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Status and Use of Tropical Hardwood Hammocks and Forested Residential Areas as Habitat for Resident and Neotropical Migr...

Permanent Link: http://ufdc.ufl.edu/UFE0021211/00001

Material Information

Title: Status and Use of Tropical Hardwood Hammocks and Forested Residential Areas as Habitat for Resident and Neotropical Migratory Birds in the Florida Keys
Physical Description: 1 online resource (61 p.)
Language: english
Creator: Karim, Annisa
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2007

Subjects

Subjects / Keywords: birds, conservation, deforestation, florida, fragmentation, gis, hammocks, hardwood, keys, migratory, tropical
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: The challenges of migration are immense and many avian species rely on stopover areas to rest and refuel before continuing migration. The availability and location of stopover habitat is vital to migratory species, and the loss of suitable stopover habitats has been implicated in the decline of migratory populations. Tropical hardwood hammocks in the Florida Keys are used by both resident and Neotropical migratory birds; however, anthropogenic activities in and around hammocks have resulted in the conversion of many of these forests to other urban uses. As a result, habitat loss and fragmentation have reduced suitable habitat. I used a Geographic Information Systems approach to quantify amount and status (protected/unprotected) of hammock forest patches in the Keys. More than 30% of hammock habitat in the upper Keys had been lost to deforestation from 1991 to 2004. Total remaining hammock habitat in the Florida Keys as of 2004 was 3,712 ha, of which 1,962 ha (53%) was located in the upper Keys, and 1,750 ha (47%) was located in the lower Keys. Of the remaining hammock habitat, approximately 37% remained unprotected throughout the Keys (46% and 27% in the upper and lower Keys, respectively). I conducted bird point counts at 86 survey sites in the northern Keys during March-May 2004 and 2005. Survey locations were established in both hammock habitat and in residential areas with varying degrees of tree canopy cover. Total hammock tree canopy had significant positive effects on presence of seven of the 35 species statistically analyzed, including the Black-whiskered Vireo (Vireo altiloquus), a bird of conservation concern. Conversely, hammock cover had significant negative effects on 12 species including common urban birds and non-native species. Cowbird species (Molothrus spp.) and feral cats (Felis catus), both of which were observed along hammock edges, may pose additional threats. Environmental education of the public on the ecological value of hammocks is needed as well as long term studies on avian-hammock interactions in the upper and lower Keys.
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.
Statement of Responsibility: by Annisa Karim.
Thesis: Thesis (M.S.)--University of Florida, 2007.
Local: Adviser: Main, Martin B.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2007
System ID: UFE0021211:00001

Permanent Link: http://ufdc.ufl.edu/UFE0021211/00001

Material Information

Title: Status and Use of Tropical Hardwood Hammocks and Forested Residential Areas as Habitat for Resident and Neotropical Migratory Birds in the Florida Keys
Physical Description: 1 online resource (61 p.)
Language: english
Creator: Karim, Annisa
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2007

Subjects

Subjects / Keywords: birds, conservation, deforestation, florida, fragmentation, gis, hammocks, hardwood, keys, migratory, tropical
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: The challenges of migration are immense and many avian species rely on stopover areas to rest and refuel before continuing migration. The availability and location of stopover habitat is vital to migratory species, and the loss of suitable stopover habitats has been implicated in the decline of migratory populations. Tropical hardwood hammocks in the Florida Keys are used by both resident and Neotropical migratory birds; however, anthropogenic activities in and around hammocks have resulted in the conversion of many of these forests to other urban uses. As a result, habitat loss and fragmentation have reduced suitable habitat. I used a Geographic Information Systems approach to quantify amount and status (protected/unprotected) of hammock forest patches in the Keys. More than 30% of hammock habitat in the upper Keys had been lost to deforestation from 1991 to 2004. Total remaining hammock habitat in the Florida Keys as of 2004 was 3,712 ha, of which 1,962 ha (53%) was located in the upper Keys, and 1,750 ha (47%) was located in the lower Keys. Of the remaining hammock habitat, approximately 37% remained unprotected throughout the Keys (46% and 27% in the upper and lower Keys, respectively). I conducted bird point counts at 86 survey sites in the northern Keys during March-May 2004 and 2005. Survey locations were established in both hammock habitat and in residential areas with varying degrees of tree canopy cover. Total hammock tree canopy had significant positive effects on presence of seven of the 35 species statistically analyzed, including the Black-whiskered Vireo (Vireo altiloquus), a bird of conservation concern. Conversely, hammock cover had significant negative effects on 12 species including common urban birds and non-native species. Cowbird species (Molothrus spp.) and feral cats (Felis catus), both of which were observed along hammock edges, may pose additional threats. Environmental education of the public on the ecological value of hammocks is needed as well as long term studies on avian-hammock interactions in the upper and lower Keys.
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.
Statement of Responsibility: by Annisa Karim.
Thesis: Thesis (M.S.)--University of Florida, 2007.
Local: Adviser: Main, Martin B.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2007
System ID: UFE0021211:00001


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6f7b5373b86e5c161a7933273c974730
256f8d6ae3f2e5ea63b4501f7b4a78873c2afaa9







STATUS AND USE OF TROPICAL HARDWOOD HAMMOCKS
AND FORESTED RESIDENTIAL AREAS AS HABITAT FOR RESIDENT AND
NEOTROPICAL MIGRATORY BIRDS INT THE FLORIDA KEYS




















By

ANNISA KARIM


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

2007




































O 2007Annisa Karim
































"In Wildness is the Preservation of the World."
-Thoreau/ Porter/ Metallica









ACKNOWLEDGMENTS

I would like to express my enormous gratitude to my parents, Muneer and Laila, and to my

sister, Farah, for their love and encouragement throughout my life. The advice and insight they

have provided and their belief in me has enabled me to achieve my goals and follow my dreams.

I am extremely fortunate to have them in my life! SHRUKRAN.

I would like to thank the chairperson of my committee, Dr. Martin B. Main for his

encouragement, support, guidance and sense of humor all of which enabled me to attain a

Master of Science Degree from the University of Florida' s Department of Wildlife Ecology and

Conservation. His passion and commitment to his work is a true inspiration. I would also like to

thank my other committee members, Dr. Mark Hostetler and Dr. Mike Moulton for their advice

and expertise. The support staff at the Department of Wildlife Ecology and Conservation

deserves a special thank you; their helpfulness, dedication and team-work is commendable.

I thank Islamorada Village of Islands for the partial funding of my research. Additionally,

I would like to thank the staff and volunteers (especially Donna Sprunt) of the Windley Key

Fossil Reef Geological State Park and the staff at the Keys Marine Laboratory for their

helpfulness during the fieldwork portion of my research.

I would like to express a special thanks to the Hennig Family...Markus, Melissa and

Robert have been extremely supportive especially during my last semester as a graduate student.

I am grateful for their encouragement, friendship and proficiency at making a great cup of coffee

(but mostly for their encouragement and friendship).












TABLE OF CONTENTS


page

ACKNOWLEDGMENT S .........__... ......._. ...............4....


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


LI ST OF FIGURE S .............. ...............7.....


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


CHAPTER


1 INTRODUCTION ................. ...............10.......... ......


2 IVETHODS ................. ...............13.......... .....


Study A rea ................. .. .............. ...............13.......
Assessment of Hammock Habitat ................. ...............14................

Bird Surveys .............. ...............15....
A analysis .............. ...............17....

3 RE SULT S .............. ...............27....


Assessment of Hammock Habitat ................. ...............27................

Bird Surveys .............. ...............29....


4 DI SCUS SSION ................. ...............45................


Assessment of Hammock Habitat ................. ...............45................
Bird Surveys ............... ...............46....
Conservation Implications ................. ...............51.......... .....


LITERATURE CITED .............. ...............55....


BIOGRAPHICAL SKETCH ............. ..............61.....










LIST OF TABLES


Table page

2-1 Number and description of 20-m, Eixed-radius point count bird survey sites including
hammock size, conservation status, and location and description (MM = mile marker
on hi ghway US- 1). ................. ................. 19.............

2-2 Description and quantifieation (mean and s.d., median and range) of habitat
variables measured at point count locations (20-m Eixed radius) where bird surveys
were conducted. ............. ...............23.....

2-3 Foraging guild and life history descriptions .............. ...............24....

3-1 Total amount (ha) of hammock habitat including percent of total in protected status
by island in the Florida Keys as of 2004 .........._._ ...... ..._ ...............34.

3-2 Partially protected hammock habitat patches in the Florida Keys including total
number of patches and area (ha) in protected and unprotected status by island and
patch size category (ha) of 2004 (islands that did not contain partially protected
patches were excluded from this table)............... ...............35.

3-3 Avian species detected during point count surveys including standard abbreviations,
common and scientific names, life history and breeding status, and foraging guild
designations............... ..............3

3-4 Influence of measured habitat variables on species presence or absence from
analyses with general linear models (data represent p-values). A negative p-value
indicates an inverse relationship between the habitat variable and species presence. .....40

3-5 Influence of measured habitat variables on life history and foraging guilds from
general linear model analyses with zero-inflated Poisson regression model (data
represent p-values). A negative p-value indicates an inverse relationship between the
habitat variable and species presence. ............. ...............41.....

3-6 Hammock habitat variables for birds that lacked sufficient data for statistical
analyses. Data reflect the range of percent cover (smallest to largest) for PCTHAM,
PCT CAN and HAM+CAN measurements where birds were observed. .........................42










LIST OF FIGURES


Figure page

2-1 Geologic delineation of the Florida Keys by substrate. ............ ......................2

2-2 Delineation of Upper Keys (consistent with Bancroft et al. 1995) study area, Lower
Keys study area and extent of avian surveys. ............. ...............26.....

3-1 Area (ha) of tropical hardwood hammock habitat patches by patch size (ha) category
and protected status in the Florida Keys in 2004 (number of habitat patches provided
above each bar). ............. ...............44.....









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

STATUS AND USE OF TROPICAL HARDWOOD HAMMOCKS
AND FORESTED RESIDENTIAL AREAS AS HABITAT FOR RESIDENT AND
NEOTROPICAL MIGRATORY BIRDS INT THE FLORIDA KEYS

By

Annisa Karim

December 2007

Chair: Martin B. Main
Major: Wildlife Ecology and Conservation

The challenges of migration are immense and many avian species rely on stopover areas to

rest and refuel before continuing migration. The availability and location of stopover habitat is

vital to migratory species, and the loss of suitable stopover habitats has been implicated in the

decline of migratory populations. Tropical hardwood hammocks in the Florida Keys are used by

both resident and Neotropical migratory birds; however, anthropogenic activities in and around

hammocks have resulted in the conversion of many of these forests to other urban uses. As a

result, habitat loss and fragmentation have reduced suitable habitat. I used a Geographic

Information Systems approach to quantify amount and status (protected/unprotected) of

hammock forest patches in the Keys. More than 30% of hammock habitat in the upper Keys had

been lost to deforestation from 1991 to 2004. Total remaining hammock habitat in the Florida

Keys as of 2004 was 3,712 ha, of which 1,962 ha (53%) was located in the upper Keys, and

1,750 ha (47%) was located in the lower Keys. Of the remaining hammock habitat,

approximately 37% remained unprotected throughout the Keys (46% and 27% in the upper and

lower Keys, respectively). I conducted bird point counts at 86 survey sites in the northern Keys

during March-May 2004 and 2005. Survey locations were established in both hammock habitat









and in residential areas with varying degrees of tree canopy cover. Total hammock tree canopy

had significant positive effects on presence of seven of the 3 5 species statistically analyzed,

including the Black-whiskered Vireo (Vireo altiloquus), a bird of conservation concern.

Conversely, hammock cover had significant negative effects on 12 species including common

urban birds and non-native species. Cowbird species (M~olothrus spp.) and feral cats (Felis

catus), both of which were observed along hammock edges, may pose additional threats.

Environmental education of the public on the ecological value of hammocks is needed as well as

long term studies on avian-hammock interactions in the upper and lower Keys.









CHAPTER 1
INTTRODUCTION

Many types of latitudinal migratory birds exist (Hayes 1995). According to Rappole

(1995), Nearctic-Neotropical migrants are birds for which all or part of their populations breed

north and winter south of the Tropic of Cancer. In the western hemisphere, there are 341 species

of Nearctic-Neotropical migratory birds, hereafter referred to as Neotropical migrants (American

Ornithologists' Union 1998; 2000).

Deforestation of wintering grounds and the fragmentation of breeding areas pose two

serious habitat constraints for the conservation of migratory passerines (Morris et al. 1996). The

challenges of migration are immense and many avian species rely on "stopover" areas to rest and

refuel energy reserves before continuing on their migrations (Kerlinger 1989; Terborgh 1989;

Moore 2000; Ydenberg et al. 2002). The loss and degradation of stopover habitats represent a

third serious habitat constraint for the conservation of these migrants. The availability, location

and quality of stopover habitat are vital to migratory species and the loss of suitable stopover

habitats has been implicated in the decline of migratory bird populations (Sauer & Droege 1989;

Moore et al 1990; Barrow et al. 2000). The combination of factors such as seasonal variability,

availability of appropriate habitat, costs of migration, and ability to navigate play an important

role in the success of migratory movements. While en route, migrants must make quick

decisions about the location and type of landscape they use.

The Florida Keys are a chain of low-lying islands that extend 354-km from the

southeastern tip of Florida and arc in a southwesterly direction to the Dry Tortugas (Florida

Department of Community Affairs 2003). These islands are located along one of the primary

migratory routes for birds that breed in tropical and temperate North America and winter in the

Caribbean and South America (Lincoln et al. 1998; US Fish and Wildlife Service 1999). The










Keys also provide habitat for Caribbean avian species that rarely appear anywhere else in North

America, such as the Mangrove Cuckoo (Coccyzus minor, MACU), Black-whiskered Vireo

(Vireo altiloquus, BWVI) and the state listed, threatened White-crowned Pigeon (Patagioena~s

leucocephala formerly Columba leucocephala, WCPI; US Fish and Wildlife Service 1999;

Banks et al. 2003).

One type of plant community used by migrant and resident birds in the Florida Keys is

tropical hardwood hammock (US Fish and Wildlife Service 1999). Historically, tropical

hardwood hammocks, hereafter referred to as hammocks, were found as far north as Cape

Canaveral on the east coast, and up to the mouth of the Manatee River on the west coast of

Florida. However, development pressure resulted in the conversion of many of these forests to

urban and agricultural uses. Today, hammocks occur primarily as remnant habitats in extreme

south Florida and in small preserves along the Atlantic coast from Miami-Dade north to Martin

County. The largest remaining tracts of extant hammocks occur in the Florida Keys. Hammocks

are closed canopy forests characterized by evergreen and semi-deciduous woody species

primarily of West Indian origin (Snyder et al. 1990; Sunquist et al. 2002; Lodge 2005).

Hammocks in the Florida Keys occur on the highest elevations where flooding rarely occurs and

are, therefore, prime areas for human habitation.

Forman (1995) defines connectivity as "spatial continuity of habitat or cover type across a

landscape". Therefore, "breaking up of a habitat or cover type into smaller, disconnected

parcels" is fragmentation. Plant communities in the Florida Keys, including hammocks, are

naturally discontinuous and fragmented due to the geologic nature of the archipelago (US Fish

and Wildlife Service 1999; Lodge 2005). Anthropogenic activities in and around hammocks

have resulted in the conversion of many of these forest patches, primarily to urban uses (Bancroft









et al. 1995). As a result, habitat loss and further fragmentation of remaining hammocks have

reduced potentially important habitat for resident and Neotropical migratory birds on the

archipelago. Furthermore, extant hammock habitat under private ownership in the Florida Keys

is under intense development pressure and loss of habitat provided by these upland forests may

have significant conservation implications for both Neotropical migrants and resident species of

birds (Wilson 1992; Russell et al. 1998; Barrow et al. 2000; Petit 2000; Simons et al. 2000). The

obj ectives of this study are to (1) determine the extent of hammock habitat throughout the

Florida Keys as of 2004, (2) quantify area and number of hammock patches under conservation

status and (3) describe bird communities by species, foraging guild, and life history that utilize

hammocks and residential areas with varying degrees of canopy cover to better understand avian

habitat preferences and needs during spring migration.









CHAPTER 2
METHOD S

Study Area

The Florida Keys (Keys) are a chain of small islands located within the subtropical region

of the Western hemisphere. The Keys have a mild tropical climate, annual rainfall averages 101

cm (40 in), temperatures vary from 10-320 C (50-900 F), and mean temperature exceeds 180C

(640 F) during all 12 months (Florida Climate Center 2005). This archipelago extends 354 km in

a southwest direction from the southeast tip of peninsular Florida (250 N, 800 W) to Key West

(240 N, 810 W; Fig. 2-1). In a geological context, the upper Keys rest on Key Largo Limestone

and extend from Soldier Key (250 N, 800 W) approximately 150 km southwest to Big Pine Key

(240 N, 810 W). The lower Keys sit on Miami oolitic limestone and extend from Big Pine Key

westward approximately 60 km to Key West (240 N, 810 W; Fig. 2-1). Recent deposits of

carbonate sands comprise the substrate of the deep, open waters between Key West, the

Marquesas Keys and the Dry Tortugas (Craighead 1971; Brown et al. 1990; Snyder et al. 1990;

Ripple 1995; Waitley 1997).

Plant communities in the Florida Keys are naturally discontinuous and fragmented due to

the geologic nature of the archipelago. Submerged portions of many islands support coral reef

communities, low-lying areas sustain mangrove, tidal marsh, and freshwater marsh communities,

while hammocks and pine (Pinus elliottii var. densa) communities thrive on the highest

elevations where flooding rarely occurs (US Fish and Wildlife Service 1999; Lodge 2005).

Hammocks are closed canopy forests and are dominated by a number of evergreen and semi-

deciduous woody species, most of which are of West Indian origin (Snyder et al. 1990; Sunquist

et al. 2002; Lodge 2005). The largest areas of hammocks occur in the northern portions of the

Keys, especially on the largest island, Key Largo. Hammocks tend to get smaller as island size









decreases south of Key Largo, with the exception of Big Pine Key, the largest island in the lower

Keys. Hammocks throughout the Keys have undergone anthropogenic influences that have

systematically reduced and further fragmented hammock habitat, particularly from residential

and urban development (US Fish and Wildlife Service 1999; Lopez 2001).

Assessment of Hammock Habitat

I used a Geographic Information Systems (ESRI GIS Software ArcMap 9.0) approach to

quantify cover, distribution, and status (protected/unprotected) of hammock forest patches in the

Florida Keys. I quantified hammock cover throughout the Keys and measured loss of hammock

cover in the upper Keys during 1991-2004 based on an earlier assessment by Bancroft et al.

(1995). For comparative purposes I defined the upper Keys consistent with Bancroft et al. (1995)

as extending from the Ragged Keys (a small chain of islands in Miami-Dade Co. north of Key

Largo, 250 N, 800 W), south to Long Key (240 N, 800 W). Lower Keys were defined as

extending from Duck Key (240 N, 800 W) to Key West (240 N, 810 W; Fig. 2-2).

I used 2004 digital orthophoto quarter quadrangles (DOQQ) from the Land Boundary

Information System (Florida Department of Environmental Protection 2003) at a 1-m resolution

as the base layer of my map. I obtained the Monroe Co. (includes Keys) boundary layer from the

Florida Geographic Data Library (University of Florida GeoPlan Center) and used the ArcMap

9.0 clip function to isolate the Monroe Co. 1995 land use/land cover data available from the

South Florida Water Management District (online: http://www.sfwmd. gov/site/index. php?id=1;

accessed December 2005). I superimposed the resulting 1995 dataset on the DOQQ and used the

Florida Land Use Land Cover Classification System (FLUCCS, level 3 code 4260) to designate

hammock habitat in the Keys (defined as tropical hardwoods by FLUCCS; Florida Department

of Transportation 1999). I created a new data layer to reflect the extent of hammock habitat

throughout the Keys (including the Ragged Keys) as of 2004. This new layer reflected









refinements to the 1995 FLUCCS code designations (e.g., new roads, water bodies, and land use

changes since 1995) based on examination of 2004 DOQQ aerial photos and ground-truthing. I

created polygons, via visual classification (1:4000), to reflect hammock habitat not recognized in

the 1995 dataset and in the Ragged Keys, which occur in Miami-Dade Co. but were included in

Bancroft et al. (1995). These additional hammock polygons were classified as commercial or

residential in the 1995 dataset because they were located within areas zoned for residential and

commercial use but had not yet been developed as of 2004.

