reproductive activities..............................................................................55

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8 LIST OF FIGURES Figure page 1-1 Map of study area. Lightly shaded areas indicate the two m anagement units in which study data were collected. Circles depict the 50km2 areas corresponding to each transect group.....................................................................................................................56

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9 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science STRUCTURE OF A LOWLAND NEOTRO PICAL GALLIFORM BIRD GUILD By Erick Hogan Baur May 2008 Chair: Ronald F. Labisky Major: Wildlife Ecology and Conservation This study (1) examined guild structure, (2) co mpared guild characteristics to predictions from potentially relevant hypotheses of guild structure regulation, and (3) assessed the impacts of hunting on a guild comprised of the Great Curassow ( Crax rubra), Crested Guan ( Penelope purpurascens ), Plain Chachalaca ( Ortalis vetula ), Ocellated Turkey ( Meleagris ocellata ), and the Great Tinamou ( Tinamus major ) in 2000-2002. The study area, located in the Maya Biosphere Reserve in northern Guatemala, included a nati onal park unit with prot ected populations of the guild and an adjacent community concession unit subject to a range of subsistence hunting pressure. Diet niches and overlap were describe d from analyses of upper digestive tract contents from 267 C. rubra 181 M. ocellata 205 O. vetula, 142 P. purpurascens and 55 T. major collected over a 24-month period. Descriptio ns of habitat niches and overlap, species abundance, and selected reproduc tive parameters were derived from line-transect sampling during a 30-month period in the park unit (1,770 km total) and over a 12-month period in the concession unit (995 km total). Descriptions of nest-sites, clutch-sizes and other reproductive characteristics were based on observations made at 24 C. rubra nests, 39 M. ocellata nests, 77 O. vetula nests, 19 P. purpurascens nests, and 66 T. major nests.

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10 Dry-mass proportions of gr it were highest and similar in the diets of M. ocellata (22%) and C. rubra (20%), lowest and similar in O. vetula (2.7%) and T. major (2.2%), and intermediate in P. purpurascens (9%). Seeds and pulp represented the greatest and second-greatest proportions respectively, of the diet of all species when grit and snail shells were excluded from analyses. Overlap in the seed component of the diet range d from 50-86% for eight of the ten species-pairs, and the average combined overlap exceeded 50% for all species except O. vetula Habitat preferences were exhibited by C. rubra and P. purpurascens for tall-forest, and by O. vetula for low-forest. T. major occurred at a greater frequency in lowforest in the two areas with the least hunting pressure, and in high-forest in the two areas with the greatest hunting pressure. The frequency of occurrence of M. ocellata did not differ among habitat t ypes. All species exhibited consistent vertical patter ns of strata occupancy. Guild characteristics were semi-consistent with structural regulation through competitive interactions with respect to habitat niches, body-size assortment, and evidence of competitive release, but were inconsistent with respect to diet niches and nu ll-pool comparisons. The variability of habitat occupancy and abundan ce within the guild we re consistent with expectations for non-equilibrium processes. Vert ical stratification of ne st-placement and other reproductive attributes of the guild were consiste nt with expectations for a guild subject to intense predation pressure. The guild appears to be a terrestrial vertebrate example of the nonequilibrium competition model of community st ructure proposed by Connell (1980). All species were most abundant in the fully-prot ected area; however, the abundance of C. rubra M. ocellata and P. purpurascens declined consistently along the incr easing gradient of hunting pressure. Guild biomass in the most heavily hunted area represented 30% of the guild biomass in the protected area.

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11 CHAPTER 1 INTRODUCTION This study exam ines the structure of a lowla nd tropical bird guild composed of the Great Curassow (Crax rubra), Crested Guan ( Penelope purpurascens), Plain Chachalaca ( Ortalis vetula ), Ocellated Turkey ( Meleagris ocellata ), and Great Tinamou ( Tinamus major ) in the El Petn department of Guatemala. These sp ecies share ecological and taxonomical or morphological similarities, and a limited size ra nge that make them likely competitors (Root 1967; MacNally 1983). Three of the species, C. rubra P. purpurascens and O. vetula, are members of the family Cracidae, a Neotropical galliform family represented by 11 genera and 50 species that reaches maximum diversity in Sout h America (Brooks & Strahl 2000). These three species are representative of the general size range of the family; Ortalis includes small cracid species, Crax large species, and Penelope intermediate-sized species. The largest species in the guild, M. ocellata is also a galliform and the only trop ical representative of the new world subfamily Meleagridinae. T. major belongs to the Neotropical fa mily, Tinamidae which is most closely related to ratites. Tinamidae is repres ented by 9 genera and 47 species and also reaches greatest diversity in South Am erica (Bertelli & Porzecanski 2 004). Although not a galliform, T. major was included in this study because tinamids ar e ecologically similar to galliforms, and its size is consistent with th e range of the guild (Thom pson 2004; Brooks et al. 2004). These species exist sympatrically in the fore st-dominated landscape of the study area and share similar feeding habits. Although cracids have been described as frugivores (Karr 1971; Thiollay 1994) or granivores (Terborgh et al. 1 990), diet studies have r ecognized the significance of seed, leaf, and animal materials (Santamari a & Franco 2000; Jimenez et al. 2001; Mamani-F 2001; Rivas & Morales 2003). Diet descriptions of M. ocellata indicate an omnivorous diet

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12 dominated by seeds (Steadman et al. 1979). T. major diet has been described as granivorous (Thiollay 1994) or omnivorous (Howell & Webb 1995). C. rubra is considered to function primarily as a seed predator on large-seeds and to a lesser degree as a disperser of small seeds (Santamaria & Fran co 2000; Rivas & Morales 2003). Guans and chachalacas are considered to have less er roles as seed predat ors and greater roles as dispersers than curassows (Brooks & Strahl 2000; Rivas & Morales 2003). M. ocellata has a muscular gizzard and consumes grit (Steadman et al. 1979) and likely functions more as a seed predator than a disperser. An assess ment of the impacts of the diet of T. major was not available; however, due to its intermediate size relative to guans and chachalacas, the species is probably a lesser seed predator and greater disper ser than the larger guild members. Guilds of large galliform species are not well represented in literature regarding bird community structure and diversit y. This under-representation may reflect less interest in generalist species, which may be less suitable for niche dimensi on research which has dominated community ecology, or may reflect vulnerability to hunting (Schoener 1983; Ricklefs 1990; Terborgh et al. 1990). Large galliform species ar e often reduced or extirpated where they are accessible, including sites where some of the most comprehensive field research of bird communities have been c onducted (Karr 1971; Karr 1976 b; Thiollay 1989: Terborgh et al. 1990). The Cracid Specialist Group of the Intern ational Union for the Conservation of Nature and Natural Resources (IUCN) desc ribes Cracidae as the most threatened avian family in the Neotropics, ranking 24 of its 50 species in need of priority conservation efforts (Brooks & Strahl 2000). Although generally less abundant than smaller bird species, large galliforms may become common where protected (Thiolla y 1989; Silva & Strahl 1991). Under natural conditions these species may represent significant proportions of the biomass of bird communities (Terborgh et

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13 al. 1990; Redford 1992), and may qualify as keystone trophic components of tropical communities (Terborgh 1992). The combination of poor scientific representation, vulnerability, threatened status, and ecological significance of large galliform species emphasizes the need for research on such guilds where they persist under unexploited conditions (Terborgh et al. 1990; Thiollay 1994; Brooks & Strahl 2000). Rahbeck and Graves (2001) cited 120 existing causal hypotheses of bird species-richness within the context of presenting yet a new model for predicting bird species-richness patterns. The tendency of tropical ecosystems to be more di verse than temperate systems, or latitudinal diversity gradient, may be the longest recogniz ed pattern of species-r ichness (Hutchinson 1959; Pianka 1966; Gill 1990; Ricklefs 1990). Many hypot heses proposed to account for the structure of ecological communities, bird species-richness pa tterns, and the latitudinal diversity gradient are potentially relevant to the di versity and structure of this guil d. Assessment of the relevance of any particular hypotheses is complicated by a variety of factors. Different hypotheses may predict similar local effects, may identify causal factors that are not mutually exclusive, or that may counteract one another. The interpretation of relationships among hypotheses varies among authors, and the same research findings are cited in support of distinct hypotheses among authors (Pianka 1966; Begon et al. 1990; Ricklefs 1990; Kricher 1997). Regional-level hypotheses of tropi cal diversity generally emphasize the roles of abiotic or historical factors operating on tempor al or spatial scales that are not amenable to testing based on the results of a single-site field study. Local -level tropical diversity hypotheses generally emphasize the causal roles of biotic factors or en vironmental variability. Biotic regulation of community structure may manifest as interspeci fic competition, external population controls, or resource heterogeneity (Begon et al. 1990; Ricklefs 1990). Lo cal-level hypotheses generally

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14 diverge with respect to variability, either regarding communities as equilibrium systems regulated by deterministic processes, or as non-equilibrium systems in which environmental factors counteract determin istic processes (Begon et al. 1990; Ricklefs 1990). Regional-Level Diversity Hypotheses The tim e hypothesis attributes greate r tropical diversity to the hi storic stability of tropical biomes relative to temperate biomes subjected to glaciation during the Pl eistocene (Begon et al. 1990; Ricklefs 1990). Evidence contrary to this hypothesis include: findings that tropical biomes were also subject to historic periods of expansion and contrac tion (Haffer 1969), that temperate biomes may be as old as tr opical biomes (Ricklefs 1990) and that speciation can occur on temporal scales that are insignificant relativ e to biome age (Remsen 1990). Support for this hypothesis includes findings of co mparisons of African and Neot ropical bird communities in similar forest and savannah biomes in which bi rd community diversity was positively correlated to the relative age and historic extension of biomes (Karr 1976 b). The climatic stability and the productivity hypotheses attribute tropical diversity to reduced variability of abiotic factors such as temperature, precipita tion, and solar radiation (Pianka 1966; Sanders 1968; Begon et al. 1990). Both hypotheses credit abiotic factors operating on regional levels with increased local diversity. Stable tropical climates are favorable to many plant and poikilothermic faunal taxa that in turn provide diverse and consistently available resources to consumers (Pianka 1966; Kricher 1997). The productivity hypothesis emphasizes the role of tropical stability in reducing physical stress on producers, permitting greater investment in non-maintenance activities such as reproduction, which results in greater abundance of producers and available res ources to consumers (Connell & Orias 1964).

