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Habitat Assessment for a Threatened Marsupial in Temperate Forest of Patagonia

Permanent Link: http://ufdc.ufl.edu/UFE0021848/00001

Material Information

Title: Habitat Assessment for a Threatened Marsupial in Temperate Forest of Patagonia
Physical Description: 1 online resource (40 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: 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: Patterns and processes that shape the distribution and abundance of animals depend on the scale at which they are evaluated. Understanding theses patterns and processes is fundamental for the study of plant-seed disperser interactions. Most plants that depend on frugivorous animals for seed dispersal are dispersed by several species of animals and, the majority of animal dispersers consume the fruit of several plant species. In contrast to the weak mutual dependence of most plants and their seed dispersers, some mutualisms in the temperate forest of Patagonia appear to be obligate. The northern portion of the temperate forest harbors a unique plant-animal interaction in which the seeds of the mistletoe Tristerix corymbosus, a proposed key species, are dispersed solely by the endemic marsupial Dromiciops gliroides. I developed a set of habitat models, in which model variables were defined a prior, to test the hypothesis that the distribution and abundance of monito del monte is associated principally with abundance of mistletoe in the temperate forest of Patagonia or that, alternatively, habitat structure and/or other plants determine the distribution and abundance of this marsupial. Results show that the distribution of monitos del monte at the broad scale was strongly influenced by bamboo cover. However, at a smaller spatial scale, the abundance of monitos del monte are explained by the abundance of mistletoe plants and fruits. The lack of dependence among fruiting plants and their seed dispersers often is attributed to the high asymmetry in these interactions and the important influence of habitat features. Conversely, my results showed that the mistletoe?monito del monte interaction is more symmetric than many other plant?frugivore interactions. This interaction is characterized by an obligate partnership of the mistletoe with the marsupial, and a strong relationship between the abundance of monitos del monte and the abundance of mistletoe.
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: Branch, Lyn C.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0021848:00001

Permanent Link: http://ufdc.ufl.edu/UFE0021848/00001

Material Information

Title: Habitat Assessment for a Threatened Marsupial in Temperate Forest of Patagonia
Physical Description: 1 online resource (40 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: 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: Patterns and processes that shape the distribution and abundance of animals depend on the scale at which they are evaluated. Understanding theses patterns and processes is fundamental for the study of plant-seed disperser interactions. Most plants that depend on frugivorous animals for seed dispersal are dispersed by several species of animals and, the majority of animal dispersers consume the fruit of several plant species. In contrast to the weak mutual dependence of most plants and their seed dispersers, some mutualisms in the temperate forest of Patagonia appear to be obligate. The northern portion of the temperate forest harbors a unique plant-animal interaction in which the seeds of the mistletoe Tristerix corymbosus, a proposed key species, are dispersed solely by the endemic marsupial Dromiciops gliroides. I developed a set of habitat models, in which model variables were defined a prior, to test the hypothesis that the distribution and abundance of monito del monte is associated principally with abundance of mistletoe in the temperate forest of Patagonia or that, alternatively, habitat structure and/or other plants determine the distribution and abundance of this marsupial. Results show that the distribution of monitos del monte at the broad scale was strongly influenced by bamboo cover. However, at a smaller spatial scale, the abundance of monitos del monte are explained by the abundance of mistletoe plants and fruits. The lack of dependence among fruiting plants and their seed dispersers often is attributed to the high asymmetry in these interactions and the important influence of habitat features. Conversely, my results showed that the mistletoe?monito del monte interaction is more symmetric than many other plant?frugivore interactions. This interaction is characterized by an obligate partnership of the mistletoe with the marsupial, and a strong relationship between the abundance of monitos del monte and the abundance of mistletoe.
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: Branch, Lyn C.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0021848:00001


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HABITAT ASSESSMENT FOR A THREATENED MARSUPIAL INT TEMPERATE FOREST
OF PATAGONIA




















BY

MARIANO ALBERTO RODRIGUEZ-CABAL


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

































O 2008 Mariano Alberto Rodriguez Cabal


































To Noelia
and to all who nurtured my curiosity and love for nature.









ACKNOWLEDGMENTS

I thank my supervisory committee (L. C. Branch, K Sieving and E. Bruna) for their

mentoring. I also thank the faculty, staff and grad students at the University of Florida (UF)

Department of Wildlife Ecology and Conservation. I thank Mauricio Failla, Director de Fauna de

la Provincia de Rio Negro, Argentina; Gustavo Iglesias and Claudio Chehebar, Nahuel Huapi

National; Park, and staff of Parque Municipal Llao Llao. Special thanks go to M. Aizen, G.

Amico, N. Barrios, G. Barrios, R. Bunge, J. M. Morales, S. Morgan, A. Martinez, M. Nufiez, E.

Perner, A. Pries, R. Reggio, and N. Tercero-Bucardo for valuable assistance in the field and for

their thoughtful comments and suggestions on previous drafts. Field work was supported by

Wildlife Conservation Society, Scott Neotropical Fund, and the Jennings Scholarship of the UF

Department of Wildlife Ecology and Conservation.












TABLE OF CONTENTS


page

ACKNOWLEDGMENTS .............. ...............4.....


LIST OF TABLES .........__.. ..... .__. ...............6....


LIST OF FIGURES .............. ...............7.....


AB S TRAC T ......_ ................. ............_........8


CHAPTER


1 HABITAT AS SES SMENT FOR MONITOS DEL MONTE ................ ................. .... 10


Introducti on ................. ...............10.................
M ethods .............. ...............13....

Study Area ................. ............ ...............13.......
Marsupial Presence Ab sence ................. ...............14........... ...
Marsupial Abundance............... ...............1
Habitat Variables ................ ...............15.................
Data Analysis............... ...............16
Re sults ................. ............ ...............19.......
Marsupial Presence Ab sence ................. ...............19........... ...
Marsupial Abundance............... ...............2
Discussion ................. ...............21.................


2 CONSERVATION IMPLICATIONS .............. ...............25....


3 CONCLUSION................ ..............3


APPENDIX


A CORRELATION ANALYSIS .............. ...............33....


LIST OF REFERENCES ................. ...............35........... ....


BIOGRAPHICAL SKETCH .............. ...............40....










LIST OF TABLES


Table page

1-1 Habitat variables for sites where monitos del monte were present or absent. ........._.......25

1-2 Logistic regression models at the 28 study sites.. ............. ...............26.....

1-3 Regression models constructed at 12 study sites.. ............ ...............27.....

A-1 Correlation coefficients between habitat variables in the presence absence survey ......33

A-2 Correlation coefficients between habitat variables in the abundance survey. ...................34










LIST OF FIGURES


Figure page

1-1 Habitat variables at sites where abundances of monito del monte were determined.. .......28

1-2 Number of monitos del monte vs. the significance predictor variables. ................... .........29









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

HABITAT ASSESSMENT FOR A THREATENED MARSUPIAL INT TEMPERATE FOREST
OF PATAGONIA

By

Mariano Alberto Rodriguez-Cabal

May 2008

Chair: Lyn C. Branch
Major: Wildlife Ecology and Conservation

Patterns and processes that shape the distribution and abundance of animals depend on the

scale at which they are evaluated. Understanding theses patterns and processes is fundamental

for the study of plant-seed disperser interactions. Most plants that depend on frugivorous

animals for seed dispersal are dispersed by several species of animals and, the majority of animal

dispersers consume the fruit of several plant species. In contrast to the weak mutual dependence

of most plants and their seed dispersers, some mutualisms in the temperate forest of Patagonia

appear to be obligate. The northern portion of the temperate forest harbors a unique plant -

animal interaction in which the seeds of the mistletoe Tristerix corymbosus, a proposed key

species, are dispersed solely by the endemic marsupial Dromiciops gliroides. I developed a set

of habitat models, in which model variables were defined a prior, to test the hypothesis that the

distribution and abundance of monito del monte is associated principally with abundance of

mistletoe in the temperate forest of Patagonia or that, alternatively, habitat structure and/or other

plants determine the distribution and abundance of this marsupial.

Results show that the distribution of monitos del monte at the broad scale was strongly

influenced by bamboo cover. However, at a smaller spatial scale, the abundance of monitos del

monte are explained by the abundance of mistletoe plants and fruits. The lack of dependence










among fruiting plants and their seed dispersers often is attributed to the high asymmetry in these

interactions and the important influence of habitat features. Conversely, my results showed that

the mistletoe-monito del monte interaction is more symmetric than many other plant-frugivore

interactions. This interaction is characterized by an obligate partnership of the mistletoe with the

marsupial, and a strong relationship between the abundance of monitos del monte and the

abundance of mistletoe.









CHAPTER 1
HABITAT ASSESSMENT FOR MONITOS DEL MONTE

Introduction

Distribution and abundance of species depends on the distribution and abundance of other

species with which they interact and habitat factors a various scales (wiens, 1989).

Understanding the patterns and processes that shape distribution and abundance is fundamental

for the study of plant-seed disperser interactions and biodiversity conservation. Seed dispersal

plays an important role in creating and maintaining diversity in plant communities (Schupp et al.,

2002). At large scales, appropriate habitat may be the most important factor limiting distribution

and abundance of frugivores (Wiens, 1989). Habitat characteristics such as availability of a

variety of food resources, nest sites, and distance to shelter, influence frugivores distribution and

could be play a critical role in regulating the abundance of these species (Fedriani, 2005). At a

small scale, habitat characteristics in which fruiting plants are surrounded may be more

important (Herrera, 1998). Previous studies have demonstrated that frugivores select plants or

good patches based on habitat features rather than on plant phenotypic traits (Herrera, 1998;

Fedriani, 2005). For example, frugivores could minimize predation risk by selecting sites with

high cover of vegetation to avoid predators during foraging (Howe, 1979). Thus, habitat factors

may limit the degree to which frugivorous animals track their food resource, and human impacts

on habitat of interacting species can have cascading effects.

Seed dispersal mutualisms influence patterns of seedling recruitment, plant demography,

and population persistence, and thus potentially have widespread effects on the rest of the

community (Janzen, 1980; Feinsinger, 1987; Bond, 1994; Levey and Benkman,1999; Herrera,

2002; Spiegel and Nathan, 2007; Rodriguez-Cabal et al., 2007). The maj ority of plants that

depend on frugivorous animals for seed dispersal are dispersed by several species of animals










and, most animal dispersers consume the fruit of several plant species (Herrera, 1998; 2002).

Mutual dependence of most plants and seed dispersers is weak, and asymmetry is common.

Asymmetry in plant-animal interactions is represented in terms of the effect of one species on

another species relative to the strength of the reciprocal effect (Vazquez and Aizen, 2004;

Vazquez et al., 2005; Bascompte et al., 2006; Vazquez et al., 2007). Asymmetry is particularly

evident for specialist plants, which tend to interact mostly with generalist animals (Vazquez and

Aizen, 2004; Bascompte et al., 2003; Bascompte et al., 2006). Generalist plants most often

interact with generalist and specialist animals (Vazquez and Aizen, 2004; Bascompte et al.,

2003; Bascompte et al., 2006).

In contrast to the weak mutual dependence of most plants and their seed dispersers, some

mutualisms in the temperate forest of Patagonia appear to be obligate. This system provides an

ideal opportunity to examine whether the distribution and abundance of an animal disperser

depends on the distribution and abundance of a specific fruit. A large proportion of the flora in

this temperate forest depends on mutualistic animals (Armesto et al., 1996; Aizen and Ezcurra,

1998; Aizen et al., 2002). This level of mutualism is one of the highest recorded for a temperate

ecosystem and is comparable to those from tropical forests (Willson et al., 1989; Willson, 1991).

Unlike the mutualism structure in tropical forests, in this temperate forest many species of flora

depend on a few mutualistic animals for their survival (Aizen et al., 2002).