The 2005 Florida Natural Areas Inventory Conservation Lands Layer was added to the

map and I used the intersect function in ArcMap to identify protected lands in Monroe Co. This

enabled me to identify hammock habitat protected for conservation under federal, state, local and

private management in the Florida Keys. I generated new layers to differentiate between

protected and unprotected status of hammock forest patches in the upper and lower Keys. I

ground-truthed 10% of the mapped hammock polygons and determined hammock coverage of

these polygons was 98% accurate. I calculated total area of extant hammock habitat (protected

and unprotected) in the upper and lower Keys by creating a new field in the attribute table for the

2004 hammock habitat layer. This included corrected data from FLUCCS code 4260

designations from the 1995 layer and the polygons I constructed. I used ArcMap's calculate

values function to calculate the area of all hammock polygons, converted results to hectares, and

compared results from the upper Keys with those of Bancroft et al. (1995).

Bird Surveys

I conducted morning bird surveys (point counts) in hammock forest habitat and residential

areas with varying levels of hammock and tree canopy cover during March-May (spring

migration) 2004 and 2005. Point counts were conducted between sunrise and 180 minutes after

sunrise as 20-m fixed-radius point counts with 5-min survey intervals (Hutto et al. 1986). I









established 86 point count locations from Key Largo to Vaca Key (Table 2-1), 65 of which were

in residential areas and 21 in protected hammock habitat. Of the 65 point count locations within

residential areas, six occurred on undeveloped parcels in private ownership that had extant

hammock habitat as of 2004. Point count locations were s parated by >250 m to maintain

independence of observations (Hutto et al.1986). I surveyed birds at each point count location

Hyve times each Hield season with the exception of one protected hammock where two point count

locations were visited four times during 2004 because permission to access the hammock was

delayed. Each 20-m Eixed radius point count assessed bird communities over an area of

approximately 0.13 ha (0.33 ac). During surveys, I recorded all birds by call, song or sight

within the 20-m radius. Species recorded as flying over/through without stopping were removed

from the dataset used for analysis. Additionally, sea birds (e.g., terns, Sterna spp.), wetland birds

(e.g., Great Egrets, Ardea alba), and domestic species (e.g., Peafowl, Pavo cristatus) were

removed from the dataset used for analysis.

Selection of point count locations for bird surveys was done separately for protected and

residential areas. Protected hammocks in the upper Keys (Key Largo to Marathon) were

identified during the GIS portion of the study. All existing protected hammocks in the upper

Keys were included in the bird surveys. All hammock habitats within the study area have been

described by the US Fish and Wildlife Service (1999) as secondary forest.

All accessible residential areas from Key Largo (city of Key Largo) to Vaca Key (city of

Marathon; 250 N, 800 W to 240 N, 810 W) were included in a pool of potential sites for bird

surveys. Most residential areas and, consequently potential bird survey sites, were located on the

larger islands. Locations dominated by the invasive non-native shrub, Brazilian pepper (Schinus

terebinthifolius), were eliminated from the pool of potential study sites because this shrub









attracts a disproportionate number of birds seeking food (Myers & Ewel 1990; Spector & Putz

2006) and its presence on some sites would confound the data. Potential survey sites were

numbered and randomly drawn to identify point count locations. Survey sites in residential areas

ranged from 0-100% hammock cover because some areas zoned as residential included

undeveloped parcels with hammock habitat.

The northern- and southern-most point count locations were separated by approximately 97

km, which necessitated a stratified random approach to sampling. I conducted surveys randomly

within groups of locations (approximately 16 point count locations/morning) and alternated the

initiation of surveys between north and south. This stratified random and alternating directional

survey scheme (first north to south then south to north) was used throughout both field seasons.

Analysis

Using a GIS approach, I was able to import GPS coordinates of all survey locations into

ArcMap and create a 20-m buffer around each survey location. Within each 20-m fixed-radius

survey location, I defined and measured eight habitat variables (Table 2-2). Variables included

hammock habitat (percent cover of native hammock tree canopy and understory cover;

PCT_~HAM) tree canopy (percent cover of native tree canopy cover with cleared understory;

PCT_CAN), ground cover (percent cover of grasses and residential lawn or managed understory

with no tree canopy cover; PCT_GROUND), and other vegetation (percent cover of non-

hammock vegetation, e.g., mangrove and pine habitats; PCT_OTHER). I also quantified

distance from survey location to the closest hammock (D2HAM; if survey location was situated

within a hammock D2HAM = 0) and the size of that hammock (SIZEHAM). I created two

combined variables, total canopy (combined sum of hammock habitat and tree canopy;

HAM+CAN) and the distance from the point count location to the nearest hammock times the

size of that hammock (DXSIZE).









Birds were analyzed on a species specific basis as well as by foraging guild and life history

traits (Table 2-3). Foraging guild (foraging technique) categories were defined by Ehrlich et al.

(1998). Migratory and non-migratory status was defined by Rappole (1995) and breeding

locations were defined by the Florida Fish and Wildlife Conservation Commission (2003).

Overall migratory and breeding status determined life history categories.

The relationships between count data and landscape attributes were regressed using general

linear models. Point count data were modeled on a per species, life history, and foraging guild

basis using eight landscape variables (Table 2-2) as covariates in Poisson regression. Species

detected fewer than 11 times were excluded from analysis due to small sample sizes. Count data

with excessive zeros were modeled using zero-inflated Poisson (ZIP) logistic regression (Martin

& McIntyre 2007). Count data without excessive zeros were modeled using the proc glimmix

command and all analyses were conducted in SAS Version 9.1.3 (Copyright 2002-2003 by SAS

Institute Inc., Cary, NC, USA).

Species richness (cumulative number of species detected at each survey location) was

calculated and modeled against the habitat variables. Additionally, the Shannon-Wiener

diversity index (Hs = -1 pi In pi; where H = diversity index; s = species richness, and pi = the

proportion of individuals found in the i.th species) was computed for all species detected (other

than seabirds and domestic species) and modeled against the habitat parameters. This index was

chosen because it is sensitive to changes in the abundance of rare species in a community (Solow

1993) and may better reflect the composition of all species found therein.









Table 2-1 Number and description of 20-m, fixed-radius point count bird survey sites including
hammock size, conservation status, and location and description (MM = mile marker
on highway US-1).


Survey Size (ha) of Protected .oato bn ecrpi
sites Hammock" Status (Y/N)



7 70.30c Y Dagny Johnson Key Largo Hammock Botanical
State Park, Key Largo
John Pennekamp Coral Reef State Park, Key
2 3.87 Y
Largo

3 N/A NResidential area (near Paradise Point Cove), Key
Largo (MM 105)

2 N/A N Residential area (Taylor Creek Village), Key
Largo (MM 105)
Located in residential area in Key Largo (In Taylor
1 17.39 N
Creek Village, MM 105)

4 N/A N Located in residential area on Key Largo (Near
Largo Sound Village and Angler's Park, MM 104)

Located in residential area on Key Largo (Near
2 N/A N Largo Sound Park, between MM 104 and MM
103)
Located in residential area on Key Largo (Near
6 N/A N Cross Key Waterway Estates and Twin Lakes,
MM 103)

6 N/A NLocated in residential area on Key Largo (Between
Lime Drive and Troup Rd, MM 102)


2 N/A NLocated in residential area on Key Largo (In Key
Largo Trailer Village, MM 101)

Located in residential area on Key Largo (Between
3 N/A N Lakeshore Drive and East Shore Drive, between
MM 101 and 100).









Table 2-1 (continued).


Survey Size (ha) of .oato bn ecrpi
sie aII1~a Protected Status (Y/N) Lcto n ecito


2 N/A N Located within a residential area on Key
Largo (Oceanside near MM 99 between
Sailfish Trail and Ocean Shores Drive)


1 86.90 N Located within a residential area on Key
Largo (Oceanside near MM 99 between
Sailfish Trail and Ocean Shores Drive)


2 N/A N Located within a residential area on Key
Largo (Northeast Papa Johns Pizza, near
MM 98)
4 N/A N Located in a residential area on Key Largo
(Near Sunset Point and the Fl. Keys
Marine Sanctuary Bldg., MM 95)
1 0.61 N Located in a residential area on Key Largo
(Between Harry Harris State Park and
Burton's Yacht Basin, MM 93)


1 N/A N Located in a residential area on Key Largo
(Between Harry Harris State Park and
Burton's Yacht Basin, MM 93)


1 4.52 N Located in a residential area on Key Largo
(Between Harry Harris State Park and
Burton's Yacht Basin, MM 93)


1 4.44 N Located in a residential area on Key Largo
(Between Harry Harris State Park and
Burton's Yacht Basin, MM 93)


1 0.31 N Located within a residential area on Key
Largo Tavernier (West of Bank of
America, MM 92)













Survey Size (a o Protected Status (Y/N) Location and Description"
sites Hammock"

3 N/A N Located within a residential area on Key
Largo Tavernier (Near Tavernier Harbor,
MM 92)
2 N/A N Located within a residential area on Key
Largo Tavernier (Near Point Lowe, MM
92)
4 N/A N Located in a residential area on Plantation
Key (Near Tavernier Creek and Sunshine
Estates, between MM 91 and MM 90)


2 4.67 Y Indigo Hammock just north of MM90
between Ocean Drive and Riviera
(Oceanside). Part of the conservation lands
within Islamorada
3 N/A N Located in residential area on Plantation
Key (Between Coral Shores High School
and Lake Harbor, MM 90)
4 N/A N Located in residential area on Plantation
Key (Between Seminole Blvd. and High
Point Rd just north of the Govt. center,
MM 89)
2 N/A N Located in residential area on Plantation
Key (Between Plantation Blvd. and Royal
lane, south of MM 88)
3 10.68 Y Windley Key Fossil Reef Geological State
Park.
2 N/A N Located in residential area on Upper
Matecumbe Key (Near Cheeca Lodge
Resort, MM 82.5)
2 N/A N Located in residential area on Upper
Matecumbe Key (Near Kaiyo Restaurant,
between MM 82 and MM 81)


Table 2-1 (continued).














Survey Size (ha) of Protected .oaio n Deciob
sites Hammock" Status (Y/N)

1 4.46 Y THH Layton Trail a part of Long Key State
Park
1 1.24 Y THH within Long Key State Park
2 12.57 Y Blue Heron Hammock West of the Marathon
Airport. Managed by FWCC.

3 14.37 Y Museums and Nature Center of Crane Point in
Marathon managed by the Florida Keys Land
and Sea Trust

a If point count locations) is/are located within a hammock, hammock size (ha) is provided. If point count
locations) is/are not located within a hammock, N/A. b MM = Mile Marker. Mile Markers are used as reference
points of locations along the Overseas Highway in the Florida Keys. Mile Marker Numbers descend from north to
south. "Dagny Johnson Key Largo Hammock Botanical State Park contains many patches of hammock, 70.3 ha
represents the amount of continuous hammock habitat within which point count surveys were conducted.


Table 2-1 (continued).










Table 2-2 Description and quantification (mean and s.d., median and range) of habitat variables measured at point count locations
(20-m fixed radius) where bird surveys were conducted.
Protected sites (%) Unprotected (%)
Habiat Vriabe DecripionMean SD Median Range Mean SD Median Range


96.0 8.4 100 6.-
100


Percent cover of native hammock (includes
native tree canopy and understory cover)

Percent cover of native hammock tree
canopy (cleared understory)
Percent cover of grasses and residential
lawns

Percent cover of non-hammock vegetation
(e.g., mangroves, pine rocklands, etc.)
Distance (m) to closest hammock from
survey location (if survey location was
located within a hammock, D2HAM = 0)
size (ha) of closest hammock from survey
location (if survey location was located
within a hammock, the size of that hammock
was recorded)

PCTHAM + PCT_CAN; percent cover of
total native hammock canopy (regardless of
understory)
D2HAM SIZEHAM; distance (m) of the
ClOsest hammock multiplied by the size of
that hammock.


18.1 30.2


0 0-100


PCT HAMa


PCT CANa


0.3 1.4 0 0-6.2 18.0 15.6 14.5 0-65.3


0.5 2.2


0 0-10.1 10.1 10.9 6.3 0- 34.5


PCT GROUNDa


PCT OTHERa


0.4 1.6 0 0-7.5 1.4 4.9


0 0-27.4


0 0 0 0-0 59.1 83.2 25.9 0- 416.5


D2HAMa


1.2-
70.3


<0.1-
86.9


SIZEHAMa



HAM+CANb


29.3 30.0 12.6


6.9 19.0 0.3


70.8-
96.3 7.3 100 10


36.2 27.9 30.9 0-100


0-
0 0 0 0-0 372.4 1828.4 9.9 143.


DXSIZEb


" Variables analyzed as covariates in first set of analyses. b Variables analyzed as covariates in second set of analyses










Table 2-3 Foraging guild and life history descriptions
Life History Description
YRa Year-round residents; non-migratory birds

aBb Migratory Breeders: Neotropical migratory birds that migrate to the northern FL,
Keys to breed.
Nb Neotropical migratory birds that use the northern FL Keys as stopover sites and
continue to migrate north for breeding purposes
Foraging Guildc Description
AF Aerial foragers: Captures flying insects while in prolonged continuous flight.
BG Bark gleaners: Gleaning prey from tree trunks and branches; includes excavating
and drilling into bark.
FG Foliage gleaners: Gleaning prey (including fruits) from foliage of vegetation and
occasionally branches.
GG Ground gleaners: Picking up items from the surface of soil, lawns, sand, forest
floor, etc.
HA Hawking: Short flights from perch to capture flying insects.
HG Hover and gleaners: Gleaning while hovering. Takes nectar, insects and/or berries
from plants above the ground while hovering.
HO Hover and pouncers: Hovering and pouncing; hovering before swooping or
dropping down on prey.
SW Swoops: Snatches up prey from ground in talons after gliding decent from perch
with wings spread.
aBirds reported to breed in the northern Keys (Florida Fish and Wildlife Conservation Commission 2003). bMigratory status of species (Rappole 1995).
c Foraging guild/foraging techniques for species defined by Ehrlich et al. (1998).






































































~~I C ~ 1.~ .~- 1.1.~.-11~ ..I~-I:L
I I I-I I~ ~ -I -I E I1 I I~ I


Figure 2-1 Geologic delineation of the Florida Keys by substrate.



































I Florida
M RonroeCounty

SUpper Keys


I 1 _I- ~ -. .. F .. I Count Locations


0 10 20 40 60


Gulf Of ewlco


Atlantic Ocean


Pt

,~ B

"' Ir-.py.~
r
:Ir a

I:-


i
I


-. I1 -1*=-a-J -I ... -us..F e-bruary2007)
'~~~~~~. I A. bers Conical EqualArea

DataSourceforF .) ...I. a IE i ...ls. .-- FGDL



Figure 2-2 Delineation of Upper Keys (consistent with Bancroft et al. 1995) study area, Lower

Keys study area and extent of avian surveys.









CHAPTER 3
RESULTS

Assessment of Hammock Habitat

Hammock habitat in the Florida Keys encompassed 3,712 ha and accounted for 19% of the

Keys landscape during 2004. Of that existing hammock habitat, 1,962 ha (53%) was located in

the upper Keys and 1,750 ha (47%) was located in the lower Keys (Table 3-1). Hammock

habitat in the upper Keys was divided among 124 habitat patches (median = 1.5 ha, range =

0.1-205.7 ha), of which 1,066 ha (54%) were under conservation status. Hammock habitat

located in the lower Keys was divided among 102 patches (median = 4.4 ha, range = 0.3-96.3

ha), of which 1,283 ha (73%) were protected under conservation status (Table 3-1). In total,

therefore, 2,349 ha (63%) of hammock habitat in the Florida Keys were protected from

development.

Key Largo, the largest and one of the northernmost islands in the archipelago, contained

the most hammock habitat (1,834 ha) and the largest amount of habitat in conservation status

(1,024 ha). These conserved patches were situated largely within the Dagny Johnson Key Largo

Hammock Botanical State Park and the Crocodile Lake National Wildlife Refuge. The largest

individual patch of hammock habitat in the entire Keys (206 ha) was also located on Key Largo.

Many islands in the Keys did not contain any hammock habitat either as a result of land use

change or unsuitable environmental conditions.

Hammock patches throughout the Keys were either fully protected (53 patches, 1062 ha),

partially protected (43 patches, 2081 ha) or in private ownership slated for development (130

patches, 569 ha). Partially protected hammocks were designated as those areas that included

protected and unprotected contiguous hammock habitat. The number and area (ha) of hammock

patches varied by patch size category (Fig. 3-1). The majority of area (892 ha, 84%) and 30% of









the total number of patches represented by fully protected hammocks were >20 ha in size.

Similar results were found for partially protected hammocks. The maj ority of area (1,956 ha,

94%) and 5 1% of the total number of patches represented by partially protected hammocks were

>20 ha in size. Twenty-five of these patches occurred in the upper Keys and 18 in the lower

Keys (Table 3-2). Conversely, the total number of unprotected hammock patches was dominated

by patches of 0-5 ha (108 patches, 83%). Seven (5%) unprotected patches >20 ha encompassed

267 ha and represented 47% of isolated, unprotected hammock habitat (i.e., not contiguous with

protected hammock habitat).

Comparisons with Bancroft et al. (1995) revealed hammock habitat declined by 31% in the

upper Keys since 1991, which represents a loss of 870 ha over a 13-year period. In 1991, the

upper Keys contained 2,832 ha of hammock divided into 1,068 patches of which, 987 were

smaller than 5 ha (Bancroft et al. 1995). By 2004, this region contained 1,962 ha of hammock

divided among 124 patches of which, 93 were smaller than 5 ha. This represents a decrease of

944 (88%) total fragments and a decrease of 894 (91%) patches under 5 ha. In 1991, Key Largo

was the only island in the upper Keys to contain hammocks >20 ha. While this remained true for

2004, the number of hammocks >20 ha went from 2-19 patches revealing fragmentation of large

tracts of hammock. This means that extremely large hammocks that fell within the >20ha size

category in 1991, were fragmented but were still, as of 2004, >20ha in size. The greatest change

in the 13 years took place in hammocks from 0-1 ha with an increase from 4-53 patches.

Between 1991 and 2004, all six size categories saw an increase in the number of hammock

patches, revealing fragmentation of hammock in the upper Keys since 1991. Historical estimates

of the number of hammock fragments and their size range was not available for the lower Keys









but as of 2004, 1 11 patches remained extant with the most (28 patches, 25.23%) occurring in the

>20.0 ha range and the least (13 patches; 11.71%) occurring in the 5-10 ha range.

Bird Surveys

During March-May 2004 and 2005, I conducted a total of 858 point count surveys over 86

point count locations. Survey locations in protected hammocks ranged from 66 100%

hammock cover (mean = 96%, sd = 8.4%, median = 100%) within the 20-m radius of the point

count survey location, with deviations from 100% being primarily due to cleared trails. Areas

zoned as residential were highly variable in nature and included hammock habitat in

undeveloped lots at one extreme, to highly manicured lawns, gardens, and cleared areas adj acent

to homes at the other extreme. The distance from the 86 survey locations to the nearest

hammock varied from 0-416 m; the size of those hammocks varied from <0. 1-87 ha.

Seventy-eight species were observed over all 86 survey locations during March-May of

2004 and 2005; these species totaled 6,400 individuals and represented three life history

categories and 11 foraging guilds (Table 3-3). Eleven species of waterbirds and domestic birds

were removed from the analysis. Thirty-five passerines were detected on fewer than 11 separate

occasions; consequently, these data were not sufficient to perform statistical analyses. Table 3-4

reveals which analyses were performed on each of the 32 remaining species and the results of the

analyses. One species detected, but not included in analyses due to lack of sufficient data was

the Brown-headed Cowbird (M~olothrus ater, BHCO), an obligate brood parasite. Many birds

commonly seen in urban environments were detected during the study including the

opportunistic nest predator, the Blue Jay (Cyan2ocitta cristata, BLJA) and two exotic species, the

Eurasian Collared-dove (Streptopelia decaocto; ECDO) and the European Starling (Sturnus

vulgaris, EUST). Five species included in these analyses were designated as Birds of

Conservation Concern by the US Fish and Wildlife Service Division of Migratory Bird










Management (U. S. Fish and Wildlife Service 2002). These included the BWVI, Common

Ground- Dove (Columbina passerine, COGD), MACU, Prairie Warbler (Dendroica discolor,

PRAW) and the state listed, threatened WCPI.