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15 Local-Level Diversity Hypotheses The regulation of community structu re through competition has been a dominant paradigm in community ecology (Schoener 1974 ; Schoener 1983; Gill 1990; Ricklefs 1990). The competition hypothesis contends that when fauna l communities are in a state of equilibrium, species will occupy distinct ecological niches (H utchinson 1959). Greater local diversity in tropical systems increases interspecific competition, which becomes the most significant determinant of community structure. When co mmunity structure is re gulated by competitive interactions, niche specialization is predicted to be prevalent among member species. Increased niche specialization permits ecological separation of competitors, which are expected to exhibit regular spacing along niche dimensions (Ricklef s 1990). According to this theory, modern tropical assemblages are divers e because increased niche speci alization has resulted in narrower niches, thus accommodating greater nu mbers of species along resource gradients (Ricklefs 1990; Begon et al. 1990). Because mode rn assemblages represent species that have resolved historic competitive conflicts via co -adaptation, competition for resources may not occur except under conditions of extreme resour ce scarcity (Wiens 1977; Fleming 1979). The significance of competition may vary among trophic le vels, with greater relevance to producers, decomposers, and predators than to herbivores, frugivores, and some omni vores (Hairston et al. 1960; Fleming 1979). An alternative role of competition in the regu lation of community structure predicts that competitive interactions in non-equilibrium system s will lead to greater niche generalization and overlap (Connell 1980). Competitive interactions are inconsistent when competitor abundance is highly variable or under conditions of diffuse-competition. Individuals of a given species may interact with distinct competitor species, or may not face consistent competition from any particular competitor species (Buckley 1983). This hypothesis contends that under such

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16 conditions, species are more li kely to develop evolutionary responses to food acquisition efficiency or mortality factors than to the niches of competing species. Biotic environmental factors that maintain populations below carrying capacity create non-equilibrium conditions that prevent competitive interactions from reaching the point of competitive exclusion (Pianka 1966; Ricklefs 1990). The tropical diversity gradient applies to predators, parasites, and pathoge ns just as it does to other sp ecies. Predators often exhibit frequency-dependent selection defined as the tendency to se lect the most abundant forms of prey. Parasites and pathogens ar e likely to increase in correlati on to increased host populations. Because competitive exclusion does not occur under such conditions, communities may become more diverse (Ricklefs 1990; Kricher 1997). In a ssemblages of populations that are subject to external biotic controls it is predicted that species will exhibit lesser niche specialization and greater resource use overlap (Pianka 1966). Abiotic environmental factors may create non-equilibrium conditions by reversing successional vegetation trends or by increasing resource variab ility. Forest-gap dynamics increase the heterogeneity of tropical forest diversity and st ructure (Connell 1978 ; Schemske & Brokaw 1981). Large rivers that flood seasonally maintain vegetation along the banks in early successional stages, increasing habitat heterogeneity and favoring greater bird diversity (Remsen & Parker 1983; Brooks et al. 2004). Highly variable precip itation patterns may reduce the availability or consistency of resources, li miting community diversity and precluding the development of niche specializations for particular resources (Karr 1976 a ). Greater local diversity in the tropics may cr eate resource gradients that are sufficiently heterogeneous to support a greater diversity of consumers (Hutchinson 1959; Kricher 1997). If resource heterogeneity is significant to communi ty structure it is predicted that niches will

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17 exhibit greater dimensionality (Ricklefs 1990). Habitat heterogeneity has been positively associated with bird species diversity in bot h tropical and temperate climates (MacArthur & MacArthur 1961; Karr & Roth 1971). The diversity of some tropical bird communities has been attributed to greater diversity of frugivorous, insectivorous, and nectivorous species, reflecting utilization of resources that are more divers e and abundant in the tr opics (Pianka 1966; Karr 1971; Terborgh 1990). Objectives The prim ary objective of this study was to examine guild structure in a manner that would permit the assessment of re levant regulatory processes to the guild. Specific objectives were (1) to determine the degree of specialization of principal niche dimensions within the guild, (2) to determine the degree of ecological separation or overl ap along corresponding resource gradients, and (3) to compare guild characteri stics to predictions de rived from potentially relevant hypotheses of structural regulation. The secondary objective of this study was to assess the impacts of hunting pressure on guild structure and composition. Niches were described along certa in dietary and habitat dimens ions. Diet is considered the most essential niche dimension among terre strial vertebrates (Schoener 1974; Schoener 1983) and dietary specialization has been documented in many tropical bird guilds (Ashmole 1968; Sherry 1984). Spatial separation along habitat resource gradients has been associated with the diversity of many bird communities (Cody 1968; Gill 1990). Habitat resources may be partitioned through preferential use, temporal vari ability of use, and ver tical stratification (Gill 1990; Pearson 1971; Karr 1971). Other guild characteristics that were co mpared to predictions based on potentially relevant hypotheses included size assortment, responsiveness of guild members to abundance variations of other guild species guild membership, variability of species abundance and habitat

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18 occupancy, and reproductive strategies. Size assortment is a mechanism of mitigating interspecific competition that occurs when combin ations of similar species with distinct sizes persist better than similar-si zed species (Case & Sidell 1983). Size assortment has been associated with the greater dive rsity of some tropical bird gu ilds (Ashmole 1968; Brooks 1998). If guild member populations are constrained by competitive interactions, then significant reductions of some species may offer an opportu nity for competitive re lease to other guild members. The use of a greater range of habitat resources in the absence of competitors has been documented in some tropical bird communities (M acArthur et al. 1966). Bird populations are not static and significant tempor al variations of abundance or habitat use may represent nonequilibrium factors that reduce the predictability of competitive interactio ns and counteract local processes of competitive exclusion (Karr & Freemark 1983; Gill 1990; Loiselle & Blake 1992). Vertical stratification of nest -placement is a mechanism of reducing nest-predation pressure through increased spatial dist ribution of nests (Martin 1988a ; Martin 1993; Schmidt & Whelan 1998). Temporal separation of reproductive activ ities may mitigate interspecific competition for nesting or brooding resources in bird communities (Ricklefs 1966). Comparisons of guild composition to null pools of potential members that are regionally available have been used to assess the significance of competitive interactions to guild structure in tropical bird communities (Pearson 1977; Cornell & Lawton 1992). It was predicted that if interspecific competition in the classic, equilibrium-system context is significant to the regul ation of guild structure that: (1 ) species would ha ve specialized niches, (2) species would exhibit regular spaci ng along niche dimensions, (3) species would be sufficiently different in size to mitigate competiti ve interactions, (4) species would exhibit some form of competitive release in response to sign ificant variations in th e abundance of other guild

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19 members, (5) guild members might stagger re productive activities to reduce competition for nesting and brooding resources, a nd (6) guild membership would include fewer similar species than are available from the regional null-pool of potential guild members. Further it was predicted that if non-equilibri um processes were significant to guild structure, species would exhibi t limited niche specialization and ecological separation, and populations would exhibit non-equilib rium properties such as signifi cant temporal variations in abundance or habitat occupancy. Also it was predic ted that if predation pr essure was significant to guild structure then guild members would exhibit differential nesting strategies or reproductive traits that are adaptive to conditions of high mortality. Conservation Threats The Maya Forest refers to m ultinational complex of protected areas corresponding to the southern Yucatan peninsula re gions of Mexico, Belize, and Guat emala. The Maya Forest is the largest Neotropical forest remnant in Mesoamerica encompassing approximately threemillion hectares of contiguous forest c over (Radachowsky & Ramos 2004; CEMEC-CONAP 2006). The principal Guatemalan component of the forest is the Maya Biosphere Reserve (MBR) created in 1990 that includes over two-m illion hectares (CONAP 2001). Despite legal restrictions on the harvest of wildlife, law enforcement in Guatemala is minimal and hunting is largely uncontrolled. Throughout most of the M BR, these species are subject to persistent hunting pressure that is positively associated with permanent settlements and human access (Radachowsky & Ramos 2004). Some of the most isolated parts of the reserve continue to support unexploited populations of the guild species. In 1960, the human population of the Pet n was approximately 60,000, but has since doubled at approximately 8-year intervals (M eyerson 1997). A census of the Petn in 2001 recorded over 700,000 inhabitants, at which ti me there were approximately 90,000 residents

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20 within the MBR (CONAP 2001). Si nce the declaration of its prot ected status, the MBR has lost over 2,000 km2 of forest cover (Ramos et al. 2007) and b een fragmented into distinct eastern and western forest blocks. Forest fires provoked by swidden (slash-and-burn) agricultural practices occur annually. The three worst fire seasons in the last 10 years were 1998, 2003, and 2005, during which 4,335 km2, 3,985 km2 and 3,752 km2 respectively were affected by forest fires (Ramos et al. 2007). The eastern forest block of the MBR retains approximately 8,000 km2 of forest cover that links the Mexican and Belizean components of the Maya Forest. Current infrastructure proposals include three major roadbuilding projects within this part of the reserve (Ramos et al. 2007). One proposed project is a highway that would connect Guatemala and Mexico through the best-conserved areas remaining in the eas tern forest block. Environmental impact assessments of that project based on models from data on existing roads in the MBR (Ramos et al. 2007) predicted severe forest-cov er loss, fragmentation into as many as three separate forests, and the loss of continuity between the Mexican and Belizean components of the Maya Forest by 2025 (Ramos et al. 2007). Study Area This study was conducted in two ad jacent management units of the MBR; the community forestry concession of Uaxactn and the Mirador-Rio Azul National Park (Figure 1-1). The Uaxactn unit includes approximately 836 km2 of the multiple-use zone and contains a single permanent settlement, the village of Uaxact n with approximately 700 residents (NPV-OMYC 1999). The Mirador-Rio Azul National Park encompasses approximately 1117 km2 and is uninhabited (CONAP-ONCA-CECON 2002). The Mirador-Rio Azul National Park is divided into two sub-units, Mirador in the west and Rio Azul in the east. Data for this study were collected in the Rio Azul unit in the northeastern part of Guatemala bordering Mexico on the