The northern portion of the temperate forest harbors a unique triangle of keystone

mutualists comprised of a hummingbird (Sephanoides sephaniodes), a mistletoe (quintral,

Tristerix corymbosus), and the marsupial (monito del monte, Dromiciops gliroides, Amico and

Aizen, 2000). This hummingbird is responsible for pollinating nearly 20% of the endemic

species of woody flora in this biome (Armesto et al., 1996; Aizen et al., 2002). The mistletoe









blooms in winter and is the only source of nectar for the hummingbird during that period (Smith-

Ramirez, 1993; Aizen and Ezcurra, 1998). The monito del monte is the only known disperser of

the mistletoe in this temperate forest (Amico and Aizen, 2000). Moreover, the passage of the

seed through the marsupial's gut is crucial to triggering germination (Amico and Aizen, 2000).

The isolation from other similar biotas since the disruption of Gondwana allowed the evolution

of monito del monte which is the only extant member from the ancient family Microbiotheriidae.

Because of the long evolutionary history of association of monitos del monte and the

mistletoe and the specific fruiting characteristics and distribution of mistletoes in this temperate

forest, I hypothesized that the distribution and abundance of monitos del monte and mistletoe

might be related closely. Mistletoe produces a large crop size every year and is the most

abundant fleshy fruit in the northern temperate forest of Patagonia (Aizen, 2003, Amico et al., in

press). Once mistletoe fruits are ripe, monitos del monte consume almost exclusively mistletoe

fruits (Rodriguez-Cabal et al., 2007). This mistletoe presents an aggregated distribution and,

therefore, may serve as high density resource patches for the monito del monte. Alternatively,

populations of monitos del monte may not track any single plant species as they are known to eat

and disperse seeds of 80% of the flora with fleshy fruit (Amico et al., in press). Because of the

diversity of fruits available, their populations maybe constrained largely by other habitat factors

such as vegetation cover and structural complexity of the habitat that influence nest site

availability, predation risk, and other aspects of their biology. In this study, I developed a set of

habitat models, in which model variables were defined a prior, to examine the hypothesis that the

distribution and abundance of monito del monte is associated principally with abundance of

mistletoe in the temperate forest of Patagonia or that, alternatively, habitat structure and/or other

plants determine the distribution and abundance of this marsupial. Although experiments are










required to fully determine the degree of mutual dependence between monitos del monte and the

mistletoe, landscape scale studies such as this provide a foundation for understanding

environmental constraints on these relationships.

Methods

Study Area

I conducted my research in Nahuel Huapi National Park (705,000 ha) and Llao-Llao

Municipal Reserve (1226 ha) in northwestern Patagonia. Nahuel Huapi National Park

encompasses a marked west-to-east rainfall gradient caused by the Andes, which form an

effective barrier to westerly air masses coming from the southern Pacific Ocean. The most xeric

forest site in my study, situated at the xeric boundary with the Patagonian steppe (Cerro Otto,

41012' S, 71019' W), has a mean annual precipitation of 1200 mm of which only 10.5 % falls

during the growing season. The western limit of my study located in Puerto Blest (410 01' S, 71o

49' W) is an area of Valdivian rainforest, one of the rainiest places in the Patagonia (mean annual

precipitation, 4000 mm). Llao-Llao Municipal Reserve (410 08' S, 71o 19' W) has intermediate

rainfall (mean annual precipitation, 1800 mm). Mean January temperature is 15oC, and mean

July temperature is 3oC.

The native forest vegetation in the study area belongs to the Subantartic biogeographical

region, which is distinctive from the Neotropical forest of the rest of South America (Cabrera,

1976). The most common trees are evergreen southern beech (Nothofagus dombeyi), a lenga

beech (Nothofagus pumilio) and a conifer (Austrocedrus chilensis). The understory is dominated

by the bamboo (Chusquea culeou) and the shrub (Aristotelia chilensis, here after A. chilensis).

That is the main host of the mistletoe in the study area (Mermoz and Martin, 1986). The two

forest layers are well differentiated with tree canopy reaching up to 40-m height and understory

reaching up to 5-m height.









I conducted presence-absence surveys of monitos del monte and examined habitat

variables at 16 sites in Nahuel Huapi National Park and 12 sites in the adj acent Llao-Llao

Municipal Reserve. In addition, trapping was conducted at the 12 sites in the Llao-Llao

Municipal Reserve to determine the relationship between abundance of monitos del monte and

abundance of mistletoe. Sites in Nahuel Huapi National Park were chosen randomly to

incorporate the range of habitats found in the study area. Potential sites in the Llao-Llao

Municipal Reserve were stratified by mistletoe density, and four sites were chosen randomly

within each of the following categories: high (>3 5 reproductive individuals of mistletoe), low

(<20 reproductive individuals), and no mistletoe. Sites were stratified on abundance of mistletoe

because mistletoe has a patchy distribution and a range of all other variables can be found within

each stratum defined by mistletoe density. I conducted most of the field work during the austral

summer (December 2006 -March 2007) because monitos del monte hibernate during winter and

the main fruiting and dispersal season of the mistletoe is summer (Aizen 2003).

Marsupial Presence Absence

In order to facilitate rapid surveys of presence absence of monitos del monte, the 28 study

sites were grouped by presence or absence of mistletoe. At sites with mistletoe, I examined all

plants < 15 cm DHB in each site for feces of monitos del monte that contained mistletoe seeds. I

concluded that monitos del monte were present if I found mistletoe seeds dispersed. If no

dispersed seeds were found in the site, I used trapping to determine the presence of monito del

monte. This technique also was used for sites with no mistletoe. Within each site, I placed a 5 x 5

grid (0.25 ha) of Tomahawk-style traps about 10-m apart. Each trap was placed between 1-2 m

above a ground in the shrub closest to the sample point. Traps were baited with apple and

banana. Because the obj ective was to document presence, once a monito del monte was trapped,

I stopped trapping at this site. Sampling was continued for four nights if no monitos del monte









were captured. Based on previous capture data for monitos del monte (Rodriguez-Cabal, 2003),

and following procedures in MacKenzie et al. (2002), probability of capture was 99.6% in four

night if monitos del monte were present. I concluded that monitos del monte were absent after

four nights of trapping if no marsupials were captured.

Marsupial Abundance

Abundance of the monito del monte was estimated using multiple capture-recapture

methods with trapping grids as described above. One trapping grid was placed at each of the 12

sites in Llao-Llao Municipal Reserve. Animals were trapped for four periods of four nights each

per site. Traps were checked daily and all trapped marsupials were marked with an individual

code based on ear perforations and released at the point of capture (for more detail see

Rodriguez-Cabal, 2003; Rodriguez-Cabal et al., 2007). Recaptures were noted in subsequent

trapping periods.

Habitat Variables

At each of the 16 sites in Nahuel Huapi National Park, a 5 x 5 grid was established with

points 10 m apart. I measured habitat variables in 5-m radius circular plots centered on every

second point (n = 13 per site). Similar sampling was conducted at sites trapped in Llao-Llao

Municipal Reserve with plots centered on every second trap. I measured the following variables

to represent the biological and physical structure of the habitat: number of reproductive

individuals and crop size of three understory plants that produce fleshy fruits (mistletoe, A.

chilensis, and Azara microphylla), cover and height of bamboo, complexity of habitat structure,

canopy cover, understory cover and the number of the two most common tree species (N.

dombeyi and Austrocedrus chilensis). Reproductive individuals were plants that presented sexual

structures (i.e., flowers or fruits). In addition for mistletoe, I counted the number of non-

reproductive individuals (2-4 years old), seedlings (those presenting the first two true leaves),









and number of seeds dispersed. To determine crop size of understory plants, I counted the

number of fruits on a subset of three branches dispersed in each crown and then estimated total

crop size by multiplying mean number of fruits per branch by total number of branches in the

crown for each plant. As with mistletoe, seeds ofA. chilensis and A. microphylla are dispersed

by monitos del monte, and passage of their seeds through the digestive tract of a monito del

monte increases chance of germination (Amico et al., in press). Cover of bamboo was estimated

visually by summing the total area occupied by bamboo stems and leaves in each plot. Also, I

measured the height of the tallest bamboo stem in each plot. As a measure of habitat complexity

within each plot, I counted the number of contacts with a vertical pole (3-m height) for branches

5 10-cm in diameter and oriented < 450 relative to the ground. Good connectivity between

plants in the understory stratum could play a key role in the mobility of this animal, and more

seeds dispersed in monito del monte feces are found on branches with these characteristics

(Amico 2000). Percent cover of the canopy and understory were estimated using a densiometer

at the center of the circular plot. Finally, I counted the total number of the most common trees for

all individuals taller than four meters.

Data Analysis

I used multiple logistic regressions to determine which habitat variables predicted monito

del monte presence-absence at the 28 study sites, and I used multiple regressions to assess the

contribution of habitat variables to explaining abundance of monitos del monte at the 12 study

sites in Llao-Llao Municipal Reserve. Data were log-transformed when necessary to achieve

normality and reduce heteroscedasticity (Table 1-1). For both analyses, I only used explanatory

variables that did not correlate significantly after a Bonferroni correction (Appendix A-1 and A-

2; Tables 1-1, 1-2 and 1-3). I did not used bamboo height and understory cover because these

measures were correlated with bamboo cover and habitat complexity, respectively, and bamboo









cover and habitat complexity may be more directly related to the biology of monitos del monte,

particular for nest material and mobility. I did not include crop size ofA. microphylla in

analyses because this understory plant did not produce fruits during the study.

Seven models were built to identify habitat variables related to presence absence and

abundance of monitos del monte. Rather than examine all possible models, models were

developed a priori, that focused on mistletoe, other food plants, and habitat structure as key

factors in the distribution and abundance of this marsupial. I examined the following models: (1)

Global -- all habitat variables, (2) Food plants -- number of reproductive individuals of maj or

food plants that produce fleshy fruits eaten by monitos del monte, (3) Fruit abundance and

bamboo -- crop size of plants with fleshy fruits and cover of bamboo, (4) Mistletoe plants and

bamboo -- number of reproductive mistletoes and bamboo cover, (5) Mistletoe plants, (6)

Bamboo, (7) Habitat structure -- all habitat variables unrelated to food plants (Table 1-2 and 1-

3). Bamboo cover was included as a separate variable because this plant is an important source

of nest material (Jimenez and Rageot 1979), provides connectivity for arboreal movement, and

may inhibit predation by aerial predators. I used an approach that compared the suite of models

using Akaike' s information criteria (AIC), which allows direct comparison of models with

different number of parameters. AIC is calculated for a suite of models and the best fitting one

has the smallest AIC (Burnham and Anderson, 2002; Whittingham et al., 2005). I applied small-

sample version of AIC (AICc), because the ratio of number of observations to number of

parameters was <40. AICc was calculated for each model with the algorithm:

a) AIC = -2 log (L) + 2 K



b) AICc = -2 log (L) + 2 K + (2 K (K + 1))/ (n K 1)









where log (L) is the log-likelihood function evaluated for the maximum likelihood parameter

estimates, K is the number of parameters in the model and n is the number of observations. AAIC

represents the difference between the AICc score for each model and the AICc score of the best

model. The absolute value of AIC is unimportant; instead AAIC indicates relative support for the

models. AAIC values <2 indicate that models are similar. When AAIC was <2, I chose the most

parsimonious model as the best model (i.e., the model that used the least number of parameters).