The percent cover of native hammock habitat (PCTHAM) was a significant predictor of

the presence of 15 species (Table 3-4). Among Neotropical migrants, PCTHAM showed a

positive predictive effect on three species and a negative effect on one species. Among migratory

breeders, effects were positive on two species including PRAW, a bird of conservation concern

and negative on one species. Among year-round residents, PCTHAM had positive predictive

effects on one species and negative effects on seven, including non-native ECDO and EUST.

The percent cover of native hammock tree canopy with cleared understory (PCT_CAN)

had significant predictive effects on 10 species. Neotropical migrants had three species with

positive and zero with negative response to this habitat variable. Migratory breeders had two

species that demonstrated positive responses, including the PRAW, and no negative responses.

Year-round residents had zero species with positive responses to this variable and five with

negative responses, including the ECDO and EUST.

Percent ground cover (PCT_GROUND) significantly influenced seven species.

Neotropical migrants had one species demonstrate a positive response and zero species with

negative responses to this variable. Migratory breeders included three species with positive

responses, including PRAW and BWVI, which are birds of conservation concern. One species of

migratory breeder demonstrated a negative response, the MACU, also a bird of conservation

concern. Year-round residents included one species that responded positively and one species

that responded negatively.










The percentage of other vegetation (PCT_OTHER; mangroves, pine habitat) showed

positive predictive effects on three species and negative predictive effects on one species.

Positive effects included one migratory breeder (PRAW), and two year-round residents,

including the exotic EUST. One year-round resident species responded negatively to

PCT OTHER.

Distance from survey locations to the nearest hammock (D2HAM) showed two positive

and two negative responses, all of which occurred among year-round resident species. The size

of the nearest hammock (SIZEHAM) showed significant effects on 10 species. Among

Neotropical migrants two species responded positively and one responded negatively. Migratory

breeders included two positive responses including the BWVI, and two negative responses

including the PRAW. Year-round residents had two species with negative responses, including

the non-native ECDO.

The combined variable HAM+CAN had significant effects on 19 species, the most of any

habitat variable. Among Neotropical migrants three species responded positively and two species

responded negatively. Migratory breeders included three positive and three negative responses.

Two birds of conservation concern, the PRAW and BWVI responded positively. Year-round

residents had one positive and seven negative responses, including the exotic ECDO and EUST,

both of which responded negatively. The other combined variable, DXSIZE, had little effect on

species. Among year-round residents one species responded positively and one negatively to this

variable.

When species were collectively analyzed based on life history and foraging guild (Table 3-

5), PCTHAM showed a negative predictive effect on one life history category (year-round

residents) and two guild categories (bark gleaners and ground gleaners), and a positive effect on









the two migratory life history categories (Neotropical migrants and migratory breeders) and one

guild (foliage gleaners). PCT_CAN also showed a negative predictive effect on year-round

residents and positive for the migratory life history categories. For guilds, PCT_CAN influenced

bark gleaners and ground gleaners negatively, and hover gleaners positively. PCT_GROUND

was a positive predictor only for the migratory breeder life history category and PCT_OTHER

had no significant influence on any life history category or guild. D2HAM showed negative

predictive effects on three guilds (bark gleaners, foliage gleaners, and hover gleaners) and no

predictive effects on life history categories. SIZEHAM showed negative predictive effects on

two life histories (Neotropical migratory birds and year-round residents). Additionally,

SIZEHAM showed negative effects on two guilds (ground gleaners and hawkers) and positive

effects on two guilds (foliage gleaners and hover gleaners). Similar to the individual species

analyses, the combined HAM+CAN variable influenced the greatest number of groups. It

showed negative predictive effects on one life history (year-round residents) and three guilds

(bark gleaners, ground gleaners, and hawkers) and positive effects on two life histories (both

migratory categories) and two guilds (foliage gleaners and hover gleaners). The combined

habitat variable DX SIZE showed no predictive effects on any of the groups analyzed.

Species richness (R2 = 0. 14, p = 0.04) and species diversity (R2 = 0.20, p < 0.01) for 67

species observed at survey locations (all species observed with the exception of the waterbirds

and domestic species) were significantly influenced by SIZEHAM. However, when birds of

conservation concern (R2 = 0. 10, p = 0. 13) or migratory species (R2 = 0.09, p = 0. 16) were

analyzed separately, or together (R2 = 0. 11, p = 0. 16), there was no relationship between

hammock size and presence.









I recorded 35 species that could not be analyzed statistically due to lack of sufficient data.

Of these, 3 1 (88%) were Neotropical migrants, two (6%) were migratory breeders, and two (6%),

were year-round residents (Table 3-3). The vast maj ority (94%), therefore, of rarely observed

birds were migratory. These species represented seven different foraging guilds, of which 40%

were foliage gleaners, 20% ground gleaners, and the remaining 40% fairly evenly divided among

five other foraging guilds. Of the rarely observed birds, eight (23%) were detected in locations

with >50% hammock cover (PCTHAM) and five of those were detected at locations with 100%

hammock cover (Table 3-6). These species included six Neotropical migrants, one migratory

breeder, and one year-round resident, none of which are identified as birds of conservation

concern. The combined variable total canopy cover (HAM+CAN) had similar results. Eleven

species (32%) were detected at locations with >50% total tree canopy cover, five of which were

detected at locations with 100% total tree canopy. These included nine Neotropical migrants, one

migratory breeder, and one year-round resident. None of these species were identified as birds of

conservation concern. Regarding guilds, these species represented two species of ground

gleaners, four foliage gleaners, one bark gleaner, three aerial foragers, and one hover gleaner.










Table 3-1 Total amount (ha) of hammock habitat including percent of total in protected status by island in the Florida Keys as of
2004.
Hammock area Hammock patches
Island No. Mean patch Median patch Range patch
total (ha) conserved (%) patches size (ha) SD size (ha) size (ha)


I


Upper Keysa
Key Largob
Plantation Key
Windley Key
Upper Matecumbe Key
Lower Matecumbe Key
Long Key
Upper Keys total


1834.2
47.8
12.4
54.2
6.3
6.7
1961.6


56
26
86
17
61
87
54

0
39
0
90
85
98
0
28
100
100
22
0
24
73
63
b includes Ragged Keys.


45
32
4
36
3
4
124

8
20
1
17
16
8
7
1
1
2
3
11
7
102
226


40.5
1.5
3.1
1.5
2.1
1.7
15.8

1.2
4.6
2.2
29.5
35.3
32.3
22.7
44.7
23.4
7.8
4.6
3.2
4.2
17.2
16.4


59.1
1.5
5.1
1.4
0.6
1.5
40.1

0.7
5.8

33.0
30.0
37.9
19.4


1.1
0.5
3.0
4.9
25.2
34.2


10.3
1.0
0.7
0.8
2.40
1.1
1.5

1.2
2.3
2.2
11.3
23.3
10.6
14.5
44.7
23.4
7.7
4.4
2.6
2.4
4.4
2.7


0.1-205.7
0.2-4.8
0.3-10.7
0.3-4.7
1.1-2.4
0.61 1.35
0.1-205.7

0.4-2.4
0.3-21.1
2.2-2.2
0.4-94.9
0.3-85.8
2.5-96.3
1.3-49.9
44.7-44.7
23.4-23.4
7.0-8.5
4.2-5.2
0.5-11.6
1.3-15.2
0.3-96.3
0.1-205.7


Lower Keys
Grassy Key 9.6
Vaca Key 91.5
Ohio Key 2.2
Big Pine Key 501.8
Middle Torch Key 564.8
Big Torch Key 258.0
Ramrod 159.2
Little Knockemdown Key 44.7
Knockemdown Key 23.4
Toptree Hammock Key 15.5
Cudjoe Key 13.8
SugarloafKey 35.8
Lower Sugarloaf Key 29.63
Lower Keys total 1749.9
Grand total 3711.5
a Delineation of upper Keys from Bancroft et al. (1995).











Table 3-2 Partially protected hammock habitat patches in the Florida Keys including total number of patches and area (ha) in
protected and unprotected status by island and patch size category (ha) of 2004 (islands that did not contain partially
protected patches were excluded from this table).
Hammock Patches by Size (ha) Category
Total
Island no.
0-1 1-5 5-10 10-20 >20
patches
#a Pb UC #o Pb Uc #o Pb Uc #o Pb Uc #o Pb Uc

Upper Keysd
Key Largoe 20 3 0.7 0.3 2 3.4 4.1 1 5.2 2.2 2 18.3 13.7 12 812.0 607.4
Plantation Key 3 3 4.4 3.0
Upper Matecumbe 2 2 0.7 0.6
Key
Upper Keys total 25 5 1.4 0.9 5 7.8 7.1 1 5.2 2.2 2 18.3 13.7 12 812.0 607.4
Lower Keys
Vaca Key 1 1 4.6 8.9
Big Pine Key 4 4 275.3 33.9
Middle Torch Key 9 2 1.0 3.00 2 26.5 5.3 5 113.3 76.3
Little Knockemdown
1 1 12.4 32.3
Key
Cudjoe Key 1 1 3.0 2.2
Lower Sugarloaf Key 2 2 6.0 0.6
Lower Keys total 18 4 7.0 3.6 1 3.0 2.2 3 31.1 14.2 10 401.0 142.5
1213.
Grand total 43 5 1.4 0.9 9 14.8 10.6 2 8.2 4.4 5 49.4 27.9 22 749.9

aNumber of partially protected hammock patches per size category. Amount (ha) of protected hammock habitat by patch size category. c Amount (ha) of
unprotected hammock habitat by patch size category. dDelineation of upper Keys from Bancroft et al. (1995). e includes the Ragged Keys.










Table 3-3 Avian species detected during point count surveys including standard abbreviations (USGS Patuxent Wildlife Research
Center Bird Banding Laboratory), common and scientific names, life history (Rappole 1995) and breeding status (Florida
Fish and Wildlife Conservation Commission 2003), and foraging guild designations (Ehrlich et al. 1998; see Table 2-3 for
definitions of life histories and foraging guilds). Frequency includes number of group observations (n) for each species and
the % of total group observations among species. Abundance includes number of individual birds recorded (n) for each
species and the % of total birds among species. Domestic and waterbirds were excluded from analyses.

Species Life Foraging Frequency Abundance
Common Name Scientific Name
Code History Guild (n) (%) (n) (%/)


GRCA
YPWA
BAWW
AMRE
REVI
OVEN
; BLPW
BGGN
COYE
EAKI
BTBW
INBU
CMWA
WEWAa
WWDO~a
MAWAa
YTWAa
BHCOa


Gray Catbird
Palm Warbler
Black-and-white Warbler
American Redstart
Red-eyed Vireo
Ovenbird
Blackpoll Warbler
Blue-gray Gnatcatcher
Common Yellowthroat
Eastern Kingbird
Black-throated Blue Warbler
Indigo Bunting
Cape May Warbler
Worm-eating Warbler
White-winged Dove
Magnolia Warbler
Yellow-throated Warbler
Brown-headed Cowbird


Dumetella carolinensis
Dendroica palmarum
M~niotilta varia
Setophaga ruticilla
Vireo olivaceus
Seiurus aurocapillus
Dendroica striata
Polioptila caerulea
Gl'lthy/pi\ trycha~s
Tyrannus tyrannus
Dendroica caerulescens
Pa~sserina cyan2ea
Dendroica tigrina
Helmitheros vermivorus
Zenaida a~siatica
Dendroica magnolia
Dendroica dominica


NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM


GG
GG
BG
HG
HG
GG
FG
FG
FG
HA
HG
FG
FG
FG
GG
HG
BG
GG


8 1 14 0
7 1 31 0


M~olothrus ater NM











Table 3-3 (continued)


Species
Code
BLBWa"
HOWAa
PABUa
YBCUa
YTVIa
AMKEa
MERLa
PIWAa
PROWa
SWWAa
EAPHa
MYWAa
NAWAa
RTHUa

SUTAa
SCTAa
WOTHa
NRWSo


Common Name

Blackburnian Warbler
Hooded Warbler
Painted Bunting
Yellow-billed Cuckoo
Yellow-throated Vireo
American Kestrel
Merlin
Pine Warbler
Prothonotary Warbler
Swainson's Warbler
Eastern Phoebe
Yellow-rumped Warbler
Nashville Warbler

Ruby-throated
Hummingbird
Summer Tanager
Scarlet Tanager
Wood Thrush
Northern Rough-winged
Swallow
Rose-breasted Grosbeak


Scientific Name

Dendroica fusca
Wilsonia citrina
Pa~sserina ciris

Coccyzus amnericanus
Vireo flavifrons
Falco sparverius
Falco sparverius
Dend oica pinus
Prothonotaria citrea
Liinnothy/pisl~i swainsonii
Sayornis phoebe
Dendroica coronata
Vermivora ruficapilla
Archilochus cohubris

Piranga olivacea
Piranga olivacea
Hylocichla mustelina
Stelgidoptelyx serripennis


Life Foraging Frequency


Abundance
(n) (%/)
15 0
6 0
4 0
4 0
6 0
2 0
2 0
2 0
2 0
2 0
3 0
3 0
4 0
4 0


History
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM

NM
NM
NM
NM


Guild
FG
FG
GG
FG
FG
HO
HA
BG
BG
GG
HA
FG
FG
HG


(%)
0
0
0
0
0
0
0
0
0
0
0
0
0
0


RBGRa


Pheucticus hidovicianus NM


FG 1 0 1 0










Table 3-3 (continued).


Species
Code
RCKla
TRESa
YBCHa
YB SAa
BLGRa
NOPAa
CEDWa
MODO
GCFL
GRAK
w PRAW
WEVI
WCPI^
BWVI
YSFL
COGD
MACU
CONIa
YWARa
NOCA
RBWO
ECDO*
COGR
BLJA


Life
History
NM
NM
NM
NM
NM
NM
NM
MB
MB
MB
MB
MB
MB
MB
MB
MB
MB
MB
MB
YR
YR
YR
YR
YR


Foraging
Guild
FG
AF
FG
BG
GG
FG
FG
GG
HA
HA
FG
FG
FG
FG
GG
GG
FG
AF
FG
GG
BG
GG
GG
GG


Frequency
(n) (%/)


Abundance


Common Name

Ruby-crowned Kinglet
Tree Swallow
Yellow-breasted Chat
Yellow-bellied Sapsucker
Blue Grosbeak
Northern Parula
Cedar Waxwing
Mourning Dove
Great-crested Flycatcher
Gray Kingbird
Prairie Warbler
White-eyed Vireo
White-crowned Pigeon
Black-whiskered Vireo
Northern Flicker
Common Ground- dove
Mangrove Cuckoo
Common Nighthawk
Yellow Warbler
Northern Cardinal
Red-bellied Woodpecker
Eurasian Collared-dove
Common Grackle
Blue Jay


Scientific Name

Regulus calendula
Tachycineta bicolor
Wilsonia citrina

Sphyrapicus varius
Guiraca caerulea
Parula amnericana

Bombycilla cedrorum
Zenaida macroura
M~yiarchus crinitus
Tyrannus dominicensis
Dendroica discolor
Vireo griseus
Patagioena~s leucocephala
Vireo altiloquus
Colaptes auratus
Columbina passerina
Coccyzus minor
Chordeiles minor
Dendroica petechia
Cardinalis cardinalis
M~elan2erpes carohinus
Streptopelia decaocto
Quiscalus quiscula
Cyanocitta cristata


(n)
1
1
1
1
2
4
45
185
133
87
66
143
53
63
15
16
13
1
1
962
533
1097
461
310


(%)
0
0
0
0
0
0
1
3
2
1
1
2
1
1
0
0
0
0
0
15
9
18
7
5










Table 3-3 (continued).


* Non-native Species. ^ State listed species Threatened. a species sited in insufficient numbers for statistical analyses. species reported not to occur in the
Florida Keys accidental or out of range sighting (Florida Fish and Wildlife Conservation Commission 2003). c removed from analysis domestic species or
waterbird species uncommon in hammocks. d data unavailable.


Species
Code
RWBL
NOMO
EUST*
RSHA
CARWa
MOPA~a


BLBU bc
COTE c
DCCO c
GHOWbc
GREGc
LAGU c
OSPR c




WHIB c


Life
History
YR
YR
YR
YR
YR
YR
YR


NM
MB



MB
MB
MB
d


MB


Foraging
Guild
GG
GG
GG
SW
GG






HD
SD
SW
SS
GG
HD
d
d

GG


Frequency
(n) (%/)
49 4
48 4
32 3
20 2
4 0
4 0
1 ----

1 ----
1 ----
1 ----
1 ----
6 ----
7 ----
4 ----
4 ----
4 ----


Abundance
(n) (%/)
206 3
195 3
250 4
38 1
6 0
14 0
3 ----

81 ----
81 ----
1 ----
1 ----
11 ----
15 ----
75 ----
9 ----
11 ----


Common Name

Red-winged Blackbird
Northern Mockingbird
European Starling
Red-shouldered Hawk
Carolina Wren
Monk Parakeet

Unidentified parrot
Blue Bunting
Common Tern
Double-crested Cormorant
Great-horned Owl

Great Egret

Laughing Gull
Osprey
Peafowl
Domestic Chicken
White Ibis


Scientific Name

Agelaius phoeniceus
M~imus polyglottos
Sturnus vulgaris
Buteo lineatus
Thr~ i yesthesiitludovicianus
M~yiopsitta monachus
Amazonia sp.

Cyan2ocompsa parellina
Sterna hirundo
Phalacrocorax auritus

Bubo virginianus
Ardea alba
Larus atricilla
Pandion haliaetus
Pavo cristatus
Gallus domesticus
Eudocimus albus











Table 3-4 Influence of measured habitat variables on species presence or absence from analyses with general linear models (data
represent p-values). A negative p-value indicates an inverse relationship between the habitat variable and species presence.
See Table 3-3 for species codes, Table 2-3 for definition of life history and foraging guild, and Table 2-2 for definition of
habitat parameters.
Statistical Species Life Foraging PCT HAMb PCT CANb PTPTb D2HAMb SIZEHAMb HM cAN DXSIZEc
Models Code History Guild --GROUND" OTHER"
ZIP RSHA YR SW <0.02
ZIP AMRE NM HG 0.05 <0.02 <0.05
ZIP REVI NM HG <0.01 <0.01
ZIP BTBW NM HG 0.05
ZIP EAKI NM HA (-) <0.02 ( -) <0.02
ZIP GCFL MB HA <0.03 <0.04 (-) <0.01
ZIP GRAK MB HA (-) <0.03 (-) 0.001
ZIP ECDO YR GG (-) <0.01 (-) <0.01 (-) <0.01 (-) <0.01
ZIP NOCA YR GG 0.05 <0.01 <0.01
ZIP COGR YR GG (-) <0.01 (-) <0.01 (-) <0.01
ZIP BLJA YR GG (-) <0.01 (-) <0.04 (-) 0.05 (-) <0.01 (-) <0.04
ZIP EUST YR GG (-) <0.01 (-) <0.01 0.05 (-) <0.01
ZIP RWBL YR GG (-) <0.02 (-) <0.01 <0.01 (-) <0.01 <0.01
ZIP NOMO YR GG (-) <0.01 <0.05 (-) <0.01
ZIP YPWA NM GG <0.01 <0.01 <0.01
ZIP GRCA NM GG <0.05 (-) <0.02
ZIP OVEN NM GG
ZIP MODO MB GG (-) <0.02 (-) <0.01
ZIP COGD MB GG
ZIP BLPW NM FG <0.04
ZIP COYE NM FG
ZIP CMWA NM FG
ZIP WEVI MB FG <0.03 <0.01 <0.01





" Statistical model used to analyze each species. ZIP = zero-inflated Poisson general linear model; GLIMIX = general linear model. b Variables analyzed as
covariates in first set of analyses. Variables analyzed as covariates in second set of analyses.