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21 northern perimeter, Belize on the eastern perime ter, and the Uaxactn un it on the southeastern perimeter. Data from the nearest meteorological stati ons indicated mean monthly temperatures ranging from 23o C to 30o C (INSIVUMEH), being lowest November-February (mean median temperature = 26o C) and highest April-July (mean median temperature = 32o C). Local average annual precipitation is approxima tely 1250 mm. The annual precip itation pattern exhibits peaks in May or June and in September, and a four-month dry season from January-April. The Uaxactn and Rio Azul units straddle the drainage basin of the Rio Tikal-Rio Hondo watershed. Elevation ranges from 100-400 m, the lowest areas corresponding to the drainage basin and the highest to a steep karst escarpment that borders the basin to the west and north. The eastern edge of the basin is characterized by isolated upla nd areas with gentle slopes. Seasonal flooding can be severe in low and poorly drained areas dur ing the rainy season. Throughout the forest, small ponds that form in depressions during the ra iny season often dry-up during the dry season, leading to seasonal scarcity of available surface water. The Rio Tikal River is an intermittent arroyo that only occasionally forms a current but retains small ponds year-round in depressions along the channel. The landscape of the study area is dominated by a forest continuum that is classified as Subtropical Moist (Holdridge et al. 1971). A classification of the forest community in Tikal National Park, which is adjacent to the Uaxact n unit, found strong correlations of forest composition and structure to the edaphic conditio ns along the topographical gradient (Schulze & Whitacre 1999). Proceeding down the slope of this gradient, canopy height and basal area decrease and canopy opening size and stem-dens ity increase (Schulze & Whitacre 1999).

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22 Although 11 predictable forest-class es were identified along the topographical gradient, species richness and diversity were consistent among forest-classes (Schulze & Whitacre 1999). The village of Uaxactn orig inated in the early 1900s as a collection center for chicle the resin of the Manilkara zapota tree that was the original ba se ingredient of chewing-gum (NPV-OMYC 1999). Although the economic importance of gum-resin extraction has since diminished greatly, the local economy continues to be dominated by the extraction of forest resources including timber, palm fronds, allspice, construction materials, and wildlife. Although small-scale agriculture is locally important, deforested agricultural areas represent only 4% of the concession area (NPV-OMYC 1 999). To accommodate extraction activities primitive camps interconnected by an extensiv e trail network are distribut ed throughout the concession. Wildlife is harvested opportunistically dur ing the course of other extractive and agricultural activ ities on a continuous basis. Community -based hunting pressure is inversely correlated to distance from the village (Polisar et al. 1998); however, camp-based extractive activities have temporary but intense impacts far from the village (McNab 1998; Morales & Morales 1998). Although subsiste nce hunting pressure is biased towards larger vertebrate species, individuals are generally harvested without regard to sex or age-class. Unpublished data provided by the Wildlife Conservation Society (WCS) in Guatemala indicated that the wildlife harvest in Uaxactn in 1997 produced approxima tely 10,400 kg of game meat, which would cost approximately $30,000 US to replace with meat produced from domestic livestock. Given that the average annual household income is less than $2,000 US (personal observation) among approximately 140 households in the village, the annual wildlife harvests represent a subsidy equivalent to more than 10% of the local economy.

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23 CHAPTER 2 METHODS Diet and Harvest Data Data were collected from birds harvested for subsistence purposes by residents of the village of Uaxactn for the purpose of describing diet, body m ass, and harvest pressure of guild species. The participating hunters provided upper-digestive tracts (gizzards and also crops when available) from harvested specimens along with re levant observations, for a reward equivalent to the value of a pound of game-meat (<1$ US). Diet samples were separated from the tracts, dried in a solar dryer, and stored in paper envelopes in the community. Samples were dried again in electric dryers in the laboratory prior to proces sing. The contents of each sample were separated into distinct components for which dry mass valu es were recorded and identified on general and specific levels. Diet compositions were based on the dry-ma ss proportions of gene ral diet components including: animal, leaf, pulp, seed, flower/stem, and grit. For composition analyses unidentifiable pulverized material in each sa mple was assigned to the identifiable items according to their relative proportions. Alte rnative diet composition descriptions were determined without grit and snail shells, due to their bias relativ e to the dry mass of plant and animal tissues. Dietary-overlap indices were calculated for all species-pairs using the equation from Pianka (1973) shown below, based on the seed component of each species diets, where Ojk is the overlap between species j and species k and Pij and Pik are the proportions of the i th resource used by the j th and k th species, respectively. Uniden tifiable pulverized material was excluded from the overlap analyses. n n Ojk = Pij Pik / Pij2 Pik2 i i

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24 The harvest data reported by local hunters were used to describe local harvest pressure on guild member populations. Adult body-mass values from the data were applied to describe harvest biomass. Harvest pressure was described by distance per 4 km interv als from the village. Birds collected by local hunters are usually cleaned in the field to avoid spoilage and reduce the burden, so access to entire specimens was limite d. Balance scales were provided to field personnel and certain local hunters in order to record body mass values from entire specimens opportunistically. Adult body-mass records were compared between species and sexes using an analysis of variance (ANOVA) and a Duncan s Multiple Range Test (alpha levels 0.05 were considered statistically significant). Size-ra tios between closest-size species-pairs were compared to predictions fr om competition theory. Line-Transect Data A system of 12 line-transects (Buckland et al. 2001) provide d data used to describe species abundance and certain ni che parameters. Transects were organized into 4 groups of three, located along a hunting pressure gradient identified a prior (Polisar et al. 1998). The Ixcan transect group was located in the core of the Rio Azul unit where no hunting pressure was expected. The Cedro group was located on the pa rk perimeter adjacent to the Uaxactn unit, approximately 25-37 km from the village where hun ting pressure was expected to be light. The Uaxactn North group was located approximately 3-15 km north of the village where hunting pressure was expected to be heavy. The Ua xactn South group include d the area surrounding the village where hunting pressure wa s expected to be heaviest. Each transect within a group was located in a stratified-random manner in order to sample all forest-classes. In the Uaxactn unit, tr ansects were oriented to avoid existing trails to reduce the probability of locals hunting on them. In the Rio Azul unit all but one of the transects were established on modified, existing trails. Eleven transects measur ed 2500 m, whereas one

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25 transect measured 2750 m to improve representati on of a forest-class in that group. Transects were flagged and labeled at 50 m in tervals and mapped by forest-class. Transects were sampled by trained observer s between 0600 and 0930 hours at intervals of 5-10 days depending on personnel capacity, weather, and logistics. Observers recorded data from detections of guild members and other specie s of interest to associated researchers (Novack 2003; Garcia & Radachowsky 2003). The transects in the Rio Azul unit were established in February 2000 and sampled over a 30-month period. Each year data collec tion in Rio Azul was interrupted when seasonal flooding prevented access to the site. The transects in the Uaxactn unit were sampled from August 2000 through Ju ly 2001. The total sampling effort on all transects was approximately 2770 km. Habitat Use Habitat use was analyzed based on species-enco unter rates by forest-class derived from the transect data. Two forest cl ass ifications based on alternative in terpretations of the results of Schulze and Whitacre (1999) were evaluated. One classification consisted of three forestclasses: Upland, representing tall forest types on the upper slopes of the topographical gradient; Scrub, representing low forest types at the bottom of the gradie nt; and Transitional, representing intermediate forest types. The second classification consolidated the Upland and Transitional classes of the former classification into a single Upl and forest-class, and used the same Scrub forest-class. Monthly species-enc ounter rates in the Upland and Transitional forest-classes of the three-cla ss system were compared using a student t-test to determine if species responded to the increase resolution of that classification. To determine if species exhibited preferential habitat occupancy, species -encounter rates by forest-class were calculated for each transect group and annua l period, converted in to relative frequencies and compared using a student t-test. Alpha levels 0.05 were considered statistically significant for both t

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26 tests. Habitat occupancy overlap was assessed using proportional simila rity indices (Schoener 1970), calculated for each species-pair in each fo rest-class. For each species the combined average overlap with all othe r species was also described. Coefficients of variation (CV) calculated fr om species-encounter rates were used to evaluate the temporal variabi lity of habitat occupation. Mont hly species-encounter rates by forest-class were plotted graphica lly and evaluated for temporal va riations of habitat occupancy. Monthly species-encounter rates by forest-class were examined for correlat ion with precipitation, temperature, and encounter rate s of other guild members using a Pearson correlation analysis. Vertical Strata Occupancy Vertical height records from the transect data were used to assess whether guild members exhibited consistent patterns of forest-strata o ccupancy. Height records were assigned to one of three categories, ground strata (0-1 m), understory strata (1m to half of the mean canopy height), or canopy strata (half of the mean canopy height and above). The relative frequencies of occurrence in the three strata were compared be tween transect groups, forest-classes, and annual periods for each species using an analysis of variance (ANOVA) and a Duncans Multiple Range Test (alpha levels 0.05 were considered statistically signifi cant). Vertical overlap in strata occupancy was estimated for each species-pair us ing proportional similarity indices (Schoener 1970). For each species the combined average overla p with all other species was also described. Species Abundance and Guild Composition Variables derived from the transect data that were used to compare guild member abundance included individual-encounter rates, density estimates by forest-class, relative population size and relative population biomass. Monthly indivi dual-encounter rates from the same annual period were compared among transect groups for each species using an analysis of variance (ANOVA) and a Duncans Multiple Range Test (alpha levels 0.05 were considered