To evaluate goodness-of-fit for the multiple logistic regressions models, I used Nagelkerke

R2. A true R2, which measures the variability in the dependent variable that is explained by a

linear regression models, cannot be calculated for logistic regression models (Hosmer and

Lemeshow, 1989). However, because deviance can be thought of as a measure of how poorly the

model fits (i.e., lack of fit between observed and predicted values), an analogy can be made to

sum of squares residual in ordinary least squares. This pseudo-R2 is designed to have similar

properties to the true R2. The pseudo-R2 is based on comparison of the likelihood of the current

model to the "null" model (one without predictors). A larger pseudo-R2 indicates that more of the

variation is explained by the model. Here, I reported the Nagelkerke R2 which is a modified

version of Cox and Snell R2 that ranges from 0 to 1 (Hosmer and Lemeshow 1989). I evaluated

the importance of significant independent variables in logistic regression using the standardized

regression coefficient p, which is the predicted change in odds for a unit increase in the

corresponding independent variable (Hosmer and Lemeshow, 1989). Values of P <1 correspond

to decreases in the odds with increases in the dependent variable; values >1 correspond to

increases in the odds with increases in the independent variable; and values close to 1 indicate

that unit changes in the independent variable do not affect the dependent variable.









To evaluate the goodness-of-fit for the multiple regression models for abundance of

monitos del monte, I calculated the R2 Of the models. In multiple regressions, I evaluated the

importance of significant independent variables by calculating the standardized regression

coefficient p (Ganas and Robbins, 2005). The standardized regression coefficient P is used to

compare the magnitude of effect of different variables by describing the change in the dependent

variable (in SD units) caused by an increase in one unit (SD) of the independent variable, while

all the other independent variables are held constant.

In addition, I compared habitat variables between sites with and without monitos del monte

and at sites where I measured marsupial abundances using an ANOVA. I performed a Chi-square

test to compare the relationship between the presence of monitos del monte and mistletoe

presence at the 28 sites. For the 12 sites where I measured abundance of monitos del monte, I

report the minimum number alive for all sites to allow comparison among sites (Wilson et al.,

1996). Because of the low number of captures in sites without mistletoe, I could not use other

methods. Finally, I performed a regression analysis to examine the relationship between

abundance of monitos del monte and habitat variables that had significant explanatory power in

the multiple regression models. These analyses were conducted using SAS 9.13 and JMP IN for

Windows Ver. 5.1 (SAS Institute, Inc., Cary, NC).

Results

Marsupial Presence-Absence

Three of the seven multiple logistic regression models received substantial support as the

best predictive models of presence-absence of monitos del monte (Table 1-2). The Global model

perfectly predicted the presence-absence of monitos del monte (i.e., Nagelkerke R2 = 1). The

most parsimonious model (Fruit abundance and bamboo) and the Habitat structure model, which

was not ranked in the best models, also explained considerable variation in presence-absence of









monitos del monte (Table 1-2). Bamboo cover was the only significant predictor variable in the

top models (Table 1-2). The seventeen sites where I found marsupials had more bamboo cover

than sites without monito del monte (Table 1-1). Monito del monte was recorded 91% of the

sites that had more than 10% bamboo cover and in only 35% of the sites with less than 10%

cover. The presence-absence of mistletoe and monito del monte were not related at the 28 sites

(Chi-square = 2.06, df = 1, P = 0. 15). Five of the 17 sites where I recorded presence of monito

del monte did not have mistletoe. Five of the eleven sites where monitos del monte were absent

had reproductive mistletoe plants, but I did not find mistletoe seeds dispersed, seedlings or non-

reproductive mistletoe (2-4 years old). The lack of seeds, seedlings and non-reproductive

individuals indicates that there has not been recruitment at least in the last four years. The other

12 sites had mistletoe seeds dispersed in monito del monte feces and seedlings.

Marsupial Abundance

The estimated minimum number alive of marsupials on study plots ranged between 0 and

45 individuals. At eleven of the 12 sites where I measured abundances, I captured monitos del

monte in the first four nights of trapping, in spite of the high variability in local abundance of

this species. Two regression models were candidates for the best predictive model for abundance

of monitos del monte, although several additional models also explained high amounts of

variation in abundance of these marsupials (Table 1-3). The number of reproductive mistletoe or

crop size of mistletoe were significant variables in five of the seven models. Habitat complexity,

bamboo cover, and crop size ofA. chilensis appear as significant variables in models that were

not supported as the best models, but P-values for these variables were lower (Table 1-3).

Marsupial abundance increased when the number of reproductive mistletoe, crop size of

mistletoe and A. chilensis, or habitat complexity increased (see regression coefficients P, Table

1-3). In contrast, the relationship between bamboo cover and monito del monte abundance was










negative. At these 12 sites, bamboo cover was lowest where the abundance of mistletoe was

highest, but the over all correlation of these variables is not significant (Figure 1-1, Appendix A-

2). Linear regression analysis of the abundance of monitos del monte and the Hyve significant

variables indicates that abundance of monitos del monte is better explained by number of

reproductive mistletoes and crop size of mistletoe than crop size ofA. chilensis or structural

variables of the habitat (Figure 1-2). Abundance of monito del monte also correlated with

abundance of non-reproductive-mi stletoe plants (R2 = 0.83; P < 0.0001) and with mistletoe

seedlings (R2 = 0.70; P < 0.001).

Discussion

High asymmetry between fleshy fruit plants and seed dispersers has been emphasized as a

mechanisms limiting coevolution of these plants and seed dispersers (Herrera, 1985; 1993;

Bascompte et al., 2003; Vazquez and Aizen, 2004). Other recent work also suggests that habitat

features constrain these species interactions (Fedriani, 2005). In this study at the scale of the

study area, habitat features played an important role in determining presence of monitos del

monte whereas presence or absence of specific food plants and mistletoe did not have an

appreciable effect. Bamboo cover was the most important habitat feature for the presence of

marsupial populations. Leaves of bamboo have been described as a principal resource for nest

building for this marsupial and because monitos del monte are arboreal, bamboo can increase the

mobility of this marsupial in forest understory where fruits and nests are located (Jimenez and

Rageot, 1979; Mann, 1955). However, presence data alone can overestimate or underestimate the

importance of habitat characteristics, because a species can be present but have very different

abundances in different sites (Royle and Nichols, 2003; Bailey et al., 2004). Although the

distribution of monitos del monte at the broad scale was influenced strongly by bamboo cover,

surveys conducted at Llao-Llao Municipal Reserve did not show a significant association









between abundance of this marsupial and cover of bamboo, except in one non-competitive model

where bamboo cover was related negatively related to abundance of monitos del monte. This

negative relationship may have occurred because the lowest amount of bamboo was in areas with

the highest amount of mistletoe (Figure 1-1). The general lack of importance of bamboo for

explaining abundance of monitos del monte in Llao-Llao Municipal Reserve could be explained

by the fact that when the minimum requirement for bamboo cover is met, which based on my

results may be about 10%, mistletoe plays a more important role than bamboo in regulating the

number of marsupials the habitat can support.

Numerous studies have addressed the link between fruit abundance and frugivore

abundance but clear examples of a single plant regulating the abundance of seed dispersers are

scarce (Levey, 1988; Herrera, 1998; 2002; Loiselle and Blake, 1991; Jordano, 1994; 2000;

Malizia, 2001; but see Moegenburg and Levey, 2003). My data suggest that the relationship

between the abundance of the monito del monte and mistletoe may be stronger and more

symmetric than in many other plant frugivory systems (Herrera, 1998; Moegenburg and Levey,

2003). Even though the monito del monte consumes and disperses 80% of the flora with fleshy

fruits, a significant correlation occurred at the Llao-Llao Municipal Reserve between the

abundance of monito del monte and the abundance of mistletoe plants and fruits. This

relationship was much stronger than with other fruiting species such as A. chilensis. Sites with

high abundance of mistletoe had from 2 to 5 times more monitos del monte that sites without

mistletoe. The strong correlation between mistletoe abundance and abundance of monito del

monte could occur because mistletoe directly affects the abundance of monitos del monte or

because both species are responding to another habitat variable not measured in this study.

Experiments are needed to resolve this issue. However, other data also support the conclusions of









the strong link between monitos del monte and mistletoe. Once mistletoe fruits are ripe, monitos

del monte change their diet from generalist to consume mistletoe fruits almost exclusively

(Rodriguez-Cabal et al., 2007).

The distribution and abundance of monitos del monte found in this study may be linked to

different habitat variables operating at different scales. This arboreal marsupial lives mainly in

the understory. At a large scale, the distribution of this marsupial may be limited by bamboo

cover, which increases nest sites and/or nest building material and connectivity to where food is

more abundant above ground. At a smaller spatial scale, the abundance of monitos del monte

tracked abundance of mistletoe. This spatial pattern may result from progressive gathering of

fine-grain foraging information by individuals in those forest sectors with high abundance of

reproductive mistletoe.

One of the most common explanations for lack of coevolution in plant-seed dispersal

interactions is the high level of asymmetry found (Bascompte et al., 2003). Specialized species

will have limited chance for coevolution because they are asymmetrically influenced by their

generalized species (Vazquez et al., 2007). Although the monito del monte could exert strong

selection on reproductive traits of this mistletoe as its only seed disperser in the temperate forest,

there is no evidence to support the coevolution of traits in this interaction. Furthermore, the

evidence points in the other direction, in which most of the relevant features of the mistletoe in

the temperate forest of Patagonia like the green color of the mature fruits, a color associated with

mammalian dispersers (van der Pijl, 1982), is driven only by environmental factors, such as

period of day light and temperatures (Amico, 2007; Amico et al., 2007). In the Chilean shrubland

this mistletoe species has fruits that are yellow when ripe, and birds disperse the seeds (Amico,

2007). As in the Australian mistletoe (Loranthaceae and Viscaceae) and mistletoebird (Dicaeum









hirundinaceum) mutualisms (Reid, 1987), the mistletoe-monito del monte interaction is an

ecological association rather than a coevolved interaction.































V


Table 1-1. Means (a SD) of habitat variables and summary ANOVA for comparison of means
for sites where monitos del monte were present or absent. Means from raw data are
shown. Variables that were log-transformed prior to analysis are indicated by "f.
Significant P-values are indicated by *.
Mean & SD


Present
(n = 17)
1.68 & 1.88
610. 17 & 725.25
4.06 & 4.05
62.11 & 57.30
0.73 A 1.01
24.97 & 27.50
3.13 A1.57
3.44 & 0.93
55.10 & 23.39
54.70 & 20.60
2.94 & 2.21


Absent
(n = 11)
1.09 & 1.56
468.68 & 927.70
1.35.63 A 1.60
47.90 & 146.57
0.69 & 2.29
3.57 & 8.23
0.98 & 1.51
2.48 & 1.36
54.24 & 36.50
46.28 & 35.79
1.12 &1.54


Habitat variables
No. reproductive mistletoe
Crop size of mistletoe
No. ofA. chilensis plants?
Crop size ofA. chilensis "
No. ofA. microphylla plants
Bamboo cover (%)Jf
Bamboo height (m)
Habitat complexity
Understory cover (%)
Canopy cover (%)Jf
Average number of trees


F dSf P-value


0.958
2.263
5.322
2.263
0.013
11.625
11.234
4.87
0.006
2.87
5.66


0.34
0.15
0.03*
0.15
0.91
0.002*
0.003*
0.04*
0.94
0.10
0.03*










Table 1-2. Summary of the seven logistic regression models constructed to determine which habitat variables predicted presence-
absence of monitos del monte at the 28 study sites. Competitive models (AAIC <2) and the significant variables in each
model are shown in bold.
Predicted correct (%)
Nagelkerke
Models Variables AIC AICc AAIC df R2 p Absent Present
Global 20.124 30.124 0.641 9 1 100 100
All variables
Food plants 39.129 40.129 10.646 3 0.28 72.7 82.4
No. of reproductive mistletoe
No. of A. chilensis plants 45.37
No. of A. microphylla plants
Fruit abundance and 28.483 29.483 0 3 0.62 81.8 88.2
bamboo Crop size of mi stletoe
Crop size of A. chilensis
Bamboo cover 13.37
Mistletoe plants and 30.817 31.297 1.814 0.49 72.7 82.4
bamboo No. of reproductive mistletoe
Bamboo cover 11.29
Mistletoe plants 40.502 40.656 11.173 1 0.05 0 100
No. of reproductive mistletoe
Bamboo 31.746 31.900 2.417 1 0.40 72.7 82.4
Bamboo cover 8.27
Habitat structure 30.517 32.256 2.773 4 0.62 81.8 88.2
Bamboo cover 10.99
Habitat complexity
Canopy cover
Average number of trees
