Table 3-5 Influence of measured habitat variables on life history and foraging guilds from general linear model analyses with zero-
inflated Poisson regression model (data represent p-values). A negative p-value indicates an inverse relationship between
the habitat variable and species presence. See Table 3-3 for species codes, Table 2-3 for definition of life history and
foraging guild, and Table 2-2 for definition of habitat parameters.
PCT PCT
PCT HAM" PCT CAN" a D2HAM" SIZEHAM" HAM+CANb DXSIZEb
-- GROUND" OTHER"
c YR (-) <0.01 (-) <0.01 (-) 0.02 (-) <0.01
-- NM <0.01 <0.01 (-) <0.01 <0.01
MB <0.01 <0.01 <0.03 <0.01
SBG (-) <0.01 (-) <0.04 (-) <0.02 (-) <0.01
SFG <0.01 (-) <0.02 <0.02 <0.01
GG (-) <0.01 (-) <0.01 (-) <0.01 (-) <0.01
HA (-) <0.04 (-) <0.01
HG <0.02 <0.04 (-) <0.04 <0.02 <0.01
SSW
aVariables analyzed as covariates in first set of analyses. Variables analyzed as covariates in second set of analyses


Table 3-4 (continued)
Statistical Species
Models Code
ZIP PRAW


Life
History
MB


Foraging PCT HAMb PCT CANb
Guild --
FG <0.01 <0.02


PCT_
GROUND
<0.01
<0.01
(-) <0.04


PCTb
OTHER~


D2HAMb SIZEHAMb HM cAN DXSIZEc


(-) 0.02


<0.01
<0.01


ZIP BWVI MB FG
ZIP MACU MB FG
ZIP RBWO YR BG (-) <0.01 (-) <0.01
ZIP BAWW NM BG


(-) <0.03 (-) <0.02 (-) <0.01


(-) <0.01


GLIMMIX YSFL
GLIMMIX BGGN
GLIMMIX INBU
GLIMMIX WCPI


<0.02
(-) <0.02


MB FG










Table 3-6 Hammock habitat variables for birds that lacked sufficient data for statistical analyses. Data reflect the range of percent
cover (smallest to largest) for PCTHAM, PCT_CAN and HAM+CAN measurements where birds were observed. See
Table 3-3 for species codes, Table 2-3 for definition of life history and foraging guild, and Table 2-2 for definition of
habitat parameters.
Survey Locations
Species Life Foraging PCT HAM PCT CAN HAM+CAN
TOTAL at which detected--
Code History Guild smallest largest smallest largest smallest largest

CEDW 45 1.16 NM FG 0 0 10.8 10.8 10.8 10.8
MAWA 31 8.14 NM HG 0 95.4 0 18.6 0 100
BLBW 15 5.81 NM FG 0 100 0 16.5 5.4 100
WWDO 14 9.30 NM GG 0 52.0 10.8 42.2 30.9 67.3
MOPA 14 4.65 YR ------ 0 29.7 0 37.6 0 67.4
WEWA 10 9.30 NM FG 6.0 100 0 6.7 12.7 100
BHCO 9 5.81 NM GG 0 0 0 25.1 0 25.2
YTWA 9 6.98 NM BG 0 100 0 46.5 0 100
CARW 6 4.65 YR GG 89.9 100 0 0 89.9 100
0 YTVI 6 3.49 NM FG 0 100 0 25.2 25.2 100
HOWA 6 4.65 NM FG 0 100 0 15.0 15.0 100
SCTA 5 2.33 NM HG 38.7 100 0 25.2 64.0 100
WOTH 5 2.33 NM GG 0 92.5 0 0 0 92.5
YBCU 4 3.49 NM FG 100 100 0 0 100 100
SUTA 4 2.33 NM FG 8.3 92.5 0 65.3 73.6 92.5
NOPA 4 1.16 NM FG 35.1 35.0 7.7 7.7 42.8 42.8
NAWA 4 2.33 NM FG 0 100 0 28.5 28.5 100
PABU 4 3.49 NM GG 0 100 0 18.6 18.6 100
RTHU 4 2.33 NM HG 0 0 5.4 38.2 5.4 38.2
MYWA 3 2.33 NM FG 0 100 0 21.0 21.0 100
EAPH 3 2.33 NM HA 0 100 0 0 0 100
SWWA 2 2.33 NM GG 94.4 100 0 0 94.4 100
PROW 2 2.33 NM BG 0 92.5 0 35.8 35.8 92.5
AMKE 2 2.33 NM HO 5.1 38.7 16.5 25.2 21.5 64.0
MERL 2 2.33 NM HA 0 0.6 13.0 35.8 13.5 35.8










Table 3-6 (continued)


Survey Locations
TOTAL at which detected


Species
Code


BLGR
CONI
RCKI
YBCH
YB SA
TRES
NRWS
YWAR
RBGR


Life
History


NM
MB
NM
NM
NM
NM
NM
MB
NM


Foraging
Guild


GG
AF
FG
FG
BG
AF
AF
FG
FG


PCT HAM
smallest largest


PCT CAN
smallest largest


HAM+CAN
smallest largest


4.9
0
0
0
0
0
30.6
6.2
0
28.5


8.0
0
100
100
100
100
77.3
70.8
35.2
28.5


100
0
100
100
100
100
77.3
70.8
35.2
28.5


1.16
1.16
1.16
1.16
1.16
1.16
1.16
1.16
1.16


0
100
100
100
100
46.6
64.6
35.2
0


0
100
100
100
100
46.6
64.6
35.2
0


0
0
0
0
0
30.6
6.2
0
28.5




















2000-

1800 -1 I Fully protected hammock patches
1 "I Partially protected hammock patches
1600 I Unprotected hammock patches
1400

1200
1016
1000




200-
47

8 2 9 65 6
61 15 9

0

0-1 1-5 5-10 10-20 >20

Hammock patch size (ha)




Figure 3-1 Area (ha) of tropical hardwood hammock habitat patches by patch size (ha) category
and protected status in the Florida Keys in 2004 (number of habitat patches provided
above each bar).









CHAPTER 4
DISCUSSION

Assessment of Hammock Habitat

Almost 3 1% of tropical hardwood hammock habitat in the upper Florida Keys was lost to

deforestation during 1991-2004. Of the remaining 1,962 ha of hammock in the upper Keys,

approximately 46% lacked protection. Of the 1,750 ha of hammock left in the lower Keys as of

2004, approximately 27% lacked protection. Therefore, of the 3,712 ha of extant hammock in

the Florida Keys, 1,375 ha (36.5%) remains unprotected and subj ect to development.

Partially protected patches of hammock exist in both the upper and lower Keys. These

patches present opportunities to increase the size of existing conservation areas. In fact, the 22

partially protected patches >20 ha encompass 750 ha of unprotected habitat, which represents

55% of all remaining unprotected hammock habitat in the Keys. If all of the unprotected portions

of these partially protected patches are lost to development, mean patch size would decrease

from 89 to 55 ha (Table 3-2).

The increasingly patchy hammocks have been and continue to be fragmented by

anthropogenic activities, the most conspicuous of which is the construction of roads. The

Overseas Highway (US Route 1) follows a trail originally formed by Henry Flagler's Florida

East Coast Railroad and stretches from Miami to Key West. The highway was completed in

1938 and today spans 182 km including 42 bridges to allow motorists access from one island to

the next. This road and the secondary roads that result from increasing urbanization are a maj or

contributor to the fragmentation of hammocks in the Keys. Although no specific studies have

been done targeting the effects of roads on hammocks in the Florida Keys, the impact of roads

has been studied extensively by a number of authors in various other systems (Forman et al.

2003).












The presence of a road in, around, or through a habitat has both direct and indirect impacts.

Direct impacts include the subdividing of natural areas into smaller habitat blocks, the

destruction and alteration of habitats, disturbance, edge effects, invasions by exotic species and,

increased roadkill (Goosem 1997; Forman & Alexander 1998). Roads have also been implicated

in attracting generalist predators (Forman et al. 2003) and qualitative observations of feral cats

(Felis catus) during bird surveys suggested a positive relationship between road density and feral

cat abundance in the Keys (A. Karim, personal obs.).

The natural island/water landscape of the archipelago lends itself to hammock habitats that

are naturally patchy in comparison to other tropical forests. The construction of roads through

these areas further increases the edge effects impacting these seasonal deciduous forests.

Consequently, resident and migratory birds that rely on these areas must contend with increasing

amounts of disturbance and other road-related impacts. Reijnen et al. (1995) found that 60% of

the 43 species of woodland species of birds that they investigated showed evidence of reduced

densities adj acent to roads. Goosem (1997) found that forest interior birds in the Amazonian

rainforest stay at least 50 m away from edges. In the Keys, 50 m is often more than half the

width of an entire hammock habitat block, which emphasizes the importance of conserving

extant large blocks.

Bird Surveys

Species specific analyses were the best way to determine habitat use by birds detected

during the study (Table 3-4). Grouping avian species by life history helped summarize the

predictive effects of some habitat variables on year-round residents, but did a poor j ob in

summarizing predictive effects on Neotropical migrants and migratory breeders (Table 3-5).

Similarly, grouping avian species by foraging guild was a poor predictor of how individual










species within those guilds responded to habitat parameters with the exception of the bark

gleaner guild. The lack of significant predictive effects of hammock variables on Neotropical

migrants and migratory breeders as collective groups is likely due in part due to the fact that

migratory species represent many different foraging guilds with unique needs and habitat

preferences. However, the lack of significant predictive effects may also be due to the fact that

many migratory birds that depend on hammocks during migration are difficult to detect because

they are in the Keys for short periods and may be difficult to detect in heavy cover typical of

hammock habitat. For example of the 35 species seen infrequently, 94% were migratory and 11

of these species were detected at locations with >50% total tree canopy cover. These analyses

suggest that when implementing conservation and management plans for resident birds (from

those examined here), grouping them together may be helpful in predicting their occupancy

within a landscape, but that implementing plans for migrants should be done on a species

specific basis.

Of the 32 species detected in sufficient numbers to be analyzed statistically, five are

designated as birds of conservation concern (BWVI, PRAW, MACU, WCPI and COGD), one is

an opportunistic nest predator (BLJA), and two are non-native (ECDO and EUST; Table 3-3).

Although the BWVI showed a significant negative response to PCT_GROUND and no positive

effects to any other habitat parameter, it has been noted that hammocks (Robertson & Kushlan

1984; Snyder et al. 1990) in addition to mangrove areas (Ehrlich et al.1988) serve as important

breeding areas. The BWVI was only detected within hammocks during this study regardless of

hammock location (residential or protected). This suggests that this species prefers hammocks to

lawns or open areas, despite a lack of significant positive response to hammock habitat

parameters.









The PRAW is usually associated with pinelands and open Hields (Ehrlich 1988; Elphick

2001) but analyses revealed that this species also responded positively to the PCTHAM,

PCT_CAN, PCT_GROUND, and HAM+CAN habitat parameters (and negatively to the

SIZEHAM variable). Both the BWVI and PRAW have experienced population declines due to

brood parasitism from the Shiny Cowbird (M~olothrus bonariensis) and the BHCO (Ehrlich 1988;

Wiley 1988; Post et al. 1990; Grzybowski 2001). Both cowbird species have been reported in the

Florida Keys (Smith & Sprunt 1987; US Fish and Wildlife Service 1999, Sibley 2000, A. Karim

personal obs.), and have been implicated in the decline of many Neotropical migratory songbirds

(Terbough 1989; Robinson et al. 1993). Specifically, the BHCO is suspected of causing the near

extirpation of the BWVI in the Tampa Bay area of Florida (Paul 1987; Paul 1988). Historically,

cowbirds were largely restricted to the mid-continental prairies where they apparently followed

herds of nomadic bison. Cowbirds mainly search for seeds and insects in short grass and on

bare ground and may have depended upon grazing by large ungulates to create suitable feeding

areas. However, since the clearing of forests for agriculture and the widespread introduction of

livestock, cowbirds have expanded their geographical range (Mayfield 1965). Additionally,

cowbirds target species that build open-cup nests (Robinson et al. 1993), which are characteristic

of the BWVI and PRAW (Ehrlich 1988). The tropical hardwood hammock multi-species

recovery plan (US Fish and Wildlife Service 1999) states that "hammocks are important for a

number of West Indian land birds including the MACU, BWVI and WCPI". The analyses

performed in this study indicated that the MACU showed a negative predictive response to

PCT_GROUND, but it did not show a significant positive response to any other habitat

parameter. Nonetheless, MACUs were only detected within protected hammocks with no urban

matrix during this study. This suggests that hammocks and native tree canopy associated with









hammocks should be considered when devising conservation plans for both the PRAW and

MACU.

The state threatened WCPI showed no negative or positive predictive effects when

modeled against the habitat variables measured within this study. While WCPI nest on small

mangrove islands, these obligate frugivores feed within native hammocks and are arguably the

most important seed dispersers of 37 species of trees and shrubs within hammocks in the Keys

(Strong and Bancroft 1994). In particular, the fruits of the blolly (Guapira discolor), strangler

fig (Ficus ureaa, and especially poisonwood (M~etopium toxiferum) are important to the survival

of this species (Bancroft & Bowman 1988; Bancroft et al. 2000). However, the strangler fig is

often removed from residential areas due to its extensive root system that interferes with septic

systems and poisonwood is removed because its sap causes contact dermatitis in some humans

(Scurlock 1987). Bancroft and Bowman (1988) state that, "breeding populations [of WCPI] are

clearly tied to the existence of sufficient hammocks for production of food". Post fledging

dispersal studies of this species show that WCPIs selectively use hammock fragments from 5-20

ha (p < 0.10) during the first 72 hours after nest dispersal; after this stage, dispersing birds show

no preference for fragment size but use hammocks more frequently than residential habitat (p <

0.10). Therefore, small hammocks are used as "stepping stones" by immature WCPIs that are

unable to fly long distances when they first leave mangrove islands to forage for food (Strong

and Bancroft 1994).

The BLJA is noted as a potentially important nest predator on open-nesting songbirds in

fragmented habitat (Wilcove 1985). Although a native bird to the Florida mainland, this species

is a fairly recent arrival to the Keys. It was first found to be breeding in the upper Keys in 1989

(Florida Fish and Wildlife Conservation Commission 2003), and has been increasing since (J.










Duquesnel, personal comm.). Engels and Sexton (1994) show a strong negative correlation

between the presence of BLJAs and the endangered Golden-cheeked Warbler (Dendroica

chrysoparia) in central Texas. During my study, the BLJA showed a negative response to three

hammock variables (PCTHAM, SIZHAM and HAM+CAN). Of the 310 BLJAs recorded, 28

were observed within hammocks and BLJAs were detected in only five of 14 hammocks

surveyed, all of which were small fragments. This suggests that this opportunistic nest predator

and typical edge species may avoid large, intact hammocks and prefer highly fragmented urban

areas.

Two non-native species were observed that deserve mention. The ECDO is noteworthy

primarily because it was the most abundant species recorded during surveys, but showed a

significant negative response to hammock and total tree canopy variables (Table 3-3). The EUST

is an aggressive competitor for cavity nesting sites and as such may be detrimental to resident

and migrant cavity nesters in the Keys. Ingold' s (1994) research in east-central Ohio shows that

Red-bellied Woodpeckers (M~elan2erpes carolinus, RBWO) "incurred the brunt of [EUST]

competition for freshly excavated nest cavities and lost 39% of their cavities to starlings. Flickers

...were significantly more aggressive than RBWOs when defending their nest cavities. Fourteen

percent of flicker cavities...were usurped by starlings." While conducting research, I observed

EUSTs competing with one RBWO pair and two pairs of Northern Flickers (Colaptes auratus)

for nest cavities. Statistical analyses suggest that the EUST did avoid hammocks; this species

showed a negative response to three hammock variables (PCTHAM, PCT_CAN and

HAM+CAN). Consequently, intact hammock habitat may provide a refuge for cavity nesting

species in the Keys.









In addition to cowbird species (M~olothrus spp.), BLJAs, and EUSTs, feral cats may pose a

substantial threat to resident and migratory birds in hammock fragments. Feral cats were

observed at 46 (54%) of survey locations and in areas that ranged from 0-100% hammock cover.

Cats were observed stalking and/or attacking resident and migratory birds. The number of feral

cats in the Keys is increasing in all probability because of lack of predators and food provided by

visitors and inhabitants of the archipelago. The Ocean Reef Club in north Key Largo has been

sustaining a colony of approximately 500 feral cats at an annual budget of approximately

$100,000 that includes a dedicated veterinarian (America Bird Conservancy 2004). In addition

to birds, feral cats prey on other species and may pose a significant threat to the recovery of two

federally listed, endangered species, the Key Largo cotton mouse (Peromyscus gossypinus

allapaticola) and Key Largo woodrat (Neotoma flori~dd~~ddana~~dd~ smallii), both of which depend on

hammocks for their survival (US Fish and Wildlife Service, 1999).

Further studies are needed in the Keys to add to the knowledge about avian-hammock

interactions. Long term surveys of the number and species of birds that use hammocks during

spring and fall migrations are needed. Comparisons of fall and spring migration will help land

managers get a better picture of the full suite of avian species that rely on hammocks in the Keys.

Studies on the impacts of roads in the Keys are needed to understand how birds and other species

that require these habitats are affected both directly and indirectly.

Conservation Implications

The study of species-environment interactions enables scientists to investigate and

potentially understand how changing landscapes effect the way species are able to respond and

ultimately survive or, alternatively, become extirpated. Bird migration often occurs over a broad

geographic scale but over a relatively short temporal scale (Russell et al. 1998, Simons et al.

2000); therefore, designing and implementing conservation strategies to help ensure the survival









of Neotropical migrants (Maurer and Heywood 1993, Keyser et al. 1998) is complicated. Since

energy demands during migrations are extreme, migrants must accumulate substantial fat stores

to fuel their j ourney. The obj ective of migrants is not only to accumulate enough fat to survive

the migration, but additionally, to arrive at a breeding ground with enough fat stores to compete

for nesting areas and therefore aid in the chance of increased population survival (Smith &

Moore 2003). Schaub and Jenni (2001) state that "the fuel deposition rate at stopover sites is the

crucial factor determining the overall speed of migration and its success." Parrish (2000)

suggests that "selection for minimization of energy expenditure during stopover has influenced

the evolution of dietary plasticity during migration. Dietary shifts to fruit allow many migrant

species to minimize the time and energy needed for foraging on 'expensive' diet types such as

insects." This "dietary opportunism" allows a migrant to handle unpredictable fluctuations in

food type and availability while en route (Levey & Stiles 1992).

As of 2004 most of the extant hammock habitat was located in the upper Keys. There

remained areas of unprotected and partially protected hammocks throughout the Keys that could

be acquired and incorporated into a network of protected hammocks that run the length of the

Florida Keys. This archipelago-wide network could benefit both resident and migratory birds

that require hammocks to survive. By simply targeting partially protected hammocks for

acquisition, the size of hammock patches >20 ha could be increased by 36% and 79% in the

lower and upper Keys, respectively. These hammocks could be utilized by migrants needing

stopover areas because of inclement weather or low fat reserves after their trans-gulf migration

and provide quality habitats for year-round residents and migratory breeders that require

hammock habitat for reproduction. Additionally, these areas would less likely be problematic









from species such as the BLJA and EUST because these species respond negatively to hammock

habitat (Table 3-4).

Acknowledging the need to acquire areas for conservation is not a sufficient end to

preserving the flora and fauna of natural plant communities; economic considerations must also

be addressed for conservation plans to work effectively. In Florida, eco-tourism contributes

millions of dollars to the state economy. Caudill (2003) reports that visitors and residents of

Florida spent over $1.5 billion dollars in activities associated with wildlife viewing, generated

more than 35,000 jobs, and created more than $76 million in state sales tax revenue in 2001.

Birds such as the BWVI, MACU and WCPI that need hammocks to persist also provide

economic benefit for the Keys. Many bird watchers visit the Florida Keys specifically to see

these native species (A. Karim pers. obs.; D. Sprunt pers. comm.).

Public costs resulting from the construction of residential neighborhoods must also be

taken into consideration. According to the Florida Department of Environmental Protection

(1997), the cost per person of moving to a previously undeveloped area ranged between $4,400

in Ocala, FL and $8,100 in Naples, FL in public expenditures subsidized by taxes paid by

existing residents to the area. In 1997, the estimated public cost for each new single-family home

in Florida ranged from $13,640-$25,110. These publicly-subsidized infrastructure costs increase

with distance to urban service centers (Florida Department of Environmental Protection 1997)

and are likely much higher in the Keys given costs associated with the transport of building

materials, limited sources of fresh water, and issues associated with trash and sewage disposal.