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27 statistically significant). Density estimates we re calculated from the transect data using the program DISTANCE version 5 (Thomas et al. 2005). Models used to estimate densities were selected by the program based on comparisons of all relevant combinations of detection functions. Data were adjusted (truncated or filtered) when necessary to control for data concentrations that were inconsistent w ith detection functions (data heaping). Spatial forest-class descriptions derived fr om Landsat 7 (USGS/EROS 2000) satellite images from the study period were provided by CEMEC-CONAP (2006). Using the program ArcView 3.2 (ESRI 2000), 4 km-radius buffers representing 50 km2 were applied to the approximate geographic center of each transect group and were used to determine habitat availability within the corres ponding areas (Figure 1) Density estimates by forest-class were extrapolated by forest-class representations within each 50 km2 area to estimate corresponding species population sizes. Guild composition descriptions for each transect group were determined based on relative population size and re lative biomass representations at each site. Guild compositions were compared between tr ansect groups and annual periods. Temporal variability of species abundance wa s assessed using coefficients of variation ca lculated from individual-encounter rates duri ng annual and 30-month periods. Nest Site Data Data collected from nest-sites of guild me mbers were used to determine reproductive characteristics of each species. Local par ticipants were compensated a days wage (approximately $7 US) to accompany data collector s to nests encountered during the course of other activities and to assist with data collection at the nest site. Clutch sizes were included in analyses if they met the minimum clutch size reported from a literature source, and did not appear abandoned. Search effort was not system atic with respect to habitat so forest-class percentages are not representative of habitat distributions for the species. Nest sites could only

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28 be visited once and some visits were made after nesting activities had concluded so nest depredation and clutch-size record s are conservative. Vertical ne st placement, tree diameter, and clutch-size were compared between species us ing an analysis of variance (ANOVA) and a Duncans Multiple Range Test (alpha levels 0.05 were considered stat istically significant). Nest placement patterns within the guild were evaluated for spatial separation. Temporal patterns of reproductive activitie s were described from field obs ervations, the line-transect and the nesting data and subsequently evaluated for temporal separation.

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29 CHAPTER 3 RESULTS Diet Composition and Overlap Diet content sam ples were derived from 267 C. rubra, 181 M. ocellata 205 O. vetula 142 P. purpurascens and 55 T. major (Table 3-1). Grit represented 20% of the diet in C. rubra 22% in M. ocellata 3% in O. vetula 9% in P. purpurascens and 2% in T. major Diet compositions without grit and snails indicat ed seed proportions of 88% of the diet in C. rubra, 77% in M. ocellata 85% in O. vetula 88% in P. purpurascens and 90% in T. major Excluding grit and snails, fruit pulp co mprised 8% of the diet in C. rubra, 11% in M. ocellata 7% in O. vetula 10% in P. purpurascens and 7% in T. major Leaf proportions without grit and snails contributed 10% of the diet in M. ocellata 7% in O. vetula and less than 2% of the diets of the remaining species. Proportions of flower/stem and animal materials varied among species, but were less than 3% of the diet of any guild member. Guild diet included 143 seed species, w ith similar quantities among guild members despite sample size differences: C. rubra (74 spp.), M. ocellata (92 spp.), O. vetula (86 spp.), P. purpurascens (72 spp.), T. major (67 spp.). Seed-species overlap values between species-pairs ranged from 33% for M. ocellata O. vetula to 86% for C. rubra T. major (Table 3-2). Average seed overlap with all other sp ecies ranged from 47-68%, with T. major ranking highest, followed in sequential order by P. purpurascens C. rubra M. ocellata and O. vetula. Harvest Description The recorded harvest of guild m embers aver aged 591 individuals representing 1625 kg of biomass annually based on average a dult, body-mass of guild members. C. rubra represented an average of 47% of the tota l harvest and 58% of the harvest biomass annually. M. ocellata represented 19% of the total harv est and 28% of the biomass the fi rst year, and declined to 13%

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30 and 22% respectively, the second year. O. vetula represented 16% of the harvest and 3% of the biomass the first year, and increased to 25% and 6%, respectively, the second year. P. purpurascens represented 7% of the harvest and 6% of the biomass the first year, and increased to 17% and 16%, respectively, the second year. T. major represented an average of 5% of the annual harvest and 2% of the bi omass. Harvest pressure on C. rubra M. ocellata and P. purpurascens was greatest in the second di stance-interval (4-8 km) from the village and declined steadily in subsequent interv als. Harvest pressure on O. vetula and T. major was greatest in the first distance interval (0-4 km) and dec lined steadily in subsequent intervals. Size Assortment Adult body-m ass records were collected from 67 C. rubra 29 M. ocellata 17 O. vetula 26 P. purpurascens and 12 T. major Average adult body-mass was 3.4 kg for C. rubra 4.16 kg for M. ocellata 0.64 kg for O. vetula 2.31 kg for P. purpurascens and 1.17 kg for T. major Adult body-mass differed between guild member s based on the results of both analyses (ANOVA, alpha = 0.0001, F = 336.91, df = 150; Duncans, alpha = 0.05, df = 146). Adult bodymass of sexes differed in M. ocellata (male = 4.81 kg; female = 3.51 kg) (ANOVA, alpha = 0.0001, F = 26.79, df = 28; Duncans, alpha = 0.05, df = 27) and P. purpurascens (male = 2.16 kg: female = 2.44 kg) (ANOVA, alpha = 0.0269, F = 5.56, df = 25; Duncans, alpha = 0.05, df = 24). Size ratios (and natural l ogarithm equivalents) between species-pairs were 1.23:1 (0.209) for C. rubra M. ocellata 1.47:1 (0.384) for C. rubra P. purpurascens 1.98:1 (0.681) for P. purpurascens T. major and 1.83:1 (0.609) for O. vetula T. major Size ratios between M. ocellata sexes and C. rubra were 1.42:1 (0.348) for males and 1.03:1 (0.032) for females. Habitat Use Monthly species-encounter rates between Upland and Transitional fo rest-classes did not differ for any guild m ember. All subsequent habitat comparisons were made using the two

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31 forest-class system. Frequencie s of occurrence between forest-c lasses were unequal (alpha = 0.0001) for O. vetula (t = -7.9141, df =10), C. rubra (t =11.681, df = 10), and P. purpurascens (t = 6.1134, df = 10) and nearly un equal (alpha = 0.0601) for T. major (t = 2.1193, df = 10). M. ocellata did not differ in fo rest-class occupancy. O. vetula occurred in Scrub forest with an average frequency of 0.77. C. rubra and P. purpurascens and T. major occurred in Upland forest with average frequencies of 0.73, 0.66, and 0.58, respectively. Habitat overlap among species-pairs ranged from 27-48% in Scrub forest and from 23-67% in Upland forest (Table 3-3). Combined habitat overlap among species-pairs ra nged from 50-94%. Average combined habitat overlap with all other speci es ranged from 61-84%, with T. major ranking highest, followed in sequential order by M. ocellata P. purpurascens C. rubra and O. vetula Ratios of CV values between Scrub fo rest and Upland forest were 1.82:1 for C. rubra 1.35:1 for M. ocellata 2.35:1 for P. purpurascens and 1.94:1 for T. major indicating greater variability in the occupancy of Scrub forest. The CV ratio between Upland forest and Scrub forest was 1.43:1 for O. vetula indicating greater variability in the occupancy of Upland forest. Graphical analyses of monthly species-encoun ter rates by forest-class did not reveal any consistent temporal trends or shifts in forest-class occupancy by guild members. The Pearson correlation analyses did not identify any (alp ha = 0.05) correlations of monthly speciesencounter rates with respect to temperature, precipitation, or en counter rates of other species. Vertical Strata Occupancy All species exhibited differential use of forest strata (ANOVA df = 35): Crax rubra (alpha = 0.0011, F = 4.68), M. oce llata (alpha = 0.0001, F = 43.07), O. vetula (alpha = 0.0018, F = 4.36), P. purpurascens (alpha = 0.0001, F = 43.41), T. major (alpha = 0.0001, F = 57). Vertical patterns of forest-strata occupancy did not differ between transect groups, forest-classes, or annual periods for any species (Duncans alpha = 0.05, df = 27) M. ocellata and T. major had

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32 average frequencies of 89% and 92%, re spectively, in the ground stratum. C. rubra had an average frequency of 59% in th e canopy stratum and an average of 32% in the ground stratum. P. purpurascens had an average frequency of 84% in the canopy stratum. O. vetula had average frequencies of 53% and 41%, respectively, in the canopy and understory strata Vertical strata overlap ranged from 14% between O. vetula T. major to 93% between M. ocellata T. major (Table 3-4). Four species-pairs exhibited vertical overlap greate r than 50%. Average combined overlap with all other speci es ranged from 40-56%, with C. rubra ranking highest, followed in sequential order by M. ocellata P. purpurascens O. vetula and T. major Species Abundance and Guild Composition Monthly individual-encounter rates differed (Duncans, alpha = 0.05, df = 42) am ong transect groups for C. rubra (ANOVA, alpha = 0.0001, F = 9.13, df = 45), O. vetula (ANOVA, alpha = 0.0012, F = 6.36, df = 45), and P. purpurascens (ANOVA, alpha = 0.0001, F = 31.49, df = 45). Encounter rates decreased alo ng the increasing hunting gradient for C. rubra M. ocellata and P. purpurascens Encounter rates were highest in the Ixcan and Uaxactn South groups for O. vetula Encounter rates were highest in the Ixcan group and stable among the remaining transect groups for T. major Density estimates for all species were highest in the protected Ixcan group (Table 3-5). Density estimates generated by DISTANCE for so me species in the less hunted areas seemed intuitively robust, however the results of analyses were ultimately accepted if they met the model selection criteria of the program, were consistent with the data and deri ved encounter-rates, and were comparable published density estimat es for these or congeneric species. C. rubra exhibited overall density declines alon g the increasing hunting gradient by 83% in both Upland and Scrub forest. M. ocellata exhibited overall declines along the increasing gradient of approximately 66% both forest-classes, with highest density estimates in the Ixcan

