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Effects of The Invasive Exotic Apple Snail (Pomacea insularum) on The Snail Kite (Rostrhamus sociabilis plumbeus) in Flo...

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Material Information

Title: Effects of The Invasive Exotic Apple Snail (Pomacea insularum) on The Snail Kite (Rostrhamus sociabilis plumbeus) in Florida, USA
Physical Description: 1 online resource (164 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: exotic, foraging, insularum, invasive, kite, paludosa, pomacea, snail, tohopekaliga, trap
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 Snail Kite (Rostrhamus sociabilis plumbeus) is an endangered raptor in the U.S. that exhibits an extreme form of dietary specialization, feeding almost exclusively on one species of freshwater snail, the Florida Apple Snail (Pomacea paludosa Say). Lake Tohopekaliga, one of the few remaining wetland fragments utilized by the snail kite in Florida, recently experienced an infestation of the invasive exotic Island Apple Snail (Pomacea insularum), which is relatively larger (length, x = 63.5 mm; weight, x = 56.8 g) than the native apple snail (length, x = 37.6 mm; weight, x = 15.9 g). This relative size difference raised questions about the ability of kites (especially juveniles) to negotiate exotic snails, and given the sensitivity of the kite population to recruitment, we conducted a comparative observational study to elucidate the effects of the exotic apple snail on snail kite foraging behavior, energetics, nest success, and survival. Relative to native snails, we found that exotic snails require longer handling times (for adults, 302 vs. 72 seconds; for juveniles, 496 vs. 97 seconds), lead to increased drop rates (for adults, 0.21 vs. 0.02; for juveniles, 0.33 vs. 0.06), and result in depressed capture rates (for adults, 1.09 vs. 3.30 snails/hour; for juveniles, 0.78 vs. 3.46 snails/hour); however, we also found that exotic snails provide more energy than natives (12.92 vs. 4.84 kcal/snail). Consequently, the effects of the exotic snail on foraging behavior do not have negative energetic repercussions for adult kites. In fact, we found that adult kites are attracted to Lake Tohopekaliga and that the relative contribution of the lake to the range-wide nesting effort increased from 6 to 33% after the invasion of the exotic snail. Conversely, the effects of the exotic snail on juvenile foraging behavior can lead to insufficient daily energy balances and may suppress juvenile survival. Given the critically endangered status of the snail kite and the propensity of the exotic apple snail to spread, this work suggests that serious management and conservation initiatives that address the exotic apple snail may be necessary to prevent further deleterious consequences for the kite population in Florida.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Kitchens, Wiley M.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0021750:00001

Permanent Link: http://ufdc.ufl.edu/UFE0021750/00001

Material Information

Title: Effects of The Invasive Exotic Apple Snail (Pomacea insularum) on The Snail Kite (Rostrhamus sociabilis plumbeus) in Florida, USA
Physical Description: 1 online resource (164 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: exotic, foraging, insularum, invasive, kite, paludosa, pomacea, snail, tohopekaliga, trap
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 Snail Kite (Rostrhamus sociabilis plumbeus) is an endangered raptor in the U.S. that exhibits an extreme form of dietary specialization, feeding almost exclusively on one species of freshwater snail, the Florida Apple Snail (Pomacea paludosa Say). Lake Tohopekaliga, one of the few remaining wetland fragments utilized by the snail kite in Florida, recently experienced an infestation of the invasive exotic Island Apple Snail (Pomacea insularum), which is relatively larger (length, x = 63.5 mm; weight, x = 56.8 g) than the native apple snail (length, x = 37.6 mm; weight, x = 15.9 g). This relative size difference raised questions about the ability of kites (especially juveniles) to negotiate exotic snails, and given the sensitivity of the kite population to recruitment, we conducted a comparative observational study to elucidate the effects of the exotic apple snail on snail kite foraging behavior, energetics, nest success, and survival. Relative to native snails, we found that exotic snails require longer handling times (for adults, 302 vs. 72 seconds; for juveniles, 496 vs. 97 seconds), lead to increased drop rates (for adults, 0.21 vs. 0.02; for juveniles, 0.33 vs. 0.06), and result in depressed capture rates (for adults, 1.09 vs. 3.30 snails/hour; for juveniles, 0.78 vs. 3.46 snails/hour); however, we also found that exotic snails provide more energy than natives (12.92 vs. 4.84 kcal/snail). Consequently, the effects of the exotic snail on foraging behavior do not have negative energetic repercussions for adult kites. In fact, we found that adult kites are attracted to Lake Tohopekaliga and that the relative contribution of the lake to the range-wide nesting effort increased from 6 to 33% after the invasion of the exotic snail. Conversely, the effects of the exotic snail on juvenile foraging behavior can lead to insufficient daily energy balances and may suppress juvenile survival. Given the critically endangered status of the snail kite and the propensity of the exotic apple snail to spread, this work suggests that serious management and conservation initiatives that address the exotic apple snail may be necessary to prevent further deleterious consequences for the kite population in Florida.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Kitchens, Wiley M.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0021750:00001


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EFFECTS OF THE INVASIVE EXOTIC APPLE SNAIL (Pomacea: insulalrum) ON THE
SNAIL KITE (Rostrhamus sociabilis plumbeus) IN FLORIDA, USA




















By

CHRISTOPHER CATTAU


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

UNIVERSITY OF FLORIDA

2008

































O 2008 Christopher Cattau






































To Florida









ACKNOWLEDGMENTS

I would like to thank my committee members (Wiley Kitchens, Phil Darby, and Ken

Meyer) for their mentorship, advice, and encouragement. Although not officially on my

committee, Julien Martin deserves the same gratitude, as his guidance and instruction, like that

of my committee, was indispensable to the development and completion of this work. I would

also like to thank all of the graduate students and field technicians who assisted with the

collection and processing of data, including Andrea Bowling, Brian Reichert, Sara Stocco,

Christina Rich, Danny Huser, Will deGravelles, Jean Olbert, Derek Piotrowicz, Nate Richardson,

Wesley Craine, Michaela Speirs, Melinda Conners, Bridget Deemer, Courtney Hooker, Andrea

Ayala, Melissa Desa, Carolyn Enloe, and all of my fellow coworkers at the Florida Fish and

Wildlife Cooperative Research Unit. I would like to express particular gratitude to Brian

Reichert for helping me with survival analyses. This proj ect would not have been possible

without the unwavering dedication that each of the aforementioned individuals expressed for

conservation of the snail kite and the challenging (and often arduous) work each performed to

advance our knowledge of this endangered species. This cooperative effort has allowed us to

elucidate threats to the snail kite population and to increase our understanding of their ecology,

which will lead to more reliable conservation strategies. I am enormously grateful to everyone

involved with this work.












TABLE OF CONTENTS


page

ACKNOWLEDGMENT S .............. ...............4.....


LI ST OF T ABLE S ........._._ ...... .__ ...............8....


LI ST OF FIGURE S .............. ...............9.....


LI ST OF AB BREVIAT IONS ........._._ ...... .... ............... 13..


AB S TRAC T ............._. .......... ..............._ 14...


CHAPTER


1 INTRODUCTION ................. ...............16.......... ......


Study Population................. ..............1
Invasion of the Exotic Snail ................. ...............19........... ...

Population Demography .............. ...............21....
Study Obj ectives and Outline ................ ...............23........... ...

2 EFFECTS OF THE EXOTIC APPLE SNAIL ON SNAIL KITE FORAGING
BEHAVIOR............... ...............27


Back ground ............ ..... ._ ...............28....
Foraging Behavior ............... .... .............2
A Note on Pomacea Measurements .............. ...............30....
The Native Apple Snail .............. ...............30....
The Exotic Apple Snail .............. ...............31....
Previous Foraging Studies .............. ...............32__. ......
Prey availability............... ... ............3
Prey negotiability and drop rate .............. ...............34....
Searching and handling time ................. .......... ...............35. ....
Age and sex effects on foraging behavior............... ...............36
Energy budgets and profitability ................. ...............37................
Economic models of foraging behavior ................. .......................... ..........38
Predictions ................ ...............39.................
Methods ............... .. .......... ..............4
Time Activity Bud gets .............. ...............40....
Sampling locations .............. ...............41....
Selection of focal individuals ................ ..................... ....................41
Timing and duration of observations .............. ...............42....
Classification of individual kites ................. ...............43................
Coll section of behavioral data .............. ...............45....
Assumptions of behavioral ob serv ations............... ..............4












A pple Snails............... ..... .... .. ..... .......4
Collection and measurement of empty snail shells .............. .....................4
Collection and processing of live snails ................ .......................... ......48
Statistical Analysis and Confidence Intervals .............. ...............49....
Analysis of Foraging Behavior ................. ...............49................
Searching and handling time ................ ...............51................
Drop rate............... ... .. .. .... .......5
Capture rate (i.e., consumption rate) ....__. ...._.._.._ ......._.... ...........5
Analysis of Energetics ........._...... .... .._._._ ....._. .. .... ..............5
Regression models and estimation of average snail weight and caloric content......53
Profitability............... .. ....... ..........5
Daily activity times and energy budgets .............. ...............54....
Re sults............ ..... .. ...............56...
D iet .............. ... ...... .... .........5
Nutritional content of apple snails .............. ...............57....
Live exotic snail measurements .............. ...............57....

Empty snail shell dimensions ............ .....___ ...............58..
Estimates of total snail weight .............. ...............58....
Foraging Behavior..................... ..............5
Comparing adults, subadults, and juveniles ....._____ ..... ...___ ........__.....59
Effects of the exotic snail on average searching times............... ..................6
Effects of the exotic snail on average handling times ....._____ ..... ......__.........60
Effects of the exotic snail on drop rates ............_...... .__ .....__ .........6
Effects of the exotic snail on capture rates............... ...............62.
Effects of the Exotic Snail on Energetics ....__ ......_____ .......___ ..........6
Estimates of caloric content .............. ...............63....
Profitability............... ... ... ...................6
Validation of time activity budget extrapolations .....__.___ ........_._ ..............64
Daily activity patterns .............. ...............65....
Daily energy balances .............. ...............65....
Discussion .........._...._. ...............66..._.... ......


3 DEMOGRAPHIC EFFECTS OF THE EXOTIC APPLE SNAIL ON THE KITE.............1 04


Introducti on ................. ...............104____.......

Reproduction .............. ............... 105...
Survival ............ ..... .. ...............105...
Predictions ............ _...... ._ ...............106...
M ethods ................ .. ....... .. ............10
Nesting Effort and Reproductive Success ............_...... ._ .....___..........10
Survival ............ ..... .. ...............111...
Re sults............ ..... .. ...............113...

Reproduction ............ _...... ._ ............... 113...
Nesting effort............. __..... ... ._ ...............113...
Nest success and productivity ............ .....___ ...............114.












Survival ................ . ....... ...............114......

Apparent annual survival ................. ...............114...............
Apparent monthly survival ................. ......... ......... ............ 1
Discussion ................. ...............116................


4 IS LAKE TOHOPEKALIGA FUNCTIONING AS AN ECOLOGICAL TRAP FOR
THE SNAIL KITE IN FLORIDA? ............. ...............137....


Introducti on ................... ...............137......... ......
Ecological Trap Theory ................. ...............137................

A Hypothetical Example .............. ...............137....
Recent Nesting History ................. ...............13. 8......... ....
Lake T ohopekal iga ..........._.._. ..............._ 13....._.. 9....
M ethods .............. ...............139....

Hypothesis 1 .............. ............... 13 9...
Hypothesis 2 ..........._.._. ...............140.._...........
Hypothesis 3 .............. ...............140....
Re sults..........._.. ....._. _. ...............141.....

Hypothesis 1 .............. ............... 141...
Hypothesis 2 ..........._.._. ...............142.._...........
Hypothesis 3 .............. ...............142....
Discussion ..............__......._. ...............143....


APPENDIX DERIVATION OF DAILY ENERGY EXPENDITURE ..........._.._.. .........._.._.....152


LIST OF REFERENCES ............. ...... ._ ...............153...


BIOGRAPHICAL SKETCH ............. ......___ ...............164...










LIST OF TABLES


Table page

2-1 Number of TABs conducted by month and location in 2003, 2004, 2005, 2006 (post-
invasion) and in 1993, 1994, 1996 (pre-invasion). ............. ...............71.....

2-2 Nutritional contents of P. insularum ........._..._._........_ ...............71...

2-3 Nutritional composition of P. insularum versus P. paludosa..................... ...............7

2-4 Average daily activity pattern of the snail kite in Florida. ........._. .... ....___...........72

3-1 The number of nests initiated range-wide and the contribution from Toho, 1995-2007.121

3-2 Model selection table for apparent annual survival of kites hatched on Toho. ...............121

3-3 Model selection table for apparent monthly survival of kites hatched on Toho. .............121










LIST OF FIGURES


Figure page

1-1 Maj or habitat fragments within the range of the snail kite in Florida .............. ..... .........._24

1-2 Confirmed locations, as of 2007, of the exotic snail (P. insularum) within the range
of the snail kite in Florida. Larger circles encompass multiple small populations in
close proximity............... ...............2

1-3 Movements of approximately 50 radio-tagged adult snail kites among wetland
fragments over a one-year period. April 1992-April 1993 .............. ....................2

2-1 Relative sizes of exotic apple snails............... ...............73.

2-2 Measurement of snail shells. ........... ........ _. ...............74....

2-3 Location of apple snail traps on Toho............... ...............74..

2-4 Linear regression models of exotic snail weight and morphology. ............. ..................75

2-5 Number of native and exotic apple snails captured per year with funnel traps on
Toho, 2005-2007............... ...............7

2-6 Proportion of native to exotic apple snails captured in five locations on Toho, 2005-
2007............... ...............77..

2-7 Average length of apple snail shells collected from feeding perches and nests in nine
wetlands throughout the range of the snail kite, 2004-2007. ........._.._..... ..._.._.........78

2-8 Average shell dimensions of native and exotic apple snails consumed by kites, 2004-
2007............... ...............79..

2-9 Estimated whole weight of native and exotic snails negotiated by foraging kites
during the post-invasion era. ............_. ...._... ...............80...

2-10 Average handling times for adult, juvenile, and subadult kites on Toho and Kiss-
WCA3A during the post-invasion era ................. ...............81........... ...

2-11 Average drop rates for adult, juvenile, and subadult kites on Toho and Kiss-WCA3A
during the post-invasion era ................. ...............82................

2-12 Average capture rates for adult, juvenile, and subadult kites on Toho and Kiss-
WCA3A during the post-invasion era ................. ...............83........... ...

2-13 Average searching times for adult, subadult, and juvenile kites on Toho and Kiss-
WCA3A during the post-invasion era ................. ...............84........... ...










2-14 Average searching times for adult male and female kites on Toho and Kiss-WCA3A
during the post-invasion era ................. ...............85................

2-15 Average searching times for adult kites on Kiss and WCA3A during the post-
invasion era and on Toho during the pre- and post-invasion eras. ............. ...................86

2-16 Average searching times for juvenile kites on Toho and Kiss-WCA3A during the
post-invasion era. .............. ...............87....

2-17 Average handling times for adult male and female kites on Toho and Kiss-WCA3A
during the post-invasion era ................. ...............88................

2-18 Average handling times for adult kites on Kiss-WCA3A during the post-invasion era
and on Toho during the pre- and post-invasion eras............... ...............89..

2-19 Average handling times for juvenile kites on Toho and Kiss-WCA3A during the
post-invasion era ................. ................ ........ ......... ........ ......... .90

2-20 Raw group drop rates for adult and juvenile kites on Toho and Kiss-WCA3A during
the post-invasion era .............. ...............9 1....

2-21 Average drop rates for adult male and female kites on Toho and Kiss-WCA3A
during the post-invasion era............... ...............92..

2-22 Average drop rates for adult and juvenile kites on Toho and Kiss-WCA3A during the
post-invasion era. .............. ...............93....

2-23 Average capture rates for adult male and female kites on Toho and Kiss-WCA3A
during the post-invasion era............... ...............94..

2-24 Average capture rates for adult kites on Kiss and WCA3A during the post-invasion
era and on Toho during the pre- and post-invasion eras. ................. .................9

2-25 Average capture rates for juvenile kites on Toho and Kiss-WCA3A during the post-
invasion era. .............. ...............96....

2-26 Estimated energetic content of native and exotic snails consumed by kites during the
post-invasion era ................. ................ ........ ......... ........ ......... .97

2-27 Average proportion of time spent flying during day-long and hourly observations by
kites on Toho, Kiss, and WCA3A in the post-invasion era. ............... ...................9

2-28 Average capture rates achieved by foraging kites during morning, afternoon, and
evening observations on Toho and Kiss-WCA3A in the post-invasion era. .....................99

2-29 Average proportion of time spent flying by all kites on Kiss, Toho, and WCA3A
during the post-invasion era............... ...............100..