Conservation and management of hammock habitat and the birds associated with them

cannot be accomplished without the education and support of the public. In south Florida,

hammocks are one of the easiest forest communities to create (US Fish and Wildlife Service










1999). Environmental education should go hand-in-hand with management activities to increase

public awareness and participation in conservation activities (Fernandez-Juricic & Jokimaki

2001) such as creation, conservation and restoration of these communities. Balmford and

Cowling (2006) state that ecologists and resource managers need to come to "the realization that

conservation is primarily not about biology but about people and the choices they make." Many

residents of the Keys appreciate the natural environment and public support will be crucial to the

future protection of hammocks. Private landowners with substantial hammock habitat may be

educated on the value of conservation easements and preservation of their land. Providing

educational materials on the ecological importance and economic benefits of hammocks can be

the starting point of a comprehensive approach to hammock conservation throughout the Keys

(Main et al. 2003; Karim & Main 2004).










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BIOGRAPHICAL SKETCH

Annisa Karim was born in Nairobi, Kenya on September 13, 1974. She moved to Florida

with her parents, Muneer and Laila, and her sister Farah, in 1980. Her lifelong curiosity of the

outdoors and her amazement of the natural world led to her to focus on ecology as a career.

Annisa received a Bachelor of Science degree in wildlife ecology and conservation and a minor

in zoology from the University of Florida' s Department of Wildlife Ecology and Conservation in

1997. She worked as a naturalist and then as a research associate in southwest Florida before she

ventured to Gainesville once again. She entered the graduate program at the University of

Florida in the Fall of 2003 and earned a Master of Science degree from the Department of

Wildlife Ecology and Conservation in 2007.





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STATUS AND USE OF TROPICAL HARDWOOD HAMMOCKS AND FORESTED RESIDENTIAL AREAS AS HABITAT FOR RESIDENT AND NEOTROPICAL MIGRATORY BIRDS IN THE FLORIDA KEYS By ANNISA KARIM 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 2007 1

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2007Annisa Karim 2

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In Wildness is the Preservation of the World. -Thoreau/ Porter/ Metallica 3

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ACKNOWLEDGMENTS I would like to express my enormous gratitude to my parents, Muneer and Laila, and to my sister, Farah, for their love and encouragement throughout my life. The advice and insight they have provided and their belief in me has enabled me to achieve my goals and follow my dreams. I am extremely fortunate to have them in my life! SHRUKRAN. I would like to thank the chairperson of my committee, Dr. Martin B. Main for his encouragement, support, guidance and sense of humor all of which enabled me to attain a Master of Science Degree from the University of Floridas Department of Wildlife Ecology and Conservation. His passion and commitment to his work is a true inspiration. I would also like to thank my other committee members, Dr. Mark Hostetler and Dr. Mike Moulton for their advice and expertise. The support staff at the Department of Wildlife Ecology and Conservation deserves a special thank you; their helpfulness, dedication and team-work is commendable. I thank Islamorada Village of Islands for the partial funding of my research. Additionally, I would like to thank the staff and volunteers (especially Donna Sprunt) of the Windley Key Fossil Reef Geological State Park and the staff at the Keys Marine Laboratory for their helpfulness during the fieldwork portion of my research. I would like to express a special thanks to the Hennig FamilyMarkus, Melissa and Robert have been extremely supportive especially during my last semester as a graduate student. I am grateful for their encouragement, friendship and proficiency at making a great cup of coffee (but mostly for their encouragement and friendship). 4

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TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES ...........................................................................................................................6 LIST OF FIGURES .........................................................................................................................7 ABSTRACT .....................................................................................................................................8 CHAPTER 1 INTRODUCTION..................................................................................................................10 2 METHODS.............................................................................................................................13 Study Area..............................................................................................................................13 Assessment of Hammock Habitat...........................................................................................14 Bird Surveys...........................................................................................................................15 Analysis..................................................................................................................................17 3 RESULTS...............................................................................................................................27 Assessment of Hammock Habitat...........................................................................................27 Bird Surveys...........................................................................................................................29 4 DISCUSSION.........................................................................................................................45 Assessment of Hammock Habitat...........................................................................................45 Bird Surveys...........................................................................................................................46 Conservation Implications......................................................................................................51 LITERATURE CITED..................................................................................................................55 BIOGRAPHICAL SKETCH.........................................................................................................61 5

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LIST OF TABLES Table page 2-1 Number and description of 20-m, fixed-radius point count bird survey sites including hammock size, conservation status, and location and description (MM = mile marker on highway US-1)..............................................................................................................19 2-2 Description and quantification (mean and s.d., median and range) of habitat variables measured at point count locations (20-m fixed radius) where bird surveys were conducted..................................................................................................................23 2-3 Foraging guild and life history descriptions......................................................................24 3-1 Total amount (ha) of hammock habitat including percent of total in protected status by island in the Florida Keys as of 2004............................................................................34 3-2 Partially protected hammock habitat patches in the Florida Keys including total number of patches and area (ha) in protected and unprotected status by island and patch size category (ha) of 2004 (islands that did not contain partially protected patches were excluded from this table)..............................................................................35 3-3 Avian species detected during point count surveys including standard abbreviations, common and scientific names, life history and breeding status, and foraging guild designations........................................................................................................................36 3-4 Influence of measured habitat variables on species presence or absence from analyses with general linear models (data represent p-values). A negative p-value indicates an inverse relationship between the habitat variable and species presence. .....40 3-5 Influence of measured habitat variables on life history and foraging guilds from general linear model analyses with zero-inflated Poisson regression model (data represent p-values). A negative p-value indicates an inverse relationship between the habitat variable and species presence................................................................................41 3-6 Hammock habitat variables for birds that lacked sufficient data for statistical analyses. Data reflect the range of percent cover (smallest to largest) for PCT_HAM, PCT_CAN and HAM+CAN measurements where birds were observed..........................42 6

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LIST OF FIGURES Figure page 2-1 Geologic delineation of the Florida Keys by substrate. ...................................................25 2-2 Delineation of Upper Keys (consistent with Bancroft et al. 1995) study area, Lower Keys study area and extent of avian surveys.....................................................................26 3-1 Area (ha) of tropical hardwood hammock habitat patches by patch size (ha) category and protected status in the Florida Keys in 2004 (number of habitat patches provided above each bar)..................................................................................................................44 7

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science STATUS AND USE OF TROPICAL HARDWOOD HAMMOCKS AND FORESTED RESIDENTIAL AREAS AS HABITAT FOR RESIDENT AND NEOTROPICAL MIGRATORY BIRDS IN THE FLORIDA KEYS By Annisa Karim December 2007 Chair: Martin B. Main Major: Wildlife Ecology and Conservation The challenges of migration are immense and many avian species rely on stopover areas to rest and refuel before continuing migration. The availability and location of stopover habitat is vital to migratory species, and the loss of suitable stopover habitats has been implicated in the decline of migratory populations. Tropical hardwood hammocks in the Florida Keys are used by both resident and Neotropical migratory birds; however, anthropogenic activities in and around hammocks have resulted in the conversion of many of these forests to other urban uses. As a result, habitat loss and fragmentation have reduced suitable habitat. I used a Geographic Information Systems approach to quantify amount and status (protected/unprotected) of hammock forest patches in the Keys. More than 30% of hammock habitat in the upper Keys had been lost to deforestation from 1991 to 2004. Total remaining hammock habitat in the Florida Keys as of 2004 was 3,712 ha, of which 1,962 ha (53%) was located in the upper Keys, and 1,750 ha (47%) was located in the lower Keys. Of the remaining hammock habitat, approximately 37% remained unprotected throughout the Keys (46% and 27% in the upper and lower Keys, respectively). I conducted bird point counts at 86 survey sites in the northern Keys during March-May 2004 and 2005. Survey locations were established in both hammock habitat 8

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and in residential areas with varying degrees of tree canopy cover. Total hammock tree canopy had significant positive effects on presence of seven of the 35 species statistically analyzed, including the Black-whiskered Vireo (Vireo altiloquus), a bird of conservation concern. Conversely, hammock cover had significant negative effects on 12 species including common urban birds and non-native species. Cowbird species (Molothrus spp.) and feral cats (Felis catus), both of which were observed along hammock edges, may pose additional threats. Environmental education of the public on the ecological value of hammocks is needed as well as long term studies on avian-hammock interactions in the upper and lower Keys. 9

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CHAPTER 1 INTRODUCTION Many types of latitudinal migratory birds exist (Hayes 1995). According to Rappole (1995), Nearctic-Neotropical migrants are birds for which all or part of their populations breed north and winter south of the Tropic of Cancer. In the western hemisphere, there are 341 species of Nearctic-Neotropical migratory birds, hereafter referred to as Neotropical migrants (American Ornithologists Union 1998; 2000). Deforestation of wintering grounds and the fragmentation of breeding areas pose two serious habitat constraints for the conservation of migratory passerines (Morris et al. 1996). The challenges of migration are immense and many avian species rely on stopover areas to rest and refuel energy reserves before continuing on their migrations (Kerlinger 1989; Terborgh 1989; Moore 2000; Ydenberg et al. 2002). The loss and degradation of stopover habitats represent a third serious habitat constraint for the conservation of these migrants. The availability, location and quality of stopover habitat are vital to migratory species and the loss of suitable stopover habitats has been implicated in the decline of migratory bird populations (Sauer & Droege 1989; Moore et al 1990; Barrow et al. 2000). The combination of factors such as seasonal variability, availability of appropriate habitat, costs of migration, and ability to navigate play an important role in the success of migratory movements. While en route, migrants must make quick decisions about the location and type of landscape they use. The Florida Keys are a chain of low-lying islands that extend 354-km from the southeastern tip of Florida and arc in a southwesterly direction to the Dry Tortugas (Florida Department of Community Affairs 2003). These islands are located along one of the primary migratory routes for birds that breed in tropical and temperate North America and winter in the Caribbean and South America (Lincoln et al. 1998; US Fish and Wildlife Service 1999). The 10

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Keys also provide habitat for Caribbean avian species that rarely appear anywhere else in North America, such as the Mangrove Cuckoo (Coccyzus minor, MACU), Black-whiskered Vireo (Vireo altiloquus, BWVI) and the state listed, threatened White-crowned Pigeon (Patagioenas leucocephala formerly Columba leucocephala, WCPI; US Fish and Wildlife Service 1999; Banks et al. 2003). One type of plant community used by migrant and resident birds in the Florida Keys is tropical hardwood hammock (US Fish and Wildlife Service 1999). Historically, tropical hardwood hammocks, hereafter referred to as hammocks, were found as far north as Cape Canaveral on the east coast, and up to the mouth of the Manatee River on the west coast of Florida. However, development pressure resulted in the conversion of many of these forests to urban and agricultural uses. Today, hammocks occur primarily as remnant habitats in extreme south Florida and in small preserves along the Atlantic coast from Miami-Dade north to Martin County. The largest remaining tracts of extant hammocks occur in the Florida Keys. Hammocks are closed canopy forests characterized by evergreen and semi-deciduous woody species primarily of West Indian origin (Snyder et al. 1990; Sunquist et al. 2002; Lodge 2005). Hammocks in the Florida Keys occur on the highest elevations where flooding rarely occurs and are, therefore, prime areas for human habitation. Forman (1995) defines connectivity as spatial continuity of habitat or cover type across a landscape. Therefore, breaking up of a habitat or cover type into smaller, disconnected parcels is fragmentation. Plant communities in the Florida Keys, including hammocks, are naturally discontinuous and fragmented due to the geologic nature of the archipelago (US Fish and Wildlife Service 1999; Lodge 2005). Anthropogenic activities in and around hammocks have resulted in the conversion of many of these forest patches, primarily to urban uses (Bancroft 11

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et al. 1995). As a result, habitat loss and further fragmentation of remaining hammocks have reduced potentially important habitat for resident and Neotropical migratory birds on the archipelago. Furthermore, extant hammock habitat under private ownership in the Florida Keys is under intense development pressure and loss of habitat provided by these upland forests may have significant conservation implications for both Neotropical migrants and resident species of birds (Wilson 1992; Russell et al. 1998; Barrow et al. 2000; Petit 2000; Simons et al. 2000). The objectives of this study are to (1) determine the extent of hammock habitat throughout the Florida Keys as of 2004, (2) quantify area and number of hammock patches under conservation status and (3) describe bird communities by species, foraging guild, and life history that utilize hammocks and residential areas with varying degrees of canopy cover to better understand avian habitat preferences and needs during spring migration. 12

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CHAPTER 2 METHODS Study Area The Florida Keys (Keys) are a chain of small islands located within the subtropical region of the Western hemisphere. The Keys have a mild tropical climate, annual rainfall averages 101 cm (40 in), temperatures vary from 10-32 C (50-90 F), and mean temperature exceeds 18C (64 F) during all 12 months (Florida Climate Center 2005). This archipelago extends 354 km in a southwest direction from the southeast tip of peninsular Florida (25 N, 80 W) to Key West (24 N, 81 W; Fig. 2-1). In a geological context, the upper Keys rest on Key Largo Limestone and extend from Soldier Key (25 N, 80 W) approximately 150 km southwest to Big Pine Key (24 N, 81 W). The lower Keys sit on Miami oolitic limestone and extend from Big Pine Key westward approximately 60 km to Key West (24 N, 81 W; Fig. 2-1). Recent deposits of carbonate sands comprise the substrate of the deep, open waters between Key West, the Marquesas Keys and the Dry Tortugas (Craighead 1971; Brown et al. 1990; Snyder et al. 1990; Ripple 1995; Waitley 1997). Plant communities in the Florida Keys are naturally discontinuous and fragmented due to the geologic nature of the archipelago. Submerged portions of many islands support coral reef communities, low-lying areas sustain mangrove, tidal marsh, and freshwater marsh communities, while hammocks and pine (Pinus elliottii var. densa) communities thrive on the highest elevations where flooding rarely occurs (US Fish and Wildlife Service 1999; Lodge 2005). Hammocks are closed canopy forests and are dominated by a number of evergreen and semi-deciduous woody species, most of which are of West Indian origin (Snyder et al. 1990; Sunquist et al. 2002; Lodge 2005). The largest areas of hammocks occur in the northern portions of the Keys, especially on the largest island, Key Largo. Hammocks tend to get smaller as island size 13

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decreases south of Key Largo, with the exception of Big Pine Key, the largest island in the lower Keys. Hammocks throughout the Keys have undergone anthropogenic influences that have systematically reduced and further fragmented hammock habitat, particularly from residential and urban development (US Fish and Wildlife Service 1999; Lopez 2001). Assessment of Hammock Habitat I used a Geographic Information Systems (ESRI GIS Software ArcMap 9.0) approach to quantify cover, distribution, and status (protected/unprotected) of hammock forest patches in the Florida Keys. I quantified hammock cover throughout the Keys and measured loss of hammock cover in the upper Keys during 1991-2004 based on an earlier assessment by Bancroft et al. (1995). For comparative purposes I defined the upper Keys consistent with Bancroft et al. (1995) as extending from the Ragged Keys (a small chain of islands in Miami-Dade Co. north of Key Largo, 25 N, 80 W), south to Long Key (24 N, 80 W). Lower Keys were defined as extending from Duck Key (24 N, 80 W) to Key West (24 N, 81 W; Fig. 2-2). I used 2004 digital orthophoto quarter quadrangles (DOQQ) from the Land Boundary Information System (Florida Department of Environmental Protection 2003) at a 1-m resolution as the base layer of my map. I obtained the Monroe Co. (includes Keys) boundary layer from the Florida Geographic Data Library (University of Florida GeoPlan Center) and used the ArcMap 9.0 clip function to isolate the Monroe Co. 1995 land use/land cover data available from the South Florida Water Management District (online: http://www.sfwmd.gov/site/index.php?id=1; accessed December 2005). I superimposed the resulting 1995 dataset on the DOQQ and used the Florida Land Use Land Cover Classification System (FLUCCS, level 3 code 4260) to designate hammock habitat in the Keys (defined as tropical hardwoods by FLUCCS; Florida Department of Transportation 1999). I created a new data layer to reflect the extent of hammock habitat throughout the Keys (including the Ragged Keys) as of 2004. This new layer reflected 14

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refinements to the 1995 FLUCCS code designations (e.g., new roads, water bodies, and land use changes since 1995) based on examination of 2004 DOQQ aerial photos and ground-truthing. I created polygons, via visual classification (1:4000), to reflect hammock habitat not recognized in the 1995 dataset and in the Ragged Keys, which occur in Miami-Dade Co. but were included in Bancroft et al. (1995). These additional hammock polygons were classified as commercial or residential in the 1995 dataset because they were located within areas zoned for residential and commercial use but had not yet been developed as of 2004. The 2005 Florida Natural Areas Inventory Conservation Lands Layer was added to the map and I used the intersect function in ArcMap to identify protected lands in Monroe Co. This enabled me to identify hammock habitat protected for conservation under federal, state, local and private management in the Florida Keys. I generated new layers to differentiate between protected and unprotected status of hammock forest patches in the upper and lower Keys. I ground-truthed 10% of the mapped hammock polygons and determined hammock coverage of these polygons was 98% accurate. I calculated total area of extant hammock habitat (protected and unprotected) in the upper and lower Keys by creating a new field in the attribute table for the 2004 hammock habitat layer. This included corrected data from FLUCCS code 4260 designations from the 1995 layer and the polygons I constructed. I used ArcMaps calculate values function to calculate the area of all hammock polygons, converted results to hectares, and compared results from the upper Keys with those of Bancroft et al. (1995). Bird Surveys I conducted morning bird surveys (point counts) in hammock forest habitat and residential areas with varying levels of hammock and tree canopy cover during March-May (spring migration) 2004 and 2005. Point counts were conducted between sunrise and 180 minutes after sunrise as 20-m fixed-radius point counts with 5-min survey intervals (Hutto et al. 1986). I 15

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established 86 point count locations from Key Largo to Vaca Key (Table 2-1), 65 of which were in residential areas and 21 in protected hammock habitat. Of the 65 point count locations within residential areas, six occurred on undeveloped parcels in private ownership that had extant hammock habitat as of 2004. Point count locations were separated by > 250 m to maintain independence of observations (Hutto et al.1986). I surveyed birds at each point count location five times each field season with the exception of one protected hammock where two point count locations were visited four times during 2004 because permission to access the hammock was delayed. Each 20-m fixed radius point count assessed bird communities over an area of approximately 0.13 ha (0.33 ac). During surveys, I recorded all birds by call, song or sight within the 20-m radius. Species recorded as flying over/through without stopping were removed from the dataset used for analysis. Additionally, sea birds (e.g., terns, Sterna spp.), wetland birds (e.g., Great Egrets, Ardea alba), and domestic species (e.g., Peafowl, Pavo cristatus) were removed from the dataset used for analysis. Selection of point count locations for bird surveys was done separately for protected and residential areas. Protected hammocks in the upper Keys (Key Largo to Marathon) were identified during the GIS portion of the study. All existing protected hammocks in the upper Keys were included in the bird surveys. All hammock habitats within the study area have been described by the US Fish and Wildlife Service (1999) as secondary forest. All accessible residential areas from Key Largo (city of Key Largo) to Vaca Key (city of Marathon; 25 N, 80 W to 24 N, 81 W) were included in a pool of potential sites for bird surveys. Most residential areas and, consequently potential bird survey sites, were located on the larger islands. Locations dominated by the invasive non-native shrub, Brazilian pepper (Schinus terebinthifolius), were eliminated from the pool of potential study sites because this shrub 16

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attracts a disproportionate number of birds seeking food (Myers & Ewel 1990; Spector & Putz 2006) and its presence on some sites would confound the data. Potential survey sites were numbered and randomly drawn to identify point count locations. Survey sites in residential areas ranged from 0-100% hammock cover because some areas zoned as residential included undeveloped parcels with hammock habitat. The northernand southern-most point count locations were separated by approximately 97 km, which necessitated a stratified random approach to sampling. I conducted surveys randomly within groups of locations (approximately 16 point count locations/morning) and alternated the initiation of surveys between north and south. This stratified random and alternating directional survey scheme (first north to south then south to north) was used throughout both field seasons. Analysis Using a GIS approach, I was able to import GPS coordinates of all survey locations into ArcMap and create a 20-m buffer around each survey location. Within each 20-m fixed-radius survey location, I defined and measured eight habitat variables (Table 2-2). Variables included hammock habitat (percent cover of native hammock tree canopy and understory cover; PCT_HAM) tree canopy (percent cover of native tree canopy cover with cleared understory; PCT_CAN), ground cover (percent cover of grasses and residential lawn or managed understory with no tree canopy cover; PCT_GROUND), and other vegetation (percent cover of non-hammock vegetation, e.g., mangrove and pine habitats; PCT_OTHER). I also quantified distance from survey location to the closest hammock (D2HAM; if survey location was situated within a hammock D2HAM = 0) and the size of that hammock (SIZEHAM). I created two combined variables, total canopy (combined sum of hammock habitat and tree canopy; HAM+CAN) and the distance from the point count location to the nearest hammock times the size of that hammock (DXSIZE). 17