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33 and Uaxactn North groups. O. vetula densities were similar a nd highest in the Ixcan and Uaxactn South groups. P. purpurascens exhibited an overall declin e along the gradient of 72% in Upland forest and 43% in Scrub forest. T. major exhibited little density variation among the Cedro, Uaxactn North and Uaxactn South transect groups. T. major exhibited higher densities in Scrub forest relative to Upland forest in the Ixcan and Cedro groups, and higher densities in Upland forest relative to Scrub forest in th e two Uaxactn groups. Co mparison of densities between years in the Ixcan and Cedro groups indi cated increased or stab le densities among all species, except for slight decreases for T. major in the Cedro group. Between years the estimated density of M. ocellata increased by 59% in Upland forest and 14% in Scrub forest in the Ixcan group, and by 159% in Upland forest a nd 569% in Scrub forest in the Cedro group. C. rubra represented proportions of the guild ranging from 24-32% in the Ixcan, Cedro, and Uaxactn North groups, but only 11% in the Uaxactn South group (Table 3-6). M. ocellata averaged 18% of the guild in the Ixcan and Uaxactn North groups, and 10% and in the Cedro and Uaxactn South. O. vetula averaged 24% of the guild in the Ixcan, Cedro, and Uaxactn North groups and represented 46% of the guild in the Uaxactn South group. P. purpurascens averaged 23% of the guild in the Ixcan and Ce dro groups, and 14% in the two Uaxactn groups. T. major averaged 10% of the guild in the Ixcan and Cedro groups and 19% in both Uaxactn groups. C. rubra accounted for the highest biomass proportion of the guild in all groups (average 40%) except Uaxactn South wher e it represented only 22%. M. ocellata averaged 31% of the guild biomass in all transect groups except the Cedro group wh ere it accounted for only 18%. The proportions of guild biomass represented by O. vetula increased steadily along the increasing hunting gradient from 6% in the Ixcan group to 18% in the Uaxactn South group. P.

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34 purpurascens accounted for 25% of guild biomass in the Cedro group, 21% in the Ixcan and Uaxactn South groups, and 14% in the Uaxactn North group. T. major accounted for 5% of guild biomass in the Ixcan and Cedro groups and an average of 12% in the two Uaxactn groups. Coefficients of variation calculated from individual-encounter ra tes during both annual and 30-month periods indicated dou ble-digit percentage changes for all species. Average CVs for annual periods were 34% for C. rubra 35% for M. ocellata 26% for O. vetula 18% for P. purpurascens and 21% for T. major CVs of variation for the 30-month period were 38% for C. rubra, 62% for M. ocellata 32% for O. vetula 22% for P. purpurascens and 22% for T. major Nest Site and Clutch Characteristics A total of 225 were observed: C. rubra nests ( n = 24), M. ocellata nests ( n = 39), O. vetula n ests ( n = 77), P. purpurascens nests ( n = 19), T. major nests ( n = 66). Mean nest height differed for O. vetula (5.7 m, range 1-18 m), and species pairs C. rubra (10 m, range 5-25 m) P. purpurascens (9.75 m, range 4-15 m), and T. major (0 m) M. ocellata (0 m) (alpha = 0.05, df = 206). Mean DBH of trees associated with ne st sites differed between the species group T. major (35.4 cm) -M. ocellata (32.3 cm) -P. purpurascens (26.3 cm) and the individual species C. rubra (20 cm) and O. vetula (16.9 cm) (alpha = 0.05, df = 151). Mean clutch sizes for M. ocellata (8.73, mode = 9) and T. major (4.49, mode = 4) differed from each other and the other guild me mbers (alpha = 0.05, df = 203). Mean clutch sizes for P. purpurascens (2.16, mode = 2) O. vetula (2.12, mode = 2) C. rubra (2.00, mode = 2). Depredation of nests, base d on a single visit, were 20% ( n = 13) for T. major 15% ( n = 6) for M. ocellata 4% ( n = 3) for O. vetula nests, 4% ( n = 1) for C. rubra and 0% for P. purpurascens With respect to habitat, 96% ( n = 23) of C. rubra nests, 95% ( n = 19) of P. purpurascens nests, 95% ( n = 23) of T. major nests, 36% ( n = 14) of M. ocellata nests, and 16% ( n = 12) of O. vetula nests were located in Upland forest Nesting in Scrub forest was only

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35 observed for O. vetula (7%), T. major (5%), and M. ocellata (3%). Only O. vetula (78%) and M. ocellata (62%) nests were observed in agricultural habitat. Specifically, all C. rubra and P. purpurascens nests were placed in trees, and all T. major nests in the roots of trees Seventy-three percent ( n = 52) of O. vetula nests were placed in trees, 20% ( n = 14) were in vines. M. ocellata placed 49% ( n = 19) and 36% ( n = 14) of their ground nests in bracken fern (Pteridium spp.) and the roots of trees, respectively. All guild members commenced nesting near th e end of the dry seas on (March-April) and continued into the wet season; however, the length of the nest ing period varied among guild members (Table 3-7). Active nests of C. rubra and M. ocellata were only observed April-June, whereas nests of P. purpurascens were observed from April-July. Nesting by T. major and O. vetula occurred during March-September and March-October, respectively.

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36 CHAPTER 4 DISCUSSION Guild Diet Characteristics Relatively high proportions of grit in the diets of C. rubra and M. ocella ta suggest that they function as seed predators. Relativ ely low proportions of grit in the diets of O. vetula and T. major suggest that they function less as seed pr edators than the rest of the guild. Diet compositions of C. rubra O. vetula and P. purpurascens were similar to previous findings (Rivas & Morales 2003). Flower and stem materials combined did not represent a significant proportion of the diet of any guild member. Flow ers have been reported in higher proportions in the diets of P. purpurascens (Rivas & Morales 2003), and M. ocellata (Steadman et al. 1979), and may be seasonally important to curassows (S antamaria & Franco 2000). Animal matter was proportionately higher in M. ocellata with snail shells included, but similar to other species when they were excluded. Diet studies based on direct observations indicate fr equent consumption of a variety of small vertebrate s by curassows (Santamaria & Franco 2000, Jimenez et al. 2001). The relatively higher proportions of leaf material in the diets of M. ocellata and O. vetula, in contrast to the other guild members, are consis tent with findings from other studies (Rivas & Morales 2003; Steadman et al. 1979 ). Excluding grit and snails, the diet of individual guild members had similar, proportions of pulp (7-11%) a nd of seeds (77-90%), thus the diet of this guild is dominated by seeds, many of which are consumed as whole fruit. Species-pairs O. vetula M. ocellata and C. rubra O. vetula displayed relatively low dietary overlap (33% and 39% re spectively), two species-pairs, P. purpurascens T. major and C. rubraT. major displayed relatively high overlap (73% and 86% respec tively), and the remaining six species-pairs had intermediate values (50-68 %). Of the 143 seed species consumed by guild members; 19% (27 seed species) were shared by all five guild member s, 13% (19) by four

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37 members, 18% (26) by three members, and 21% ( 30) were shared by two guild members. Seed species shared by the entire guild represented significant proportions of the diets of individual species: 82% of the seed dry mass in C. rubra 76% in M. ocellata 57% in O. vetula 79% in P. purpurascens and 83% in T. major The fruits Brosimum alicastrum and Psuedolmedia spp. were among the top-five ranking seeds in the diet s of four of the guild members and ranked 15th and 8th, respectively, in O. vetula Rivas & Morales (2003) analyzed diets of C. rubra O. vetula and P. purpurascens collected from the study area be tween January 2002 and July 2003. In that study the two most significant seed species in the diet of C. rubra were from the fruits Pouteria amygdalina (47%) and Protium copal (8%); the two most significant seeds in the diet of P. purpurascens were P. amygdalina (17%) and Vitex gaumeri (12%); and the most signifi cant seed in the diet of O. vetula was Psuedolmedia spp. (19%). In the current study P. amygdalina represented 9% and P. copal 1% in C. rubra ; P. amygdalina represented 1% and V. gaumeri 5% in P. purpurascens ; and Psuedolmedia spp. represented 3% in O. vetula In the current study, the most signifi cant seed species in the diet of C. rubra were B. alicastrum (20%) and Calophyllum brasiliensis (19%), and the most significant seed species in the diet of P. purpurascens was Chrysophyla argentea (14%). Rivas and Morales (2003) did not detect B. alicastrum in the diet of C. rubra and C. brasiliensis represented 6% of the diet furthermore they found that C. argentea represented 5% of the diet in P. purpurascens Those authors noted only m inor representation of B. alicastrum which represented (3%) in O. vetula and (0.05%) in P. purpurascens Rivas (1995) analyzed diet contents of C. rubra from the same area in 1995 and identified B. alicastrum as the most important diet species.