2-30 Average proportion of time spent flying by adult and juvenile kites on Toho and
Kiss-WCA3A during the post-invasion era. ................ .......... .......................101

2-3 1 Estimated gross daily energetic gains for adult and juvenile kites foraging on Toho,
Kiss, and WCA3A during the post-invasion era ................. ..............................10

2-32 Estimated daily energy balances for adult and juvenile kites on Toho, Kiss, and
WCA3A during the post-invasion era ................. ...............103........... ...

3-1 Annual number of snail kite nests initiated range-wide, 1995-2007. ............. .................122

3-2 Average number of nests initiated annually on Toho during the pre- and post-
invasion eras. Post-invasion estimate excludes 2004.. ............ ...............123.....

3-3 Number of nests initiated annually on Toho, 1995-2007. ................ ........... ..........124

3-4 Relative annual contribution of Toho to the total nesting effort range-wide, 1995-
2007............... ...............125.

3-5 Trend model expressing the increasing contribution of Toho to the total population
nesting effort over time (includes all years 1995-2007). ................... ............... 12

3-6 Trend model expressing the increasing contribution of Toho to the total population
nesting effort over time (excludes drought years 2001, 2007 and drawdown year
2004) .............. ...............127....

3-7 Relative contribution of Toho to the total population nesting effort during the pre-
and post-invasion eras............... ...............128.

3-8 Annual nest success on Toho, 1995-2007............... ...............12

3-9 Average nest success on Toho during the pre- and post-invasion eras............................130

3-10 Annual number of young fledged per successful nest on Toho, 1995-2007. .................. 13 1

3-11 Average nest productivity on Toho during the pre- and post-invasion eras. ...................132

3-12 Average number of young fledged per successful nest on Toho vs. all other wetlands
during the post-invasion era............... ...............133..

3-13 Apparent annual survival of juveniles hatched on Toho during the pre- and post-
invasion eras............... ...............134.

3-14 Apparent annual survival of juveniles hatched on Toho, 1992-2006. ............. ................135

3-15 Apparent monthly survival of juveniles hatched on Toho during the pre- and post-
invasion eras.............. .............136...











4-1 Average number of nests initiated annually on nine wetlands during the post-
invasion era. .............. ...............147....

4-2 Annual nest success on nine wetlands during the post-invasion era. ............. ................148

4-3 Annual nest productivity on nine wetlands during the post-invasion era. ................... ....149

4-4 Total number of nests initiated annually on nine wetlands during the post-invasion
era. ............ ...... ._ ...............150...

4-5 Total number of young fledged per year (uncorrected counts) on nine wetlands
during the post-invasion era. ........._.._.. ...._... ...............151...









LIST OF ABBREVIATIONS

Ninety-five percent confidence interval

American Ornithologists' Union

Degrees of freedom

Florida Fish and Wildlife Conservation Commission

Lake Kissimmee

Sample size

Standard Deviation

South Florida Water Management District

Time Activity Budget

Lake Tohopekaliga

United States Army Corp of Engineers

United States Fish and Wildlife Service

Water Conservation Area 3A

Sample mean


95% CI

AOU

df

FFWCC

KISS (or Kiss)



SD

SFWMD

TAB

TOHO (or Toho)

USACE

USFWS

WCA3A









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

EFFECTS OF THE INVASIVE EXOTIC APPLE SNAIL (Ponmacea: insulalrun2) ON THE
SNAIL KITE (Rostrhanzus sociabilis plunmbeus) IN FLORIDA, USA

By

Christopher Cattau

May 2008

Chair: Wiley Kitchens
Major: Wildlife Ecology and Conservation

The Snail Kite (Rostrha~nus sociabilis phenbeus) is an endangered raptor in the U.S. that

exhibits an extreme form of dietary specialization, feeding almost exclusively on one species of

freshwater snail, the Florida Apple Snail (Pontacea pahedosa Say). Lake Tohopekaliga, one of

the few remaining wetland fragments utilized by the snail kite in Florida, recently experienced an

infestation of the invasive exotic Island Apple Snail (Pontacea insularunt), which is relatively

larger (length, x = 63.5 mm; weight, x = 56.8 g) than the native apple snail (length, x = 37.6 mm;

weight, x = 15.9 g). This relative size difference raised questions about the ability of kites

(especially juveniles) to negotiate exotic snails, and given the sensitivity of the kite population to

recruitment, we conducted a comparative observational study to elucidate the effects of the

exotic apple snail on snail kite foraging behavior, energetic, nest success, and survival.

Relative to native snails, we found that exotic snails require longer handling times (for

adults, 302 vs. 72 seconds; for juveniles, 496 vs. 97 seconds), lead to increased drop rates (for

adults, 0.21 vs. 0.02; for juveniles, 0.33 vs. 0.06), and result in depressed capture rates (for

adults, 1.09 vs. 3.30 snails/hour; for juveniles, 0.78 vs. 3.46 snails/hour); however, we also found

that exotic snails provide more energy than natives (12.92 vs. 4.84 kcal/snail). Consequently, the

effects of the exotic snail on foraging behavior do not have negative energetic repercussions for









adult kites. In fact, we found that adult kites are attracted to Lake Tohopekaliga and that the

relative contribution of the lake to the range-wide nesting effort increased from 6% to 33% after

the invasion of the exotic snail. Conversely, the effects of the exotic snail on juvenile foraging

behavior can lead to insufficient daily energy balances and may suppress juvenile survival.

Given the critically endangered status of the snail kite and the propensity of the exotic apple snail

to spread, this work suggests that serious management and conservation initiatives that address

the exotic apple snail may be necessary to prevent further deleterious consequences for the kite

population in Florida









CHAPTER 1
INTTRODUCTION

Species dependent upon specialized niches are particularly subj ect to environmental

perturbations (Brown & Maurer 1989; Owens & Bennett 2000; Purvis et al. 2000): narrow

dietary breadth is one such specialization (Real & Caraco 1986; Begon et al. 1996). In fact,

niche specialization is positively correlated with extinction risk across a wide range of taxa

(Owens & Bennett 2000; Purvis et al. 2000), including birds (Hughes et al. 2000). The snail kite

(Rostrha~nus sociabilis phenbeus) is a wetland-dependent raptor that displays an extreme form of

dietary specialization, feeding almost exclusively on a single species of apple snail (Pontacea

pahedosa Say) (Howell 1932; Stieglitz & Thompson 1967), the only species of this genus native

to Florida (Rawlings et al. 2007). After several decades of landscape fragmentation and

hydroscape alteration, the kite population is now confined to a patchwork of freshwater wetlands

that remain within its historical range, and the viability of the population rests entirely on the

conditions and dynamics of these wetland fragments (Sykes 1979, 1987a; Bennetts & Kitchens

1992, 1997; Martin 2007). However, many of the remaining wetlands are no longer sustained by

the natural processes under which they evolved (USFWS 1999; RECOVER 2005), and hence,

are not necessarily characteristic of the historical ecosystems that once supported the kite

population (Bennetts & Kitchens 1992, 1997, 1999; Martin 2007). Snail kites now face another

potential threat, the recently established populations of the invasive exotic apple snail (Pontacea

insularunt) in Florida.

Evidence suggests that the exotic apple snail negatively affects the food handling ability of

snail kites (Darby et al. 2007), and there is speculation as to what the consequences for the kite

population may be (Darby et al. 2007; Rawlings et al. 2007), but many unanswered questions

remain. The snail kite already faces a high risk of extinction (Martin 2007), so the complications










experienced by kites attempting to exploit exotic apple snails are of particular concern, especially

given that populations of the native apple snail have been declining throughout the kites' range

(Darby et al. 2005) while populations of the exotic snail in Florida have been spreading

(Rawlings et al. 2007). Native snail populations have declined largely in the absence of exotic

snails; hence, these population trends are likely coincidental, but they are concerning

nonetheless. Both snail species have characteristics that may influence their utility to snail kites.

The elucidation of these differences, and of their respective effects on snail kite foraging

behavior, energetic, and population demography, is an essential prerequisite to future

conservation planning.

Study Population

Snail kite (Rostrha~nus sociabilis) populations occur in North, Central, and South America.

Based on apparent geographical variations in body size, distinctions are drawn among three

subspecies (i.e., R. s. major, R .s. sociabilis, and R. s. phenabeus), which are recognized by the

American Ornithologists' Union (AOU) (Amadon 1975). However, only a small number of

morphometric measurements were used to separate these subspecies, and the tenuous

methodology has been called into question by numerous authors (e.g., Beissinger 1988, Bennetts

& Kitchens 1997, Martin 2007). While the AOU states that R. s. phenabeus occurs in both the

United States and Cuba (Amadon 1975), there is no documentation of movement between these

two geographic locations (Bennetts & Kitchens 1997; Martin 2007). Furthermore, the

aforementioned classification scheme of subspecies, which pools the Florida and Cuba

populations together, has no genetic basis (Beissinger 1988; Martin 2007). Although no study

has directly addressed the relationship between the Florida and Cuba snail kite populations,

multiple studies (e.g., Sykes 1979; Beissinger et al. 1983; Bennetts & Kitchens 1997; Dreitz et

al. 2002; Martin et al. 2006a) of movement and population dynamics suggest that the snail kite









population in Florida is isolated. This study addresses the Florida snail kite population only, and

for all practical conservation purposes, we also consider this population geographically isolated.

The snail kite (Rostrhantus sociabilis phenbeus) is an endangered raptor (Federal Register

1967, 2007) whose range in the U.S. is confined to the freshwater wetlands of central and

southern Florida (Sykes 1984; Bennetts & Kitchens 1992, 1997; Martin 2007). As an extreme

dietary specialist, dependent almost entirely on a single species of freshwater apple snail

(Pontacea pahedosa, from here on referred to as the native snail or native apple snail) for food,

the snail kite is a wetland-dependent species (Howell 1932; Stieglitz & Thompson 1967; Snyder

& Snyder 1969; Sykes 1987a). In addition to foraging, nesting is also tied directly to wetland

habitats. Snail kites always build nests in vegetation surrounded by standing water, which aids

in the deterrence of terrestrial predators (Stieglitz & Thompson 1967; Sykes 1987b; Beissinger

1988). Over the past several decades, landscape fragmentation and hydroscape alteration have

severely j eopardized the quantity and quality of the historically contiguous wetland habitats that

once comprised the range of the snail kite in Florida (Bennetts & Kitchens 1997, 2000; Kitchens

et al. 2002; Martin 2007). The major remaining wetland fragments used by the snail kite are

depicted in Figure 1-1 and include the following: Everglades National Park (ENP), Big Cypress

National Preserve (BICY), Water Conservation Areas (WCA) 1A, 2A, 2B, 3A, 3B, Lake

Okeechobee (OKEE), Grassy Waters Preserve (GW), Saint John's Marsh (SJM), Lake

Kissimmee (KISS), Lake Tohopekaliga (TOHO), and East Lake Tohopekaliga (ETOHO)

(Bennetts & Kitchens 1997; Dreitz 2000; Kitchens et al. 2002; and Martin et al. 2006a). As of

2007, only one of the maj or wetlands utilized by the snail kite, Lake Tohopekaliga (from here on

referred to simply as Toho), has suffered a major invasion ofP. insularunt (from here on referred

to as the exotic snail or exotic apple snail) (Rawlings et al. 2007; Darby et al. 2007); however,









confirmed exotic snail populations exist in close proximity to many of the other primary

wetlands used by the kite in Florida (Rawlings et al. 2007) (Figure 1-2).

Within Florida, the kite population is described as nomadic, and monthly movement

probabilities among wetland fragments may be as high as 0.25 (Bennetts & Kitchens 1997,

2000). More recent analyses, which include multiple levels of spatial and temporal resolution,

suggest that the monthly movement probability among contiguous fragments differs significantly

from that among isolated fragments (0.29 vs. 0. 10, respectively) (Martin et al. 2006a, 2007b).

However, numerous studies, of both movement (Sykes 1979; Rodgers et al. 1988; Bennetts &

Kitchens 1992, 1997; Martin 2007) and genetics (Rodgers & Stangel 1996), confirm that the

spatial distribution of kites in Florida shifts temporally and that sufficient individual movement

among wetlands occurs, thus uniting the entire Florida population (Figure 1-3). Even though the

exotic snail has thus far only infested one of the maj or wetlands utilized by the snail kite, the kite

population in Florida does not function as a metapopulation (Bennetts & Kitchens 1997, 2000;

Martin et al. 2006b, 2007b; Martin 2007); therefore, the scope of our proj ect covers the entire

kite population.

Invasion of the Exotic Snail

The exotic apple snail (P. insularunt), commonly known as the island apple snail, is native

to Argentina, Brazil, and Bolivia. The invasion history of the exotic apple snail is somewhat

unclear (Rawlings et al. 2007). P. insularunt, along with other members of the Pontacea genus

that have invaded the U.S., have historically been misidentified as P. canaliculata or lumped

together with other species in what was called the Canaliculata (or, channeled apple snail)

complex (Thompson 1997; Howells et al. 2006; Rawlings et al. 2007). Reliable genetic analysis

confirming the presence ofP. insularunt in Florida dates back to only 2002 (Rawlings et al.

2007); however, exotic snails observed on Toho in 2001 were later identified as P. insularunt










(Darby, personal communication). Many new populations of P. insularum were observed in

Florida from 2004 to 2006, suggesting that it may be a relatively new invasive species to the

state (Rawlings et al. 2007).

The exotic apple snail is known to feed on the eggs of congeneric snails, and some

anecdotal evidence suggests that native apple snail populations appear to decline or even

disappear after the introduction of the exotic snail (Rawlings et al. 2007); however, native snail

populations have declined largely in the absence of exotic snails, and the declining native snail

populations have been attributed to other sources, such as altered hydrologic regimes and shifts

in vegetative communities (Kushlan 1975; Turner 1996; Darby et al. 2003).

The ecology of Toho has undergone drastic anthropogenic changes recently. As part of a

management strategy aimed at improving habitat conditions for fish populations, a drawdown of

the lake stage occurred between late-2003 and early-2004, and an intensive scraping treatment of

littoral vegetation followed. After this treatment, water levels on Toho remained low until June

2004. This course of action significantly altered the physical and biological conditions of the

lake's littoral zone (Welch 2004; Williams et al. 2005). The emergent vegetation in the littoral

zone is a critical component of suitable snail kite foraging habitat (Sykes 1987a; Bennetts &

Kitchens 1997), and once it was removed via mechanical scraping, kites temporarily abandoned

Toho (Martin et al. 2003; Kitchens et al. 2005), subsequently returning in late-2004 (Kitchens et

al. 2005; Martin et al. 2006b). In the interim, the distribution and abundance of exotic apple

snails on Toho grew dramatically (Kitchens et al. 2005). Prior to the drawdown and scraping

treatment, native snails dominated the littoral habitats on Toho, while the exotic snails were

largely confined to the pelagic habitats on the interior side of the lake. However, the drawdown

and scraping treatment, as well as the prolonged dry conditions that followed it, greatly









suppressed the population of native snails on Toho. When the lake stage rose, the exotic snail

population spread out and invaded the littoral zone (Darby, personal communication). Since

2003, the exotic snail has been the most abundant apple snail in portions of the lake utilized by

snail kites (Kitchens et al. 2007).

Population Demography

Since 1992, the snail kite population has been monitored annually via robust population

surveys, which include extensive mark-recapture techniques. Reliable estimates of abundance,

survival, and movement are attainable from these data by utilizing multi-state models, such as

Cormack-Jolly-Seber models (CJS), which incorporate detection probabilities and spatial

variation (Bennetts & Kitchens 1997; 1999; Dreitz et al. 2002; Martin et al. 2006a, 2007a).

Estimates of the snail kite population size and growth rate for the years 1992 to 2005 show

several alarming trends. During the period 2000 to 2002 the snail kite population essentially

halved, falling from around 3500 to around 1700 individuals (Martin et al. 2007a). The most

recent demographic analyses show no signs of a significant population rebound. In fact, the

estimated stochastic population growth rate for the post-population-decline period (i.e., 2003-

2006) is less than one (Martin 2007; Martin et al. 2007a). As indicated by a life table response

experiment (LTRE), over 80% of the reduction in the stochastic population growth rate after

1998 is attributable to adult fertility (Martin 2007). Finally, the most recent snail kite population

viability analysis (PVA) shows that extinction within the next 60 years is highly probable if

current conditions are representative of future conditions. (For a detailed discussion of snail kite

population viability analyses see Martin 2007). Since the snail kite population is at risk of

extinction and because adult fertility plays such an overwhelming role in the population growth

rate, it is critical to identify and attempt to remedy all factors that negatively affect snail kite

fertility in order to properly manage for the conservation of the species









Adult fertility is simply the product of 1) the number of juveniles produced per adult

during the breeding season and 2) the probability that these juveniles survive until the next

breeding season (i.e., juvenile survival) (Martin 2007). Using the aforementioned demographic

trends, the recent history of the snail kite population in Florida can be divided into two time

periods: 1) the pre-1998 period (including 1998), which is representative of a stable population

in terms of demographic trends and environmental conditions, and 2) the post-1998 period (not

including 1998), which is representative of the population decline and subsequent suppression

(Martin 2007). The snail kite population, which was in a steady state of decline throughout the

period 1999 to 2002, has since stabilized. Adult and juvenile survival have, on average, resumed

levels comparable to the pre-1998 period, but nesting efforts and fecundity remain significantly

lower. One anomaly is the unprecedented flurry of nesting attempts that have taken place on

Toho since the invasion of the exotic apple snail. In spite of these nesting efforts, preliminary

analyses show less than expected recruitment from kites on Toho during the post-invasion era

(Martin et al. 2007c), suggesting that the exotic snail may negatively affect nest success or

juvenile survival.