Global 44.623 134.623 86.154 9 0.87
No. of reproductive mistletoe 1.50 <0.0001
Habitat complexity 2.02 0.05
Food plants 48.070 51.070 2.600 3 0.80
No. of reproductive mistletoe 0.72 <0.0001
Fruit abundance and 48.292 51.292 2.823 3 0.83
bamboo Crop size of mistletoe 0.75 0.0005
Crop size ofA. chilensis 0.45 0.04
Mistletoe plants and 48.065 49.398 0.929 2 0.83
bamboo No. of reproductive mistletoe 0.95 <0.0001
Mi stl etoe 48.069 48.469 0 1 0.80
No. of reproductive mistletoe 0.89 <0.0001
Bamboo 65.311 65.711 17.242 1 0.01
Habitat structure 54.068 64.068 15.599 5 0.67
Bamboo cover -2.02 0.01


Table 1-3.Summary of the seven regression models constructed to determine which habitat variables predicted abundances of monito
del monte at 12 study sites. Competitive models (AIC <2) and the significant variables in each model are shown in bold.
Model variables are the same as in Table 1-2 except for habitat structure. For this analysis, number of N. dombeyi and
Austrocedrus chilensis were used instead of average number of trees.


Models


Significant variables


AIC AICc AAIC df R2 P P-value





JcJe


30000 -
20 -
20000 -


mHigh abundance of mistletoe
I Low abundance of mistletoe
I No-mistletoe


."3~
~J\
~L~


Figure 1-1.Means (a SD) of the habitat variables measured at the 12 sites where abundances of
monito del monte were determined. Means from raw data are shown. Variables log
transformed prior to analysis are indicated by +. Significant differences are shown as
*** P < 0.0001, ** P < 0.01, P < 0.05.












SU
R2 =0.22, P= 0.13*
40 1

30



20*


0.0 0.5 1.0 1.5 2.0 0 1 2 3 4 5


No. of reproductive mistletoe (log)


Crop size of A. chilensis (log)

R2 = 0.01, P= 0.80 *



--------*


012345678

Crop size of mistletoe (log)

R2 = 0.26, P= 0.09*


1 2 3 4


Bamboo cover (log)


0 1 2 3 4 5

Habitat complexity




Figure 1-2. Simpler linear regression analysis of the number of monitos del monte vs. the
significance predictor variables of the multiple regression models. The coefficient of
determination (R2) and P-value are shown.









CHAPTER 2
CONSERVATION IMPLICATIONS

The interaction between this mistletoe and its marsupial disperser may have consequences

at the community and ecosystem levels and habitat changes that impact either of these species

could have indirect cascading effects throughout different interaction webs. This mistletoe is

considered a keystone species due to the production of sugar-rich nectar and protein-rich fruits

during periods of food scarcity. This mistletoe also increases mortality of host trees and could

play an important role in plant community diversity. As the only known disperser for the

mistletoe, the monito del monte is important in maintaining these processes. Habitat

fragmentation disrupts the interaction between the mistletoe and monitos del monte (Rodriguez-

Cabal et al., 2007). Fragmentation negatively affects marsupial abundance, fruit removal, seed

dispersal, and seedling recruitment. The local extinction of monitos del monte has been

associated with the complete disruption of mistletoe seed dispersal (Rodriguez-Cabal et al.,

2007). The reason for the local extinction of monitos del monte in fragmented habitats is

unknown. One possible mechanism is reduction of bamboo in fragments results in loss of nest

sites and connectivity. Grazing by cattle and other introduced ungulates significantly alters

habitat structure and composition of the forest, including reduction of bamboo (Veblen et al.,

1992). Effects of grazing are particularly severe in forest fragments where introduced ungulates

have easy access to entire patch.

In this temperate forest, the bamboo blooms and dies at approximately 40-60 years

intervals. Although this is a natural process, the die-off of the bamboo also likely has important

consequences for populations of the monito del monte by reducing nest sites and connectivity.

Loss of bamboo within habitat fragments may have particularly severe consequences because









these patches are isolated, reducing the probability of demographic rescue and recolonization

following regeneration of bamboo within the patch.









CHAPTER 3
CONCLUSION

In the temperate forest of southern South America, the northern portion (to 41oS) has the

highest biodiversity and number of endemic species, the lowest proportion of protected areas,

and the highest rate of disturbance in this forest (Armesto et al., 1998, Rodriguez-Cabal et al., in

press). The biodiversity of this temperate forest currently is threatened by the highest rates of

destruction, fragmentation, and degradation ever experienced by this system, with more than

120,000 ha of native forest destroyed or degraded annually (Lara et al., 1996).

The alteration of natural habitats through human activities is the principal cause of the

decline and extinction of natural populations (Gaston and Spicer, 2004). Long-term plans to

conserve biodiversity must not only focus in maintaining the elements that form biodiversity but

also on the interactions among these elements (Vazquez and Simberloff, 2003). Understanding

the species and processes that shape ecosystems is fundamental for effective conservation

(Wayne et al., 2006). No previous study has investigated explicitly factors that determine this

austral marsupial's distribution and abundance or the causes of its present decline. This study

constitutes the first detailed ecological research focused on the identification of the habitat

variables that play the most important role in maintaining the marsupial's populations. My work

demonstrates that both habitat factors (bamboo) and interacting species (mistletoe) are important

in determining the distribution and abundance of this species.










APPENDIX
CORRELATION ANALYSIS


Table A-1. Correlation coefficients (r) between habitat variables in the presence-absence survey (n = 28). Significant correlations
after the Bonferroni correction are indicated by (P < 0.0009).
Crop size of No. ofA. Crop size ofA. No. ofA. Bamboo cover


microphylla plants
0.17
0.18
0.42
0.10


(%)
-0.19
-0.17
0.29
0.39
0.33


mistl etoe
0.89*


chilensis plants
0.56
0.50


chilensis
0.15
0.10
0.66*


No. of reproductive mistletoe
Crop size of mistletoe
No. of A. chilensis plants
Crop size of A. chilensis
No. of A. microphylla plants
Bamboo cover (%)
Bamboo height (m)
Habitat complexity
Understory cover (%)
Canopy cover (%)


Table A-1.


Bamboo height
(m)
0.11
0.02
0.47
0.40
0.36
0.88*


Habitat
complexity
0.29
0.01
0.02
0.19
-0.06
0.35
0.20


Understory
cover (%)
-0.004
-0.01
-0.23
-0.004
0.39
-0.11
-0.29
0.73*


Average no.
trees
-0.24
-0.28
0.13
0.16
0.004
0.43
0.40
0.44
0.15
0.88*


Canopy cover (%)
0.38
0.44
-0.07
-0.06
-0.05
0.37
0.29
0.43
0.23


No. of reproductive mistletoe
Crop size of mistletoe
No. of A. chilensis plants
Crop size of A. chilensis
No. of A. microphylla plants
Bamboo cover (%)
Bamboo height (m)
Habitat complexity
Understory cover (%)
Canopy cover (%)










Table A-2. Correlation coefficients (r) between habitat variables in the abundance survey (n = 12). Significant correlations after the
Bonferroni correction are indicated by (P < 0.0006).
Crop size of No. of A. Crop size of A. No. of A. Bamboo cover
mistletoe chilensis plants chilensis microphylla plants (%)
No. of reproductive mistletoe 0.94* 0.78 0.30 0.09 -0.28
Crop size of mistletoe 0.70 0.16 0.12 -0.13


0.41


0.33
0.05


-0.32
0.46
0.10


No. of A. chilensis plants
Crop size of A. chilensis
No. of A. microphylla plants
Bamboo cover (%)
Bamboo height (m)
Habitat complexity
Understory cover (%)
Canopy cover (%)
N. dombeyi


Table A-2.


Bamboo
height (m)
0.33
0.39
0.63
0.38
0.32
0.65


Habitat
complexity
0.28
0.34
0.39
0.70
0.00
0.70
0.67


Understory
cover (%)
0.13
0.18
0.11
0.67
-0.23
0.71
0.50
0.89*


Canopy cover
(%)
0.21
0.28
0.56
0.36
0.10
0.68
0.96*
0.58
0.40


A.
chilensis
0.46
0.55
0.53
-0.42
0.09
-0.26
0.30
-0.08
-0.35
0.27
0.17


N. dombeyi
-0.35
0.20
-0.02
-0.14
0.18
0.46
0.33
-0.16
-0.16
0.51


No. of reproductive mistletoe
Crop size of mistletoe
No. of A. chilensis plants
Crop size of A. chilensis
No. of A. microphylla plants
Bamboo cover (%)
Bamboo height (m)
Habitat complexity
Understory cover (%)
Canopy cover (%)
N. dombevi










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BIOGRAPHICAL SKETCH

Mariano Alberto Rodriguez Cabal was born in Buenos Aires, Argentina in 1976. His

interest in biology began when he very young. Nature has always exhibited a powerful attraction

for him. He studied in a religious high school and obtained a degree in economics. When he

finished his secondary studies, he moved to San Carlos de Bariloche in the foothills of the

Andes, in western Argentina. The magnificent environment around this city, which is within a

national park, led him unequivocally to pursue university studies in biology.


In 1997, he began studying Biology at Universidad Nacional del Comahue. In July 2003,

he graduated as a "Licenciado en Ciencias Biol6gicas." Under the guidance of Dr. Marcelo

Aizen and Dr. Andres Novaro, he conducted his undergraduate thesis research on the effects of

habitat fragmentation on a dispersal mutualism in the temperate forest of southwestern

Argentina. After two years of Hield work, he completed and defended his thesis, obtaining the

highest grade.

In 2004, he received a Fulbright Scholarship to study abroad. In the next year, he moved

to Florida, USA, where he started his master' s studies at the University of Florida in the

Department of Wildlife Ecology and Conservation. During those years worked under the

supervision of Professor Lyn C. Branch. This work was recognized with three grants. Twice he

earned the Outstanding Academic Achievement Award at UF. In December 2007, he defended

his master' s thesis. He made a firm decision to devote himself to scientific research. He hopes to

continue his doctoral education in the United States of America. On completing his studies, he

will return to his country to contribute to the welfare of his people and the conservation of

Argentina' s biodiversity through teaching and research.





PAGE 1

1 HABITAT ASSESSMENT FOR A THREATENED MARSUPIAL IN TEMPERATE FOREST OF PATAGONIA BY MARIANO ALBERTO RODRIGUEZ-CABAL A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORI DA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2008

PAGE 2

2 2008 Mariano Alberto Rodriguez Cabal

PAGE 3

3 To Noelia and to all who nurtured my curiosity and love for nature.

PAGE 4

4 ACKNOWLEDGMENTS I thank my supervisory committee (L. C. Br anch, K Sieving and E. Bruna) for their mentoring. I also thank the faculty, staff and grad students at the Univer sity of Florida (UF) Department of Wildlife Ecology and Conservation. I thank Mauricio Failla, Director de Fauna de la Provincia de Rio Negro, Arge ntina; Gustavo Iglesias and Claudio Chehebar, Nahuel Huapi National; Park, and staff of Par que Municipal Llao Llao. Special thanks go to M. Aizen, G. Amico, N. Barrios, G. Barrios, R. Bunge, J. M. Morales, S. Morgan, A. Martnez, M. Nuez, E. Perner, A. Pries, R. Reggio, and N. Tercero-Buca rdo for valuable assistance in the field and for their thoughtful comments and suggestions on previous drafts. Fiel d work was supported by Wildlife Conservation Society, Sc ott Neotropical Fund, and the Je nnings Scholarship of the UF Department of Wildlife Ecology and Conservation.