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38 In summary members of this guild members consume many of the same seed species, some of which are important to the diet of most members of the guild. Common usage of important seed species by guild members sugge sts that food resources were not limited. The contrasting results of these two st udies indicate that the importance of seed species varies with time. Temporal variability in the importance of some seeds may reflect variability in production due to fluctuations in annual precipitation. To illustrate, Peters (1989) found that fruit production of B. alicastrum is strongly correlated with precipitation. Local meteorological data indicated mean annual variations in precipitation of 200mm (16% of annual mean) with extremes of as much as 800mm (64% of annual mean). Guild Habitat Characteristics Overall guild occupancy was greatest in Up land f orest, which may reflect a greater volume of habitat spac e relative low-canopy Scrub forest. Habitat occupancy patterns of O. vetula C. rubra and P. purpurascens were consistent between transect groups and annual periods, and highly skewed towards a particular fo rest-class, indicating ha bitat selection by these species. Preferential habitat use was exhibited by all six cracid species in a similar guild in Bolivia (Wallace et al. 2001). Pref erential occupancy of Scrub forest by O. vetula and of Upland forest by C. rubra and P. purpurascens were consistent with habitat-use descriptions for the species (Gonzalez et al. 1998; Howell & Webb 1995; Brooks et al. 2004). Open, grassy clearings have been identified as important habitat for reprodu ctive activities and feeding for M. ocellata and are known to be important brooding habitat for M. gallopavo in North America (Steadman et al. 1977; Gonzalez et al. 1998; Willia ms & Austin 1988). Open, grassy clearings are almost nonexistent in the study area theref ore local habitat may not be optimal for M. ocellata

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39 Overlap values calculated usi ng the proportional similarity index are expected to be conservative (Krebs 1989). The two species that exhibited the strongest and most divergent habitat preferences were O. vetula and C. rubra which subsequently had the lowest combined habitat overlap among species-pair with 50%. The species-pairs w ith the most similar patterns of habitat use, M. ocellata T. major and C. rubra P. purpurascens shared the highest overlap values in the guild at 94%. Based on that range, three other species-pairs ( P. purpurascens T. major M. ocellata P. purpurascens and C. rubra T. major ) had high combined habitat overlap (85-91%), two species-pairs ( M. ocellata O. vetula, C. rubra M. ocellata ) had intermediate overlap (71-79%), and two species-pairs ( O. vetulaP. purpurascens O. vetula T. major ) had low overlap (56-65%). All species that exhibited habitat preferen ces also exhibited greater variability of occurrence in the alternative forest-class. M. ocellata exhibited the most variability in habitat occupancy and the least. C. rubra O. vetula, and P. purpurascens exhibited similar variability of habitat occupancy. Despite variability of habitat occupanc y among all species, consistent temporal patterns of differential habitat occ upancy were not exhibited by any guild member. The latter finding is in contrast with seasonal habitat shifts reported for M. ocellata by Gonzalez et al. (1998). Consistent and distinct patterns of vertical strata occupation indicate vertical stratification within the guild, which is considered to be impor tant in tropical bird communities (Karr & Roth 1971; Pearson 1971). The vertical patterns of strata occupation exhibited by guild members were consistent with descriptions for thes e species (Howell & Webb 1995; Brooks et al. 2004). Relevance of Competition to Guild Structure Diet com positions, high divers ity of dietary resources, a nd significant dietary overlap among most species-pairs indicate generalist diet niches for all guild members. The hypothesis

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40 that guild members have specialized dietary ni ches is not supported by the data. The only evidence of differential dietary resource use in the guild were the proportionally greater leaf material in the diets of M. ocellata and O. vetula, and the low overlap values between speciespairs O. vetula M. ocellata and C. rubra O. vetula. The hypothesis that the guild members exhibit regular spacing along a gr adient of dietary resources is not supported by the comparisons of guild member diets. Differential habitat preferences between O. vetula and the two larger cracids indicate specialized use of hab itat by these three species. The limited habitat preference exhibited by T. major and the lack of preference exhibited by M. ocellata suggest generalist habitat resource use by these species. The hypothesis that the guild members have specialized habitat niches is only partially supported by the data. Less selective use of habitat by M. ocellata and T. major increased the representation of th e guild in Scrub forest, and ma y represent deliberate spatial separation from C. rubra and P. purpurascens populations in Upland fo rest. Overall habitat occupancy patterns among all species indicate sp atial separation along a gradient of habitat resources. The hypothesis that th e guild exhibits regular spaci ng along the gradient of habitat resources is supported by comparisons of habitat occupancy patterns. The hypothesis that guild members would have specialized niches with respect to the vertical occupation of forest st rata is supported by the consiste nt patterns exhibited by all species. The hypothesis that the guild exhibits regul ar spacing along a vertical gradient of forest strata is supported by the vertical stratification observed within the guild. Size ratios in a community structured by comp etitive interactions are predicted to be either constant, have minimum values from 1.6-2.0:1 (natural logarithm values between 0.405 and 0.683), or else to increase with species size (Hutchinso n 1959; Diamond 1962; Oskanen et

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41 al. 1979). Ratios of size differences between guild members are not constant and decrease among the largest three species. The si ze differences between species-pairs P. purpurascens T. major (0.681) and T. major O. vetula (0.609), were similar and also consistent with predicted minimum differences. The size differences between species-pairs M. ocellata C. rubra (0.209) and C. rubra P. purpurascens (0.384), are not similar to one a nother or to the size differences between other species-pairs, and are less than predicted minimums in assemblages structured by competitive interactions. The hypothesis that size a ssortment of the guild is sufficient to mitigate competitive interactions is supported by size diffe rences among half of the guild species. C. rubra M. ocellata and P. purpurascens populations exhibited significant reductions along the increasing hunting pressure gradient. O. vetula and T. major abundance did not increase significantly nor did they exhibit changes in ve rtical strata occupancy in the transect group where C. rubra M. ocellata and P. purpurascens populations were most reduced. T. major exhibited a relative shift in habitat occupancy from Scrub fo rest in the two groups where hunting pressure was light, to Upland forest in the two groups where hunting pressure heavy. Considering that T. major had the lowest relative abundance in the transect groups where hunting pressure was light and exhibited the highest overlap with all other species in dietary and habitat resources, it may experience the greatest comp etitive pressure from the other species. T. major was the only species that exhibited an inter-annu al density decrease in the Cedro group, where M. ocellata concurrently exhibited extraordinar y density increases, which may indicate competitive inhibition of T. major by M. ocellata These two species had the highest overlap in the guild along diet and vertical niche dimensions as well as relatively high dietary overlap. The hypothesis that guild members would exhibit comp etitive release in response to the decreased abundance of other species is partially supported by the data.

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42 Although guild members exhibited variable ne sting period lengths, reproductive activities peaked during the same time of year for all gui ld members. The hypothesis that guild members may stagger reproductive activities to avoid comp etition was not supported from the temporal nesting patterns exhibited by the guild. Aside from the five existing guild members, the only potential guild members available within a 200 km radius are the Horned Guan ( Oreophasis derbianus ), the Highland Guan ( Penelopina nigra), and the White-bellied Chachalaca ( Ortalis luecogastra ). The two guans are not legitimate null pool species because they are highland species with lower distribution limits at elevations of 1000 m for Penelopina and 2000 m for Oreophasis Although O. luecogastra exists at low elevations, regional Ortalis species were previously c onsidered con-specific, and current taxonomic descriptions reflect distinct geographic distributions (Howell & Webb). Because regional Ortalis species are geographically separate, they may not qualify as legitimate null pool species capable of coloni zing a distinct geographic area. O. luecogastra is confined to the Pacific slope of the isthmus, which is se parated from the study area by mountain ranges along the entire length of the species distribution, so the abse nce of the species in the study area is not likely to be the result of competit ive exclusion. A regional null pool including O. luecogastra would consist of only six species, in which case the local absence of O. luecogastra could be attributable to competitive exclusion by O. vetula If O. luecogastra is excluded from the regional null pool, then all available potential guild member s are represented locally. Competitive interactions within the guild do not a ppear sufficient to exclude regionally available guild members except possibly in the case of O. luecogastra. The hypothesis that the guild would include fewer similar species than are re gionally available is not supported based on null pool comparisons.

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43 Guild characteristics are partially consistent with predictions based on competition theory in an equilibrium system. Guild members exhibite d niche specialization with respect to vertical occupancy of forest strata, and to some degree w ith respect to habitat use, but did not exhibit niche specialization with respect to the use of di etary resources. Guild members appeared to be ecologically separated along habita t resource and vertical forest strata gradients, but not along the gradient of dietary resources. Size differences appeared to be sufficient to mitigate competitive interactions for O. vetula T. major and P. purpurascens but not for C. rubra M. ocellata and P. purpurascens Overlap between C. rubra and P. purpurascens was relatively high along all niche dimensions. Overlap between M. ocellata with both C. rubra and P. purpurascens was relatively high with respect to hab itat and dietary reso urces and low with respect to vertical stra ta use. If competitive interactio ns were significant within the guild, C. rubra, M. ocellata and P. purpurascens would be expected to exhi bit greater specialization and separation along the other niche dimensions. T. major exhibited inverse responses to variations of the abundance of other guild members. Guild membership was not notably different from the regional null pool. Although some guild characte ristics are consistent with an assemblage structured by competitive interactions, the degree is less than would be expected based on predictions from competition theory in an equilibrium system. The limited evidence of competitive interactions displaye d by the guild may be consistent with the non-equilibrium competition model proposed by Connell (1980). Relevance of Non-Equilibrium Factors to Guild Structure Niches should evolve in response to mortal ity factors and dietary resources as well as com petition, therefore some niche specialization and ecological separation would be expected whether or not competitive interactions are impor tant to the guild. Guild member niches were generalist with respect to diet, semi-specialized with respect to habitat, and specialized with