The invasion of Toho by the exotic apple snail has created unique conditions, in terms of

available prey species, for kites. Any deleterious effects that exotic snails have on snail kite

foraging behavior may, in turn, have severe demographic repercussions. Notwithstanding the

very real possibility that the exotic snail will spread into other wetlands utilized by the snail kite,

the fact that the kite population in Florida does not function as a metapopulation implies that

negative effects experienced by kites on Toho may have population-wide consequences,

particularly given the sensitivity of the population growth rate to fecundity and recruitment.









Study Objectives and Outline

The government agencies responsible for managing natural resources (e.g., FFWCC,

SFWMD, USFWS, USACE) exhibit tremendous influence over the remaining wetlands that

comprise the range of the snail kite in Florida (Bennetts & Kitchens 1997; USFWS 1999;

RECOVER 2005). Water regulation schedules are used to manage the composition and

dynamics of wetland ecosystems on a regional scale. The hydrology is dictated via a complex

network of levees, canals, and other water control structures; and smaller-scale management

actions targeted at specific flora and fauna are often conducted via mechanical and chemical

controls (USFWS 1999; Welch 2004). It is the goal of the multi-billion dollar restoration

initiative, the Comprehensive Everglades Restoration Plan (CERP), to create a management

paradigm that mimics the historical, natural-flow patterns and community compositions that once

drove and maintained the integrity and biodiversity of Florida' s natural wetland ecosystems

(USFWS 1999).

Understanding the effects that the exotic apple snail has on snail kite foraging and

demography precludes our ability to strategically manage Florida's wetlands in a way that

sustains overall ecosystem health and alleviates any detrimental consequences to the kite

population. As stated by the CERP, "The desired restoration condition for the Everglade snail

kite is to restore and maintain a network of snail kite foraging habitats and promote habitat that

supports primary prey (apple snails) recruitment throughout the South Florida ecosystem"

(USFWS 1999; RECOVER 2005). To help reach these goals, we conducted a comparative

observational study focused on discerning the effects of the exotic apple snail on the snail kite

population. We hope that this study will have important applications in both species

management and conservation biology:








































Figure 1-1. Maj or habitat fragments within the range of the snail kite in Florida
(modified from Dreitz 2000).


Southern Range









Northern Range


1. Everglades National Park (ENP)
2. Big Cypress National Preserve (BICY)
3. Water Conservation Area 1A (WCA1A)
4. Water Conservation Area 2A (WCA2A)
5. Water Conservation Area 2B (WCA2B)
6. Water Conservation Area 3A (WCA3A)
7. Water Conservation Area 3B (WCA3B)
8. Lake Okeechobee (OKEE)
9. Grassy Waters Preserve (GW)
A. Saint John's Marsh (SJM)
B. Lake Kissimmee (KISS)
C. Lake Tohopekaliga (TOHO)
D. East Lake Tohopekaliga (ETOHO)


B wam



































Figure 1-2. Confirmed locations, as of 2007, of the exotic snail (P. insularum) within the range
of the snail kite in Florida. Larger circles encompass multiple small populations in
close proximity (modified from Rawlings et al. 2007).














































Figure 1-3. Movements of approximately 50 radio-tagged adult snail kites among wetland
fragments over a one-year period. April 1992-April 1993 (modified from Bennetts &
Kitchens 1997).









CHAPTER 2
EFFECTS OF THE EXOTIC APPLE SNAIL ON SNAIL KITE FORAGINTG BEHAVIOR

An animal's fitness is inextricably linked to its diet. The ability to meet certain energetic

and nutritional thresholds is a requisite for survival, growth, and reproduction. The options

available to an individual attempting to meet these energetic requirements are dictated by its

physical and behavioral traits and by the environment in which it lives. An animal's dietary

breadth is therefore constrained by its genetic makeup and by the resources presently available

(Fox 1981; Pyke 1984; Stevens & Krebs 1986; Ehlinger 1990). In ecology, animal species are

often broadly classified as generalists and specialists. Generalists typically incorporate a wide

range of prey types into their diets and utilize varying tactics to attack, subdue, and consume

different prey. Dietary specialists, on the other hand, have narrow dietary breadths and often

utilize acute skill sets that facilitate the efficient exploitation of specific prey types (McArthur

1972; Pyke 1984; Stevens & Krebs 1986). Adaptations facilitating the ability of a species to

exploit specific food resources may hinder the ability of that species to exploit alternative prey

types. Antagonistic pleiotropy, the positive genetic feedback loop through which such inhibition

may occur, can lead to extreme dietary specialization in some instances (Futuyma & Moreno

1988; Cooper & Lenski 2000). Thus, an organism's current behavioral traits and dietary options

are restricted, to an extent, by morphological or physiological characteristics consequent of past

adaptations (McArthur 1972; Pyke 1984; Futuyma & Moreno 1988; Fry 1990; Holt 1996). The

snail kite (Rostrhamnus sociabilis phembeus) is one such extreme dietary specialist, and its unique

physical traits and foraging behaviors are tied to its nearly exclusive prey, the Florida apple snail

(Pomacea pahedosa Say) (Cottom & Knappen 1939; Stieglitz & Thompson 1967; Snyder &

Snyder 1969; Sykes 1987a). The extreme form of dietary specialization observed in the snail

kite has led some authors to question the ability of the kite to efficiently incorporate a newly









introduced exotic apple snail, Pomacea insularum, into its diet (Rawlings et al. 2007; Darby et

al. 2007).

Background

Foraging Behavior

Numerous authors (e.g., Holgerson 1967; Snyder & Snyder 1969; Haverschmidt 1970;

Collett 1977; Beissinger 1983; Sykes 1987a) have described the details of snail kite foraging

behavior, and here we provide a brief synopsis of those details pertinent to our study. Snail kites

exercise two methods of foraging: course-hunting and perch-hunting. While course-hunting,

kites usually fly between 1.5 to 10.0 meters above the surface of the water with their heads

angled downward looking for snails. These flights follow a slow flap-glide pattern and are

usually directed into the wind, when present. Upon sighting a snail, kites will make an abrupt

turn and descend toward the water, hovering briefly just above the surface while reaching into

the water with the talons to capture the snail. Snails are always captured with the talons, but they

may be transferred into the bill during flight (Snyder & Snyder 1969; Beissinger 1983; Sykes

1987a). Kites will then return to a feeding perch in order to manipulate and consume the snail.

Using the talons, kites position the snail shell so that the aperture is facing away from the perch

and the spire is pointed downward. Kites pry open the hard operculum of the snail using the

torque generated by their upper mandible being pressed against the shell wall. The sharp tip of

the upper mandible is then used to cut the muscle attachments of the operculum. After the

operculum is removed, kites must further negotiate the shell, repositioning it so that the aperture

is facing upward then inserting their curved upper mandible into the spiraling aperture to cut the

columellar muscle that connects the snail's body to the shell. The soft body tissue of the snail is

removed and consumed either whole or in torn parts. The albumen gland of female snails is

often discarded, and the viscera of snails is sometimes eaten and sometimes not. The shell is









usually dropped after extraction, but sometimes kites will hold onto the shell until consumption

is complete. Shells of consumed snails frequently accumulate under feeding perches (Snyder &

Snyder 1969; Snyder & Kale 1983; Sykes 1987a). While employing the second form of

foraging, perch-hunting, kites remain perched while scanning the surrounding water (1 to 12

meters away) for snails (Sykes 1987a). Upon detection, the kite will swoop down from the

perch, and just as in course-hunting, hover directly above the surface of the water, grabbing the

snail with the talons. The kite will then return to the perch of departure in order to consume the

snail. After every successful foraging bout, kites perform the same meticulous process of

negotiating, extracting, and eating the snail (Snyder & Snyder 1969; Sykes 1987a).

It is important to note that snail kites in Florida coevolved with the Florida apple snail.

Therefore, certain morphological attributes of the kite seem specifically tailored to exploiting

native snails that fall within a particular size range (around 42.6 mm or "golf ball-sized")

(Snyder & Snyder 1969; Sykes 1987a). The bill of the snail kite is especially adapted to extract

the soft tissue from these shells. The deeply-hooked upper mandible of the snail kite has a

curvature that closely mimics that of the inner spiral of the apple snail shell (Snyder & Snyder

1969), and the bill width of adult kites ranges from 26-34 mm, restricting the depth into the shell

that a kite can reach to cut the columellar muscle (Sykes et al. 1995). Other physical features of

snail kites may also affect their ability to handle alternative prey items. Middle toe plus claw

widths for adult kites range from 49 to 68 mm (Sykes et al. 1995). Average weighs of adult

females, adult males, and juveniles are 446, 394 (Sykes et al. 1995), and 389 grams (Valentine-

Darby et al. 1997) respectively. These attributes potentially impose limitations on the sizes,

weights, and types of prey items that can be successfully captured, handled, and consumed by

kites.









On rare occasions (e.g., during drought conditions or other instances of food scarcity),

kites in Florida have been observed attempting to feed on alternative (i.e., non-snail) prey items

(e.g., small turtles, crawfish, a dead bird, a snake) (Sykes & Kale 1974; Woodin & Woodin

1981; Sykes 1987a; Beissinger 1990; Bennetts et al. 1994); however, they are rarely successful,

and certainly not efficient, at extracting edible body parts (Sykes & Kale 1974; Beissinger 1988,

1990). Many reported cases of non-snail foraging have involved juvenile birds (Sykes 1987a).

In another isolated incident, three to four snail kites were observed feeding on another exotic

snail, Pontacea bridgesii, in a flooded agricultural field during a drought (Takekawa &

Beissinger 1983). Small, isolated populations of P. bridgesii have been present in south Florida

since at least 1966. However, unlike the newly introduced P. insularunt, P. Bridgesii have not

spread to any of the maj or wetlands utilized by the snail kite and show no evidence of being an

invasive threat (Rawlings et al. 2007).

A Note on Pomacea Measurements

The field of malacology has yet to adopt standardized measurement protocols for Pontacea

(Youens & Burks 2007); therefore, throughout much of the literature, identical dimensional

measurements have been described using inconsistent terminology. We adopted the

measurement protocols for linear dimensions of length and width as presented in Darby et al.

(2007). In this study, we standardized the use of these terms, and for all of the results from

previous Pontacea studies described within, we relabeled the dimensional measurements in terms

of length and width by assessing the measurement methods reported by the respective authors.

The Native Apple Snail

Of the many morphological and physiological factors that can influence prey selection and

suitability, prey size may be the most pertinent when dealing with extreme dietary specialists

(Stevens & Krebs 1986; Sih 1987). The Florida apple snail (Pontacea paludosa Say), which is









the only apple snail native to Florida, effectively constitutes the entirety of the snail kite' s diet

and fulfills all of the necessary energetic and nutritional requirements for kites (Cottom &

Knappen 1939; Stieglitz & Thompson 1967; Snyder & Snyder 1969; Sykes 1987a). Adult snail

shells have a normal range of 40 to 70 mm in length (Thompson 1984). Sykes (1987a) collected

empty shells (n=697) from beneath feeding perches used by snail kites. He found that kites

selected snails ranging from 25.2 to 71.3 mm in width (x=42.8, SD=4.9) and from 27.4 to 82.4

mm in length (x=45.8, SD=5.1), with 98.5% of the snails falling in the 30 to 60 mm range for

both measures and 70% falling in the 40 to 50 mm range. Several other authors (e.g., Beissinger

1983; Bourne 1983, 1985a; Tanaka et al. 2006) have also noted that kites rarely eat snails greater

than 60 mm in length. Tanaka et al. (2006) recorded the largest native snail (86 mm in length) to

have been consumed by a kite. Sykes (1987a) found that the average wet weight (whole snail

with shell) of live native apple snails ranged from 12.7 to 38. 1 grams (x=22.3, SD=6.1i, n=24).

Darby et al. (2007) reported that the wet weight of an average-size native apple snail collected on

Lake Kissimmee during the fall of 2004 was 3 5 grams (n=1).

The Exotic Apple Snail

From 2003 to 2007, exotic apple snails were dominant throughout Toho in the areas of the

lake utilized most heavily by kites (Kitchens et al. 2007; Darby, personal communication).

Exotic apple snails commonly exceed 90 mm and can reach 150 mm in length (Benson 2007;

Darby et al. 2007), possibly creating a conflict with the life history strategy of the snail kite.

Darby et al. (2007) sampled exotic apple snails in Goblets Cove on Toho via throw traps and dip

nets. Shell widths of these snails ranged from 67 to 97 mm (x=81, SD=6, n=64) with estimated

lengths (calculated using a 1.15 width to length ratio) ranging from 77 to 112 mm (x=95, SD=7,

n=64). The wet weight (whole snail with shell) of an exotic snail with average width and length

dimensions was 175 grams (n=1).









On average, the exotic snail is large relative to the native snail, and some size classes of the

exotic apple snail may not constitute suitable prey types for snail kites (Figure 2-1). The largest

exotic snails may be rendered unavailable if kites cannot capture them or if kites cannot properly

negotiate them while attempting to feed. Relative to the native snails, exotic snails may also

require longer handling times, which translate into increased energy expenditures for kites.

Useful information alluding to how the size disparity between the native and exotic apple snail

may affect snail kite foraging behavior recently emerged (Darby et al. 2007), but the issue has

yet to be fully addressed, especially in the context of energetic repercussions.

Previous Foraging Studies

Most snail kite foraging studies in Florida pre-date the invasion of Toho by the exotic

apple snail, and given the snail kite's extreme nature of dietary specialization, previous authors

were largely unconcerned with prey type or snail size as influential factors of foraging success.

Nevertheless, several aspects of snail kite foraging behavior have been quantified (e.g., average

searching times, handling times, and capture rates) (Cary 1985; Sykes 1987a; Beissinger 1990;

Bennetts & Kitchens 1997, 2000). Most early studies of snail kite foraging in Florida focus on

modeling capture rates (as measured by the number of snails captured divided by the time spent

hunting) as a function of environmental variables (e.g., water depth, temperature, vegetative

community) and attempt to make inferences about prey availability and relative habitat quality

(Darby et al. 2006; Karunaratne et al. 2006; Bennetts et al. 2006).

Prey availability

Foraging snail kites are limited by the availability of apple snails (Sykes et al. 1995).

Some authors have used prey density as a proxy to prey availability in avian foraging studies, but

this may result in biased inferences (Kushlan 1989; Gawlik 2002). In the case of the snail kite,

snail densities below 0.14 snails/m2 do not support foraging kites (Darby et al. 2006), but snail









density is only one of the factors that influences prey availability (Beissinger 1983; Bourne

1985a, 1985b; Sykes 1987a; Bennetts & Kitchens 1997, Darby et al. 2006, Bennetts et al. 2006).

To be available, snails must not only be present; they must also be visible and accessible to

foraging kites. For example, snail densities may be suitably high in certain vegetative

communities (e.g., sawgrass, cattail) but inaccessible due to visual or physical obstruction

(Beissinger 1983; Sykes 1987a; Bennetts et al. 2006). In other cases, snails may be present and

visible (e.g., open water habitats), but without sufficient emergent vegetation, snails cannot

surface (Hanning 1979; Turner 1996) and may remain too deep (>16 cm) for kites to capture

(Sykes 1987a).

A number of authors have identified other factors that may influence the visibility and

accessibility of snails to kites. Apple snails are poikilothermic and become less active as water

temperature decreases. As temperatures fall, the metabolism of apple snails slows down, which

lowers their oxygen requirement, and at the same time, dissolved oxygen in the water column

increases. Thus, snails visit the surface much less frequently when water temperatures are low

(Hanning 1979; Stevens et al. 2002), and temperature is negatively correlated with the capture

rates achieved by foraging kites (Cary 1985; Bennetts & Kitchens 1997). In extreme cases

(<10 oC), snails may be present but completely inactive, and thus unavailable (Cary 1985;

Stevens et al. 2002). Snails also become inactive during low water events (<10 cm) (Darby et al.

2002), often burying themselves in the substrate, presumably in preparation for aestivation

(Snyder and Snyder 1969; Kushlan 1975, 1989; Darby & Percival 2000; Darby et al. 2002),

again creating a situation where snails are present but undetectable or inaccessible for kites

(Stevens et al. 2002; Darby et al. 2003, 2005). Rain and heavy wind have also been implicated

as factors that reduce kites ability to detect snails (Cary 1985; Bennetts & Kitchens 1997). All of









these studies were conducted before the invasion of Toho by the exotic snail, and none truly

address the final component of apple snail availability: negotiability.