PAGE 5

5 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........6 LIST OF FIGURES................................................................................................................ .........7 ABSTRACT....................................................................................................................... ..............8 CHAPTER 1 HABITAT ASSESSMENT FOR MONITOS DEL MONTE................................................10 Introduction................................................................................................................... ..........10 Methods........................................................................................................................ ..........13 Study Area..................................................................................................................... ..13 Marsupial Presence Absence.........................................................................................14 Marsupial Abundance......................................................................................................15 Habitat Variables.............................................................................................................15 Data Analysis.................................................................................................................. .16 Results........................................................................................................................ .............19 Marsupial Presence Absence.........................................................................................19 Marsupial Abundance......................................................................................................20 Discussion..................................................................................................................... ..........21 2 CONSERVATION IMPLICATIONS....................................................................................25 3 CONCLUSION.......................................................................................................................32 APPENDIX A CORRELATION ANALYSIS...............................................................................................33 LIST OF REFERENCES............................................................................................................. ..35 BIOGRAPHICAL SKETCH.........................................................................................................40

PAGE 6

6 LIST OF TABLES Table page 1-1 Habitat variables for sites where mon itos del monte were present or absent....................25 1-2 Logistic regression mode ls at the 28 study sites................................................................26 1-3 Regression models constr ucted at 12 study sites...............................................................27 A-1 Correlation coefficients betw een habitat variables in the presence absence survey ......33 A-2 Correlation coefficients between habita t variables in the abundance survey....................34

PAGE 7

7 LIST OF FIGURES Figure page 1-1 Habitat variables at sites where abundan ces of monito del monte were determined.........28 1-2 Number of monitos del monte vs. the significance predictor variables.............................29

PAGE 8

8 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Master of Science HABITAT ASSESSMENT FOR A THREATENED MARSUPIAL IN TEMPERATE FOREST OF PATAGONIA By Mariano Alberto Rodriguez-Cabal May 2008 Chair: Lyn C. Branch Major: Wildlife Ecology and Conservation Patterns and processes that sh ape the distribution and abunda nce of animals depend on the scale at which they are evaluated. Understanding theses patterns and pr ocesses is fundamental for the study of plant-seed disperser interac tions. Most plants th at depend on frugivorous animals for seed dispersal are dispersed by several species of animals and, the majority of animal dispersers consume the fruit of several plant spec ies. In contrast to the weak mutual dependence of most plants and thei r seed dispersers, some mutualisms in the temperate forest of Patagonia appear to be obligate. The northern portion of the temperate forest harbors a unique plant animal interaction in which the seeds of the mistletoe Tristerix corymbosus a proposed key species, are dispersed solely by the endemic marsupial Dromiciops gliroides I developed a set of habitat models, in which model variables were defined a prior, to test the hypothesis that the distribution and abundance of monito del monte is associated principally with abundance of mistletoe in the temperate forest of Patagonia or that, alternatively, habita t structure and/or other plants determine the distribution and abundance of this marsupial. Results show that the distribution of monito s del monte at the broa d scale was strongly influenced by bamboo cover. However, at a sma ller spatial scale, the ab undance of monitos del monte are explained by the abundan ce of mistletoe plants and fruits. The lack of dependence

PAGE 9

9 among fruiting plants and their seed dispersers often is attributed to the high asymmetry in these interactions and the important influence of habi tat features. Conversely, my results showed that the mistletoemonito del monte interaction is more symmetric than many other plantfrugivore interactions. This interaction is characterized by an obligate partnership of the mistletoe with the marsupial, and a strong relationship between the abundance of monitos del monte and the abundance of mistletoe.

PAGE 10

10 CHAPTER 1 HABITAT ASSESSMENT FO R MONITOS DEL MONTE Introduction Distribution and abundance of species depends on the distri bution and abundance of other species with which they interact and ha bitat factors a various scales (Wiens,1989). Understanding the patterns and processes that shape distribution and abundance is fundamental for the study of plant-seed disperser interactio ns and biodiversity cons ervation. Seed dispersal plays an important role in creating and maintain ing diversity in plant communities (Schupp et al., 2002). At large scales, appropriate habitat may be the most important factor limiting distribution and abundance of frugivores (Wiens, 1989). Habitat characteristics such as availability of a variety of food resources, nest si tes, and distance to shelter, in fluence frugivores distribution and could be play a critical role in regulating the abundan ce of these species (Fedriani, 2005). At a small scale, habitat character istics in which fruiting plants are surrounded may be more important (Herrera, 1998). Previous studies have demonstrated that frugivores select plants or good patches based on habitat feat ures rather than on plant ph enotypic traits (Herrera, 1998; Fedriani, 2005). For example, fr ugivores could minimize predation risk by selecting sites with high cover of vegetation to avoi d predators during foraging (How e, 1979). Thus, habitat factors may limit the degree to which frugivorous animal s track their food resource, and human impacts on habitat of interact ing species can have cascading effects. Seed dispersal mutualisms influence patterns of seedling recruitment, plant demography, and population persistence, and thus potentially have widespread effects on the rest of the community (Janzen, 1980; Feinsinger, 1987; Bond, 1994; Levey and Benkman,1999; Herrera, 2002; Spiegel and Nathan, 2007; Rodriguez-Cabal et al., 2007). The majority of plants that depend on frugivorous animals for seed dispersal are dispersed by several species of animals

PAGE 11

11 and, most animal dispersers consume the fru it of several plant speci es (Herrera, 1998; 2002). Mutual dependence of most plants and seed dispersers is weak, and asymmetry is common. Asymmetry in plantanimal interactions is repres ented in terms of the ef fect of one species on another species relative to the strength of the reciprocal effect (Vazquez and Aizen, 2004; Vazquez et al., 2005; Bascompte et al., 2006; Vaz quez et al., 2007). Asymme try is particularly evident for specialist plants, which tend to interact mostly with generalist animals (Vazquez and Aizen, 2004; Bascompte et al., 2003 ; Bascompte et al., 2006). Gene ralist plants most often interact with genera list and specialist anim als (Vazquez and Aizen, 2004; Bascompte et al., 2003; Bascompte et al., 2006). In contrast to the weak mutual dependence of most plants and their seed dispersers, some mutualisms in the temperate forest of Patagonia a ppear to be obligate. This system provides an ideal opportunity to examine whether the distri bution and abundance of an animal disperser depends on the distribution and abundance of a speci fic fruit. A large pro portion of the flora in this temperate forest depends on mutualistic an imals (Armesto et al., 1996; Aizen and Ezcurra, 1998; Aizen et al., 2002). This leve l of mutualism is one of the hi ghest recorded for a temperate ecosystem and is comparable to those from tropical forests (Willson et al., 1989; Willson, 1991). Unlike the mutualism structure in tropical forests, in this temperate forest many species of flora depend on a few mutualistic animals for th eir survival (Aizen et al., 2002). The northern portion of the temperate forest harbors a unique tr iangle of keystone mutualists comprised of a hummingbird ( Sephanoides sephaniodes ), a mistletoe (quintral, Tristerix corymbosus ), and the marsupial (monito del monte, Dromiciops gliroides Amico and Aizen, 2000). This hummingbird is responsible for pollinating nearly 20% of the endemic species of woody flora in this biome (Armesto et al., 1996; Aizen et al., 2002). The mistletoe

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12 blooms in winter and is the only source of nect ar for the hummingbird during that period (SmithRamrez, 1993; Aizen and Ezcurra, 1998). The m onito del monte is the only known disperser of the mistletoe in this temperat e forest (Amico and Aizen, 2000). Moreover, the passage of the seed through the marsupials gut is crucial to triggering germination (Amico and Aizen, 2000). The isolation from other similar biotas since the disruption of Gondwan a allowed the evolution of monito del monte which is the only extant me mber from the ancient family Microbiotheriidae. Because of the long evolutionary history of association of monitos del monte and the mistletoe and the specific fruiting characteristics and distribution of mistle toes in this temperate forest, I hypothesized that the distribution a nd abundance of monitos de l monte and mistletoe might be related closely. Mistletoe produces a large crop size every year and is the most abundant fleshy fruit in the north ern temperate forest of Patag onia (Aizen, 2003, Amico et al., in press). Once mistletoe fruits are ripe, monitos del monte consume almost exclusively mistletoe fruits (Rodriguez-Cabal et al., 2007). This mistletoe presents an aggregated distribution and, therefore, may serve as high dens ity resource patches for the m onito del monte. Alternatively, populations of monitos del monte ma y not track any single plant spec ies as they are known to eat and disperse seeds of 80% of the flora with fleshy fruit (Amico et al., in press). Because of the diversity of fruits available, their populations maybe constraine d largely by other habitat factors such as vegetation cover and structural complexi ty of the habitat that influence nest site availability, predation risk, and other aspects of their biology. In this study, I developed a set of habitat models, in which model variables were de fined a prior, to examine the hypothesis that the distribution and abundance of monito del monte is associated principally with abundance of mistletoe in the temperate forest of Patagonia or that, alternatively, habita t structure and/or other plants determine the distribution and abundance of this marsupial. Although experiments are

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13 required to fully determine the degree of mutu al dependence between monitos del monte and the mistletoe, landscape scale studies such as this provide a foundation for understanding environmental constraints on these relationships. Methods Study Area I conducted my research in Nahuel Huapi National Park (705,000 ha) and Llao-Llao Municipal Reserve (1226 ha) in northwester n Patagonia. Nahuel Huapi National Park encompasses a marked west-to-east rainfall gr adient caused by the Andes, which form an effective barrier to westerly air masses coming fr om the southern Pacific Ocean. The most xeric forest site in my study, situated at the xeric boundary with the Patagonian steppe (Cerro Otto, 41' S, 71' W), has a mean annual prec ipitation of 1200 mm of which only 10.5 % falls during the growing season. The west ern limit of my study located in Puerto Blest (41 01' S, 71 49' W) is an area of Valdivian rainforest, one of the rainiest places in the Patagonia (mean annual precipitation, 4000 mm). Llao-Llao Municipal Reserve (41 08' S, 71 19' W) has intermediate rainfall (mean annual precipitation, 1800 mm). M ean January temperatur e is 15C, and mean July temperature is 3C. The native forest vegetation in the study area belongs to the Subantartic biogeographical region, which is distinctive from the Neotropical forest of the rest of South America (Cabrera, 1976). The most common trees are evergreen southern beech ( Nothofagus dombeyi ) a lenga beech ( Nothofagus pumilio ) and a conifer ( Austrocedrus chilensis ). The understory is dominated by the bamboo ( Chusquea culeou ) and the shrub ( Aristotelia chilensis here after A. chilensis ). That is the main host of the mistletoe in th e study area (Mermoz and Martn, 1986). The two forest layers are well differentiated with tr ee canopy reaching up to 40-m height and understory reaching up to 5-m height.