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Birds were analyzed on a species specific basis as well as by foraging guild and life history traits (Table 2-3). Foraging guild (foraging technique) categories were defined by Ehrlich et al. (1998). Migratory and non-migratory status was defined by Rappole (1995) and breeding locations were defined by the Florida Fish and Wildlife Conservation Commission (2003). Overall migratory and breeding status determined life history categories. The relationships between count data and landscape attributes were regressed using general linear models. Point count data were modeled on a per species, life history, and foraging guild basis using eight landscape variables (Table 2-2) as covariates in Poisson regression. Species detected fewer than 11 times were excluded from analysis due to small sample sizes. Count data with excessive zeros were modeled using zero-inflated Poisson (ZIP) logistic regression (Martin & McIntyre 2007). Count data without excessive zeros were modeled using the proc glimmix command and all analyses were conducted in SAS Version 9.1.3 (Copyright 2002-2003 by SAS Institute Inc., Cary, NC, USA). Species richness (cumulative number of species detected at each survey location) was calculated and modeled against the habitat variables. Additionally, the Shannon-Wiener diversity index (H s = p i ln p i ; where H = diversity index; s = species richness, and p i = the proportion of individuals found in the i th species) was computed for all species detected (other than seabirds and domestic species) and modeled against the habitat parameters. This index was chosen because it is sensitive to changes in the abundance of rare species in a community (Solow 1993) and may better reflect the composition of all species found therein. 18

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Table 2-1 Number and description of 20-m, fixed-radius point count bird survey sites including hammock size, conservation status, and location and description (MM = mile marker on highway US-1). Survey sites (#) Size (ha) of Hammock a Protected Status (Y/N) Location and Description b 7 70.30 c Y Dagny Johnson Key Largo Hammock Botanical State Park, Key Largo 2 3.87 Y John Pennekamp Coral Reef State Park, Key Largo 3 N/A N Residential area (near Paradise Point Cove), Key Largo (MM 105) 2 N/A N Residential area (Taylor Creek Village), Key Largo (MM 105) 1 17.39 N Located in residential area in Key Largo (In Taylor Creek Village, MM 105) 4 N/A N Located in residential area on Key Largo (Near Largo Sound Village and Angler's Park, MM 104) 2 N/A N Located in residential area on Key Largo (Near Largo Sound Park, between MM 104 and MM 103) 6 N/A N Located in residential area on Key Largo (Near Cross Key Waterway Estates and Twin Lakes, MM 103) 6 N/A N Located in residential area on Key Largo (Between Lime Drive and Troup Rd, MM 102) 2 N/A N Located in residential area on Key Largo (In Key Largo Trailer Village, MM 101) 3 N/A N Located in residential area on Key Largo (Between Lakeshore Drive and East Shore Drive, between MM 101 and 100). 19

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Table 2-1 (continued). Survey sites (#) Size (ha) of Hammock a Protected Status (Y/N) Location and Description b 2 N/A N Located within a residential area on Key Largo (Oceanside near MM 99 between Sailfish Trail and Ocean Shores Drive) 1 86.90 N Located within a residential area on Key Largo (Oceanside near MM 99 between Sailfish Trail and Ocean Shores Drive) 2 N/A N Located within a residential area on Key Largo (Northeast Papa Johns Pizza, near MM 98) 4 N/A N Located in a residential area on Key Largo (Near Sunset Point and the Fl. Keys Marine Sanctuary Bldg., MM 95) 1 0.61 N Located in a residential area on Key Largo (Between Harry Harris State Park and Burton's Yacht Basin, MM 93) 1 N/A N Located in a residential area on Key Largo (Between Harry Harris State Park and Burton's Yacht Basin, MM 93) 1 4.52 N Located in a residential area on Key Largo (Between Harry Harris State Park and Burton's Yacht Basin, MM 93) 1 4.44 N Located in a residential area on Key Largo (Between Harry Harris State Park and Burton's Yacht Basin, MM 93) 1 0.31 N Located within a residential area on Key Largo Tavernier (West of Bank of America, MM 92) 20

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Table 2-1 (continued). Survey sites (#) Size (ha) of Hammock a Protected Status (Y/N) Location and Description b 3 N/A N Located within a residential area on Key Largo Tavernier (Near Tavernier Harbor, MM 92) 2 N/A N Located within a residential area on Key Largo Tavernier (Near Point Lowe, MM 92) 4 N/A N Located in a residential area on Plantation Key (Near Tavernier Creek and Sunshine Estates, between MM 91 and MM 90) 2 4.67 Y Indigo Hammock just north of MM90 between Ocean Drive and Riviera (Oceanside). Part of the conservation lands within Islamorada 3 N/A N Located in residential area on Plantation Key (Between Coral Shores High School and Lake Harbor, MM 90) 4 N/A N Located in residential area on Plantation Key (Between Seminole Blvd. and High Point Rd just north of the Govt. center, MM 89) 2 N/A N Located in residential area on Plantation Key (Between Plantation Blvd. and Royal lane, south of MM 88) 3 10.68 Y Windley Key Fossil Reef Geological State Park. 2 N/A N Located in residential area on Upper Matecumbe Key (Near Cheeca Lodge Resort, MM 82.5) 2 N/A N Located in residential area on Upper Matecumbe Key (Near Kaiyo Restaurant, between MM 82 and MM 81) 21

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22 Table 2-1 (continued). Survey sites (#) Size (ha) of Hammock a Protected Status (Y/N) Location and Description b 1 4.46 Y THH Layton Trail a part of Long Key State Park 1 1.24 Y THH within Long Key State Park 2 12.57 Y Blue Heron Hammoc k West of the Marathon Airport. Managed by FWCC. 3 14.37 Y Museums and Nature Center of Crane Point in Marathon managed by the Florida Keys Land and Sea Trust a If point count location(s) is/are located within a hamm ock, hammock size (ha) is provided. If point count location(s) is/are not located within a hammock, N/A. b MM = Mile Marker. Mile Markers are used as reference points of locations along the Overseas Highway in the Florida Keys. Mile Marker Numbers descend from north to south. c Dagny Johnson Key Largo Hammock Botanical State Park contains many patches of hammock, 70.3 ha represents the amount of continuous hammock habitat within which point count surveys were conducted.

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Table 2-2 Description and quantification (mean and s.d., median and range) of habitat variables measured at point count locations (20-m fixed radius) where bird surveys were conducted. Habitat Variable Description Protected sites (%) Mean SD Median Range Unprotected (%) Mean SD Median Range PCT_HAM a Percent cover of native hammock (includes native tree canopy and understory cover) 96.0 8.4 100 64.6 100 18.1 30.2 0 0 PCT_CAN a Percent cover of native hammock tree canopy (cleared understory) 0.3 1.4 0 0.2 18.0 15.6 14.5 0.3 PCT_GROUND a Percent cover of grasses and residential lawns 0.5 2.2 0 0.1 10.1 10.9 6.3 0 34.5 PCT_OTHER a Percent cover of non-hammock vegetation (e.g., mangroves, pine rocklands, etc.) 0.4 1.6 0 0.5 1.4 4.9 0 0.4 D2HAM a Distance (m) to closest hammock from survey location (if survey location was located within a hammock, D2HAM = 0) 0 0 0 0 59.1 83.2 25.9 0 416.5 SIZEHAM a size (ha) of closest hammock from survey location (if survey location was located within a hammock, the size of that hammock was recorded) 29.3 30.0 12.6 1.270.3 6.9 19.0 0.3 <0.186.9 HAM+CAN b PCT_HAM + PCT_CAN; percent cover of total native hammock canopy (regardless of understory) 96.3 7.3 100 70.8100 36.2 27.9 30.9 0 DXSIZE b D2HAM SIZEHAM; distance (m) of the closest hammock multiplied by the size of that hammock. 0 0 0 0 372.4 1828.4 9.9 014013.5 23 a Variables analyzed as covariates in first set of analyses. b Variables analyzed as covariates in second set of analyses

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24 Table 2-3 Foraging guild and life history descriptions Life History Description YR a Year-round residents; non-migratory birds MB ab Migratory Breeders: Neotropi cal migratory birds that mi grate to the northern FL Keys to breed. NM b Neotropical migratory birds that use th e northern FL Keys as stopover sites and continue to migrate nor th for breeding purposes Foraging Guild c Description AF Aerial foragers: Captures flying inse cts while in prolonged continuous flight. BG Bark gleaners: Gleaning prey from tree trunks and branches; includes excavating and drilling into bark. FG Foliage gleaners: Gleaning prey (including fruits) from foliage of vegetation and occasionally branches. GG Ground gleaners: Picking up items from th e surface of soil, lawns, sand, forest floor, etc. HA Hawking: Short flights from perch to capture flying insects. HG Hover and gleaners: Gleaning while hovering. Takes nectar, insects and/or berries from plants above the ground while hovering. HO Hover and pouncers: Hovering and pounc ing; hovering before swooping or dropping down on prey. SW Swoops: Snatches up prey from ground in ta lons after gliding decent from perch with wings spread. a Birds reported to breed in the northern Keys (Florida Fish and Wildlife Conservation Commission 2003). b Migratory status of species (Rappole 1995). c Foraging guild/foraging techniques for species defined by Ehrlich et al. (1998).

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25 Figure 2-1 Geologic delineation of the Florida Keys by substrate. 25

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Figure 2-2 Delineation of Upper Keys (consistent with Bancroft et al. 1995) study area, Lower Keys study area and extent of avian surveys. 26

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CHAPTER 3 RESULTS Assessment of Hammock Habitat Hammock habitat in the Florida Keys encompassed 3,712 ha and accounted for 19% of the Keys landscape during 2004. Of that existing hammock habitat, 1,962 ha (53%) was located in the upper Keys and 1,750 ha (47%) was located in the lower Keys (Table 3-1). Hammock habitat in the upper Keys was divided among 124 habitat patches (median = 1.5 ha, range = 0.1-205.7 ha), of which 1,066 ha (54%) were under conservation status. Hammock habitat located in the lower Keys was divided among 102 patches (median = 4.4 ha, range = 0.3-96.3 ha), of which 1,283 ha (73%) were protected under conservation status (Table 3-1). In total, therefore, 2,349 ha (63%) of hammock habitat in the Florida Keys were protected from development. Key Largo, the largest and one of the northernmost islands in the archipelago, contained the most hammock habitat (1,834 ha) and the largest amount of habitat in conservation status (1,024 ha). These conserved patches were situated largely within the Dagny Johnson Key Largo Hammock Botanical State Park and the Crocodile Lake National Wildlife Refuge. The largest individual patch of hammock habitat in the entire Keys (206 ha) was also located on Key Largo. Many islands in the Keys did not contain any hammock habitat either as a result of land use change or unsuitable environmental conditions. Hammock patches throughout the Keys were either fully protected (53 patches, 1062 ha), partially protected (43 patches, 2081 ha) or in private ownership slated for development (130 patches, 569 ha). Partially protected hammocks were designated as those areas that included protected and unprotected contiguous hammock habitat. The number and area (ha) of hammock patches varied by patch size category (Fig. 3-1). The majority of area (892 ha, 84%) and 30% of 27

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the total number of patches represented by fully protected hammocks were >20 ha in size. Similar results were found for partially protected hammocks. The majority of area (1,956 ha, 94%) and 51% of the total number of patches represented by partially protected hammocks were >20 ha in size. Twenty-five of these patches occurred in the upper Keys and 18 in the lower Keys (Table 3-2). Conversely, the total number of unprotected hammock patches was dominated by patches of 0-5 ha (108 patches, 83%). Seven (5%) unprotected patches >20 ha encompassed 267 ha and represented 47% of isolated, unprotected hammock habitat (i.e., not contiguous with protected hammock habitat). Comparisons with Bancroft et al. (1995) revealed hammock habitat declined by 31% in the upper Keys since 1991, which represents a loss of 870 ha over a 13-year period. In 1991, the upper Keys contained 2,832 ha of hammock divided into 1,068 patches of which, 987 were smaller than 5 ha (Bancroft et al. 1995). By 2004, this region contained 1,962 ha of hammock divided among 124 patches of which, 93 were smaller than 5 ha. This represents a decrease of 944 (88%) total fragments and a decrease of 894 (91%) patches under 5 ha. In 1991, Key Largo was the only island in the upper Keys to contain hammocks >20 ha. While this remained true for 2004, the number of hammocks >20 ha went from 2-19 patches revealing fragmentation of large tracts of hammock. This means that extremely large hammocks that fell within the >20ha size category in 1991, were fragmented but were still, as of 2004, >20ha in size. The greatest change in the 13 years took place in hammocks from 0-1 ha with an increase from 4-53 patches. Between 1991 and 2004, all six size categories saw an increase in the number of hammock patches, revealing fragmentation of hammock in the upper Keys since 1991. Historical estimates of the number of hammock fragments and their size range was not available for the lower Keys 28

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but as of 2004, 111 patches remained extant with the most (28 patches, 25.23%) occurring in the >20.0 ha range and the least (13 patches; 11.71%) occurring in the 5-10 ha range. Bird Surveys During March-May 2004 and 2005, I conducted a total of 858 point count surveys over 86 point count locations. Survey locations in protected hammocks ranged from 66 100% hammock cover (mean = 96%, sd = 8.4%, median = 100%) within the 20-m radius of the point count survey location, with deviations from 100% being primarily due to cleared trails. Areas zoned as residential were highly variable in nature and included hammock habitat in undeveloped lots at one extreme, to highly manicured lawns, gardens, and cleared areas adjacent to homes at the other extreme. The distance from the 86 survey locations to the nearest hammock varied from 0-416 m; the size of those hammocks varied from <0.1-87 ha. Seventy-eight species were observed over all 86 survey locations during March-May of 2004 and 2005; these species totaled 6,400 individuals and represented three life history categories and 11 foraging guilds (Table 3-3). Eleven species of waterbirds and domestic birds were removed from the analysis. Thirty-five passerines were detected on fewer than 11 separate occasions; consequently, these data were not sufficient to perform statistical analyses. Table 3-4 reveals which analyses were performed on each of the 32 remaining species and the results of the analyses. One species detected, but not included in analyses due to lack of sufficient data was the Brown-headed Cowbird (Molothrus ater, BHCO), an obligate brood parasite. Many birds commonly seen in urban environments were detected during the study including the opportunistic nest predator, the Blue Jay (Cyanocitta cristata, BLJA ) and two exotic species, the Eurasian Collared-dove (Streptopelia decaocto; ECDO) and the European Starling (Sturnus vulgaris, EUST). Five species included in these analyses were designated as Birds of Conservation Concern by the US Fish and Wildlife Service Division of Migratory Bird 29

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Management (U.S. Fish and Wildlife Service 2002). These included the BWVI, Common GroundDove (Columbina passerine, COGD), MACU, Prairie Warbler (Dendroica discolor, PRAW) and the state listed, threatened WCPI. The percent cover of native hammock habitat (PCT_HAM) was a significant predictor of the presence of 15 species (Table 3-4). Among Neotropical migrants, PCT_HAM showed a positive predictive effect on three species and a negative effect on one species. Among migratory breeders, effects were positive on two species including PRAW, a bird of conservation concern and negative on one species. Among year-round residents, PCT_HAM had positive predictive effects on one species and negative effects on seven, including non-native ECDO and EUST. The percent cover of native hammock tree canopy with cleared understory (PCT_CAN) had significant predictive effects on 10 species. Neotropical migrants had three species with positive and zero with negative response to this habitat variable. Migratory breeders had two species that demonstrated positive responses, including the PRAW, and no negative responses. Year-round residents had zero species with positive responses to this variable and five with negative responses, including the ECDO and EUST. Percent ground cover (PCT_GROUND) significantly influenced seven species. Neotropical migrants had one species demonstrate a positive response and zero species with negative responses to this variable. Migratory breeders included three species with positive responses, including PRAW and BWVI, which are birds of conservation concern. One species of migratory breeder demonstrated a negative response, the MACU, also a bird of conservation concern. Year-round residents included one species that responded positively and one species that responded negatively. 30

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The percentage of other vegetation (PCT_OTHER; mangroves, pine habitat) showed positive predictive effects on three species and negative predictive effects on one species. Positive effects included one migratory breeder (PRAW), and two year-round residents, including the exotic EUST. One year-round resident species responded negatively to PCT_OTHER. Distance from survey locations to the nearest hammock (D2HAM) showed two positive and two negative responses, all of which occurred among year-round resident species. The size of the nearest hammock (SIZEHAM) showed significant effects on 10 species. Among Neotropical migrants two species responded positively and one responded negatively. Migratory breeders included two positive responses including the BWVI, and two negative responses including the PRAW. Year-round residents had two species with negative responses, including the non-native ECDO. The combined variable HAM+CAN had significant effects on 19 species, the most of any habitat variable. Among Neotropical migrants three species responded positively and two species responded negatively. Migratory breeders included three positive and three negative responses. Two birds of conservation concern, the PRAW and BWVI responded positively. Year-round residents had one positive and seven negative responses, including the exotic ECDO and EUST, both of which responded negatively. The other combined variable, DXSIZE, had little effect on species. Among year-round residents one species responded positively and one negatively to this variable. When species were collectively analyzed based on life history and foraging guild (Table 3-5), PCT_HAM showed a negative predictive effect on one life history category (year-round residents) and two guild categories (bark gleaners and ground gleaners), and a positive effect on 31

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the two migratory life history categories (Neotropical migrants and migratory breeders) and one guild (foliage gleaners). PCT_CAN also showed a negative predictive effect on year-round residents and positive for the migratory life history categories. For guilds, PCT_CAN influenced bark gleaners and ground gleaners negatively, and hover gleaners positively. PCT_GROUND was a positive predictor only for the migratory breeder life history category and PCT_OTHER had no significant influence on any life history category or guild. D2HAM showed negative predictive effects on three guilds (bark gleaners, foliage gleaners, and hover gleaners) and no predictive effects on life history categories. SIZEHAM showed negative predictive effects on two life histories (Neotropical migratory birds and year-round residents). Additionally, SIZEHAM showed negative effects on two guilds (ground gleaners and hawkers) and positive effects on two guilds (foliage gleaners and hover gleaners). Similar to the individual species analyses, the combined HAM+CAN variable influenced the greatest number of groups. It showed negative predictive effects on one life history (year-round residents) and three guilds (bark gleaners, ground gleaners, and hawkers) and positive effects on two life histories (both migratory categories) and two guilds (foliage gleaners and hover gleaners). The combined habitat variable DXSIZE showed no predictive effects on any of the groups analyzed. Species richness (R 2 = 0.14, p = 0.04) and species diversity (R 2 = 0.20, p < 0.01) for 67 species observed at survey locations (all species observed with the exception of the waterbirds and domestic species) were significantly influenced by SIZEHAM. However, when birds of conservation concern (R 2 = 0.10, p = 0.13) or migratory species (R 2 = 0.09, p = 0.16) were analyzed separately, or together (R 2 = 0.11, p = 0.16), there was no relationship between hammock size and presence. 32

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33 I recorded 35 species that could not be analyzed statistically due to lack of sufficient data. Of these, 31 (88%) were Neotropical migrants, two (6%) were migratory breeders, and two (6%), were year-round residents (Table 3-3). The vast ma jority (94%), therefore, of rarely observed birds were migratory. These species represented seven different foraging guilds, of which 40% were foliage gleaners, 20% ground gleaners, and the remaining 40% fairly evenly divided among five other foraging guilds. Of the rarely observed birds, eight (23%) were detected in locations with >50% hammock cover (PCT_HAM) and five of those were detected at locations with 100% hammock cover (Table 3-6). These species in cluded six Neotropical migrants, one migratory breeder, and one year-round resident, none of which are identified as birds of conservation concern. The combined variable total canopy cover (HAM+CAN) had similar results. Eleven species (32%) were detected at locations with >50% total tree ca nopy cover, five of which were detected at locations with 100% total tree canopy. These included nine Neotropical migrants, one migratory breeder, and one year-roun d resident. None of these species were identified as birds of conservation concern. Regarding guilds, these species represented two species of ground gleaners, four foliage gleaners, one bark glea ner, three aerial foragers, and one hover gleaner.