Prey negotiability and drop rate

Apple snails can be present, detectable, and accessible, yet still unavailable if they are not

within the size range that can be captured and successfully negotiated by kites to the point of

consumption. If a snail is dropped (and not recovered) during any phase of the handling process

before it is consumed, that snail effectively becomes unavailable. Cary (1985) observed 1% and

9% drop rates for kites feeding on native snails while course-hunting and perch-hunting

respectively. However, the author noted that shells from previously consumed snails may

remain suspended near the surface of the water in close proximity to feeding perches and that

accidental captures of these empty shells may have contributed to the higher drop rates observed

for kites employing the perch-hunting strategy. Nonetheless, Cary (1985) did not distinguish

between live snails that were dropped due to mishandling and empty shells that were accidentally

captured and subsequently rej ected, nor did he implicate size as the causal factor of snail

dropping.

In a recent study, Darby et al. (2007) reported a 44% drop rate for kites foraging on the

exotic snail on Toho and did, in fact, verify that no empty shells were counted as dropped live

snails. The authors also noted that an average-size exotic apple snail may weigh 37% to 45% as

much as a snail kite, and due to their relative sizes, many exotic snails may be unavailable even

though they exist in high densities (Darby et al. 2007). It is this latter component of availability

involving present and accessible, yet nonnegotiable, snails that raises questions about the

suitability of the exotic apple snail as prey for the snail kite in Florida.









Searching and handling time

Flight is one of the most energetically expensive activities in which birds engage (Masman

& Klaassen 1987; Krebs & Davies 1997; Harrison & Roberts SP 2000). Given that snail kites in

Florida rely heavily on a course-hunting strategy (94% of all foraging bouts) (Cary 1985), the

time spent searching for snails can significantly affect the net energetic gains of foraging.

As described in Sykes (1987a), who studied the foraging behavior of kites intermittently from

1967 to 1980, the average duration of a course-hunting foraging bout was 72 seconds for adult

males (SD=1.2, range=30-180, n=18) and 150 seconds for adult females (SD=2.9, range=30-720,

n=34), which the author notes is a significant difference. This conclusion should be viewed with

caution considering the relatively small sample sizes used and the fact that spatiotemporal

heterogeneity was ignored. No significant difference between the sexes was found for the mean

interval between successive captures, 1338 seconds (range=120 to 5400, n=109), or for the

average captures per hour, 2.5 snails/h (range=1.7 to 3.4, n=109) (Sykes 1987a). Taking

temperature effects into consideration, Cary (1985) reported average capture rates of 5.28 snails

per hour above 30oC, 2.68 between 21-30oC, 0.72 between 11-20oC, and 0.0 below 10oC.

The primary prey of an extreme dietary specialist must be profitable enough to provide

sufficient net energy for survival, growth, and reproduction. Handling time also directly affects

the profitability of selected prey items and is one of the central determinants of diet breadth

(Werner & Hall 1974; Stevens and Krebs 1986). Sykes (1987a) reported the mean combined

handling and eating time for adult kites to be 162 seconds (SD=84, range 60-420, n=16 males

and 53 females), with no significant difference between the sexes. Another study of adult kites

consuming native apple snails, Beissinger (1990) found the mean combined handling and eating

time to be 95.7 seconds (SD=37.3, n=197).









More recently, kites foraging on exotic snails on Toho were compared with those foraging

on native snails on Kiss, and snail size was implied to affect handling time. On Toho, the

average extraction time for the exotic snail was 333 seconds (SD=178, n=10), but the handling

times for snails captured on Kiss were not recorded (nor were the searching times for either

location). However, as mentioned above, kites on Toho were observed capturing 25 exotic

snails, of which 11 were dropped while handling. Kites on Kiss were observed capturing 136

native snails, of which none were dropped (Darby et al. 2007). If one calculates the cumulative

handling time (and likewise, searching time) by including all of the failed handling attempts that

occur between successful foraging bouts, then these instances of snail dropping could greatly

increase the time and energy spent by kites foraging for exotic apple snails. Darby et al. (2007)

did not test for age or sex effects on handling times or drop rates but did identify experience and

relative body weights as likely influential factors of prey handling.

Age and sex effects on foraging behavior

Learning and experience have been recognized as important factors contributing to

foraging behavior. Animals develop search images and improve handling techniques for familiar

prey types (Krebs & Davies 1997). Several studies have confirmed that adult snail kite survival

remains relatively constant from year to year, while juvenile survival varies widely (Bennetts &

Kitchens 1997; Dreitz et al. 2004; Martin et al. 2006a). Experience, and in turn, foraging

efficiency may play a large role in this disparity. Juveniles of most vertebrate species lack the

experience accumulated by adults and are thus more sensitive to environmental variation

(Stephens & Krebs 1986; Stearns 1986; Martin et al. 2007a). Sykes et al. (1995) notes that

compared to adults, recently-fledged kites do not demonstrate the same adept negotiation skills

when extracting native apple snails, and observations of frequent snail dropping by juvenile kites

on Toho during the post-invasion era suggest the same (personal observation). In addition to









lacking experience, juvenile kites also weigh roughly 10% less than adults (males and females

combined) (Sykes et al. 1995; Valentine-Darby et al. 1997); therefore, the ability of juveniles to

negotiate exotic snails may be further limited by their relative physical proportions.

Adult kites may also experience handling complications due to the decreased predator to

prey weight ratios associated with exotic snails. Snail kites are sexually dimorphic, with males

ranging from 360 to 440 grams (x=394, SD=22.8, n=28) and females ranging from 350-570

(x=446, SD=47.8, n=29) (Sykes et al 1995); therefore, sex may play a role in the ability of adult

snail kites to efficiently exploit exotic apple snails.

Energy budgets and profitability

Many of the components necessary for the computation of energy budgets for snail kites

are present in the literature, including daily activity budgets (Beissinger 1984; Cary 1985; Sykes

1987a), energetic costs of behavior (Beissinger 1984, 1986, 1987; Beissinger & Snyder 1987),

and energetic contents of native apple snails (Beissinger 1984; Sykes 1987a), but none have

holistically addressed energetic balances for kites in Florida in the context of alternative prey

types.

Snail kites in South America (i.e., R. s. sociabilis) have been the focus of many prey

selection and energetic studies (Beissinger 1983, 1990; Beissinger et al. 1994; Bourne 1983,

1985a, 1985b, 1993; Tanaka et al. 2006). Of note, the ranges of P. insularum and R. s. sociabilis

overlap, but no documentation of kites feeding on P. insularum in South America exists (Sykes

et al. 1995; Darby et al. 2007). The diet of R. s. sociabilis commonly includes one species of

apple snail (P. dolioides) and one species of crab (Dilocarcinus dentatus), each with varying size

classes, creating an ideal environment in which to test certain predictions of optimal foraging

theory (Bourne 1983; Beissinger et al. 1994). Handling time was found to be positively related

to snail size for kites in Venezuela (Beissinger et al. 1994). Further study of negotiability as a









limiting factor of prey availability and suitable prey size is necessary in order to draw

meaningful conclusions for the conservation of the snail kite if Florida. Research methods aimed

at elucidating the effects of different prey types (as used in these South American studies) are

now applicable to Florida snail kites foraging on native and exotic apple snails.

The calculation of profitability values will demonstrate the utility that each prey species

has to snail kites. Searching time, handling time, and energetic content may vary among prey

types, and profitability is a meaningful quantitative scale on which the costs and benefits of

foraging on alternative prey items can be compared (MacArthur & Pianka 1966; Stevens &

Krebs 1986). Such an approach was used to compare dietary choices and foraging site selection

of snail kites (R. s. sociabilis) in Guyana (Beissinger 1983, 1990; Boumne 1985a, 1985b).

However, kites in South America remain idle during long periods of the day (periods that could

be spent foraging), and profitability may not be the sole determinant of foraging decisions in

snail kites (Beissinger 1983; Bourne 1985a, 1985b). Therefore, a more holistic measure of the

costs and benefits associated with alternative foraging conditions may be more useful. The

calculation of daily energy balances for kites foraging on alternative prey types will shed light on

the potential effects the exotic snail may have on the kite population. As stated above,

negotiability has been largely ignored until recently, and unfortunately, data collected in the

recent study (i.e., Darby et al. 2007) is insufficient to calculate profitability values and daily

energy balances for kites foraging on native versus exotic snails.

Economic models of foraging behavior

In the study of foraging behavior, economic models that compare the costs and benefits of

alternative food choices are commonplace (Krebs & Davies 1997). A general assumption of all

studies that use this cost-benefit approach to compare alternative decisions is that it pays for an

animal to be efficient. That is, natural selection will act in favor of individuals that maximize









their benefits while minimizing their costs (MacArthur 1972; Staddon 2003). Using economic

models in the study of foraging behavior requires investigators to define a set of costs and

benefits that can be quantified and compared using a common currency. The ideal currency with

which to weigh the costs and benefits of decisions would be fitness (Krebs & Davies 1997).

Glimcher (2002) points out that "Darwinian fitness provides a natural and genuinely one-

dimensional scale." Unfortunately, directly quantifying the effects that certain foraging

behaviors have on fitness can be difficult; therefore, more easily quantifiable currencies (e.g.,

time and energy) are often employed by these economic models. An underlying assumption of

these surrogate currencies is that they correlate directly with fitness (Krebs & Kacelnik 1991;

Brown et al., 1993). Time and energy were the first two currencies used in the development of

foraging and diet selection models (MacArthur & Pianka 1966; Emlen 1966), and they have

remained the dominant measures in these behavioral studies ever since (Cuthill & Houston 1997;

Sih & Christensen 2001).

Predictions

Based on our knowledge of snail kite ecology, we expect the exotic apple snail to have

deleterious effects on kite foraging behavior, and we expect these effects to vary in magnitude

relative to foraging experience and relative body weight; therefore, we make the following

predictions:

* Prediction 1. If the average drop rate for exotic snails is significantly greater than the
average drop rate for natives (i.e., if Prediction 3 is true), then the average searching time for
exotic snails will be significantly longer than the average searching time for native snails

* Prediction 2. The average handling time for exotic snails will be significantly longer than
the average handling time for native snails.

* Prediction 3. The average drop rate for exotic snails will be significantly greater than the
average drop rate for native snails.









* Prediction 4. When feeding on exotic snails, juvenile kites will have significantly greater a)
handling times and b) drop rates than adults.

* Prediction 5. When feeding on exotic snails, adult males will have significantly greater a)
handling times and b) drop rates than adult females.

* Prediction 6. The average capture rate for exotic snails will be significantly lower than the
average capture rate for native snails.

* Prediction 7. The average profitability of the exotic snail will be significantly lower than the
average profitability of the native snail.

* Prediction 8. Daily energy balances will be significantly lower for kites feeding on the
exotic snail relative to kites feeding on native snails.

Methods

To address the above predictions, we conducted a comparative observational study. First,

we quantified the necessary set of measures that pertain to foraging behavior and prey items.

Then we used these data to address the anticipated behavioral differences between/among groups

of kites. After teasing out the effects of the exotic apple snail on foraging behavior, we assessed

the energetic repercussions for kites by calculating prey profitabilities, deriving activity budgets,

and proj ecting daily energy balances.

Time Activity Budgets

In order to obtain quantitative behavioral data related to foraging activities, we conducted

time-activity budget (TAB) observations on wild snail kites from 2003-2007 using a

modification of focal animal sampling (Altman 1974). Such a method was used to collect TAB

data on kites in 1993, 1994, and 1996 (Bennetts & Kitchens 1997, 2000), and so that we could

make meaningful comparisons between datasets, we followed similar protocols. During each

TAB we observed a focal kite continuously for an allotted period of time and recorded the

frequency and duration of all pertinent events and behaviors (see Collection of Behavioral Data).










Sampling locations

Most of our TAB sampling effort was focused in the following wetlands: Toho, Kiss,

WCA3A (Table 2-1; Figure 1-1). Together, these fragments represent much of the remaining

core of snail kite habitat in Florida. In recent years, the snail kite population has relied heavily

on Toho, Kiss, and WCA3A during the breeding season. During our study, these were the only

wetlands that produced significant numbers of observable juveniles. Kites also utilize Toho,

Kiss, and WCA3A during the non-breeding season; therefore, numerous kites can be found

foraging in these wetlands throughout most of the year (Bennetts & Kitchens 1997; Martin 2007;

Martin et al. 2007c). We also collected data, at least opportunistically, throughout the remainder

of the kite' s range, but much less effort was expended conducting TABs in the following non-

focal wetlands: WCA3B, WCA2B, ENP, OKEE, GW, and SJM (Table 2-1; Figure 1-1).

Selection of focal individuals

Given the extensive range of the snail kite and the high expense of Hield operations, on any

given day the wetlands in which we made observations were determined systematically to

optimize travel and funding efficiency. However, we randomly selected the individual birds that

were observed within each wetland fragment. To do so, a block grid with numbered cells was

overlaid on a map of the wetland area, and we used a random number generator to select the cell

in which to start our observations. Using an airboat and GPS unit for transportation and

navigation, we traveled to the selected cell and observed the first snail kite detected upon our

arrival. Observations started as soon as the boat was parked and necessary equipment set up. If

no snail kite was detected in the given cell, we randomly moved to one of the eight surrounding

cells by blindly spinning a compass and driving to the adj acent cell at which it pointed. This

process was repeated until a snail kite was located for observation. The latitude and longitude

(measured in UTMs, Datum NAD83) of all observation locations were recorded with a hand-









held GPS unit. After completing a TAB, the random number generator was used to select a cell

for the next kite observation, and the process started again.

Timing and duration of observations

TABs were conducted during all months of the year (Table 2-1). Months were assigned to

one of three seasons as follows: Spring included January, February, March and April; Summer

included May, June, July and August; and Fall included September, October, November and

December (Bennetts & Kitchens 2000). Spring coincides loosely with the peak of nest initiation

during the breeding season and with the annual dry season (Bennetts & Kitchens 1997), although

both of these measures show some overlap with Summer (Sykes et al. 1995, Martin 2007).

Summer connects the post-breeding period (i.e., when kites tend to disperse) with the onset of

the wet season (Bennetts & Kitchens 1997; Martin et al. 2006a), while Fall encompasses a period

of occasional late-season breeding and the onset of the dry season (Bennetts & Kitchens 2000).

Depending on the sex and reproductive stage of a breeding snail kite, a high degree of

variability may be observed in its behavior (Sykes et al. 1995). Since we could not collect

enough TAB data on actively breeding individuals during different reproductive stages to treat

such data differently, such variability would have only confounded our analyses; therefore, we

did not conduct TABs on individuals that were actively engaged in any stage of the reproductive

process, and if any breeding behavior was observed (e.g., courtship, snail delivery to a mate, nest

building, nest defense, provisioning of young) then the TAB was discontinued and the data

censored from our analyses. Thus, our inferences are limited to snail kites that are foraging

independently and cannot necessarily be extrapolated to kites that are provisioning nestlings or a

mate.

Observations were made during all hours of the day, as kites forage throughout the day in

Florida (Cary 1985; Sykes 1987a); however, in following with previous studies (e.g., Cary 1985;









Sykes 1987a; Bennetts & Kitchens 2000), we recorded the exact time of day during which

observations took place. We assigned observations to one of three daytime categories: morning

(sunrise plus three hours), evening (sunset minus three hours), and afternoon (everything in

between). To avoid potential confounding effects of inclement weather conditions on foraging

behaviors, observations were not made during times of rain or heavy wind (Cary 1985; Bennetts

& Kitchens 1997).

We conducted a combination of one hour (n=713), two hour (n=48), and day-long

observations (n=21). The intended duration of the observation was always determined prior to

locating a focal individual. Due to unpredictable weather events and equipment malfunctions,

observations were sometimes stopped prematurely. Observations lasting at least eight hours

were classified as day-long observations, and those less than 30 minutes were censored from our

analyses.

Classification of individual kites

Whenever possible we determined the sex and relative age of the focal kite. Snail kites are

sexually dichromatic, and mature adults of each sex have very distinct plumage coloration

(Sykes et al. 1995). We visually identified the sex of all mature adults by their plumage.

However, full adult plumage may not be achieved until the breeding season of the 3rd year (i.e.,

until 36 months old), and the sex of younger kites, especially those from 6 to 24 months old

cannot be reliably distinguished by plumage cues (Bennetts & Kitchens 1999). Fortunately,

during annual breeding season population surveys, which were conducted by the Florida Fish

and Wildlife Cooperative Research Unit from 1992 to 2007, numerous kites were banded as

juveniles and had feather samples taken for DNA analysis (Bennetts & Kitchens 1997, 1999;

Martin 2007). Therefore, the sex of many young birds (i.e., all those with specific alphanumeric

bands and corresponding DNA results) could be determined, despite their ambiguous plumage.









To avoid inducing bias by systematically bypassing non-banded adults and/or old juveniles of

ambiguous sex, we still observed these individuals when they were encountered, though birds of

unknown sex were censored from all analyses in which sex was a covariate.