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14 I conducted presenceabsence surveys of m onitos del monte and examined habitat variables at 16 sites in Nahuel Huapi National Park and 12 sites in th e adjacent Llao-Llao Municipal Reserve. In addition, trapping was conducted at the 12 sites in the Llao-Llao Municipal Reserve to determine the relationshi p between abundance of monitos del monte and abundance of mistletoe. Sites in Nahuel Hu api National Park were chosen randomly to incorporate the range of habita ts found in the study area. Pote ntial sites in the Llao-Llao Municipal Reserve were stratified by mistletoe de nsity, and four sites were chosen randomly within each of the following categories: high (> 35 reproductive individuals of mistletoe), low (<20 reproductive individuals), a nd no mistletoe. Sites were stra tified on abundance of mistletoe because mistletoe has a patchy distribution and a ra nge of all other variables can be found within each stratum defined by mistletoe density. I conduc ted most of the field work during the austral summer (December 2006 March 2007) because mon itos del monte hibernate during winter and the main fruiting and dispersal season of the mistletoe is summer (Aizen 2003). Marsupial Presence Absence In order to facilitate rapid surveys of presence absen ce of monitos del monte, the 28 study sites were grouped by presence or absence of mistle toe. At sites with mistletoe, I examined all plants < 15 cm DHB in each site for feces of monitos del monte th at contained mistletoe seeds. I concluded that monitos del monte were presen t if I found mistletoe seeds dispersed. If no dispersed seeds were found in the site, I used tr apping to determine the pr esence of monito del monte. This technique also was used for sites with no mistletoe. Within e ach site, I placed a 5 x 5 grid (0.25 ha) of Tomahawk-style traps about 10-m apart. Each trap was placed between 1-2 m above a ground in the shrub closest to the sa mple point. Traps were baited with apple and banana. Because the objective was to document presence, once a monito del monte was trapped, I stopped trapping at this site. Sampling was cont inued for four nights if no monitos del monte

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15 were captured. Based on previous capture data for monitos del monte (Rodriguez-Cabal, 2003), and following procedures in MacKenzie et al. (2002), probability of capture was 99.6% in four night if monitos del monte were present. I conc luded that monitos del monte were absent after four nights of trapping if no marsupials were captured. Marsupial Abundance Abundance of the monito del monte was estimated using multiple capture-recapture methods with trapping grids as described above. On e trapping grid was placed at each of the 12 sites in Llao-Llao Municipal Reserve. Animals we re trapped for four periods of four nights each per site. Traps were checked daily and all tra pped marsupials were mark ed with an individual code based on ear perforations and released at the point of capture (for more detail see Rodriguez-Cabal, 2003; Rodriguez-Cabal et al., 2007). Recaptures were noted in subsequent trapping periods. Habitat Variables At each of the 16 sites in Nahuel Huapi National Park, a 5 x 5 grid was established with points 10 m apart. I measured habitat variables in 5-m radius circular plots centered on every second point (n = 13 per site). Similar sampli ng was conducted at sites trapped in Llao-Llao Municipal Reserve with plots ce ntered on every second trap. I m easured the following variables to represent the biological a nd physical structure of the habitat: number of reproductive individuals and crop size of th ree understory plants that produce fleshy fruits (mistletoe A. chilensis, and Azara microphylla ), cover and height of bamboo, co mplexity of habitat structure, canopy cover, understory cover and the numbe r of the two most common tree species ( N. dombeyi and Austrocedrus chilensis ). Reproductive individuals were plants that presented sexual structures (i.e., flowers or fruits). In additi on for mistletoe, I count ed the number of nonreproductive individuals (2-4 year s old), seedlings (those presenti ng the first two true leaves),

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16 and number of seeds dispersed. To determine cr op size of understory plants, I counted the number of fruits on a subset of three branches dispersed in each crown and then estimated total crop size by multiplying mean number of fruits per branch by total number of branches in the crown for each plant. As with mistletoe, seeds of A. chilensis and A. microphylla are dispersed by monitos del monte, and passage of their seed s through the digestive tr act of a monito del monte increases chance of germinat ion (Amico et al., in press). Cover of bamboo was estimated visually by summing the total area occupied by bamboo stems and leaves in each plot. Also, I measured the height of the tallest bamboo stem in each plot. As a measure of habitat complexity within each plot, I counted the num ber of contacts with a vertical pole (3-m height) for branches 5 10-cm in diameter and oriented < 45 relative to the ground. Good connectivity between plants in the understory stratum could play a key role in the mobility of this animal, and more seeds dispersed in monito del monte feces are found on branches with these characteristics (Amico 2000). Percent cover of the canopy and understory were estimated using a densiometer at the center of the circular plot Finally, I counted the total numb er of the most common trees for all individuals taller than four meters. Data Analysis I used multiple logistic regressions to determ ine which habitat variables predicted monito del monte presence-absence at the 28 study sites, and I used multiple regressions to assess the contribution of habitat variable s to explaining abundance of m onitos del monte at the 12 study sites in Llao-Llao Municipal Reserve. Data we re log-transformed when necessary to achieve normality and reduce heteroscedasticity (Table 1-1). For both analyses, I only used explanatory variables that did not correlat e significantly after a Bonferroni correction (Appendix A-1 and A2; Tables 1-1, 1-2 and 1-3). I did not used ba mboo height and understory cover because these measures were correlated with bamboo cover an d habitat complexity, respectively, and bamboo

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17 cover and habitat complexity may be more direct ly related to the biology of monitos del monte, particular for nest material and mobility. I did not include crop size of A. microphylla in analyses because this understory plan t did not produce fruits during the study. Seven models were built to identify habitat variables related to presence absence and abundance of monitos del monte. Rather than examine all possible models, models were developed a priori, that focused on mistletoe, other food plants, and ha bitat structure as key factors in the distribu tion and abundance of this marsupial. I examined the following models: (1) Global -all habitat variables, (2) Food plants -numb er of reproductive i ndividuals of major food plants that produce fleshy fruits eaten by monitos del monte, (3) Fruit abundance and bamboo -crop size of plants with fleshy fruits and cover of bamboo, (4) Mistletoe plants and bamboo -number of reproductive mistletoes a nd bamboo cover, (5) Mistletoe plants, (6) Bamboo, (7) Habitat structure -a ll habitat variables unrelated to food plants (Table 1-2 and 13). Bamboo cover was included as a separate variab le because this plant is an important source of nest material (Jimenez and Rageot 1979), pr ovides connectivity for arboreal movement, and may inhibit predation by aerial predators. I used an approach that compared the suite of models using Akaikes information criteria (AIC), whic h allows direct comparison of models with different number of parameters. AIC is calculated for a suite of models and the best fitting one has the smallest AIC (Burnham and Anderson, 2002; Whittingham et al., 2005). I applied smallsample version of AIC (AICc), because the ra tio of number of observations to number of parameters was <40. AICc was calculated for each model with the algorithm: a) AIC = -2 log ( L ) + 2 K b) AICc = -2 log ( L ) + 2 K + (2 K (K + 1))/ (n K 1)

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18 where log ( L ) is the log-likelihood function evaluate d for the maximum likelihood parameter estimates, K is the number of parameters in the model and n is the number of observations. AIC represents the difference between the AICc score for each model and the AICc score of the best model. The absolute value of AIC is unimportant; instead AIC indicates relative support for the models. AIC values <2 indicate that models are similar. When AIC was <2, I chose the most parsimonious model as the best model (i.e., the mode l that used the least number of parameters). To evaluate goodness-of-fit for the multiple logist ic regressions models, I used Nagelkerke R2. A true R2, which measures the variability in the de pendent variable that is explained by a linear regression models, cannot be calculated for logistic regression models (Hosmer and Lemeshow, 1989). However, because deviance can be thought of as a measure of how poorly the model fits (i.e., lack of fit be tween observed and predicted valu es), an analogy can be made to sum of squares residual in ordina ry least squares. This pseudoR2 is designed to have similar properties to the true R2. The pseudoR2 is based on comparison of the likelihood of the current model to the null model (one wi thout predictors). A larger pseudoR2 indicates that more of the variation is explained by the model. Here, I reported the Nagelkerke R2 which is a modified version of Cox and Snell R2 that ranges from 0 to 1 (Hosmer and Lemeshow 1989). I evaluated the importance of significant independent variables in logistic regression using the standardized regression coefficient which is the predicte d change in odds for a unit increase in the corresponding independent variable (Hos mer and Lemeshow, 1989). Values of <1 correspond to decreases in the odds with increases in th e dependent variable; va lues >1 correspond to increases in the odds with increa ses in the independent variable; and values close to 1 indicate that unit changes in the inde pendent variable do not affect the dependent variable.

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19 To evaluate the goodness-of-fit for the multiple regression models for abundance of monitos del monte, I calculated the R2 of the models. In multiple regressions, I evaluated the importance of significant independent variable s by calculating the standardized regression coefficient (Ganas and Robbins, 2005). The sta ndardized regression coefficient is used to compare the magnitude of effect of different va riables by describing the ch ange in the dependent variable (in SD units) caused by an increase in one unit (SD) of the independent variable, while all the other independent vari ables are held constant. In addition, I compared habitat variables betw een sites with and wit hout monitos del monte and at sites where I measured marsupial abundan ces using an ANOVA. I performed a Chi-square test to compare the relationship between the presence of monitos del monte and mistletoe presence at the 28 sites. For the 12 sites wher e I measured abundance of monitos del monte, I report the minimum number alive for all sites to allow comparison among sites (Wilson et al., 1996). Because of the low number of captures in sites without mistletoe, I could not use other methods. Finally, I performed a regression anal ysis to examine the relationship between abundance of monitos del monte and habitat variab les that had significant explanatory power in the multiple regression models. These analyses were conducted using SAS 9.13 and JMP IN for Windows Ver. 5.1 (SAS Institute, Inc., Cary, NC). Results Marsupial Presence-Absence Three of the seven multiple logistic regressi on models received substantial support as the best predictive models of presence-absence of monitos del monte (Table 1-2). The Global model perfectly predicted the presence-absence of monitos del monte (i.e., Nagelkerke R2 = 1). The most parsimonious model (Fruit abundance and ba mboo) and the Habitat structure model, which was not ranked in the best models, also e xplained considerable variation in presence-absence of

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20 monitos del monte (Table 1-2). Bamboo cover was the only significant pred ictor variable in the top models (Table 1-2). The seventeen sites where I found marsupials had more bamboo cover than sites without monito del monte (Table 11). Monito del monte wa s recorded 91% of the sites that had more than 10% bamboo cover a nd in only 35% of the s ites with less than 10% cover. The presence-absence of mistletoe and monito del mo nte were not related at the 28 sites (Chi-square = 2.06, df = 1, P = 0.15). Five of the 17 sites where I recorded presence of monito del monte did not have mistletoe. Five of the el even sites where monitos del monte were absent had reproductive mistletoe plants but I did not find mi stletoe seeds dispersed, seedlings or nonreproductive mistletoe (2-4 year s old). The lack of seeds, seedlings and non-reproductive individuals indicates that there has not been recrui tment at least in the last four years. The other 12 sites had mistletoe seeds dispersed in monito del monte feces and seedlings. Marsupial Abundance The estimated minimum number alive of ma rsupials on study plots ranged between 0 and 45 individuals. At eleven of the 12 sites wher e I measured abundances, I captured monitos del monte in the first four nights of trapping, in sp ite of the high variability in local abundance of this species. Two regression models were candida tes for the best predic tive model for abundance of monitos del monte, although several additi onal models also expl ained high amounts of variation in abundance of these marsupials (Table 1-3). The number of re productive mistletoe or crop size of mistletoe were signif icant variables in five of the seven models. Habitat complexity, bamboo cover, and crop size of A. chilensis appear as significant vari ables in models that were not supported as the best models, but P -values for these variables were lower (Table 1-3). Marsupial abundance increased when the number of reproductive mistle toe, crop size of mistletoe and A. chilensis or habitat complexity increased (see regression coefficients Table 1-3). In contrast, the relationship between ba mboo cover and monito del monte abundance was