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44 respect to vertical occupation of forest strata. The guild did not exhibit ecological separation with respect to dietary resources, but did appear to be distributed along habitat and vertical strata dimensions. The hypothesis that guild species would exhibit limited niche specialization and separation along niche dimensions is supported by the niche characte ristics of the guild. Significant temporal variability of gu ild member abundance would constitute environmental non-equilibrium. CVs of indivi dual-encounter rates du ring annual and 30-month periods represented double-digit percentages for all species. Although an objective basis of comparison of the coefficient values is not avai lable, these percentages appear to represent significant variation in guild member abundan ce. CVs were consistently highest for C. rubra and M. ocellata which may reflect greater movement in foraging strategies. Significant local temporal variation of curassow abundance has been attributed to movement between patchy fruit resources (Thiollay 1989). CVs were relativ ely consistent betwee n inter-annual and the 30month periods for all species except M. ocellata which had a much higher coefficient for the 30month period. This observation suggests that inter-annual variation may be greater than intraannual variation in this species. Inter-annual changes of guild member densities in the two, Rio Azul transect groups also indi cated significant variation of a bundance for all species. The hypothesis that guild members may exhibit signifi cant variation in abundan ce is supported by the variability exhibited by individual-en counter rates and density estimates. Temporal shifts in habitat us e by bird species in response to microhabitat variability may result in dynamic assemblages representing non-equilibrium environmental factors (Karr & Freemark 1983). The CVs of species-encounter rates in Upland and Scrub forests indicated considerable variation of hab itat occupancy. The hypothesis th at guild members may exhibit significant variation in habitat occupancy is s upported by the variation observed from species-

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45 encounter rates in each forest-class. In summary the combination of limited niche specialization and ecological separation and the high variati on of species abundance and habitat occupancy, are consistent with predictions of non-equilibrium systems. Relevance of Predation to Guild Structure If guild m embers are subject to significant or differential pr edation pressure, it should be reflected by reproductive ch aracteristics (Martin 1988 a). Predation is the primary cause of nest failure in most bird species (C ody 1971). Nest losses to predation have been reported as high as 80% for T. major (Brennan 2004) and almost 50% for M. ocellata (Gonzalez et al. 1998). Potential mammalian nest predators observed on the line-transects included coatimundi ( Nasua narica ), raccoon ( Procyon lotor ), collared peccary (Tayassu tajacu ), white-lipped peccary ( Tayassu pecari ), and spider monkey ( Ateles geoffroyii ). Other potential mammalian nest predators occurring locally in clude: two large opossums ( Didelphis spp.), nine-banded armadillo ( Dasypus novemcinctus), ring-tailed cat ( Bassariscus sumichrasti ), and striped hog-nosed skunk ( Conepatus semistriatus ) (Garcia & Radachowsky 2003; pers onal observation). Although avian nest predators may be less significant for large species, corvids have been reported to be minor nest predators on M. gallopavo (Williams & Austin 1988). Also curassow predation on tinamou nests has been documented (Santamaria & Fr anco 2000). Although evidence of avian nest predation is not available loca lly, the study area supports divers e and abundant populations of corvids and rhamphastids (toucans). Spatial separation of nest-sites through diffe rential vertical or habitat placement may favor bird community diversity by mitig ating predation pressure (Martin 1988 a). The combination of terrestrial nest placement by M. ocellata and T. major arboreal nest placement by C. rubra and P. purpurascens and shrub-arboreal placement by O. vetula indicates vertical nest-separation within the guil d. Although nest-site da ta collection was no t systematic with

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46 respect to habitat, there is evidence of select ive habitat use within the guild. All species had significant but variable proportions of their nests in Upland forest. C. rubra and P. purpurascens nests were almost exclus ively in Upland forest. M. ocellata O. vetula and T. major were the only species that nested in Scrub forest. Only M. ocellata and O. vetula exhibited substantial use of agricultural habitat. Predation on ground nests may be greater than in other strata in tropi cal forests (Loiselle & Hoppes 1983; Martin 1993; Gibbs 1991). This phenomenon is supported by the relatively higher proportions of nests lost to predation by M. ocellata and T. major in this study. The perception of greater predation on ground nests in the tropics may be attributable to greater general predation pressure in the ground stratum, and therefor e on terrestrial species (Martin 1988a). Gonzalez et al. (1998) reported that 40% of the M. ocellata hens that had successfully nested were killed by predators within 15 days of leaving the nest, and poult survival for the breeding season was only 15%. Predation on pou lts during the first tw o-weeks after hatching may represent the greatest limit to population size in M. gallopavo (Williams & Austin 1988). Increased clutch sizes of some terrestrial bi rds are adaptive traits for conditions of high natural mortality (Martin 1988 b). The significantly larger clutch sizes of M. ocellata and T. major are consistent with the predicti on of greater mortality on terres trial birds. The diversity of mating systems within the guild reflects the vari ation observed in clutch sizes. Curassows, guans and chachalacas not only share similar, small clutch sizes but also, as monogamous breeders, exhibit biparental care of young. T. major is polygynandrous; both males and females have multiple partners, but uniparental care is provided by males (Brennan 2004). M. ocellata is polygamous, with uniparental car e provided by females. Polygynandry and polygamy, which are

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47 life history traits consistent with high populatio n turnover, may increase re productive capacity or may improve population response to environmental variabil ity, and are life history. Relative to other general categories of birds, galliforms have lower annual survival rates and a higher rate of population turnover (Gill 1990). Although local im pacts of predation on guild populations could not be directly asse ssed, existing evidence suggests that natural predation pressure on the guild may be significant. Potential predator species observed on the line-transects included jaguar ( Panthera onca ), puma ( Puma concolor ), ocelot ( Leopardus pardalis ), tayra ( Eira barbara), and grey fox ( Urocyon cineoargenteus ). Other locally occurring predators include margay ( Leopardus weidii ), jaguarundi ( Herpailurus yagouaroundi ), ornate hawk-eagle (Spizaetus ornatus ), black hawk-eagle ( Spizaetus tyrannus ), crested eagle ( Morphnus guianensis ), and solitary eagle (Harpyhaliaetus solitarus) (Garcia & Radachowsky 2003, personal observation). The only information regarding the local contri bution of guild members to predator diets is from research on jaguar and puma conducted on the same study area (Novack 2003). The reported biomass proportions of mostly cracids were approximately 4% and 6% of jaguar and puma diet, respectively. Based on the daily caloric requirements of jaguar reported in that study, and applying a 5% correction for non-edible parts, the average jaguar or puma consumes approximately 28 kg of guild biomass annuall y, or the equivalent of about 8 adult C. rubra Applying the density estimates for jaguar and pu ma from Novack (2003) to the current study area of approximately 1000 km2, the local populations of these two predator species consume approximately 1,400 kg of mostly cracids annuall y. Considering that th e human off-take from the guild in the study area averaged 1,625 kg of biomass annually and appeared to have a

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48 significant impact on local populations the natural predation pressure on the guild appears to be significant. Alternative Influences on Guild Structure The influence of other potentially relevant biotic fact ors whose impacts on the guild could not be directly assessed included resour ce heterogeneity, disease, and parasitism. If heterogeneity is significant to the guild, resource gradients should be dive rse and species niches should have greater dimensionality. The heterogeneity of the forest habitat and the broad diet of the guild were consistent with predictions of dive rse resource gradients. One potentially relevant niche dimension that was not addressed was specia lized daily-activity pattern s of guild members. Brooks et al. (2004) reported s ubstantive nocturnal activity for T. major in a diverse tinamou community, indicating that other niche dimensions may be important to the guild. Anecdotal and circumstantial support for the potential relevanc e of disease comes from local reports of an historic regional crash in M. ocellata populations, and region al population crashes of M. gallopavo in North America (Williams 1981; personal observation). Parasitism may be especially relevant in tropical environments (Kricher 1997), especially for terrestrial species, which due to increased contact with the soil, of ten have greater paras ite loads than arboreal species (Gill 1990). Guild Response to Hunting Pressure The recorded harvest is not re presentative of all subsisten ce hunting pressure in the study area. Although local participation in the collecti on effort was significant, there is no evidence that all local hunters partic ipated or that regular participants did so all the tim e. In particular, harvest pressure from camp-based activities far from the village is unlikely to be well represented in the study data. During th e initial stages of specimen collection, O. vetula and T. major were poorly represented. Through the delibera te efforts of some regularly participating

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49 hunters, collection rates for O. vetula were increased significa ntly, although rates for T. major remained low throughout the collection period. Relative to the actua l impacts of local subsistence hunting, the number of individuals harvested is expected to be slightly conservative for C. rubra M. ocellata P. purpurascens and T. major and robust for O. vetula Because the proportion of sub-adult birds harves ted could not be determined, the application of average adult body-mass values to the recorded harvest for each guild species probably overestimated the recorded harvest biomass. However, additional harvest pressure attributable to camp-based extraction activities (McNab 1998; Morales & Morales 1998) may compensate for any overestimation. The harvest data indicated a disproportionately greater mean annual harvest of C. rubra (47%) than of other guild member s, which are consistent with pr eviously findings from the area (Morales & Morales 1998). Mean annual harvest proportions were intermediate and relatively similar for O. vetula (20%), M. ocellata (16%), and P. purpurascens (12%), and were consistently low for T. major (5%). These harvest levels we re inconsistent with previous findings, which indicated that the harvest of P. purpurascens exceeded that of M. ocellata by a factor of more than 2:1, and the harvest of M. ocellata exceeded that of O. vetula by a factor of more than 5:1 (Morales & Morales 1998). Selectivity for C. rubra M. ocellata and P. purpurascens by local hunters is consistent with population trends along the hunting gradient Based on the changes observed in abundance estimates and guild composition between transect groups, local hunting pr essure appeared to have significant impacts on populations of these three species. Although O. vetula and T. major abundance was greatest in the protected Ixcan transect group, abundance among the other three groups did not appear to be related to distance from the village of Uaxactn.