We also identified the relative age of the focal kite. Kites with mature adult plumage (see

above) were classified as adults. The second age class, juveniles, also has definitive plumage

coloration, although the sexes cannot be distinguished. The distinct buff brown and cinnamon

juvenile plumage is retained until about the 5th month of life (Sykes et al. 1995). Juveniles also

have dark brownish black eyes, in contrast to the bright red eyes of adults. Eye color begins to

change between 4 and 6 months of age, and juveniles become hard to distinguish from second

year birds after 6 months of age (personal observation). Therefore, birds from 6 to 36 months

old are hard to age with visual cues. The equipping of juveniles with unique alphanumeric bands

during the annual population surveys helped us keep track of many birds' ages through the

period of ambiguous plumage coloration. The ages of all kites that were banded as nestlings

could be estimated to the day ( /- 7 days) (Sykes et al. 1995).

We still observed kites of ambiguous age that we encountered during TABs. These kites

(along with banded kites of know ages falling from 6 to 36 months old) were placed into a third

category: subadults. Even though there is no evidence for a subadult age class in snail kites, as

juvenile survival parallels that of adults by the fifth or sixth month of age (Bennetts & Kitchens

1999; Bennetts et al. 1999) and kites can begin breeding as young as 9 months of age (Synder et

al. 1989), it was useful to define a third age class. Birds whose age could not be positively

identified in the Hield (i.e., non-banded individuals from 6 to 36 months old) were, at first,

categorized as subadults, but upon comparing measures of foraging behavior among the three










age classes we found evidence that justified combining adults and subadults into a single adult

age class (see Comparison ofAdults, Subadults, and Juveniles).

Collection of behavioral data

TAB observations were made from an airboat with the use of 10x binoculars and a 15x-

60x spotting scope. Two observers were always present so that observation and data recording

could occur simultaneously. Using a twenty-four hour digital clock, we recorded the time of day

to the second that the focal kite engaged in each of the following behaviors: started flying,

captured a snail, stopped flying (i.e., returned to a perch), extracted a snail, Einished eating a

snail, or dropped a snail.

Each flight was categorized as either a foraging bout or a non-foraging flight. Each

foraging bout was categorized as either a course-hunt or perch-hunt. These distinctions were

determined by telltale behavioral cues (Snyder & Snyder 1969; Beissinger 1983; Sykes 1987a).

Foraging bouts were marked successful if the kite returned to the perch with a snail and

consumed it, and they were marked unsuccessful if the kite returned to a perch without a snail or

if the snail was dropped at the perch before consumption. In cases where a snail was dropped in

flight but a kite made multiple attempts during a single foraging flight, the bout was considered

successful if the kite eventually returned to a perch with a snail and consumed it, and it was

marked unsuccessful if it did not. Even if a snail was dropped, it was still tallied as a capture,

but the associated foraging bout was classified as unsuccessful unless a successful capture

followed by successful consumption occurred. We distinguished between dropped snails and

empty shells that were picked up and rej ected by visual cues (e.g., the splash on the water; the

lag time before sinking) (Darby et al. 2007).

While the kites were perched, we distinguished between handling time and idle perch time.

Handling time was defined as the time that elapsed between arriving at a perch with a snail and









finishing the consumption of a snail. Upon arriving at a feeding perch, kites usually began

negotiating the captured snail immediately and continued through the extraction and

consumption process without interruption (Sykes 1987a). Handling time included negotiating,

extracting, and consuming a snail. The remaining time spent perched (i.e., all but the handling

time) was classified as idle perch time. Although kites occasionally preened or engaged in intra-

and inter-species interactions while perched, the times of these behaviors were not recorded

during our observations unless such a behavior interrupted the handling time of the focal kite or

induced the focal kite to take flight.

Missing data is one of the inescapable shortcomings associated with field observation as a

mode of scientific inquiry (Morrison 2001). In all circumstances we did our best to record every

behavior to the second, but oftentimes, due to conditions beyond our control, particular actions

were unobservable, resulting in unknown starting or ending times for given events.

Assumptions of behavioral observations

A maj or assumption using data collected during TABs is that our observation actions did

not affect the observed individual's behavior. Snail kites are relatively docile birds and are not

easily disturbed by the presence of humans or airboats from distances greater than 49+18 meters

(Rodgers & Schwikert 2003; Bennetts et al. 2006). To minimize potential disruption to the

natural behavior of the birds, we always stayed at least 70 meters away when making

observations. It is highly unlikely that our presence at such a distance altered the focal bird's

normal behavior, unless an active nest was present. In spite of the care taken by observers, on

occasion human activities did affect the behavior of the snail kite under observation, resulting in

the bird flushing evasively. There were two common scenarios in which this occurred. First,

when we temporarily lost sight of a bird and were forced to move the boat in order to locate it so

that the observation could continue, we sometimes approached too closely before getting a visual









on the bird. The second scenario occurred when a third party (e.g., fishing boats, tour boats)

passed through our observation area unknowingly close to the snail kite under observation.

Instances of human intrusion, observer induced or otherwise, were recorded.

Apple Snails

Collection and measurement of empty snail shells

After the completion of a TAB, empty snail shells were collected from the feeding perches

that were used by the focal kite. Snail shells often accumulate beneath frequently used feeding

perches, and snails consumed by kites can be distinguished from those consumed by other

predators (Snyder & Snyder 1969; Collett 1977; Sykes 1987a). Snail shells may also accumulate

on and around snail kite nests (personal observation). Shells were also collected from nests as

part of the nest monitoring protocol under the systematic annual breeding season population

surveys (Bennetts & Kitchens 1997; Martin et al. 2007c). To ensure that collection dates were

representative of consumption dates, only fresh shells, as determined by coloration and smell

(Bourne 1993), were collected from feeding perches and nests. All collected shells were bagged

and labeled with the date, GPS coordinates, wetland, and specific place of collection (i.e.,

feeding perch or nest). We also recorded the TAB during which the shells were collected;

however, it is common for some feeding perches to be used by several individual kites

intermittently or with close temporal proximity (Sykes 1987a), so we could not be certain that all

of the shells we collected during TABs were from the focal individual. As in Sykes (1987a), the

linear widths and lengths (dimensions renamed) of these shells were measured to the nearest

millimeter using vernier calipers (Figure 2-2a, 2-2b respectively), and these dimensions were

used to estimate the whole weight and caloric content of the snails that once occupied the

collected shells (see Regression models and estimation of average snail weight and caloric

content) .









Collection and processing of live snails

Funnel traps were used to collect apple snails on Toho (For details on and a discussion of

this collection method see Darby et al. 2001). We sampled from 18 traps that were deployed by

an ongoing study (Kitchens et al. 2005, 2007) and from 6 additional traps, which we deployed in

an area used heavily by kites (i.e., Grassy Island) that was not covered in the scope of the

overlapping study (Figure 2-3).

Traps were checked every one to two weeks. All snails caught in the traps located on the

east, south, and west shores were identified to species (Rawlings et al. 2007) and tallied. Then,

for each trap, individual snails were pooled together by species and the cumulative species

weights were recorded to the nearest gram using a spring scale. The snails were subsequently

released at the location of capture. Snails caught in traps located in Goblets Cove and Grassy

Island (Figure 2-3), the two areas of the lake used heavily by snail kites (Martin et al. 2007c),

were removed, placed in a plastic bags labeled with the date and trap location, and immediately

frozen. In the lab, we identified each individual to species, took the dimensional measurements

of width and length (see Collection and' measurement of empty snail shells), and measured the

whole wet weight (i.e., whole snail with shell) to the nearest tenth of a gram using a digital scale.

A subset of these frozen snails was randomly selected and the snails removed from their

shells. The wet weight of each individual's internal body parts (i.e., no shell and no operculum)

was recorded. Female snails have an albumen gland, which is the only internal tissue commonly

discarded by kites prior to consumption (Snyder & Snyder 1969; Sykes 1987a). The internal

bodies of female snails were weighted with and without their albumen glands, and then the gland

was discarded before nutritional analyses were conducted. Three pooled samples (consisting of

20, 30, and 30 snails respectively) were packed on dry ice and shipped overnight to Silliker

Laboratory in Chicago, IL for biochemical analysis. The samples were homogenized with a









laboratory blender, and then the caloric content was determined as well as the percent moisture,

ash, protein, lipid, and carbohydrate (Table 2-2).

Statistical Analysis and Confidence Intervals

All confidence intervals (95% CI) around our derived estimates were approximated as

follows: CI = x +- ta/ SE, where x is the sample estimate (i.e., mean), t is the test statistic

(evaluated at a= 0.05 on n-1 degrees of freedom), and SE is the standard error of the mean. Such

estimation is valid for positive sample estimates of count, measurement, or proportion that

generally follow a lognormal distribution (Burnham & Anderson 2002). We also report standard

deviations (SD) of some measures for comparison with previous studies. All statistical analyses

were performed using the software package R version 2.5.1.

Analysis of Foraging Behavior

Using the observational data that we collected, we quantified the following foraging

behaviors for each TAB: average searching time, average handling time, drop rate, and capture

rate (i.e., consumption rate). These measures were used to compare the behavioral differences

between kites foraging on exotic versus native snails. Most of our conclusions were drawn from

observations made on Toho (in which exotic snails are prevalent) and on Kiss and WCA3A

(neither of which harbors the exotic apple snail). In order to account for small sample sizes, we

pooled data from Kiss and WCA3A for many of our analyses; however, if the measure of

comparison differed significantly between Kiss and WCA3A or if small sample sizes were not an

issue, then Kiss and WCA3A were treated independently. Data collected in the non-focal

wetlands (i.e., WCA3B, WCA2B, ENP, OKEE, GW, SJM) were, in many cases, not evenly

distributed (e.g., observations were only made during certain seasons or years; unbalanced

numbers of observations were made on a particular age group or a particular sex), and to avoid









biasing our results via nonrandom effects, these data were excluded from our analyses (unless

otherwise noted).

Exotic snails were first discovered on Toho in 2001 and were likely predominant on by

Toho 2003 (Darby, personal conanunication), yet native snails were still relatively abundant in

the littoral zone of the lake prior to the commencement of the drawdown and scraping treatment

in late-2003 (Kitchens et al., unpublished data; Darby, personal conanunication). Reliable data

on the selection of snails by kites during this transitional period (2001-2003) is lacking, but

anecdotal evidence suggests that kites were foraging heavily on exotics by 2003: kites were

commonly seen course-hunting in deeper regions of the lake (i.e., habitats that do not harbor

native snails) and piles of exotic snail shells were observed below kite feeding perches (Darby &

Welch, personal conanunication). During the pilot year of our study (i.e., 2003), only 23 TABs

were conducted, and all of these observations were of juvenile kites, none of which were on

Toho. By 2004 kites on Toho were feeding almost exclusively on exotic snails (see Results), and

although our inferences related to foraging on exotic snails were drawn from TABs made during

the period 2004 to 2007, we defined 2003-2007 as the post-invasion era based on the confirmed

presence and likely abundance of the exotic snail on Toho during 2003. And in order to increase

the sample size of observations made on juvenile kites, we included the 2003 data for juveniles

on Kiss and WCA3A in our analyses concerning the post-invasion era.

We also incorporated data from Bennetts and Kitchens (1997, 2000) into some of our

analyses. These data were collected in 1993, 1994, and 1996. We used these observations to

represent the pre-invasion era. None of the aforementioned foraging behaviors differed

significantly among Toho, Kiss, and ETOHO during the pre-invasion era, so to increase the

sample size, we pooled these data. We then compared measures of handling time and drop rate









for adults foraging on Toho during the pre- and post-invasion eras. Data on juveniles from the

pre-invasion era was lacking; therefore, we could not compare juvenile foraging behavior before

and after the invasion of the exotic snail.

Searching and handling time

The time that elapsed during a foraging bout between takeoff and capture was measured as

searching time; however, the calculation of the searching time per foraging bout was contingent

on the foraging bout being successful (i.e., it resulted in the consumption of a snail). Since not

all foraging bouts were successful yet kites still expended time and energy searching during these

bouts, we calculated the searching time for each successful foraging bout as the cumulative time

spent searching between successful bouts. Therefore, for a given successful foraging bout, we

took the search time of that bout and added the total flight time of all the previous unsuccessful

bouts (i.e., foraging bouts in which no snail was captured or in which the snail was dropped

before consumption) that had occurred since the last successful one. Non-foraging flights that

occurred between successful foraging bouts were not included in the calculation of searching

times. Searching time (ts) was calculated as follows: ts = tf t, + (ts), where tf is the total flight

time of the successful foraging bout, tr is the returning flight time after the capture was made,

and (t)S is the sum of the total flight times for all preceding unsuccessful foraging bouts that

occurred since the last successful bout.

Handling time was measured as elapsed time from landing on a perch with a snail to the

end of consumption. In unusual cases where the focal kite was interrupted (e.g., by a boat or

another bird), we recorded the duration of the alternative behavior and subtracted it from the

actual handling time. Handling time (th) WaS calculated as follows: th Te-Tp- t,, where Te is the

time at which the kite finishes eating, Tp is the time at which the kite perches with the snail, and t,

is the total time of any interruptions that may have occurred during the handling process.









We only analyzed foraging bouts that had complete time documentation. Foraging bouts

with missing capture times or handling times were censored from respective analyses. In order

to avoid problems associated with pseudoreplication that may have resulted from treating every

foraging bout independently, we calculated the average searching and handling times for each

TAB (i.e., each individual bird) and used these averages for comparative analyses.

Drop rate

We calculated the drop rate for each TAB as the number of snails dropped divided by the

total number of snails captured by the focal kite. For this analysis, we calculated the total

number of snails captured as the sum of dropped snails and successfully consumed snails.

Empty shells that were picked up and rej ected were not included in this calculation. Drop rate

(D) was calculated as follows: D = sd (s,+Sc d, where Sd is the number of snails dropped during

a TAB and so is the number of snails consumed during a TAB.

We also calculated raw drop rates for adults and juveniles on Toho and Kiss-WCA3A so

that we could compare our results with those reported by Darby et al. (2007). The raw drop rate

(D,) was calculated as follows: D, = Sd, (Sc,, Sd;), where Sd; is the total number of snails

dropped during TABs by all individuals in group i and Sc, is the total number of snails consumed

during TABs by all individuals in group i.

Capture rate (i.e., consumption rate)

Capture rates were measured as the number of snails captured and successfully consumed

per elapsed time. While snails that were captured and subsequently dropped were included in the

total number of captures used in our calculation of drop rates (see Drop rate), these mishandled

snails were censored from the calculation of capture rates. We calculated the capture rate for

each TAB as the total number of snails successfully consumed during a TAB divided by the total

length of the TAB in seconds (the rate was later converted to snails per hour). If the focal









individual was lost from sight for greater than one minute during the TAB, we subtracted the

time it was lost from the total observation time before calculating the capture rate. Capture rate

(C) was calculated as follows: C = sc / to, where so is the number of snails consumed during a

TAB and tois the total length of the TAB in seconds.

Analysis of Energetics

Regression models and estimation of average snail weight and caloric content

Using the measurements from the live exotic snails that were collected in funnel traps, we

ran a linear regression of the total snail weight (i.e., shell, operculum, all soft body parts) as a

function of shell length (as done on native snails in Sykes 1987a), which yielded the following

equation: total snail weight = (-0. 08495465 0. 01279404 shell length) ^\ 2, (p<0.001, adjusted

R-squared= 0.78, df= 887) (Figure 2-4a). We used this regression model to derive the original

weights of the empty exotic snail shells that were collected from feeding perches after being

handled by kites on Toho.

We also used the live exotic snails from Toho to run a linear regression of the wet weight

of edible soft body parts (i.e., no shell, no operculum, no albumen) as a function of shell length

(as done on native snails in Sykes 1987a), which yielded the following equation: wet weight of

soft body parts = (0.8143635 shell length 8. 08201), (p<0.001, adjusted R-squared= 0.76,

df=362) (Figure 2-4b). Then we converted the wet weights from this model to dry weights by

multiplying the output by 0. 1215 (i.e., the proportion of dry weight to wet weight of edible soft

body parts) (Table 2-3). Finally, in order to derive the gross caloric value of exotic snails

consumed by kites on Toho, we multiplied the derived dry weights by the caloric value of the

exotic snail, 3.30 kcal/g (Table 2-2 and 2-3), and took the average. Our models and respective

coefficients of determination were comparable to those in Sykes (1987a).









We estimated weights and caloric values for the empty native snail shells that were

collected during TABs using the following linear models from Sykes (1987a): total snail weight

= (0. 6769 shell length- 20. 3448) / 0.48, where 0.48 is the proportional weight of wet edible

soft body parts to total snail weight; caloric content = (0. 6769 shell length- 20. 3448) 0. 145

* 4.60, where 0. 145 is the proportion of dry weight to wet weight of edible soft body parts of the

native snail and 4.60 kcal/g is the caloric value of the native apple (for both models, adjusted

R^'2= 0.75) (Table 2-3).