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21 negative. At these 12 sites, bamboo cover was lowest where the abunda nce of mistletoe was highest, but the over all correlati on of these variables is not si gnificant (Figure 1-1, Appendix A2). Linear regression analysis of the abundance of monitos del monte and the five significant variables indicates that abunda nce of monitos del monte is better explained by number of reproductive mistletoes and crop size of mistletoe than crop size of A. chilensis or structural variables of the habitat (Figur e 1-2). Abundance of monito de l monte also correlated with abundance of non-reproductive-mistletoe plants ( R2 = 0.83; P < 0.0001) and with mistletoe seedlings ( R2 = 0.70; P < 0.001). Discussion High asymmetry between fleshy fruit plants and seed dispersers has been emphasized as a mechanisms limiting coevolution of these plants and seed dispersers (Herrera, 1985; 1993; Bascompte et al., 2003; Vazquez and Aizen, 2004). Ot her recent work also suggests that habitat features constrain these species interactions (Fed riani, 2005). In this stud y at the scale of the study area, habitat features play ed an important role in dete rmining presence of monitos del monte whereas presence or abse nce of specific food plants a nd mistletoe did not have an appreciable effect. Bamboo cover was the most important habitat featur e for the presence of marsupial populations. Leaves of bamboo have been described as a principal resource for nest building for this marsupial and because monitos del monte are arboreal, bamboo can increase the mobility of this marsupial in forest understory where fruits and nests are located (Jimenez and Rageot, 1979; Mann, 1955). However, presence data alone can overestimate or underestimate the importance of habitat characteristics, because a species can be present but have very different abundances in different sites (Royle and Nic hols, 2003; Bailey et al ., 2004). Although the distribution of monitos del monte at the broad s cale was influenced strongly by bamboo cover, surveys conducted at Llao-Llao Municipal Re serve did not show a significant association

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22 between abundance of this marsupial and cove r of bamboo, except in one non-competitive model where bamboo cover was related negatively relate d to abundance of monitos del monte. This negative relationship may have occurred because th e lowest amount of bamboo was in areas with the highest amount of mistletoe (Figure 1-1). The general lack of importance of bamboo for explaining abundance of monitos del monte in Ll ao-Llao Municipal Reserve could be explained by the fact that when the minimum requirement for bamboo cover is met, which based on my results may be about 10%, mistletoe plays a more important role than bamboo in regulating the number of marsupials the habitat can support. Numerous studies have a ddressed the link between fr uit abundance and frugivore abundance but clear examples of a single plant re gulating the abundance of seed dispersers are scarce (Levey, 1988; Herrera, 1998; 2002; Lo iselle and Blake, 1991; Jordano, 1994; 2000; Malizia, 2001; but see Moegenburg and Levey, 2003) My data suggest that the relationship between the abundance of the monito del mont e and mistletoe may be stronger and more symmetric than in many other plant frugivor y systems (Herrera, 1998; Moegenburg and Levey, 2003). Even though the monito del monte consumes and disperses 80% of the flora with fleshy fruits, a significant correlation occurred at the Llao-Llao Municipal Reserve between the abundance of monito del monte and the abundanc e of mistletoe plants and fruits. This relationship was much stronger than w ith other fruiting species such as A. chilensis Sites with high abundance of mistletoe had from 2 to 5 times more monitos del monte that sites without mistletoe. The strong correlation between mis tletoe abundance and abund ance of monito del monte could occur because mistletoe directly a ffects the abundance of monitos del monte or because both species are responding to another habitat variable not measured in this study. Experiments are needed to resolve this issue. Howe ver, other data also support the conclusions of

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23 the strong link between monitos del monte and mistletoe. Once mis tletoe fruits are ripe, monitos del monte change their diet from generalist to consume mistle toe fruits almost exclusively (Rodriguez-Cabal et al., 2007). The distribution and abundance of monitos del monte found in this study may be linked to different habitat variables operati ng at different scales. This arbor eal marsupial lives mainly in the understory. At a larg e scale, the distribution of this marsupial may be limited by bamboo cover, which increases nest site s and/or nest building material and connectivity to where food is more abundant above ground. At a smaller spatia l scale, the abundance of monitos del monte tracked abundance of mistletoe. This spatial pa ttern may result from progressive gathering of fine-grain foraging information by individuals in those forest sectors with high abundance of reproductive mistletoe. One of the most common explanations for lack of coevolution in plant-seed dispersal interactions is the high level of asymmetry found (Bascompte et al., 2003). Specialized species will have limited chance for coevolution because they are asymmetrically influenced by their generalized species (Vazquez et al., 2007). Although the monito del monte could exert strong selection on reproductive traits of th is mistletoe as its only seed di sperser in the temperate forest, there is no evidence to support the coevolution of traits in this interaction. Furthermore, the evidence points in the other direction, in which mo st of the relevant featur es of the mistletoe in the temperate forest of Patagonia lik e the green color of the mature fruits, a color associated with mammalian dispersers (van der Pijl, 1982), is dr iven only by environmental factors, such as period of day light and temperat ures (Amico, 2007; Amico et al., 2007). In the Chilean shrubland this mistletoe species has fruits that are yellow when ripe, and birds disperse the seeds (Amico, 2007). As in the Australian mistletoe (Lor anthaceae and Viscaceae) and mistletoebird ( Dicaeum

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24 hirundinaceum ) mutualisms (Reid, 1987), the mistletoemonito del monte interaction is an ecological association rather than a coevolved interaction.

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25 Table 1-1. Means ( SD) of habitat variable s and summary ANOVA for comparison of means for sites where monitos del monte were present or absent. Means from raw data are shown. Variables that were log-transforme d prior to analysis are indicated by Significant P -values are indicated by *. Mean SD Habitat variables Present (n = 17) Absent (n = 11) F df P -value No. reproductive mistletoe 1.68 1.88 1.09 1.560.958 1 0.34 Crop size of mistletoe 610.17 725.25468.68 927.702.263 1 0.15 No of A. chilensis plants 4.06 4.051.35.63 1.605.322 1 0.03* Crop size of A. chilensis 62.11 57.3047.90 146.572.263 1 0.15 No of A. microphylla plants 0.73 1.010.69 2.290.013 1 0.91 Bamboo cover (%) 24.97 27.503.57 8.2311.625 1 0.002* Bamboo height (m) 3.13 1.570.98 1.5111.234 1 0.003* Habitat complexity 3.44 0.932.48 1.364.87 1 0.04* Understory cover (%) 55.10 23.3954.24 36.500.006 1 0.94 Canopy cover (%) 54.70 20.6046.28 35.792.87 1 0.10 Average number of trees 2.94 2.211.12 1.545.66 1 0.03*

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26Table 1-2. Summary of the seven logistic re gression models constructed to determine wh ich habitat variables predicted presenceabsence of monitos del monte at the 28 study sites. Competitive models ( AIC <2) and the significant variables in each model are shown in bold. Predicted correct (%) Models Variables AIC AICc AIC df Nagelkerke R2 Absent Present Global 20.12430.1240.641 9 1 100 100 All variables Food plants 39.12940.12910.646 3 0.28 72.7 82.4 No. of reproductive mistletoe No. of A. chilensis plants 45.37 No. of A. microphylla plants Fruit abundance and 28.48329.4830 3 0.62 81.8 88.2 bamboo Crop size of mistletoe Crop size of A. chilensis Bamboo cover 13.37 Mistletoe plants and 30.81731.2971.814 0.49 72.7 82.4 bamboo No. of reproductive mistletoe Bamboo cover 11.29 Mistletoe plants 40.50240.65611.173 1 0.05 0 100 No. of reproductive mistletoe Bamboo 31.74631.9002.417 1 0.40 72.7 82.4 Bamboo cover 8.27 Habitat structure 30.51732.2562.773 4 0.62 81.8 88.2 Bamboo cover 10.99 Habitat complexity Canopy cover Average number of trees

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27Table 1-3.Summary of the seven regression models constructed to determine which habitat variable s predicted abundances of monit o del monte at 12 study sites. Competitive models (AIC <2) and the significant variables in each model are shown in bold. Model variables are the same as in Table 1-2 except for habitat structure. For this analysis, number of N. dombeyi and Austrocedrus chilensis were used instead of average number of trees. Models Significant variables AIC AICc AIC df R2 Pvalue Global 44.623134.623 86.1549 0.87 No. of reproductive mistletoe 1.50 <0.0001 Habitat complexity 2.02 0.05 Food plants 48.07051.070 2.6003 0.80 No. of reproductive mistletoe 0.72 <0.0001 Fruit abundance and 48.29251.292 2.8233 0.83 bamboo Crop size of mistletoe 0.75 0.0005 Crop size of A. chilensis 0.45 0.04 Mistletoe plants and 48.06549.398 0.9292 0.83 bamboo No. of reproductive mistletoe 0.95 <0.0001 Mistletoe 48.06948.469 01 0.80 No. of reproductive mistletoe 0.89 <0.0001 Bamboo 65.31165.711 17.2421 0.01 Habitat structure 54.06864.068 15.5995 0.67 Bamboo cover -2.020.01

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28 N o o f r e p r o d u c t i v e m i s t l e t o e + C r o p s i z e o f m i s t l e t o e + N o A c h i l e n s i s p l a n t s + C r o p s i z e o f A c h i l e n s is + N o A m i c r o p h y l l a p l a n t s B a m b o o c o v e r + B a m b o o h e i g h t H a b i t a t c o m p l e x i t y C a n o p y c o v e r + U n d e r s t o r y c o v e rN o N d o m b e y i N o A c h i l e n s i s Mean and SD 0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 10000 20000 30000 High abundance of mistletoe Low abundance of mistletoe No-mistletoe *** ** Figure 1-1.Means ( SD) of the habitat variable s measured at the 12 sites where abundances of monito del monte were determined. Means from raw data are shown. Variables log transformed prior to analysis are indicated by +. Significant diffe rences are shown as *** P < 0.0001, ** P < 0.01, P < 0.05.

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29 Bamboo cover (log) 01234 0 10 20 30 40 50 R2 = 0.01, P = 0.80 No. of reproductive mistletoe (log) 0.00.51.01.52.0 No. monitos del monte 0 10 20 30 40 50 R2 = 0.80, P < 0.001 Crop size of A. chilensis (log) 012345 0 10 20 30 40 50 R2 = 0.22, P = 0.13 Crop size of mistletoe (log) 012345678 0 10 20 30 40 50 R2 = 0.72, P < 0.001 Habitat complexity 012345 0 10 20 30 40 50 R2 = 0.26, P = 0.09 Figure 1-2. Simpler linear regression analysis of the number of monitos del monte vs. the significance predictor variables of the multiple regression models. The coefficient of determination (R2) and P -value are shown.

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30 CHAPTER 2 CONSERVATION IMPLICATIONS The interaction between this mistletoe and its marsupial disperser may have consequences at the community and ecosystem levels and habita t changes that impact either of these species could have indirect cascading e ffects throughout different interac tion webs. This mistletoe is considered a keystone species du e to the production of sugar-rich nectar and protein-rich fruits during periods of food scarcity. Th is mistletoe also increases mo rtality of host trees and could play an important role in plant community diversity. As the only known disperser for the mistletoe, the monito del monte is important in maintaining these processes. Habitat fragmentation disrupts the interaction between the mistletoe and monitos del monte (RodriguezCabal et al., 2007). Fragmentation negatively affects marsupial abundance, fruit removal, seed dispersal, and seedling recruitment. The lo cal extinction of monito s del monte has been associated with the complete disruption of mi stletoe seed dispersal (Rodriguez-Cabal et al., 2007). The reason for the local extinction of mon itos del monte in fragmented habitats is unknown. One possible mechanism is reduction of bam boo in fragments results in loss of nest sites and connectivity. Grazing by cattle and other introduced ungulates significantly alters habitat structure and composition of the forest, including reduc tion of bamboo (Veblen et al., 1992). Effects of grazing are particularly severe in forest fragments where introduced ungulates have easy access to entire patch. In this temperate forest, the bamboo bloom s and dies at approximately 40-60 years intervals. Although this is a natural process, th e die-off of the bamboo also likely has important consequences for populations of the monito de l monte by reducing nest sites and connectivity. Loss of bamboo within habitat fragments may have particularly severe consequences because

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31 these patches are isolated, reduc ing the probability of demogra phic rescue and recolonization following regeneration of bamboo within the patch.