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50 The combined population size of the guild in the heavily hunted Uaxactn South group (1,930 individuals) represented 45% of the combined guild population size in the protected Ixcan group (4,332 individuals). Estimated population sizes for Uaxactn South e quated to 17% of the Ixcan population for C. rubra 28% for M. ocellata 89% for O. vetula 31% for P. purpurascens and 79% for T. major Population biomass estimates were calculated usin g average adult body mass values and were therefore robust; however, these estimates should adequately reflect the relative differences between populations and sites. The combined biomass of the guild populations in the Uaxactn South group (3,215 kg) represented 30% of the combined biomass in the Ixcan group (10,587 kg). Based on the cha nges observed in the guild along the gradient, local subsistence hunting pressure has signi ficantly reduced local guild populations. Conclusions Guild char acteristics that were consistent with predictions of structural regulation by competitive interactions included: separation along habitat dimens ions, vertical stratification, limited size assortment, and apparent competitive rel ease by T. major. Guild characteristics that were inconsistent with regulation by competition included: lack of diet-niche specialization, considerable dietary overlap, lim ited specialization of habitat nich es, and the similarity of guild membership to the regional null pool. Additiona l evidence contrary to regulation by competition derives from the high abundance and inter-annual in crease all guild members in the protected Ixcan transect group. If the fitness of indivi dual guild members is constrained by competitive interactions with other guild memb ers, then it is counterintuitiv e that all species would exhibit greatest abundance in the same area at the sa me time, and that all species would exhibit abundance increases in the same area during the same time period. Guild characteristics that were consistent wi th non-equilibrium pro cesses included: limited niche specialization, limited ecologi cal separation, and the high va riability of species abundance

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51 and habitat occupancy. Guild char acteristics that were consistent with external limitations by predation included: spatial separati on of nest sites and high reproduc tive potential of M. ocellata and T. major. Based on limited evidence of competition, evidence of non-e quilibrium processes, and evidence of the importance of predation, the characteristics of the guild are most consistent with the non-equilibrium model of competition proposed by Connell (1980). The limited niche specializations and separation along resource dimens ions may be sufficient to minimize direct interactions among species. The unpredictabl e local abundance of competitor species and external limitation of populations by predat ion create conditions under which dietary generalization is more adaptive than dietar y specialization. Guild-member populations are unlikely to reach the carrying capac ity of the environment, therefor e, competitive interactions are held below levels that would lead to the co mpetitive exclusion of potential guild members.

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52 Table 3-1. General diet compositions of guild members based on dry-mass proportions of contents from upper-digestive tract specime ns collected in Uaxactn, Flores, El Petn, from January 2000 through December 2001. Diet compositions with all sample contents Crax Meleagris Ortalis Penelope Tinamus ( n = 267) ( n = 181) ( n = 205) ( n = 142) ( n = 55) Flower/Stem 0.004 0.008 0.013 0.002 0.001 Animal 0.015 0.121 0.007 0.011 0.024 Leaf 0.015 0.65 0.063 0.014 0.007 Pulp 0.066 0.07 0.068 0.087 0.065 Grit 0.204 0.222 0.027 0.093 0.022 Seed 0.696 0.514 0.822 0.793 0.881 Diet compositions with grit and snail contents excluded Crax Meleagris Ortalis Penelope Tinamus Flower/Stem 0.005 0.012 0.013 0.002 0.001 Animal 0.016 0.011 0.006 0.012 0.024 Leaf 0.019 0.098 0.064 0.015 0.007 Pulp 0.083 0.105 0.07 0.096 0.066 Seed 0.877 0.774 0.847 0.875 0.902 Proportions include pulverized material detected in samples. The dry-mass of pulverized material was assigned to the values of separable items in each sample according to their relative proportions. Table 3-2. Overlap of dietar y seed components between spec ies-pairs and average seed component overlap of individual spec ies with all other guild members. Species-pair Seed component Overlapa Species Average Overlapb Crax Meleagris 57.9% Crax 62.6% Crax Penelope 67.9% Crax Ortalis 39.1% Meleagris 53% Crax Tinamus 85.6% Meleagris Penelope 57.4% Ortalis 46.8% Meleagris Ortalis 32.9% Meleagris Tinamus 63.6% Penelope 66% Penelope Ortalis 65.4% Penelope Tinamus 73.3% Tinamus 68.1% Ortalis Tinamus 49.8% a Calculated using equation five from Pianka (1973), pulverized material in samples was not included. b Average seed component overlap with all other guild members.

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53 Table 3-3. Overlap of habitat occupancy between species-pairs and average combined overlap of individual species with all other guild members. Species-pair Habitat Overlap a Species Average Overlap b Crax Meleagris 79.5% Crax 77.3% Crax Penelope 94.2% Crax Ortalis 50.3% Meleagris 82.6% Crax Tinamus 85% Meleagris Penelope 85.4% Ortalis 60.7% Meleagris Ortalis 70.8% Meleagris Tinamus 94.5% Penelope 81.7% Penelope Ortalis 56.2% Penelope Tinamus 90.9% Tinamus 83.9% Ortalis Tinamus 65.3% a Calculated using proportional similarity index equation from Schoener (1970). b Average combined habitat occupancy ove rlap with all other guild members. Table 3-4. Overlap of vertical strata occupancy between species-pairs and average combined vertical overlap of individual spec ies with all other guild members. Species-pair Vertical Overlap a Species Average Overlap b Crax Meleagris 43.5% Crax 56.4% Crax Penelope 73.8% Crax Ortalis 68.3% Meleagris 55.6% Crax Tinamus 40% Meleagris Penelope 17.3% Ortalis 41.8% Meleagris Ortalis 17.1% Meleagris Tinamus 93.1% Penelope 43.3% Penelope Ortalis 68.3% Penelope Tinamus 13.8% Tinamus 40.1% Ortalis Tinamus 13.6% a Calculated using proportional similarity index equation from Schoener (1970). b Average combined vertical overlap in both forest-classes with all other guild members.

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54 Table 3-5. Density estimates for Crax rubra Meleagris ocellata Ortalis vetula Penelope purpurascens and Tinamus major by forest-class in the Maya Biosphere Reserve, Guatemala. Transect group Ixcan Ixcan Cedro Cedro Uaxactn North Uaxactn South Sampling period 20002001 20012002 20002001 20012002 2000-2001 2000-2001 Upland forest densities (km2) Crax 31.19 31.05 21.37 26.27 14.44 5.46 Meleagris 15.51 24.63 6.44 16.7 8.04 4.7 Ortalis 11.25 12.56 14.51 22.91 3.37 18.51 Penelope 24.31 28.29 16.43 15.78 7.27 6.73 Tinamus 8.44 14.64 5.46 5.16 10.96 9.13 Scrub forest densities (km2) Crax 14.17 19.54 6.12 9.94 6.18 2.42 Meleagris 15.19 17.41 3.4 22.73 7.87 5.2 Ortalis 37.22 56.03 24.19 25.29 19.03 25.01 Penelope 11.11 15.81 6.12 6.28 4.56 6.32 Tinamus 10.37 8.97 6.8 6.54 6.39 4.51 Density estimates calculated using DI STANCE version 5, release Beta 5. Table 3-6. Guild compositions based on proportional population size and relative biomass at each transect group. Transect group Ixcan Ixcan Cedro Cedro Uaxactn North Uaxactn South Sampling period 20002001 20012002 20002001 20012002 2000-2001 2000-2001 Proportional population size Crax 0.28 0.24 0.317 0.289 0.236 0.106 Meleagris 0.172 0.167 0.098 0.197 0.181 0.109 Ortalis 0.227 0.247 0.248 0.264 0.25 0.455 Penelope 0.219 0.213 0.246 0.168 0.135 0.15 Tinamus 0.102 0.133 0.091 0.082 0.198 0.18 Proportional population biomass Crax 0.391 0.351 0.464 0.399 0.355 0.216 Meleagris 0.294 0.302 0.177 0.336 0.334 0.274 Ortalis 0.059 0.068 0.068 0.069 0.071 0.175 Penelope 0.207 0.212 0.245 0.157 0.138 0.208 Tinamus 0.049 0.067 0.046 0.039 0.102 0.127 Population sizes were based on density estimates by forest-class extrapolated by the availability of each forest-class within the 50km2 area corresponding to each transect group. Population biomass estimates were based on population size and average adult body mass of each species.

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55 Table 3-7. Temporal reproductive patterns of Crax rubra, Meleagris ocellata Ortalis vetula Penelope purpurascens and Tinamus major based on the number of monthly observations of repr oductive activities. Category Jan. Feb. Mar. Apr. May Jun. Jul. Aug.Sep. Oct. Nov.Dec. Crax rubra Calls 8 14 38 36 46 30 8 4 1 Nests 8 1 5 Young 2 5 7 10 1 1 2 Meleagris ocellata Calls 1 21 43 38 2 1 Nests 3 9 4 Young 1 1 4 1 2 2 2 Ortalis vetula Nests 1 7 30 31 Young 3 5 7 5 3 3 1 Penelope purpurascens Nests 3 3 4 3 Young 3 5 8 1 Tinamus major Nests 1 6 9 12 1 1 Young 2 3 6 8 3 Calls include records of vocalizations specific to re productive behavior which were recorded only for C. rubra and M. ocellata Nests include only observations of nests with laying or incubating adults present. Young include observations of dependent young and hatched nests observed during the nesting period.

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56 Figure 1-1. Map of study area. Lightly shaded ar eas indicate the two management units in which study data were collected. Circles depict the 50km2 areas corresponding to each transect group.

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64 BIOGRAPHICAL SKETCH At his point I have lived r oughly equal proportions of m y lif e in Texas, New Jersey, and Guatemala. I received a B. S. in zoology and a nother in wildlife management from Texas Tech University in Lubbock, Texas. After graduating from Texas Tech I entered the U. S. Peace Corps and was stationed in Guatemala where I worked with non-governmental organizations working in the two largest prot ected areas in that nation, firs t in the Sierra de las Minas Biosphere Reserve and later in the Maya Biosph ere Reserve. My interests in terrestrial vertebrate ecology and sustainable wildife resource-u se lead me to pursue graduate studies at the Department of Wildlife Ecology and Conservation at the University of Florida. Since 1999 I have co-developed and managed the Guatemal an operations of a community-based, sporthunting, conservation project for the Ocellated Turkey working in community forestry concessions in the multiple-use zone of the Maya Biosphere Reserve.


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