Profitability

Profitability (i.e., the energy gained from a prey item per unit time invested in obtaining

and consuming the prey item) is a common measure used to quantify the benefit that an

individual receives from a foraging bout. The costs associated with a foraging bout include the

time and energy spent searching for and handling a prey item. Search time is taken as a function

of encounter rate (which itself depends of prey density and detection probability), while handling

time is a function of prey size and shape (i.e., prey type).

We derived estimates of profitability for the following four groups: adults feeding on

native snails, adults feeding on exotic snails, juveniles feeding on native snails, and juveniles

feeding on exotic snails. We calculated profitability (P,,) as follows: P,, = c, (0.9) (ts,, ts,,)

where c, is the average caloric content of snail species j, 0.9 is the approximated digestion

coefficient for kites (Beissinger 1984), and ts,, is the average searching time and to,, the average

handling time for individuals in age class i feeding on snail species j.

Daily activity times and energy budgets

Daily time-activity budgets can be constructed with incomplete observations by using the

proportional activity times observed during TABs to extrapolate the actual time spent conducting

each behavior on a daily basis (Pearson 1954; Wolf et al. 1975; Ashkenazie & Safriel 1979;









Goldstein 1988). We employed this method to estimate average daily time-activity budgets for

groups of kites. An obvious assumption is that the activity patterns observed during TABs are

representative of normal daily behavior. Previous studies do not report significant differences in

behavior patterns among kites observed during different times of the day (Beissinger 1983; Cary

1985; Sykes 1987a). We validated this assumption with our own data by comparing the relative

proportion of time spent flying between day-long and hourly observations. Moreover, capture

rates are known to be affected by temperature (Cary 1985), and it has been suggested by some

authors (e.g., Cary 1885; Sykes 1987a) that there is a lull in foraging activity during mid-day,

although these same authors have also shown that there is no statistically significant difference in

capture rates throughout the day (Cary 1885; Sykes 1987a). Nonetheless, we also validated that

the average capture rates achieved by kites did not differ among observations made during the

morning, afternoon, and evening (see Validation of time activity budget exturapolation).

After validating our assumptions, we calculated the relative proportion of time that the

following five groups of kites spent flying and perched: adults on Toho, adults on Kiss, adults on

WCA3A, juveniles on Toho, and juveniles on Kiss-WCA3A. Using these estimates, we derived

the average daily energy expenditure (DEE,) for each of the five groups by employing a series of

equations from Koplin et al. (1980) that were adapted for the snail kite by Beissinger (1984) (see

Appendix). Then we estimated the average daily energy gain (DEG,) for each of the five groups

as follows: DEG, = C, c, (0.9) dl, where C, is the average capture rate for individuals in

group i, c, is the average caloric content of snail species j, 0.9 is the approximated digestion

coefficient for kites (Beissinger 1984), and dl is the day length. Since we had insufficient data

to subdivide each of the aforementioned group by season, we used an overall average day length

of 12 hours for our calculations. The annual average day length for WCA3A was 12.16 hours










(SD= 1.25) and for Kiss and Toho was 12.15 hours (SD= 1.12). Finally, we calculated the

average daily energy balance (DEB,) for each of these five groups by subtracting DEE, from

DEG,. This is the most common approach in such foraging studies (Goldstein 1988), and

variations of this approach have been applied to the snail kite previously (Beissinger 1983, 1984;

Bourne 1985b; Sykes 1987a).

Results

Diet

We collected 336 piles of empty snail shells (ranging from 1 to 121 shells each) from snail

kite feeding perches and nests from 2004 to 2007 in the following areas: WCA3A (n=4108),

WCA3B (n=532), WCA2B (n=27), ENP (n=379), Okee (n=154), GW (n=61), SJM (n=161),

Kiss (n=999), and Toho (n=1486). Toho was the only wetland in which we found shells from

the exotic, P. insularum; however, both native and exotic snails were found on Toho throughout

our study. The annual frequencies of each species as they appeared in our funnel traps on Toho

appear in Figure 2-5, and the proportions of native to exotic snails captured in each of the five

maj or sampling locations on Toho throughout the study appear in Figure 2-6; however, these

years and trap locations do not represent equal sampling efforts, and neither relative species

proportions nor trends should be interpreted from these graphs.

During the nearly 790 hours spent observing kite behavior, we observed kites consuming

2697 snails (360 exotics and 2367 natives). We also observed 15 cases of kites eating turtles, all

of which involved adult kites on Toho during January, February, and November of 2006. We

observed one female kite feeding on Marisa sp. shells in WCA3A during late summer of 2004,

and several Marisa shells were found mixed in with native apple snail shells below two well-

used feeding perches in the same location. Snails of the genus Pomacea accounted for 99.5% of

the snail kite's diet during our observations.









During our TABs, we also observed five occasions of juvenile kites on Toho consuming

vegetation. Although anecdotal, all five instances involved juveniles that had not been observed

eating for over an hour and that may have been starving. We observed these individuals

rummaging around in vegetation near the surface of the water and picking up empty shells and

pieces of water lily and hydrilla. There were two instances in which the juveniles picked apart

and swallowed entire water lily seed heads. The other three observations involved the juveniles

ingesting large chunks of plant material, including hydrilla, water lily stems, and an unidentified

woody twig. After searching the literature, we believe that this is the first documentation of kites

feeding on vegetation.

Nutritional content of apple snails

The nutritional content of exotic snails as reported by Silliker Laboratories, appears in

Figure 2-2. We found that exotic snails store significantly fewer calories per gram of dry weight

than do native snails (3.30 versus 4.52 kcal/g) but that other nutritional measures for the exotic

fall within the range of reported values for the native snail (Figure 2-3).

Live exotic snail measurements

Live exotic snails (n=890) collected in funnel traps from Toho from 2005 to 2007 ranged

from 11.0 to 73.1 mm in width (x=37.7, SD=12.5) and from 16 to 81 mm in length (x=45.5,

SD=10.9). Weights of whole snails (with shell) ranged from 1.1 to 117.9 grams (x=27.1,

SD=16.2). However, this sample cannot be considered an accurate representation of the pool of

available snails for several reasons. Firstly, the physical design of the funnel traps used by

Kitchens et al. (2005, 2007) systematically restricted snails over 80 mm from entering. In

addition to this drawback, snails of different size classes also have different probabilities of

being captured, escaping, and being eaten by other species while in the trap (Darby et al. 2001;

Kitchens et al. 2005, 2007). Nonetheless, 70% of the empty shells collected from kite feeding









perches on Toho fell within the 45.5 to 81.0 mm range (see Empty snail shell dimensions), and

the funnel traps that we used to capture live snails covered this range.

Empty snail shell dimensions

We found that kites on Toho are eating exotic snails (n=1486) ranging from 17.0 to 92.1

mm (x=58.2, SD=13.6) in width and from 19.0 to 103.5 mm in length (x=63.5, SD=15.5), with

the lengths of 98.5% of the snails falling in the 26.3 to 92.9 mm range and 70% falling in the

45.5 to 81.0 mm range. Empty native snails shells (n=6421) discarded by kites after feeding

ranged from 8.50 to 65.8 mm (x=37.6, SD=5.6) in width and from 12.0 to 67.1 mm in length

(x=40.2 mm, SD=6.3), with the lengths of 98.5% of the snails falling in the 17.5 to 54. 1 mm

range, and 70% falling in the 35.5 to 45.7 mm range. We found that the average shell

dimensions of native snails did not significantly differ among wetlands from which adequately

sized samples were collected (Figure 2-7), but a significant difference was found between the

dimensions of the native shells found in these areas and the exotic snail shells collected from

Toho (Figure 2-8).

Estimates of total snail weight

With the length measurements from the empty exotic shells, we predicted the weights

(whole snail with shell and operculum) of exotic snails captured by kites on Toho using the

following linear model: total snail weight = (-0. 08465614 0. 012791 72 shell length) ^\ 2. Our

estimated weights ranged from 2.5 to 153.6 grams, and we found that, on average, kites on Toho

are negotiating exotic snails that weigh 56.8 grams (95% CI= 53.9-59.7).

With the length measurements from the empty native shells, we predicted the weights

(whole snail with shell and operculum) of native snails captured by kites using the following

linear model from Sykes (1987a): total snail weight = (0. 6769 shell length -20. 3448) / 0. 48.

Our predicted native snail weights ranged from 0.1 to 52.2 grams, and we found that the average










weight of native apple snails negotiated by kites was 15.9 grams (95% CI=15.1-16.7). Thus, the

exotic snails consumed by kites are significantly heavier (over three times heavier) than the

natives (Figure 2-9).

Foraging Behavior

Comparing adults, subadults, and juveniles

We compared measures of foraging behavior among the three age classes defined by our

TAB protocol (i.e., adults, subadults, juveniles), and consistent with previous literature, we

found no supporting evidence for a subadult age class. Average handling times on Kiss-WCA3A

were similar for adults (x=72 seconds, 95% CI= 68-76, n= 259) and subadults (x= 72 seconds,

95% CI= 65-79, n =65) and their confidence intervals barely included the average handling time

for juveniles, 96 seconds (95% CI= 71-122, n= 22). On Toho, we found that the average

handling times for adults (x= 303 seconds, 95% CI= 245-360, n= 78) and subadults (x= 205

seconds, 95% CI= 109-301, n= 7) were similar and that they differed significantly from

juveniles (x= 496 seconds, 95% CI= 376-617, n= 44) (Figure 2-10). Likewise, the average drop

rates for adults (on Toho, x= 0.17; on Kiss-WCA3A, x=0.02) were comparable to those for

subadults (on Toho, x=0.18; on Kiss-WCA3A, x= 0.02) and differed significantly from the

average drop rates of juveniles (on Toho, x= 0.33; on Kiss-WCA3A, x= 0.06) (Figure 2-11).

Average capture rates for adults (on Toho, x= 1.09, 95% CI= 1.05-1.14, n= 171; on Kiss-

WCA3A, x= 3.14, 95% CI= 2.85-3.43, n= 313) were comparable to those for subadults (on

Toho, x= 1.05, 95% CI= 0.92-1.17, n= 15; on Kiss-WCA3A, x= 3.35, 95% CI= 2.66-4.05, n=

65) and differed significantly from the average capture rates of juveniles (on Toho, x= 0.77, 95%

CI= 0.69-0.85, n= 110; on Kiss-WCA3A, x= 3.46 snails/hour, 95% CI= 2.49-4.44, n= 37)

(Figure 2-12). Average searching times were similar among all age classes (Figure 2-13). As a









result of these findings, we reclassified all subadults as adults and used only two age classes (i.e.,

adults, juveniles) in all of the following analyses.

Effects of the exotic snail on average searching times

We found no evidence that average searching times differ between adult males and adult

females (Figure 2-14). After pooling the sexes, we found that during the post-invasion era, the

average searching time for adults on Toho (x= 67 seconds, 95% CI= 49-85, n= 117) was

significantly less than the average search times on Kiss (x= 111 seconds, 95% CI= 95-125, n=

118), and WCA3A (x= 76 seconds, 95% CI= 68-83, n= 270). The average searching time for

adults on Toho during the pre-invasion era (x= 32 seconds, 95% CI= 18-44, n= 79) was

significantly less than the average searching time on Toho during the post-invasion era (Figure 2-

15). For juveniles, we found that the average searching time during the post-invasion era was

61 seconds (95% CI= 43-78, n= 50) on Toho and was 75 seconds (95% CI= 58-93, n= 46) on

Kiss-WCA3A (Figure 2-16). Barring the pre-invasion average from Toho, our results showed a

trend opposite from that in Prediction 1, and considering all of the data we found no evidence

that the exotic snail affects average searching times.

Effects of the exotic snail on average handling times

We compared handling times between the adults of each sex and found that the average

handling times for adult males (on Toho, x=261 seconds, 95% CI= 198-323, n= 27; on Kiss-

WCA3A, x= 71, 95% CI= 65-77, n= 153) and females (on Toho, x= 319 seconds, 95% CI=

245-392, n= 56; on Kiss-WCA3A, x= 75, 95% CI= 70-81, n= 132) were not significantly

different (Figure 2-17). Thus, we found no evidence for Prediction 5a; however, these results

did support Prediction 2.

After combining males and females we found that the average handling time for adults on

Toho during the post-invasion era was 302 seconds (95% CI= 255-349, n= 85), while it was 78









seconds (95% CI= 75-123, n= 130) on Toho during the pre-invasion era and 72 seconds (95%

CI= 68-75, n= 324) on Kiss-WCA3A during the post-invasion era and (Figure 2-18).

We found that the average handling time for juveniles eating exotic snails on Toho was

496 seconds (95% CI= 376-617, n=44). In contrast, we found that the average handling time for

juveniles eating native snails on Kiss-WCA3A was 96 seconds (95% CI= 71-122, n=22) (Figure

2-19). Thus, we found that the handling times for exotic snails were significantly higher for each

age class relative to the handling times for native snails, which also supported Prediction 2.

Additionally, juveniles spent significantly longer than adults handling exotic (but not native)

snails, which supported Prediction 4a.

Effects of the exotic snail on drop rates

Under normal foraging conditions (e.g., no heavy wind or rain) on Toho, we observed

adults capturing 309 snails and juveniles capturing 199 snails, with raw group drop rates of

20.3% and 42. 1% respectively. Under normal foraging conditions in Kiss-WCA3A, we

observed adults capturing 1796 snails and juveniles capturing 3 86 snails, with raw group drop

rates of 2.7% and 5.3% respectively (Figure 2-20). The raw drop rate we found for juveniles on

Toho (i.e., 42. 1%) was comparable to the raw drop rate reported for unidentified kites on Toho

(i.e., 44%) by Darby et al. (2007).

These raw group drop rates did not account for individual variation and were inappropriate

for our comparisons between/among groups; therefore we used the drop rates from all of the

individuals within each group to calculate average group drop rates. Doing so, we found a

significant difference between the average drop rates of adult males and adult females on Toho.

Adult males (x= 0. 14, 95% CI= 0. 11-0. 17, n= 70) had a lower average drop rate than adult

females (x= 0. 19, 95% CI= 0. 15-0.22, n=109), and this trend was opposite from that which we

expected; therefore, we found no support for Prediction 5b. On Kiss-WCA3A, we found that the










average drop rates for adult males (x= 0.02, 95% CI= 0.01-0.03, n= 186) and adult females (x=

0.03, 95% CI= 0.01-0.04, n= 175) were similar (Figure 2-21), which, when compared to Toho,

lent support to Prediction 3.

We found that the average drop rate for juveniles on Toho was 0.33 (95% CI= 0.28-0.39,

n= 110) and that on Kiss-WCA3A it was 0.06 (95% CI= 0.03-0.09, n= 39). After pooling adult

males and females, we found the average drop rates for adults were 0. 17 (95% CI= 0. 15-0. 19, n=

187) on Toho and 0.02 (95% CI= 0.01-0.03, n= 405) on Kiss-WCA3A (Figure 2-22). Hence,

we found that drop rates for adults and juveniles increased significantly when handling exotic

snails, which supported Prediction 3. We also found support for Prediction 4b in that the

average drop rate of adults was lower than that of juveniles when foraging on the exotic snail.

Effects of the exotic snail on capture rates

We found that capture rates did not differ significantly between adult males (x= 3.13

snails/hour, 95% CI= 2.75-3.51, n= 179) and adult females (x= 3.18 snails/hour, 95% CI= 2.78-

3.58, n= 169) when foraging on native snails from Kiss-WCA3A, but when foraging on exotic

snails from Toho their capture rates did differ (males, x= 1.27 snails/hour, 95% CI= 1.13-1.41,

n= 70; females, x= 0.98. snails/hour, 95% CI= 0.95-1.00, n= 109) (Figure 2-23). However, we

did not feel that the difference between the average capture rates of adult males and adult

females on Toho justified separating the adult age class by sex before making comparisons with

juveniles. First of all, average adult capture rates did not differ between sexes in Kiss-WCA3A,

from which we had a larger sample size. Additionally, capture rates are more sensitive to

proximate conditions (e.g., temperature, season specific behavioral patterns) than are handling

times and drop rates, so the variation we observed could have resulted from such conditions.

We found that during the post-invasion era the average capture rate of adult kites was

significantly lower on Toho (x= 1.09 snails/hour, 95% CI= 1.04-1.13, n= 187) than it was on









Kiss (x=3.22 snails/hour, 95% CI= 2.76-3.69, n= 129) or on WCA3A (x= 3.38 snails/hour, 95%

CI= 3.03-3.73, n= 259). We also found that adult capture rates on Toho were lower in the post-

invasion era (see above) relative to the pre-invasion era (1.95 snails/hour, 95% CI= 1.61-2.29,

n= 65) (Figure 2-24). We found similar trends when comparing the average capture rates of

juvenile kites foraging on Toho (x= 0.77 snails/hour, 95% CI= 0.69-0.85, n= 110) with those on

Kiss-WCA3A (x= 3.46 snails/hour, 95% CI= 2.49-4.44, n= 37) (Figure 2-25). These results

provided strong support for Prediction 6.