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32 CHAPTER 3 CONCLUSION In the temperate forest of southern South America, the northern portion (to 41oS) has the highest biodiversity and number of endemic species, the lowest proportion of protected areas, and the highest rate of disturbanc e in this forest (Armesto et al., 1998, Rodriguez-Cabal et al., in press). The biodiversity of this te mperate forest currently is thr eatened by the highest rates of destruction, fragmentation, and de gradation ever experienced by th is system, with more than 120,000 ha of native forest destroyed or degraded annually (L ara et al., 1996). The alteration of natural hab itats through human ac tivities is the principal cause of the decline and extinction of natu ral populations (Gaston and Sp icer, 2004). Long-term plans to conserve biodiversity must not only focus in main taining the elements that form biodiversity but also on the interactions among these elements (Vazquez and Simberloff, 2003). Understanding the species and processes that shape ecosyste ms is fundamental for effective conservation (Wayne et al., 2006). No previous study has inves tigated explicitly factors that determine this austral marsupials distribution and abundance or the causes of its present decline. This study constitutes the first detailed ecological research focused on the identification of the habitat variables that play the most important role in maintaining the marsupials populations. My work demonstrates that both habitat factors (bamboo) and interacting species (mistletoe) are important in determining the distribution and abundance of this species.

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33APPENDIX CORRELATION ANALYSIS Table A-1. Correlation coefficients (r) betw een habitat variables in the presence-abs ence survey (n = 28). Significant correla tions after the Bonferroni corr ection are indicated by ( P < 0.0009). Crop size of mistletoe No. of A. chilensis plants Crop size of A. chilensis No. of A. microphylla plants Bamboo cover (%) No. of reproductive mistletoe 0.89* 0.56 0.15 0.17 -0.19 Crop size of mistletoe 0.50 0.10 0.18 -0.17 No. of A. chilensis plants 0.66* 0.42 0.29 Crop size of A. chilensis 0.10 0.39 No. of A. microphylla plants 0.33 Bamboo cover (%) Bamboo height (m) Habitat complexity Understory cover (%) Canopy cover (%) Table A-1. Bamboo height (m) Habitat complexity Understory cover (%) Canopy cover (%) Average no. trees No. of reproductive mistletoe 0.11 0.29 -0.004 0.38 -0.24 Crop size of mistletoe 0.02 0.01 -0.01 0.44 -0.28 No. of A. chilensis plants 0.47 0.02 -0.23 -0.07 0.13 Crop size of A. chilensis 0.40 0.19 -0.004 -0.06 0.16 No. of A. microphylla plants 0.36 -0.06 0.39 -0.05 0.004 Bamboo cover (%) 0.88* 0.35 -0.11 0.37 0.43 Bamboo height (m) 0.20 -0.29 0.29 0.40 Habitat complexity 0.73* 0.43 0.44 Understory cover (%) 0.23 0.15 Canopy cover (%) 0.88*

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34Table A-2. Correlation coefficients (r) between habitat variables in the abundance survey (n = 12). Significant correlations af ter the Bonferroni correction are indicated by ( P < 0.0006). Crop size of mistletoe No. of A. chilensis plants Crop size of A. chilensis No. of A. microphylla plants Bamboo cover (%) No. of reproductive mistletoe 0.94* 0.78 0.30 0.09 -0.28 Crop size of mistletoe 0.70 0.16 0.12 -0.13 No. of A. chilensis plants 0.41 0.33 -0.32 Crop size of A. chilensis 0.05 0.46 No. of A. microphylla plants 0.10 Bamboo cover (%) Bamboo height (m) Habitat complexity Understory cover (%) Canopy cover (%) N. dombeyi Table A-2. Bamboo height (m) Habitat complexity Understory cover (%) Canopy cover (%) N. dombeyi A. chilensis No. of reproductive mistletoe 0.33 0.28 0.13 0.21 -0.35 0.46 Crop size of mistletoe 0.39 0.34 0.18 0.28 0.20 0.55 No. of A. chilensis plants 0.63 0.39 0.11 0.56 -0.02 0.53 Crop size of A. chilensis 0.38 0.70 0.67 0.36 -0.14 -0.42 No. of A. microphylla plants 0.32 0.00 -0.23 0.10 0.18 0.09 Bamboo cover (%) 0.65 0.70 0.71 0.68 0.46 -0.26 Bamboo height (m) 0.67 0.50 0.96* 0.33 0.30 Habitat complexity 0.89* 0.58 -0.16 -0.08 Understory cover (%) 0.40 -0.16 -0.35 Canopy cover (%) 0.51 0.27 N. dombeyi 0.17

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37 Lara, A., C. Donoso, Aravena, J. C., 1996. La conservacin del bosque nativo en Chile: problemas y desafos. In: J.J. Armesto, C. Villagrn and M.T.K. Arroyo, (Eds.), Ecologa de los bosques nativos de Chile. Editorial Universitaria, Santiago de Chile pp. 335-362. Levey, D. J., 1988. Spatial and temporal variatio n in Costa Rican fruit and fruit-eating bird abundance. Ecological Monograph 58, 251-269. Levey, D. J., Benkman, C. W., 1999. Fruit-seed disperser interactions: timely insights from a long-term perspective. Trends in Ecology and Evolution 14, 41-43. Loiselle, B. A., Blake, J. G., 1991. Potential conse quences of local frugivore extinction for plant population in neotropical lowland wet forest. In : Levey D. J., Silva W. R. and Galetti M. (Eds.), Seed dispersal and frugivory: ecology, evolution and conservation. CAB International Press, Oxfordshire, UK pp. 397-406. MacKenzie, D. I., Nichols, J. D., Hines, J. E., Knutson, M. G., Franklin, A. B., 2002. Estimating site occupancy, colonization, and local extincti on when a species is detected imperfectly. Ecology 84, 2200-2207. Malizia, L. R., 2001. Seasonal fluctuations of birds, fruits, and flowers in a subtropical forest of Argentina. Condor 103, 45-61. Mann, G. F., 1955. Monito del monte Dromiciops australis Philippi. Investigacin Zoolgica Chilenas 2, 159-166. Mermoz, M., Martn, C., 1986. Mapa de vegetacin del Parque y la Reserva Nacional Nahuel Huapi Secretara de Ciencia y Tcnica de la Nacin, Delega cin Regional Patagonia, S. C. de Bariloche, Argentina. Moegenburg, S. M., Levey, D. J. ,2003. Do frugivores respond to fruit harvest? An experimental study of short-term responses. Ecology 84,2600-2612. Reid, N., 1987. The mistletoebird and Australian mistletoe: co-evolution or coincidence? Emu 87, 130-131. Rodrguez-Cabal, M. A., 2003. Habitat fragmentati ons effect on a mutualism of dispersal in the temperate forest of South America. Univer sidad Nacional del Coma hue, Bariloche, RN, Argentina. Rodrguez-Cabal, M. A., Aizen, M. A., Novar o, A. J., 2007. Habitat fragmentation disrupts a plant-disperser mutualism in the temperat e forest of South America. Biological Conservation 139, 195-202. Royle, J. A., Nichols, J. D., 2003. Estimating abu ndance from repeated presence-absence data or point counts. Ecology 84, 777-790.

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38 Schupp, E.W., Milleron, T., Russo, S.E., 2002. Dissemination limitation and the origin of species-rich tropical forests. In: Levey D.J., Silva W.R. and Galetti M. (eds.) Seed Dispersal and Frugivory: Ecology, Evoluti on, and Conservation. CAB International, Wallingford, Oxon, UK, pp. 19-43. Smith-Ramirez, C., 1993. Los picaflores y su recurso floral en el bosque te mplado de la isla de Chilo, Chile. Revista Chilena de Historia Natural 66, 65-73. Spiegel, O., Nathan, R., 2007. Incorpating disper sal distance into the disperser effectiveness framework: frugivorous birds provide comple mentary dispersal to plants in a patchy environment. Ecology Letters 10, 718-728. van der Pijl, L., 1982 Principles of dispersal in higher plants. Springer, Berlin, Germany. Vazquez, D. P., Simberloff, D,. 2003. Change s in interaction biodiversity induced by an introduced ungulate. Ecology Letters 6, 1077-1083. Vazquez, D. P., Aizen, M. A., 2004. Degree distri bution in plant-animal mutualistic networks: forbidden links or random interactions. Oikos 108, 421-426. Vazquez, D. P., Morris, W. F., Jordano, P., 2005. In teraction frequency as surrogate for the total effect of animal mutualists on plants. Ecology Letters 8, 1088-1094. Vazquez, D. P., Melian, C. J., Williams, N. M., Bluthgen, N., Krasnov, R., Poulin, R., 2007. Species abundance and asymmetric interacti on strength in ecological networks. Oikos 116, 1120-1127. Veblen, T V., Mermoz, M., Martin, C., Kitzbe rger, T., 1992. Ecological impacts of introduced animals in Nahuel Huapi National Park, Argentina. Conservation Biology 6, 71-83. Wayne, A. F., Cowling, A., Lindenmayer, D. B., Ward, C. G., Vellios, C. V., Donnelly, C. F., Calver, M. C., 2006. The abundance of a threat ened arboreal marsupial in relation to anthropogenic disturbance at lo cal and landscape scales in Me diterranean-type forest in south-western Australia. Bi ological Conservation 127, 463-476. Wiens, J. A., 1989. Spatial scaling in ecology. Functional Ecology 3, 385-397. Whittingham, M. J., Swetnam, R. D., Wilson, J. D., Chamberlain, D. E., Freckleton, R. P., 2005. Habitat selection by yellowhammers Emberiza citrinella on lowland farmland at two spatial scales: implications for conservati on management. Journal of Applied Ecology 42, 270-280. Wilson, D. E., Cole, F. R., Nichols, J. D ., Rudran, R., Foster, M., 1996. Measuring and monitoring biological diversity. Standard me thods for mammals. Smithsonian Institution Press, Washington, D. C, USA.

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40 BIOGRAPHICAL SKETCH Mariano Alberto Rodriguez Cabal was born in Buenos Aires, Argentina in 1976. His interest in biology began when he very young. Natu re has always exhibited a powerful attraction for him. He studied in a reli gious high school and obtained a degree in economics. When he finished his secondary studies, he moved to San Carlos de Bariloche in the foothills of the Andes, in western Argentina. The magnificent e nvironment around this city, which is within a national park, led him unequivocally to pursue university studies in biology. In 1997, he began studying Biol ogy at Universidad Nacional de l Comahue. In July 2003, he graduated as a Licenciado en Ciencias Bi olgicas. Under the guidance of Dr. Marcelo Aizen and Dr. Andrs Novaro, he conducted his unde rgraduate thesis resear ch on the effects of habitat fragmentation on a dispersal mutualism in the temperate forest of southwestern Argentina. After two years of field work, he completed and defended his thesis, obtaining the highest grade. In 2004, he received a Fulbright Scholarship to study abroad. In the next year, he moved to Florida, USA, where he started his masters studies at the University of Florida in the Department of Wildlife Ecology and Conserva tion. During those years worked under the supervision of Professor Lyn C. Branch. This wo rk was recognized with three grants. Twice he earned the Outstanding Academic Achievemen t Award at UF. In December 2007, he defended his masters thesis. He made a firm decision to devote himself to scientific research. He hopes to continue his doctoral education in the United Stat es of America. On completing his studies, he will return to his country to contribute to th e welfare of his people and the conservation of Argentinas biodiversity thr ough teaching and research.