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Table 3-1 Total amount (ha) of hammock habitat including percent of total in protected status by island in the Florida Keys as of 2004. Island Hammock area total (ha) conserved (%) Hammock patches No. Mean patch Median patch Range patch patches size (ha) SD size (ha) size (ha) Upper Keys a Key Largo b 1834.2 56 45 40.5 59.1 10.3 0.1-205.7 Plantation Key 47.8 26 32 1.5 1.5 1.0 0.2-4.8 Windley Key 12.4 86 4 3.1 5.1 0.7 0.3.7 Upper Matecumbe Key 54.2 17 36 1.5 1.4 0.8 0.3.7 Lower Matecumbe Key 6.3 61 3 2.1 0.6 2.40 1.1.4 Long Key 6.7 87 4 1.7 1.5 1.1 0.61 1.35 Upper Keys total 1961.6 54 124 15.8 40.1 1.5 0.1.7 Lower Keys Grassy Key 9.6 0 8 1.2 0.7 1.2 0.4.4 Vaca Key 91.5 39 20 4.6 5.8 2.3 0.3.1 Ohio Key 2.2 0 1 2.2 2.2 2.2.2 Big Pine Key 501.8 90 17 29.5 33.0 11.3 0.4.9 Middle Torch Key 564.8 85 16 35.3 30.0 23.3 0.3.8 Big Torch Key 258.0 98 8 32.3 37.9 10.6 2.5-96.3 Ramrod 159.2 0 7 22.7 19.4 14.5 1.3.9 Little Knockemdown Key 44.7 28 1 44.7 44.7 44.7.7 Knockemdown Key 23.4 100 1 23.4 23.4 23.4.4 Toptree Hammock Key 15.5 100 2 7.8 1.1 7.7 7.0.5 Cudjoe Key 13.8 22 3 4.6 0.5 4.4 4.2.2 Sugarloaf Key 35.8 0 11 3.2 3.0 2.6 0.5.6 Lower Sugarloaf Key 29.63 24 7 4.2 4.9 2.4 1.3.2 Lower Keys total 1749.9 73 102 17.2 25.2 4.4 0.3.3 Grand total 3711.5 63 226 16.4 34.2 2.7 0.1.7 34 a Delineation of upper Keys from Bancroft et al. (1995). b includes Ragged Keys.

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Table 3-2 Partially protected hammock habitat patches in the Florida Keys including total number of patches and area (ha) in protected and unprotected status by island and patch size category (ha) of 2004 (islands that did not contain partially protected patches were excluded from this table). Hammock Patches by Size (ha) Category Island Total no. patches 0-1 # a P b U c 1-5 # a P b U c 5-10 # a P b U c 10-20 # a P b U c >20 # a P b U c Upper Keys d Key Largo e 20 3 0.7 0.3 2 3.4 4.1 1 5.2 2.2 2 18.3 13.7 12 812.0 607.4 Plantation Key 3 3 4.4 3.0 Upper Matecumbe Key 2 2 0.7 0.6 Upper Keys total 25 5 1.4 0.9 5 7.8 7.1 1 5.2 2.2 2 18.3 13.7 12 812.0 607.4 Lower Keys Vaca Key 1 1 4.6 8.9 Big Pine Key 4 4 275.3 33.9 Middle Torch Key 9 2 1.0 3.00 2 26.5 5.3 5 113.3 76.3 Little Knockemdown Key 1 1 12.4 32.3 Cudjoe Key 1 1 3.0 2.2 Lower Sugarloaf Key 2 2 6.0 0.6 Lower Keys total 18 4 7.0 3.6 1 3.0 2.2 3 31.1 14.2 10 401.0 142.5 Grand total 43 5 1.4 0.9 9 14.8 10.6 2 8.2 4.4 5 49.4 27.9 22 1213.0 749.9 35 a Number of partially protected hammock patches per size category. b Amount (ha) of protected hammock habitat by patch size category. c Amount (ha) of unprotected hammock habitat by patch size category. d Delineation of upper Keys from Bancroft et al. (1995). e includes the Ragged Keys.

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Table 3-3 Avian species detected during point count surveys including standard abbreviations (USGS Patuxent Wildlife Research Center Bird Banding Laboratory), common and scientific names, life history (Rappole 1995) and breeding status (Florida Fish and Wildlife Conservation Commission 2003), and foraging guild designations (Ehrlich et al. 1998; see Table 2-3 for definitions of life histories and foraging guilds). Frequency includes number of group observations (n) for each species and the % of total group observations among species. Abundance includes number of individual birds recorded (n) for each species and the % of total birds among species. Domestic and waterbirds were excluded from analyses. Species Code Common Name Scientific Name Life History Foraging Guild Frequency (n) (%) Abundance (n) (%) GRCA Gray Catbird Dumetella carolinensis NM GG 70 6 251 4 YPWA M G 6090 Palm Warbler Dendroica palmarum NM GG 64 5 468 7 BAWW Black-and-white Warbler Mniotilta varia NM BG 34 3 97 2 AMRE American Redstart Setophaga ruticilla NM HG 34 3 107 2 REVI Red-eyed Vireo Vireo olivaceus NM HG 25 2 48 1 OVEN Ovenbird Seiurus aurocapillus NM GG 20 2 31 0 BLPW Blackpoll Warbler Dendroica striata NM FG 20 2 41 1 BGGN Blue-gray Gnatcatcher Polioptila caerulea NM FG 17 1 41 1 COYE Common Yellowthroat Geothylpis trychas NM FG 14 1 31 0 EAKI Eastern Kingbird Tyrannus tyrannus NM HA 13 1 17 0 BTBW Black-throated Blue Warbler Dendroica caerulescens NM HG 10 1 16 0 INBU Indigo Bunting Passerina cyanea NM FG 10 1 33 1 CMWA Cape May Warbler Dendroica tigrina NM FG 9 1 27 0 WEWA a Worm-eating Warbler Helmitheros vermivorus NM FG 8 1 10 0 WWDO* a White-winged Dove Zenaida asiatica NM GG 8 1 14 0 MAWA a Magnolia Warbler Dendroica magnolia NM HG 7 1 31 0 YTWA a Yellow-throated Warbler Dendroica dominica N B BHCO a Brown-headed Cowbird Molothrus ater NM GG 5 0 9 0 36

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Table 3-3 (continued) Species Code Common Name Scientific Name Life History Foraging Guild Frequency (n) (%) Abundance (n) (%) BLBW ab Blackburnian Warbler Dendroica fusca NM M G 4060 M G 3040M G 3060 M G 2020M G 2020 M G 2030M G 2040 M G 2040 M G 1010 FG 5 0 15 0 HOWA a Hooded Warbler Wilsonia citrina N F PABU a Painted Bunting Passerina ciris NM GG 3 0 4 0 YBCU a Yellow-billed Cuckoo Coccyzus americanus N F YTVI a Yellow-throated Vireo Vireo flavifrons N F AMKE a American Kestrel Falco sparverius NM HO 2 0 2 0 MERL a Merlin Falco sparverius NM HA 2 0 2 0 PIWA a Pine Warbler Dendroica pinus N B PROW a Prothonotary Warbler Prothonotaria citrea N B SWWA a Swainson's Warbler Limnothylpis swainsonii NM GG 2 0 2 0 EAPH a Eastern Phoebe Sayornis phoebe NM HA 2 0 3 0 MYWA a Yellow-rumped Warbler Dendroica coronata N F NAWA a Nashville Warbler Vermivora ruficapilla N F RTHU a Ruby-throated Hummingbird Archilochus colubris NM HG 2 0 4 0 SUTA a Summer Tanager Piranga olivacea N F SCTA a Scarlet Tanager Piranga olivacea NM HG 2 0 5 0 WOTH a Wood Thrush Hylocichla mustelina NM GG 2 0 5 0 NRWS a Northern Rough-winged Swallow Stelgidopteryx serripennis NM AF 1 0 1 0 RBGR a Rose-breasted Grosbeak Pheucticus ludovicianus N F 37

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Table 3-3 (continued). Species Code Common Name Scientific Name Life History Foraging Guild Frequency (n) (%) Abundance (n) (%) RCKI a Ruby-crowned Kinglet Regulus calendula N M 0 M G 1010M G 1010 M G 1040 B F 1010B G 1010 F G 1 01 TRES a Tree Swallow Tachycineta bicolor NM AF 1 0 1 0 YBCH a Yellow-breasted Chat Wilsonia citrina N F YBSA a Yellow-bellied Sapsucker Sphyrapicus varius N B BLGR a Blue Grosbeak Guiraca caerulea NM GG 1 0 2 0 NOPA a Northern Parula Parula americana N F CEDW a Cedar Waxwing Bombycilla cedrorum NM FG 1 0 45 1 MODO Mourning Dove Zenaida macroura MB GG 59 5 185 3 GCFL Great-crested Flycatcher Myiarchus crinitus MB HA 46 4 133 2 GRAK Gray Kingbird Tyrannus dominicensis MB HA 36 3 87 1 PRAW Prairie Warbler Dendroica discolor MB FG 26 2 66 1 WEVI White-eyed Vireo Vireo griseus MB FG 25 2 143 2 WCPI^ White-crowned Pigeon Patagioenas leucocephala MB FG 24 2 53 1 BWVI Black-whiskered Vireo Vireo altiloquus MB FG 22 2 63 1 YSFL Northern Flicker Colaptes auratus MB GG 11 1 15 0 COGD Common Grounddove Columbina passerina MB GG 11 1 16 0 MACU Mangrove Cuckoo Coccyzus minor MB FG 7 1 13 0 CONI a Common Nighthawk Chordeiles minor M A YWAR a Yellow Warbler Dendroica petechia M F NOCA Northern Cardinal Cardinalis cardinalis YR GG 85 7 962 15 RBWO Red-bellied Woodpecker Melanerpes carolinus YR BG 80 7 533 9 ECDO* Eurasian Collared-dove Streptopelia decaocto YR GG 75 6 1097 18 COGR Common Grackle Quiscalus quiscula YR GG 68 6 461 7 BLJA BlueJay Cyanocitta cristata YR GG 60 5 310 5 38

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Table 3-3 (continued). Species Code Common Name Scientific Name Life History Foraging Guild Frequency (n) (%) Abundance (n) (%) RWBL Red-winged Blackbird Agelaius phoeniceus YR GG 49 4 206 3 NOMO Northern Mockingbird Mimus polyglottos YR GG 48 4 195 3 EUST* European Starling Sturnus vulgaris YR GG 32 3 250 4 RSHA Red-shouldered Hawk Buteo lineatus YR SW 20 2 38 1 CARW a Carolina Wren Thryothorus ludovicianus YR GG 4 0 6 0 MOPA* a Monk Parakeet Myiopsitta monachus YR --d 4 0 14 0 ---c Unidentified parrot Amazonia sp. YR --d 1 ---3 ---BLBU bc Blue Bunting Cyanocompsa parellina --b --d 1 ---1 ---COTE c Common Tern Sterna hirundo NM HD 1 ---81 ---DCCO c Double-crested Cormorant Phalacrocorax auritus MB SD 1 ---1 ---GHOW bc Great-horned Owl Bubo virginianus --b SW 1 ---1 ---GREG c Great Egret Ardea alba MB SS 1 ---1 ---LAGU c Laughing Gull Larus atricilla MB GG 6 ---11 ---OSPR c Osprey Pandion haliaetus MB HD 7 ---15 -----cd Peafowl Pavo cristatus --d --d 4 ---7 -----cd Domestic Chicken Gallus domesticus --d --d 4 ---9 ---WHIB c White Ibis Eudocimus albus MB GG 4 ---11 ---39 Non-native Species. ^ State listed species Threatened. a species sited in insufficient numbers for statistical analyses. b species reported not to occur in the Florida Keys accidental or out of range sighting (Florida Fish and Wildlife Conservation Commission 2003). c removed from analysis domestic species or waterbird species uncommon in hammocks. d data unavailable.

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Table 3-4 Influence of measured habitat variables on species presence or absence from analyses with general linear models (data represent p-values). A negative p-value indicates an inverse relationship between the habitat variable and species presence. See Table 3-3 for species codes, Table 2-3 for definition of life history and foraging guild, and Table 2-2 for definition of habitat parameters. Statistical Model a Species Code Life History Foraging Guild PCT_HAM b PCT_CAN b PCT_ GROUND b PCT_ OTHER b D2HAM b SIZEHAM b HAM+CAN c DXSIZE c ZIP RSHA YR SW <0.02 ZIP AMRE NM HG 0.05 <0.02 <0.05 ZIP REVI NM HG <0.01 <0.01 ZIP BTBW NM HG 0.05 ZIP EAKI NM HA (-) <0.02 ( -) <0.02 ZIP GCFL MB HA <0.03 <0.04 (-) <0.01 ZIP GRAK MB HA (-) <0.03 (-) 0.001 ZIP ECDO YR GG (-) <0.01 (-) <0.01 (-) <0.01 (-) <0.01 ZIP NOCA YR GG 0.05 <0.01 <0.01 ZIP COGR YR GG (-) <0.01 (-) <0.01 (-) <0.01 ZIP BLJA YR GG (-) <0.01 (-) <0.04 (-) 0.05 (-) <0.01 (-) <0.04 ZIP EUST YR GG (-) <0.01 (-) <0.01 0.05 (-) <0.01 ZIP RWBL YR GG (-) <0.02 (-) <0.01 <0.01 (-) <0.01 <0.01 ZIP NOMO YR GG (-) <0.01 <0.05 (-) <0.01 ZIP YPWA NM GG <0.01 <0.01 <0.01 ZIP GRCA NM GG <0.05 (-) <0.02 ZIP OVEN NM GG ZIP MODO MB GG (-) <0.02 (-) <0.01 ZIP COGD MB GG ZIP BLPW NM FG <0.04 ZIP COYE NM FG ZIP CMWA NM FG ZIP WEVI MB FG <0.03 <0.01 <0.01 40

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Table 3-4 (continued) Statistical Model a Species Code Life History Foraging Guild PCT_HAM b PCT_CAN b PCT_ GROUND b PCT_ OTHER b D2HAM b SIZEHAM b HAM+CAN c DXSIZE c ZIP PRAW MB FG <0.01 <0.02 <0.01 0.05 (-) 0.02 <0.01 ZIP BWVI MB FG <0.01 <0.01 <0.01 ZIP MACU MB FG (-) <0.04 ZIP RBWO YR BG (-) <0.01 (-) <0.01 (-) <0.03 (-) <0.02 (-) <0.01 (-) <0.01 ZIP BAWW NM BG GLIMMIX YSFL MB GG GLIMMIX BGGN NM FG <0.02 GLIMMIX INBU NM FG (-) <0.02 GLIMMIX WCPI MB FG a Statistical model used to analyze each species. ZIP = zero-inflated Poisson general linear model; GLIMIX = general linear model. b Variables analyzed as covariates in first set of analyses. c Variables analyzed as covariates in second set of analyses. 41 Table 3-5 Influence of measured habitat variables on life history and foraging guilds from general linear model analyses with zero-inflated Poisson regression model (data represent p-values). A negative p-value indicates an inverse relationship between the habitat variable and species presence. See Table 3-3 for species codes, Table 2-3 for definition of life history and foraging guild, and Table 2-2 for definition of habitat parameters. PCT_HAM a PCT_CAN a PCT_ GROUND a PCT_ OTHER a D2HAM a SIZEHAM a HAM+CAN b DXSIZE b YR (-) <0.01 (-) <0.01 (-) 0.02 (-) <0.01 NM <0.01 <0.01 (-) <0.01 <0.01 Life History MB <0.01 <0.01 <0.03 <0.01 BG (-) <0.01 (-) <0.04 (-) <0.02 (-) <0.01 FG <0.01 (-) <0.02 <0.02 <0.01 GG (-) <0.01 (-) <0.01 (-) <0.01 (-) <0.01 HA (-) <0.04 (-) <0.01 HG <0.02 <0.04 (-) <0.04 <0.02 <0.01 Foraging Guild SW a Variables analyzed as covariates in first set of analyses. b Variables analyzed as covariates in second set of analyses

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Table 3-6 Hammock habitat variables for birds that lacked sufficient data for statistical analyses. Data reflect the range of percent cover (smallest to largest) for PCT_HAM, PCT_CAN and HAM+CAN measurements where birds were observed. See Table 3-3 for species codes, Table 2-3 for definition of life history and foraging guild, and Table 2-2 for definition of habitat parameters. Species Code TOTAL Survey Locations at which detected (%) Life History Foraging Guild PCT HAM smallest largest PCT CAN smallest largest HAM+CAN smallest largest CEDW 100 45 1.16 NM FG 0 0 10.8 10.8 10.8 10.8 MAWA 31 8.14 NM HG 0 95.4 0 18.6 0 100 BLBW 15 5.81 NM FG 0 100 0 16.5 5.4 WWDO 14 9.30 NM GG 0 52.0 10.8 42.2 30.9 67.3 MOPA 14 4.65 YR -----0 29.7 0 37.6 0 67.4 WEWA 10 9.30 NM FG 6.0 100 0 6.7 12.7 100 BHCO 9 5.81 NM GG 0 0 0 25.1 0 25.2 YTWA 9 6.98 NM BG 0 100 0 46.5 0 100 CARW 6 4.65 YR GG 89.9 100 0 0 89.9 100 YTVI 6 3.49 NM FG 0 100 0 25.2 25.2 100 HOWA 6 4.65 NM FG 0 100 0 15.0 15.0 100 SCTA 5 2.33 NM HG 38.7 100 0 25.2 64.0 100 WOTH 5 2.33 NM GG 0 92.5 0 0 0 92.5 YBCU 4 3.49 NM FG 100 100 0 0 100 100 SUTA 4 2.33 NM FG 8.3 92.5 0 65.3 73.6 92.5 NOPA 4 1.16 NM FG 35.1 35.0 7.7 7.7 42.8 42.8 NAWA 4 2.33 NM FG 0 100 0 28.5 28.5 100 PABU 4 3.49 NM GG 0 100 0 18.6 18.6 100 RTHU 4 2.33 NM HG 0 0 5.4 38.2 5.4 38.2 MYWA 3 2.33 NM FG 0 100 0 21.0 21.0 100 EAPH 3 2.33 NM HA 0 100 0 0 0 100 SWWA 2 2.33 NM GG 94.4 100 0 0 94.4 100 PROW 2 2.33 NM BG 0 92.5 0 35.8 35.8 92.5 AMKE 2 2.33 NM HO 5.1 38.7 16.5 25.2 21.5 64.0 MERL 2 2.33 NM HA 0 0.6 13.0 35.8 13.5 35.8 42

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43 Table 3-6 (continued) Species Code TOTAL Survey Locations at which detected (%) Life History Foraging Guild PCT HAM smallest largest PCT CAN smallest largest HAM+CAN smallest largest 4.9 8.0 100 BLGR 2 1.16 NM GG 0 0 0 0 0 0 CONI 1 1.16 MB AF 100 100 0 0 100 100 RCKI 1 1.16 NM FG 100 100 0 0 100 100 YBCH 1 1.16 NM FG 100 100 0 0 100 100 YBSA 1 1.16 NM BG 100 100 0 0 100 100 TRES 1 1.16 NM AF 46.6 46.6 30.6 30.6 77.3 77.3 NRWS 1 1.16 NM AF 64.6 64.6 6.2 6.2 70.8 70.8 YWAR 1 1.16 MB FG 35.2 35.2 0 0 35.2 35.2 RBGR 1 1.16 NM FG 0 0 28.5 28.5 28.5 28.5

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Figure 3-1 Area (ha) of tropical hardwood hammock habitat patches by patch size (ha) category and protected status in the Florida Keys in 2004 (number of habitat patches provided above each bar). 44

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CHAPTER 4 DISCUSSION Assessment of Hammock Habitat Almost 31% of tropical hardwood hammock habitat in the upper Florida Keys was lost to deforestation during 1991-2004. Of the remaining 1,962 ha of hammock in the upper Keys, approximately 46% lacked protection. Of the 1,750 ha of hammock left in the lower Keys as of 2004, approximately 27% lacked protection. Therefore, of the 3,712 ha of extant hammock in the Florida Keys, 1,375 ha (36.5%) remains unprotected and subject to development. Partially protected patches of hammock exist in both the upper and lower Keys. These patches present opportunities to increase the size of existing conservation areas. In fact, the 22 partially protected patches >20 ha encompass 750 ha of unprotected habitat, which represents 55% of all remaining unprotected hammock habitat in the Keys. If all of the unprotected portions of these partially protected patches are lost to development, mean patch size would decrease from 89 to 55 ha (Table 3-2). The increasingly patchy hammocks have been and continue to be fragmented by anthropogenic activities, the most conspicuous of which is the construction of roads. The Overseas Highway (US Route 1) follows a trail originally formed by Henry Flagler's Florida East Coast Railroad and stretches from Miami to Key West. The highway was completed in 1938 and today spans 182 km including 42 bridges to allow motorists access from one island to the next. This road and the secondary roads that result from increasing urbanization are a major contributor to the fragmentation of hammocks in the Keys. Although no specific studies have been done targeting the effects of roads on hammocks in the Florida Keys, the impact of roads has been studied extensively by a number of authors in various other systems (Forman et al. 2003). 45