Effects of the Exotic Snail on Energetics

Estimates of caloric content

With the length measurements from the empty exotic shells collected during TABs, we

predicted the weights of the edible soft body parts (no shell, no operculum, and no albumen

gland) of exotic snails captured by kites on Toho using the following linear model: wet weight of

soft body parts = (0.8143635 shell length 8. 08201). We converted wet weight to dry

weight by multiplying by 0. 1215 (the proportion of dry weight to wet weight of soft body parts),

and then we took the product of dry weight and caloric content, 3.30 kcal/g dry weight (Table 2-

3). We found that kites obtain an average of 12.92 kcal (95% CI= 11.62-12.27) per exotic apple

snail.

We predicted the energetic content of the native shells collected during TABs using the

following linear model from Sykes 1987(a): caloric content = (0. 6769 shell length- 20. 3448)

* 0.145 4. 60. We found that kites are obtaining an average of 4.84 kcal (95% CI= 4.60-5.07)

per native snail. Thus, kites obtain significantly more gross energy from exotic apple snails than

from native snails (Figure 2-26).









Profitability

Using average searching times and handling times for each group of kites and the average

caloric content of each snail species, we found the following profitabilities: adults feeding on the

exotic snail, 2.10 kcal/min; adults feeding on the native, 1.75 kcal/min; juveniles feeding on the

exotic snail, 1.39 kcal/min; juveniles feeding on the native, 1.47 kcal/min. We found that exotic

snails are more profitable than natives for adults but that native snails are more profitable than

exotics for juveniles. Hence, Prediction 7 holds true for juveniles but not for adults.

Validation of time activity budget extrapolations

We found evidence validating our assumption that day-long time activity budgets could be

extrapolated from shorter observations throughout the day. On Toho, the average proportion of

time spent flying during day-long (x= 0.108, 95% CI= 0.086-0.131) and hourly (x= 0.99, 95%

CI= 0.077-0. 121) TABs was almost identical. We found a similar level of agreement between

day-long (x= 0. 148, 95% CI= 0. 112-0. 174) and hourly (x= 0. 158, 95% CI= 0. 131-0. 184) TABs

on WCA3A. On Kiss, the average proportion of time spent flying during day-long TABs was

0. 158 (95% CI= 0. 113-0. 185), while during hourly TABs it was 0. 192 (95% CI= 0. 164-0.221);

however, the confidence intervals still overlapped (Figure 2-27).

We found additional evidence that behavioral patterns observed during hourly observations

were representative day-long activity patterns when we compared the average capture rates

among morning, afternoon, and evening TABs. For this analysis, we combined Kiss and

WCA3A because of the similar average capture rates observed for each area (Figure 2-24). On

Toho, average capture rates in the morning and evening were higher than in the afternoon, but

there was no significant difference between morning and evening or between afternoon and

evening. On Kiss-WCA3A, we found that average capture rates decreased throughout the day

from morning to afternoon to evening, but all confidence intervals overlapped so this trend was









not significant (Figure 2-28). Since we found no clear pattern of average capture rates varying

significantly among different times of the day, which agreed with the findings presented in the

literature (Cary 1985; Sykes 1987a), we felt comfortable with the assumption that hourly TABs

were representative of day-long TABs.

Daily activity patterns

On average, we found that kites spent 85.6% of their diurnal activity time perched and

14.4% flying (SD=14.4), which agrees with the patterns recorded by previous behavioral studies

(e.g., Cary 1985; Bennetts and Kitchens 1997) (Table 2-4). However, we found that theses

proportions differed significantly between Toho (flying, x= 0. 104, 95% CI= 0.0761-0. 133)

compared to Kiss (flying, x= 0. 157, 95% CI= 0. 124-0. 190) and WCA3A (flying, x= 0. 142, 95%

CI= 0. 111-0. 174) (Figure 2-29). This made biological sense given that we found no difference

in the average search times for Toho, Kiss, and WCA3A but that we did find that capture rates

on Toho were much lower that on Kiss-WCA3A (see E~ffects of the exotic snail on average

handling times and Effects of the exotic snail on drop rates). In addition, we found a trend across

areas that juveniles spend more time flying than adults, but this trend was not significant (Figure

2-30).

Daily energy balances

Assuming a 12 hour photoperiod, we used the average capture rate from each group of

kites (see E~ffects of the exotic snail on capture rates) to estimate the number of snails captured

per day and to derive the gross daily energetic gain from such consumption. We estimated that

adults on Toho consumed between 18-20 snails/day (i.e., 157 kcal/day, 95% CI= 150-164) and

that they consumed 41-48 snails/day on Kiss (i.e., 174 kcal/day, 95% CI= 168-180) and

between 44-52 snails/day on WCA3A (i.e., 166 kcal/day, 95% CI= 158-174). We estimated

that juveniles consumed between 15-18 snails/day on Toho (i.e., 110 kcal/day, 95% CI= 105-










116) and between 37-57 snails/day on Kiss-WCA3A (i.e., 178 kcal/day, 95% CI = 170-186)

(Figure 2-31).

We found that kites on Toho expended approximately 133 kcal/day while those on Kiss-

WCA3A expended approximately 138 kcal/day. Therefore, the net daily energy gain for adults

on Toho was 24 kcal/day and, while it was 36 on Kiss and 28 on WCA3A. The daily energy

balance for juveniles was 40 kcal/day on Kiss-WCA3A, while, on average, there was a net loss

of 23 kcal/day for juveniles on Toho (Figure 2-32).

Discussion

It is evident that the exotic apple snail affects several aspects of snail kite foraging

behavior. When compared to the average handling times for native snails, handling times for

exotic snails are extremely inflated, with adults and juveniles respectively taking three and five

times longer to negotiate, extract, and consume exotic snails. Kites also experience elevated

drop rates when foraging for exotic snails. On average, adults drop over eight times more exotic

snails than native snails (17% versus 2%). Juveniles, already dropping around 6% of the native

snails that they handle, experience an average drop rate of 33% when foraging for exotic snails.

We found that, on average, kites are consuming native apple snails that fall between 35 to 45 mm

in length and weigh around 16 grams, while the exotic snails that they typically consume range

from 45 to 81 mm and weigh between 50 to 60 grams. It is likely that the larger relative

proportions of the exotic snail are underlying, at least in part, the increased handling times and

drop rates. However, we believe that experience may affect handling times and drop rates more

so than does the relative size/weight ratio of predator to prey. While handling times and drop

rates increase significantly for all groups of kites when foraging on exotic apple snails, there is

an obvious discrepancy in the magnitude of the effect between adults and juveniles, but this

discrepancy cannot be attributed to relative size/weight differences alone. Adult kites display










reverse sexual dimorphism, with males weighing up to 10% less than females, yet while foraging

on exotic snails, adult males maintain significantly lower drop rates than adult females. This

contradicts our expectations that were based on relative weight ratios alone and suggests that the

lack of experience in juveniles exacerbates the problems they have with negotiating exotic apple

snails.

While the negative effects of the exotic apple snail on the aforementioned foraging

behaviors of the snail kite may seem debilitating, they cannot be viewed, substantively, outside

the context of energetic ramifications. By dry weight, the edible body tissue of the exotic snail

contains 3.30 kcal/g, while that of the native snail contains 4.60 kcal/g. Interestingly, our results

for the exotic snail (i.e., P. insularum) are similar to those reported for P. canaliculata, 3.37

kcal/g (Catalma et al. 1991; PhilRice 2001). Although the edible tissues of exotic apple snails

contain less energy per gram, whole exotic snails still contain more total energy than whole

native snails, and this is due to the relative difference in their average sizes. The average exotic

snail contains 12.92 kcal, while the average native snail contains 4.84 kcal. In light of these

findings, other effects of the exotic snail on snail kite foraging behavior seem to make biological

sense.

Adult kites on Toho achieve an average capture rate of 1.09 snails/h, which is significantly

lower than their average capture rates on Kiss (3.22 snails/h) and WCA3A (3.38 snails/h). The

average capture rates of juveniles display a similar trend, 0.77 snails/h on Toho and 3.46 snails/h

on Kiss-WCA3A. While the difference in capture rates between adults and juveniles on Toho

may be a consequence of juveniles' increased negotiability problems with the exotic snail, the

overall trend in capture rates between Toho and Kiss-WCA3A is likely due to the fact that

similar energetic benefits can be obtained by consuming fewer exotic snails. The exotic snail has









no direct measurable affect on searching time. Even though the elevated drop rates associated

with exotic snails result in kites making more attempts before successfully capturing and

consuming exotic snails, both adults and juveniles have shorter average searching times per

success on Toho than they do on Kiss and WCA3A. Therefore, the trend we observe in capture

rates is not a product of increased searching times for exotic snails.

Searching time, however, is strongly affected by the abundance and availability of apple

snails. Even though some of the exotic snails on Toho may be unavailable due to the inability of

kites to negotiate them, kites may still maintain short searching times if exotic snails exist in a

high enough abundance. Estimates of exotic snail abundance specific to the vegetative

communities exploited by foraging kites on Toho preclude our ability to disentangle the

relationship between exotic snails, searching times, and capture rates. If the short searching

times on Toho are being maintained by a superabundance of exotic apple snails, then under

different conditions (i.e., lower snail densities) the drop rates and presumably longer searching

times associated with the exotic snail could translate into much greater energetic expenditures for

the snail kite, potentially causing the profitability of exotic snails to fall below levels that are

energetically sustainable for snail kites. Elucidating the affects of exotic snail density on

average snail kite searching times and capture rates should receive immediate attention given

that populations of the native apple snail have been declining throughout the kites' range (Darby

et al. 2005) while populations of the exotic snail have been spreading (Rawlings et al. 2007) and

that exotic snail densities will likely vary among wetlands. In South America, the range of the

snail kite (R. s. sociabilis) overlaps the natural range of P. insularum (i.e., what we call the

exotic snail) but there is no documentation that kites feed on these snails (Sykes et al. 1995;

Darby et al. 2007), which provides reason for further concern if one considers the possibility that










in its natural habitat, P. insularum, may exist in densities too low to provide a sustainable

wforaging base for kites.

Our estimates of DEE are comparable to those reported in Beissinger (1984). Assuming

that the conditions experienced by each group of kites on Toho, Kiss, and WCA3A during the

post-invasion era are representative of average conditions and that the searching times, handling

times, capture rates, and activity patterns of each respective group are representative of average

responses to such conditions, then adults kites can maintain comparable daily energy balances

whether feeding on the exotic or the native snail; however, juveniles, while maintaining energy

balances similar to those of adults on Kiss-WCA3A, are on average experiencing net energy

losses of around 20 kcal/day when foraging for exotic snails on Toho. This is concerning given

that multiple authors (e.g., Kirkwood 1981; Newton 1991) have shown that raptors successively

failing to meet daily energy requirements may face a high risk of mortality within several days to

a week.

Our analyses of energy balances are crude and we present the findings here to demonstrate

the possible energetic consequences of the effects that the exotic apple snail has on snail kite

foraging behavior. We utilized some parameters (e.g., rates of energetic expenditure for flying

and food handling; the caloric contents of native apple snails) that were reported in previous,

possibly outdated, literature (e.g., Beissinger 1984; Sykes 1987a). In order to conclude

unequivocally that juvenile kites on Toho fail, on average, to meet their daily energetic

requirements, we recommend that more advanced methods of energetic analysis be employed.

We provide here a valuable starting point and suggest that serious efforts should be made to

apply recent work by Porter et al. (1994, 2000, 2002, 2003) to the snail kite. Using

spatiotemporally explicit models that can account for variations in the local microclimate as well









as in the biophysical aspects of individual snail kites and their prey will provide more robust and

reliable estimates of daily energy balances.










Table 2-1. Number of TABs conducted by month and location in 2003, 2004, 2005, 2006 (post-
invasion) and in 1993, 1994, 1996 (pre-invasion).
Spring Summer Fall
Era Location JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
TOHO 4 15 0 2 13 5 53 51 53 36 29 25
KSS 17 26 8 2 7 23 12 4 5 2 12 2
WCA3A 36 53 8 2 34 2 41 23 11 38 46 54
SWA2B 0 2 0 0 0 0 2 1 9 0 0 0
WC A3B 0 0 8 0 0 0 0 0 0 0 0 0
.=EP 0 0 0 0 0 0 0 0 0 0 4 0
E OEE0 0 0 0 0 0 0 0 0 0 0 3
SJM0 0 5 0 4 2 2 0 0 0 0 3
EOHO 0 0 0 0 0 0 0 0 0 0 0 0
TOHO 4 9 0 2 9 42 48 49 47 33 29 20
SKSS 14 22 5 2 6 21 11 4 4 18 12 19
uJ WA3A 32 36 7 2 25 20 33 21 8 32 38 41
0 WA2B 0 2 0 0 0 0 1 1 9 0 0 0
SWA3B 0 0 4 0 0 0 0 0 0 0 0 0
-EP 0 0 0 0 0 0 0 0 0 0 3 0
8 KE 0 0 0 0 0 0 0 0 0 0 0 2
SJM0 0 0 0 4 0 2 0 0 0 0 3
EOHO 2 2 2 0 0 2 3 0 1 2 0 7






Table 2-2. Nutritional contents of P. insularum.
Nutritional Measurement S1 S2 S3 Average SD
Calories 49.5 36.8 43.3 43.2 6.35
Calories from Fat 4.2 3 3.2 3.47 0.64
Total Fat (g) 0.47 0.33 0.35 0.38 0.08
Saturated Fat (g) 0.19 0.12 0.13 0.15 0.04
Total Monounsaturated Fat (g) 0.08 0.09 0.08 0.08 0.01
Total Polyunsaturated Fat (g) 0.16 0.1 0.11 0.12 0.03
Trans Fat (g) 0.01 0.01 0.01 0.01 0
Cholesterol (mg) 94.2 66.8 55.7 72.23 19.82
Sodium (mg) 71.6 97.2 89.7 86.17 13.16
Total Carbohydrate (g) 1.5 0.7 2.3 1.5 0.8
Protein (g) 9.82 7.76 7.74 8.44 1.2
Calcium (mg) 464 535 681 560 110.64
Iron (mg) 59 43.6 43.3 48.63 8.98
Moisture (g) 84.21 89 86.7 86.64 2.4
Ash (g) 3.99 2.25 2.89 3.04 0.88
Note: Nutritional content is measured per 100g of wet weight of edible soft body tissues.











Table 2-3. Nutritional composition of P. insularum versus P. paludosa.
P. insularum P. paludosa P. paludosa
Cattau et al. 2007 Beissinger 1984 Sykes 1987a
Kcal/g 3.25 +/- 0.11 4.52 +/- 0.25 4.60 +/- 0.18
% Fat 2.87 +/- 0.21 3.11 +/- 0.49 3.4
% Carbohydrate 11.05 +/- 5.63 NA 27.4
% Protein 63.63 +/- 6.28 61.5 +/- 5.2 36.3
% Moisture 13.37 +/- 2.40 NA 14.5
% Ash 22.48 +/- 2.50 9.59 +/- 1.16 32.1
Note: Nutritional Contents of apple snails reported as % dry weight of edible soft body parts, mean +/-
standard deviation.


Table 2-4. Average daily activity pattern of the snail kite in Florida.
Time Standard
Investigator Years Covered by Study Perched Time Flying Deviation
Cary 1985 1977, 1978, 1979 85.30% 14.70% 15.60%
Bennetts and Kitchens 1997 1993, 1994, 1996 86.10% 13.90% 14.00%
Cattau et al. 2007 2004, 2005, 2006, 2007 85.60% 14.40% 14.40%

























I I I I II eF I


Figure 2-1. Relative sizes of exotic apple snails. A) 50, 63, and 80 mm in length. B) 50 and 80
mm in length.





































Figure 2-2. Measurement of snail shells. A) Shell width. B) Shell length.


I slundl TIaps


Gonhless Corer Tlips


shones
TaysP


Ealss shourI Traps

m


Tip


South sholr


*. .


Figure 2-3. Location of apple snail traps on Toho.
















OO


ogo oo
o oo o ono
a -l o o
OO O~ OOO
oooo o At
a o 8 I 8 ooo a o
c~~~ o 0oEs o5g
5 0 0 o e 0g i 0 g g 0
o o o ao
a, o aa o o
oo g o o oo

o Be


oo8 oo

oo
I- I I


20 30 40 50 60 70 80

length (Rn)




O



Oi O

o oo oi
go O O O 80 O
c~ 80

oS eoo oo
o o~ o8
8 8 a0

a gge8 co,
ao e
20 30 40 580 60 708





functio ofsel egh


















o Native Apple Snail
SExotic Apple Snail


o
-


O
o


O
0-
Ln







0-


2005


2006


2007


year



Figure 2-5. Number of native and exotic apple snails captured per year with funnel traps on
Toho, 2005-2007.

















O native
Exotic


o


ai

d-

a


E.shore Gob.Cove


L.grassy


S.shore W.shore


Lake Location



Figure 2-6. Proportion of native to exotic apple snails captured in five locations on Toho, 2005-
2007.