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The presence of a road in, around, or through a habitat has both direct and indirect impacts. Direct impacts include the subdividing of natural areas into smaller habitat blocks, the destruction and alteration of habitats, disturbance, edge effects, invasions by exotic species and, increased roadkill (Goosem 1997; Forman & Alexander 1998). Roads have also been implicated in attracting generalist predators (Forman et al. 2003) and qualitative observations of feral cats (Felis catus) during bird surveys suggested a positive relationship between road density and feral cat abundance in the Keys (A. Karim, personal obs.). The natural island/water landscape of the archipelago lends itself to hammock habitats that are naturally patchy in comparison to other tropical forests. The construction of roads through these areas further increases the edge effects impacting these seasonal deciduous forests. Consequently, resident and migratory birds that rely on these areas must contend with increasing amounts of disturbance and other road-related impacts. Reijnen et al. (1995) found that 60% of the 43 species of woodland species of birds that they investigated showed evidence of reduced densities adjacent to roads. Goosem (1997) found that forest interior birds in the Amazonian rainforest stay at least 50 m away from edges. In the Keys, 50 m is often more than half the width of an entire hammock habitat block, which emphasizes the importance of conserving extant large blocks. Bird Surveys Species specific analyses were the best way to determine habitat use by birds detected during the study (Table 3-4). Grouping avian species by life history helped summarize the predictive effects of some habitat variables on year-round residents, but did a poor job in summarizing predictive effects on Neotropical migrants and migratory breeders (Table 3-5). Similarly, grouping avian species by foraging guild was a poor predictor of how individual 46

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species within those guilds responded to habitat parameters with the exception of the bark gleaner guild. The lack of significant predictive effects of hammock variables on Neotropical migrants and migratory breeders as collective groups is likely due in part due to the fact that migratory species represent many different foraging guilds with unique needs and habitat preferences. However, the lack of significant predictive effects may also be due to the fact that many migratory birds that depend on hammocks during migration are difficult to detect because they are in the Keys for short periods and may be difficult to detect in heavy cover typical of hammock habitat. For example of the 35 species seen infrequently, 94% were migratory and 11 of these species were detected at locations with >50% total tree canopy cover. These analyses suggest that when implementing conservation and management plans for resident birds (from those examined here), grouping them together may be helpful in predicting their occupancy within a landscape, but that implementing plans for migrants should be done on a species specific basis. Of the 32 species detected in sufficient numbers to be analyzed statistically, five are designated as birds of conservation concern (BWVI, PRAW, MACU, WCPI and COGD), one is an opportunistic nest predator (BLJA), and two are non-native (ECDO and EUST; Table 3-3). Although the BWVI showed a significant negative response to PCT_GROUND and no positive effects to any other habitat parameter, it has been noted that hammocks (Robertson & Kushlan 1984; Snyder et al. 1990) in addition to mangrove areas (Ehrlich et al.1988) serve as important breeding areas. The BWVI was only detected within hammocks during this study regardless of hammock location (residential or protected). This suggests that this species prefers hammocks to lawns or open areas, despite a lack of significant positive response to hammock habitat parameters. 47

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The PRAW is usually associated with pinelands and open fields (Ehrlich 1988; Elphick 2001) but analyses revealed that this species also responded positively to the PCT_HAM, PCT_CAN, PCT_GROUND, and HAM+CAN habitat parameters (and negatively to the SIZEHAM variable). Both the BWVI and PRAW have experienced population declines due to brood parasitism from the Shiny Cowbird (Molothrus bonariensis) and the BHCO (Ehrlich 1988; Wiley 1988; Post et al. 1990; Grzybowski 2001). Both cowbird species have been reported in the Florida Keys (Smith & Sprunt 1987; US Fish and Wildlife Service 1999, Sibley 2000, A. Karim personal obs.), and have been implicated in the decline of many Neotropical migratory songbirds (Terbough 1989; Robinson et al. 1993). Specifically, the BHCO is suspected of causing the near extirpation of the BWVI in the Tampa Bay area of Florida (Paul 1987; Paul 1988). Historically, cowbirds were largely restricted to the mid-continental prairies where they apparently followed herds of nomadic bison. Cowbirds mainly search for seeds and insects in short grass and on bare ground and may have depended upon grazing by large ungulates to create suitable feeding areas. However, since the clearing of forests for agriculture and the widespread introduction of livestock, cowbirds have expanded their geographical range (Mayfield 1965). Additionally, cowbirds target species that build open-cup nests (Robinson et al. 1993), which are characteristic of the BWVI and PRAW (Ehrlich 1988). The tropical hardwood hammock multi-species recovery plan (US Fish and Wildlife Service 1999) states that hammocks are important for a number of West Indian land birds including the MACU, BWVI and WCPI. The analyses performed in this study indicated that the MACU showed a negative predictive response to PCT_GROUND, but it did not show a significant positive response to any other habitat parameter. Nonetheless, MACUs were only detected within protected hammocks with no urban matrix during this study. This suggests that hammocks and native tree canopy associated with 48

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hammocks should be considered when devising conservation plans for both the PRAW and MACU. The state threatened WCPI showed no negative or positive predictive effects when modeled against the habitat variables measured within this study. While WCPI nest on small mangrove islands, these obligate frugivores feed within native hammocks and are arguably the most important seed dispersers of 37 species of trees and shrubs within hammocks in the Keys (Strong and Bancroft 1994). In particular, the fruits of the blolly (Guapira discolor), strangler fig (Ficus aurea), and especially poisonwood (Metopium toxiferum) are important to the survival of this species (Bancroft & Bowman 1988; Bancroft et al. 2000). However, the strangler fig is often removed from residential areas due to its extensive root system that interferes with septic systems and poisonwood is removed because its sap causes contact dermatitis in some humans (Scurlock 1987). Bancroft and Bowman (1988) state that, breeding populations [of WCPI] are clearly tied to the existence of sufficient hammocks for production of food. Post fledging dispersal studies of this species show that WCPIs selectively use hammock fragments from 5-20 ha (p < 0.10) during the first 72 hours after nest dispersal; after this stage, dispersing birds show no preference for fragment size but use hammocks more frequently than residential habitat (p < 0.10). Therefore, small hammocks are used as stepping stones by immature WCPIs that are unable to fly long distances when they first leave mangrove islands to forage for food (Strong and Bancroft 1994). The BLJA is noted as a potentially important nest predator on open-nesting songbirds in fragmented habitat (Wilcove 1985). Although a native bird to the Florida mainland, this species is a fairly recent arrival to the Keys. It was first found to be breeding in the upper Keys in 1989 (Florida Fish and Wildlife Conservation Commission 2003), and has been increasing since (J. 49

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Duquesnel, personal comm.). Engels and Sexton (1994) show a strong negative correlation between the presence of BLJAs and the endangered Golden-cheeked Warbler (Dendroica chrysoparia) in central Texas. During my study, the BLJA showed a negative response to three hammock variables (PCT_HAM, SIZHAM and HAM+CAN). Of the 310 BLJAs recorded, 28 were observed within hammocks and BLJAs were detected in only five of 14 hammocks surveyed, all of which were small fragments. This suggests that this opportunistic nest predator and typical edge species may avoid large, intact hammocks and prefer highly fragmented urban areas. Two non-native species were observed that deserve mention. The ECDO is noteworthy primarily because it was the most abundant species recorded during surveys, but showed a significant negative response to hammock and total tree canopy variables (Table 3-3). The EUST is an aggressive competitor for cavity nesting sites and as such may be detrimental to resident and migrant cavity nesters in the Keys. Ingolds (1994) research in east-central Ohio shows that Red-bellied Woodpeckers (Melanerpes carolinus, RBWO) incurred the brunt of [EUST] competition for freshly excavated nest cavities and lost 39% of their cavities to starlings. Flickers were significantly more aggressive than RBWOs when defending their nest cavities. Fourteen percent of flicker cavitieswere usurped by starlings. While conducting research, I observed EUSTs competing with one RBWO pair and two pairs of Northern Flickers (Colaptes auratus) for nest cavities. Statistical analyses suggest that the EUST did avoid hammocks; this species showed a negative response to three hammock variables (PCT_HAM, PCT_CAN and HAM+CAN). Consequently, intact hammock habitat may provide a refuge for cavity nesting species in the Keys. 50

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In addition to cowbird species (Molothrus spp.), BLJAs, and EUSTs, feral cats may pose a substantial threat to resident and migratory birds in hammock fragments. Feral cats were observed at 46 (54%) of survey locations and in areas that ranged from 0-100% hammock cover. Cats were observed stalking and/or attacking resident and migratory birds. The number of feral cats in the Keys is increasing in all probability because of lack of predators and food provided by visitors and inhabitants of the archipelago. The Ocean Reef Club in north Key Largo has been sustaining a colony of approximately 500 feral cats at an annual budget of approximately $100,000 that includes a dedicated veterinarian (America Bird Conservancy 2004). In addition to birds, feral cats prey on other species and may pose a significant threat to the recovery of two federally listed, endangered species, the Key Largo cotton mouse (Peromyscus gossypinus allapaticola) and Key Largo woodrat (Neotoma floridana smallii), both of which depend on hammocks for their survival (US Fish and Wildlife Service, 1999). Further studies are needed in the Keys to add to the knowledge about avian-hammock interactions. Long term surveys of the number and species of birds that use hammocks during spring and fall migrations are needed. Comparisons of fall and spring migration will help land managers get a better picture of the full suite of avian species that rely on hammocks in the Keys. Studies on the impacts of roads in the Keys are needed to understand how birds and other species that require these habitats are affected both directly and indirectly. Conservation Implications The study of species-environment interactions enables scientists to investigate and potentially understand how changing landscapes effect the way species are able to respond and ultimately survive or, alternatively, become extirpated. Bird migration often occurs over a broad geographic scale but over a relatively short temporal scale (Russell et al. 1998, Simons et al. 2000); therefore, designing and implementing conservation strategies to help ensure the survival 51

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of Neotropical migrants (Maurer and Heywood 1993, Keyser et al. 1998) is complicated. Since energy demands during migrations are extreme, migrants must accumulate substantial fat stores to fuel their journey. The objective of migrants is not only to accumulate enough fat to survive the migration, but additionally, to arrive at a breeding ground with enough fat stores to compete for nesting areas and therefore aid in the chance of increased population survival (Smith & Moore 2003). Schaub and Jenni (2001) state that the fuel deposition rate at stopover sites is the crucial factor determining the overall speed of migration and its success. Parrish (2000) suggests that selection for minimization of energy expenditure during stopover has influenced the evolution of dietary plasticity during migration. Dietary shifts to fruit allow many migrant species to minimize the time and energy needed for foraging on expensive diet types such as insects. This dietary opportunism allows a migrant to handle unpredictable fluctuations in food type and availability while en route (Levey & Stiles 1992). As of 2004 most of the extant hammock habitat was located in the upper Keys. There remained areas of unprotected and partially protected hammocks throughout the Keys that could be acquired and incorporated into a network of protected hammocks that run the length of the Florida Keys. This archipelago-wide network could benefit both resident and migratory birds that require hammocks to survive. By simply targeting partially protected hammocks for acquisition, the size of hammock patches >20 ha could be increased by 36% and 79% in the lower and upper Keys, respectively. These hammocks could be utilized by migrants needing stopover areas because of inclement weather or low fat reserves after their trans-gulf migration and provide quality habitats for year-round residents and migratory breeders that require hammock habitat for reproduction. Additionally, these areas would less likely be problematic 52

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from species such as the BLJA and EUST because these species respond negatively to hammock habitat (Table 3-4). Acknowledging the need to acquire areas for conservation is not a sufficient end to preserving the flora and fauna of natural plant communities; economic considerations must also be addressed for conservation plans to work effectively. In Florida, eco-tourism contributes millions of dollars to the state economy. Caudill (2003) reports that visitors and residents of Florida spent over $1.5 billion dollars in activities associated with wildlife viewing, generated more than 35,000 jobs, and created more than $76 million in state sales tax revenue in 2001. Birds such as the BWVI, MACU and WCPI that need hammocks to persist also provide economic benefit for the Keys. Many bird watchers visit the Florida Keys specifically to see these native species (A. Karim pers. obs.; D. Sprunt pers. comm.). Public costs resulting from the construction of residential neighborhoods must also be taken into consideration. According to the Florida Department of Environmental Protection (1997), the cost per person of moving to a previously undeveloped area ranged between $4,400 in Ocala, FL and $8,100 in Naples, FL in public expenditures subsidized by taxes paid by existing residents to the area. In 1997, the estimated public cost for each new single-family home in Florida ranged from $13,640-$25,110. These publicly-subsidized infrastructure costs increase with distance to urban service centers (Florida Department of Environmental Protection 1997) and are likely much higher in the Keys given costs associated with the transport of building materials, limited sources of fresh water, and issues associated with trash and sewage disposal. Conservation and management of hammock habitat and the birds associated with them cannot be accomplished without the education and support of the public. In south Florida, hammocks are one of the easiest forest communities to create (US Fish and Wildlife Service 53

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1999). Environmental education should go hand-in-hand with management activities to increase public awareness and participation in conservation activities (Fernandez-Juricic & Jokimaki 2001) such as creation, conservation and restoration of these communities. Balmford and Cowling (2006) state that ecologists and resource managers need to come to the realization that conservation is primarily not about biology but about people and the choices they make. Many residents of the Keys appreciate the natural environment and public support will be crucial to the future protection of hammocks. Private landowners with substantial hammock habitat may be educated on the value of conservation easements and preservation of their land. Providing educational materials on the ecological importance and economic benefits of hammocks can be the starting point of a comprehensive approach to hammock conservation throughout the Keys (Main et al. 2003; Karim & Main 2004). 54

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LITERATURE CITED American Bird Conservancy. 2004.Managed cat colonies in Florida. American Bird Conservancy. Washington, D.C. Available at: http://www.abcbirds.org/cats/states/florida_intro.htm (accessed November 2004). American Ornithologists Union. 1998. The A.O.U. Checklist of North American Birds, seventh edition. American Ornithologists Union and Allen Press, Inc., Lawrence, Kansas. American Ornithologists Union. 2000. Forty-second supplement to the American Ornithologists Union Checklist of North American Birds. The Auk 117:847-858. Bancroft, G. T. and R. Bowman. 1988. Relationship between the reproductive ecology of White-crowned pigeons and fruiting phenology of tropical hardwood hammock trees: Second Annual Repot. 1988. National Audubon Society, Research Department. Tavernier, Florida. Balmford, A. and R. M. Cowling. 2006. Fusion or failure? The future of conservation biology. Conservation Biology 20:692-695. Bancroft, G. T., A. M. Strong, and M. Carrington. 1995. Deforestation and its effects on forest-nesting birds in the Florida Keys. Conservation Biology 9: 835-844. Bancroft, G. T., R. Bowman, and R .J. Sawicki. 2000. Rainfall, fruiting phenology, and the nesting Season of White-crowned pigeons in the upper Florida Keys. The Auk 117: 416-426. Banks, R. C., C. Cicero, J. L Dunn, A. W. Kratter, P. C. Rasmussen, J. V. Remsen Jr., J. D. Rising, and D. F. Stotz. 2003. Forty-fourth supplement to the American Ornithologists Union check-list of North American birds. The Auk 120:923-931. Barrow, W. C. Jr., C. Chen, R. B. Hamilton, K. Ouchley, and T. J. Spengler. 2000. Disruption and restoration of en route habitat, a case study: the Chenier Plain. Pages 71-87 in Moore, F. R. editor. Stopover ecology of Nearctic-Neotropical landbird migrants: habitat relations and conservation implications. Studies in Avian Biology. Cooper Ornithological Society. Brown, R. B., E. L. Stone, V. W. Carlisle. 1990. Soils. Pages 35-69 in R. L. Myers and J. J. Ewel editors. Ecosystems of Florida. University of Central Florida Press; Orlando, Florida. Caudill, J. 2003. 2001 National and state economic impacts of wildlife watching: addendum to the 2001 national survey of fishing, hunting and wildlife-associated recreation. Report 2001-2. US Fish and Wildlife Service, Arlington, Virginia. Craighead, F. C. Sr., 1971. The trees of south Florida Volume 1: the natural environments and their succession. University of Miami Press, Coral Gables, FL. Ehrlich, P. R., D. S. Dobkin, D. Wheye. 1988. The Birder's Handbook: A field guide to the natural history of North American birds. Simon & Schuster Inc., New York. 55

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Elphick, C., J. B. Dunning Jr, and D. A. Sibley 2001. The Sibley guide to bird life and behavior. Chanticleer Press; New York, NY. Engels, T. M. and C. W. Sexton. 1994. Negative correlation of Blue Jays and Golden-cheeked Warblers near an urbanizing area. Conservation Biology 8:286-290. Fernandez-Juricic, E., and J. Jokimaki. 2001. A habitat island approach to conserving birds in urban landscapes: case studies from southern and northern Europe. Biodiversity and Conservation 10:2023-2043. Florida Climate Center. 2005. The Center for Ocean-Atmospheric Prediction Studies. Florida State University, Tallahassee, FL. Available from http://coaps.fsu.edu/climate_center/ (accessed January 2007). Florida Department of Environmental Protection. 1997. State lands economics of acquisition excerpt from the Florida Preservation 2000 Program remaining needs and priorities study. Tallahassee, FL. Available from http://www.dep.state.fl.us/lands/acquisition/P2000/ECONOMIC.htm (accessed August 2007). Florida Department of Community Affairs. 2003. Final Report of the Florida Keys Carrying Capacity Study Implementation Rule 28-20 Work Group. Tallahassee, FL. Available from http://www.dca.state.fl.us/FDCP/DCP/acsc/final.pdf (accessed May 2006). Florida Department of Environmental Protection. 2003. Land Boundary Information System. Tallahassee, FL. Available from http://data.labins.org/2003/ (accessed May 2005). Florida Fish and Wildlife Conservation Commission. 2003, January 6. Florida's breeding bird atlas: A collaborative study of Florida's birdlife. Available from http://www.myfwc.com/bba/ (accessed August 2004). Forman, R. T. T. 1995. Land Mosaics: The Ecology of Landscapes and Regions. Cambridge University Press, Cambridge, UK. Forman, R. T. T., and L. E. Alexander. 1998. Roads and their major ecological effects. Annual Review of Ecology and Systematics, 29: 207-231. Forman, R. T. T., D. Sperling, J. A. Bissonette, A.P. Clevenger, C.D. Cutshall, V.H. Dale, L. Fahrig, R. France, C.R. Goldman, K. Heanue, J.A. Jones, F.J. Swanson, T. Turrentine, and T.C. Winter. 2003. Road Ecology: Science and Solutions. Island Press, Washington, D.C. Goosem, M. 1997. Internal Fragmentation: The effects of roads, highways, and powerline clearings on movements and mortality of Rainforest vertebrates. Pages 241 255 in W.F. Lawrence and R.O. Bierrgarrd Jr. editors. Tropical Forest Remnants Ecology, Management, and Conservation of Fragmented Communities. University of Chicago Press. Chicago, IL. 56

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BIOGRAPHICAL SKETCH Annisa Karim was born in Nairobi, Kenya on September 13, 1974. She moved to Florida with her parents, Muneer and Laila, and her sister Farah, in 1980. Her lifelong curiosity of the outdoors and her amazement of the natural world led to her to focus on ecology as a career. Annisa received a Bachelor of Science degree in wildlife ecology and conservation and a minor in zoology from the University of Floridas Department of Wildlife Ecology and Conservation in 1997. She worked as a naturalist and then as a research associate in southwest Florida before she ventured to Gainesville once again. She entered the graduate program at the University of Florida in the Fall of 2003 and earned a Master of Science degree from the Department of Wildlife Ecology and Conservation in 2007. 61