0

o

o o


o


o
0


o


ENP WCA3A WCA3B WCA2B


OKEE GW


SJM KISS TOHO


LOCATAlON


Figure 2-7. Average length of apple snail shells collected from feeding perches and nests in nine
wetlands throughout the range of the snail kite, 2004-2007. Shaded boxes represent
95% CIs. Lower and upper whiskers represent first and third quartiles respectively.
Open circles represent extreme outliers.


TT
o A


a





























O_


exotic shell width


exotic shell length


native shell width


native shell length


Figure 2-8. Average shell dimensions of native and exotic apple snails consumed by kites, 2004-
2007. Shaded boxes represent 95% CIs. Lower and upper whiskers represent first
and third quartiles respectively. Open circles represent extreme outliers.

















































I


CO



O











CO

-


Native Snails


Exotic Snails


Figure 2-9. Estimated whole weight of native and exotic snails negotiated by foraging kites
during the post-invasion era. Error bars represent 95% CIs.





















80





I I


TOHO.adults KISS-WCA3A.adults


I
III
I


TOHO.subadults


TOHO.juveniles KISS-WCA3A.juveniles


KISS-WCA3A.subadults


LOCATION.age



Figure 2-10. Average handling times for adult, juvenile, and subadult kites on Toho and Kiss-
WCA3A during the post-invasion era. Shaded boxes represent 95% CIs. Lower and

upper whiskers represent first and third quartiles respectively.

































o
TOOa OOsbd THOu ISWAa ISWAsbd KS-C~u


LOAIOUg


Fiue21.Aeaedo aesfraut ueie n ubdl ie nTh n is
WC 3 duigteps-nainea

















LO


CV


O


TOHO.juy KISS-WCA.adul


KISS-WCA.subad


KISS-WCA.juy


TOHO.adult TOHO.subad


LOCAllON.age


Figure 2-12. Average capture rates for adult, juvenile, and subadult kites on Toho and Kiss-
WCA3A during the post-invasion era. Error bars represent 95% CIs.


























I




















88





I
I


I


I


I



I
I


I


I



TOHO.male


KISS-WCA3A.female


TOHO.female


KISS-WCA3A.male


LOCATION.sex




Figure 2-14. Average searching times for adult male and female kites on Toho and Kiss-WCA3A
during the post-invasion era. Shaded boxes represent 95% CIs. Lower and upper

whiskers represent first and third quartiles respectively.























O









I II

Too(otivso) Th peivso) W AA(otivso) Ks ps-nain
LOAIN ea



Fiue21.Aeaesacigtmsfrautktso isadW AAdrn h ot
inaineaado ooduigtepe n otinainea.Sae oe
rereen 95 .Lwradupr hsesrpeetfrs n hr urie
repciey

















O



O
O-


CY

O









cn-i
UI I

TOOKS-C3

LOATO

Fiure21.Aeaesacigtmsfrjvnl ie nTh n isW AAdrn h
potivso r.Sae bxsrpeet9%Cs Lwraduprwikr
rersn is adtidqatle epciey











































I
I
I
I
I
I
I
I
I
I
I


KISS-WCA3A.male


KISS-WCA3A.female


I



TOHO.male


TOHO.female


LOCATION.sex





Figure 2-17. Average handling times for adult male and female kites on Toho and Kiss-WCA3A

during the post-invasion era. Shaded boxes represent 95% CIs. Lower and upper

whiskers represent first and third quartiles respectively.









































88





WCA3A-KISS (post-invasion)


Toho (pre-invasion)


LOCATION (era)


Figure 2-18. Average handling times for adult kites on Kiss-WCA3A during the post-invasion
era and on Toho during the pre- and post-invasion eras. Shaded boxes represent 95%
CIs. Lower and upper whiskers represent first and third quartiles respectively.






















89


Toho (post-invasion)





I I


O
0-
hi



0-


TOHO


KISS-WCA3A


LOCATION



Figure 2-19. Average handling times for juvenile kites on Toho and Kiss-WCA3A during the
post-invasion era. Shaded boxes represent 95% CIs. Lower and upper whiskers
represent first and third quartiles respectively.























o


O





O,


o

c5 -
TOOaut OOaueie IS-C~dlsKSSWAjvnls









Figre2-0.Ra goHaup ro ats o adO~uvnlts an ueiekites on uToh n Kiss-W CA3A during



the post-invasion era





























o


O







c5,

TOOfmls THoae ISWAfmls KS-C~ae

LOAIOre


Figur2-1AvrgdrprtsfrautmlanfeaektsoTooadKs-C3
duigteps-naso r.Errbrsrpeet9%Cs





























o


O








o
c5 -
TOOjvnls THaaut ISWAjvnie ISWAaut











Figure2-22.Averae dopraes for aultandut KIWjuvenile ieso oh n Kiss-WCA3A during




the post-invasion era. Error bars represent 95% CIs.


































































m-


L
O

V)

I~
V)
r\l-


I


0-


Kiss-WCA3A.males


Kiss-WCA3A.females


TOHO.females


TOHO.males


LOCATION.sex





Figure 2-23. Average capture rates for adult male and female kites on Toho and Kiss-WCA3A

during the post-invasion era. Error bars represent 95% CIs.






































KISS (post-invasion)


TOHO (post-i nvasi on) WCA3A (post-i nvasi on) TOHO (pre-i nvasi on)


LOCATION (era)


Figure 2-24. Average capture rates for adult kites on Kiss and WCA3A during the post-invasion
era and on Toho during the pre- and post-invasion eras. Error bars represent 95% CIs.





O








KISWAATH








KISS-WLOCATION




Figure 2-25. Average capture rates for juvenile kites on Toho and Kiss-WCA3A during the post-
invasion era. Error bars represent 95% CIs.
























































I


hi




O


CO


hi




0-


Native Snails


Exotic Snails


Figure 2-26. Estimated energetic content of native and exotic snails consumed by kites during
the post-invasion era. Error bars represent 95% CIs.





























































I I I I


O
(3
O




LO
CI~
O


rn
o
i. ~u
c
o
c
a,
a
m
a,
LO
E
c o
c
o

o

o o

o o
a


I


TOHO (daylong) TOHO (houdly) KISS (daylong) KISS (hourly) WCA3A (daylong)


WCA3A (hourly)


LOCATION (length of observation)





Figure 2-27. Average proportion of time spent flying during day-long and hourly observations by

kites on Toho, Kiss, and WCA3A in the post-invasion era. Error bars represent 95%

CIs.















LO


~-I II II








Toho.morning Toho.afternoon Toho.emening Kiss-WCA.morning Kiss-WCA.afternoon Kiss-WCA.emening


Location.daytime.


Figure 2-28. Average capture rates achieved by foraging kites during morning, afternoon, and
evening observations on Toho and Kiss-WCA3A in the post-invasion era. Error bars
represent 95% CIs.


















O



C3l
O








On

OO



O-

Os

O
O-h
d
KISTOOWC3

LOATO


Fiue22.Aeaepooto ftm pn fyn yalktso is oo n C 3
duigteps-naso r.Errbrsrpeet9%Cs


















I10
















I


~


KISS.adult WCA3A.adult


TOHO.ad ult


TOHO.juy KISS-WCA3A.juy


LOCATION.age

Figure 2-30. Average proportion of time spent flying by adult and juvenile kites on Toho and
Kiss-WCA3A during the post-invasion era. Error bars represent 95% CIs.












101





O


WCA3A.adults


KISS.adults


TOHO.adults


TOHO.juveniles KISS-WCA3A.juveniles


LOCATION.age



Figure 2-31. Estimated gross daily energetic gains for adult and juvenile kites foraging on Toho,
Kiss, and WCA3A during the post-invasion era. Error bars represent 95% CIs.
































102





I
TOHO.adults


I
WCA3A.adults


I
KISS.adults


I I
TOHO.juveniles KISS-WCA3A.juveniles


LOCATION.age


Figure 2-32. Estimated daily energy balances for adult and juvenile kites on Toho, Kiss, and
WCA3A during the post-invasion era. Error bars represent 95% CIs.


T









CHAPTER 3
DEMOGRAPHIC EFFECTS OF THE EXOTIC APPLE SNAIL ON THE KITE

Introduction

Compared to other vertebrate species, birds, especially those that employ active foraging

strategies, have high metabolic rates and internal temperatures. Physiological conditions of such

species can only be maintained through frequent energy acquisition (Blem 1976; Krebs &

Kacelnik 1991; Carey 1996). Many species, such as the snail kite (Beissinger & Snyder 1987;

Bennetts & Kitchens 2000), face the risk of starvation daily, as failing to meet minimum

energetic thresholds over the course of several days to a week will likely result in mortality

(Kirkwood 1981; Newton 1991). Given the vital necessity for kites to maintain proper energy

balances, foraging takes precedence over other behaviors. Thus, foraging behavior directly

influences the amount of time and energy an individual can afford to allocate toward growth and

reproduction (Krebs & Kacelink 1991; Carey 1996; Newton 1991). As shown, the exotic apple

snail has significant effects on the foraging behavior of snail kites, especially juveniles (Chapter

2), and thus it is important to determine what demographic consequences the kite population may

face as a result. Martin (2007) urges that "under the current system, adult fertility appears to be

crucial [to the viability of the snail kite population], and factors likely to have large effects on

adult reproduction and juvenile survival should receive more attention." To determine whether

the exotic apple snail constitutes an environmental stressor that has negative demographic

consequences for the kite population, we compare measures of reproduction (including nesting

effort, nest success, and nest productivity), along with estimates of survival, from Toho between

the pre- and post-invasion eras.










Reproduction

The reproductive activity of snail kites is limited, in large part, by the availability of food

resources (Nichols et al. 1980; Snyder et al. 1989). The height of the breeding season stretches

from March through June, but kites may breed during any month of the year (Beissinger 1988;

Snyder et al. 1989; Bennetts & Kitchens 1997). Kites are sensitive to proximate environmental

conditions throughout the breeding process. In most circumstances, adult kites attempt to breed

at least once a year and, on occasion, have been observed successfully fledging two broods in the

same breeding season (Snyder et al. 1989; Bennetts & Kitchens 1992, 1997). On the other hand,

kites may forgo nesting completely in years of severe droughts (Sykes 1979b; Nichols et al.

1980; Beissinger 1986; Snyder et al. 1989). Kites may also limit reproductive effort during other

times of food scarcity (Beissinger 1986; Bennetts et al. 1988; Snyder et al. 1989). In addition to

influencing the decision of whether or not to commence nesting, environmental conditions can

also affect kite behavior after nesting has begun. Adults may abandon initiated nests or

provision fewer young in times of food stress, resulting in lower nest productivity (Beissinger &

Snyder 1987; Sykes 1987b; Bennetts et al. 1988). Nest failures not resulting from parental

abandonment are usually the result of either depredation or structural collapse (Sykes 1979a,

1987b; Bennetts et al. 1988; Snyder et al. 1989; Bennetts & Kitchens 1997).

Survival

Kite survival has been linked to a number of factors, but prey availability plays a critical

role, as starvation is thought to be the leading cause of mortality (Sykes et al. 1995). After

reaching independence, juvenile kites commonly become emaciated, especially when dispersing

through unfamiliar territory in which adequate prey is not available (Bennetts & Kitchens 1992,

1997; Martin et al. 2007b). Likewise, most adult mortalities are linked to starvation during

severe droughts (Beissinger 1988; Bennetts & Kitchens 1997; Martin et al. 2006a, 2007b).










Demographic evidence suggests that the snail kite population can be divided into two age classes

(juveniles and adults) and that after approximately 5 months of age, the survival probability of

juveniles effectively parallels that of adults (Bennetts and Kitchens 1999; Bennetts et al. 1999;

Dreitz et al. 2004; Martin et al. 2006a).

Predictions

Relative to adult kites feeding on native snails, adults feeding on exotic snails spend more

time handling prey, capture fewer snails per day, and achieve lower daily energy balances

(Chapter 2). If these behavioral discrepancies translate into significantly less available time or

energy for adults on Toho during the post-invasion era, then we would expect to see a decrease

in the occurrence and/or success of behaviors requiring additional energy reserves, such as

reproduction. It must also be considered that the foraging behaviors of adults and juveniles are

differentially affected by the exotic apple snail (Chapter 2). Given that adult kites maintain

positive daily energy balances while feeding on exotic snails, we assume that the exotic snail

does not represent a threat to adult survival. The foraging behavior of juvenile kites, on the other

hand, is severely impacted by the exotic snail, and preliminary estimates indicate that juveniles

on Toho may not be achieving sustainable daily energy balances (Chapter 2). In light of our

previous findings, we made the following predictions:

* Prediction 1. Average annual nesting effort on Toho is significantly lower during the post-
invasion era than it was during the pre-invasion era.

* Prediction 2. The average nest success on Toho is significantly lower during the post-
invasion than it was during the pre-invasion era.

* Prediction 3. Nest productivity (i.e., the number of young produced per successful nest) is
significantly lower during the post-invasion than it was during the pre-invasion era.

* Prediction 4. Nest productivity during the post-invasion era is significantly lower on Toho
relative to other wetlands utilized by the snail kite.










* Prediction 5. The survival of juveniles hatched on Toho is significantly lower during the
post-invasion era relative to the pre-invasion era.

* Prediction 6. The survival of adult kites that were hatched on Toho does not differ
significantly between the pre- and post-invasion eras.

Methods

Robust breeding-season population surveys, along with other long-term studies of

demography and movement of the snail kite population, began in 1992. Protocols utilized

systematic surveys throughout the range of the kite that coupled counts with capture-mark-

recapture methods, including band-resighting and radio-telemetry. A wealth of demographic

data (including survival estimates of juveniles and adults) and movement data have been

accumulated. Range-wide snail kite nesting data (including nesting attempts, nest success, and

nest productivity) has also been collected in conjunction with the annual population monitoring

project. However, on Toho, not all data types were available for all years (see below) (For

detailed methods, findings, and discussions see Bennetts & Kitchens 1992, 1997, 2000; Dreitz et

al. 2001, 2004; Martin et al. 2006a, 2006b, 2007a, 2007b, 2007c). Yet, comparisons of

demographic parameters with specific regard to the invasion of Toho by the exotic apple snail

have not been made previously.

Based on recent demographic trends of the snail kite population and on the invasion

history of the of the exotic apple snail in Florida (Chapter 1), we defined 1992-1998 as the pre-

invasion era and 2003-2007 as the post-invasion era. These years also corresponded roughly

with our pre- and post-invasion TAB data (Chapter 2). The period 1999 to 2002 corresponds

with the severe decline of the snail kite population (Martin et al. 2006a; Martin 2007) and with

the first occurrences of the exotic snail on Toho (Darby, personal communication). To avoid

confounding effects associated with the population decline and with the unknown relative










proportion of native and exotic snails making up the diet of the snail kite on Toho, we defined

1999-2002 as the decline era and treated it independently from the pre- and post-invasion eras.

Nesting Effort and Reproductive Success

All nests observed with at least one egg or one chick during any monitoring visit were

classified as initiated (i.e., active). Kites commonly build nests that are never initiated (Chandler

& Anderson 1974; Beissinger 1984; Snyder et al. 1989; Bennetts & Kitchens 1997); therefore,

nests in which no eggs or young were ever observed were censored from our analysis (from here

on nests refers to initiated nests only). All nests in which at least one chick was observed to

reach potential fledging age, 24 days old (Snyder et al. 1989), were classified as successful, and

all chicks that reached at least 24 days old were assumed to fledge. Nests that failed before

fledging young were classified as unsuccessful. Nests of unknown fate were censored from our

analyses. The average fledging age of kites is 28.7 days (Sykes 1987b), but it ranges from 24 to

35 days (Synder 1989). We used this 24-day benchmark to reduce the likelihood of missing

fledging events, which would result in the misclassification of these nests as failures or as nests

of unknown fate. We do not feel that using the 24-day benchmark biased our estimates of nest

success or nest productivity, as mortality during this nesting stage is minimal for kites (Steenhof

& Kochert 1982; Bennetts & Kitchens 1997; Dreitz et al. 2001).

Measures commonly used to assess the relative reproductive effort of snail kites in specific

wetland units during a given year include 1) the number of breeding individuals present and 2)

the number of nests initiated. The latter is less ambiguous and provides a more tangible measure

of nesting effort; therefore, we used the average number of nests initiated annually on Toho to

compare the pre- and post-invasion eras (2004 was excluded from the post-invasion average).

Nest abundance can vary radically between 'wet' and 'dry' years (Sykes et al. 1995), making it

hard to draw definitive conclusions when comparing two eras; therefore, we also looked at the









relative annual abundance of kite nests on Toho in contrast to other wetlands. We calculated the

relative contribution of Toho to the annual nesting effort of the entire kite population for the

years 1995 to 2007 by dividing the number of nests on Toho by the total number of nests found

range-wide in a given year (no nesting data was available for 1992-1994). Then we developed

two trend models to assess whether the contribution of Toho to the overall nesting effort has

changed over time. In the first trend model we included all years from 1995-2007. In the second

trend model we removed anomalous years, in which documented environmental phenomenon

may have drastically altered kite behavior. We censored 2001 and 2007, two years in which

severe droughts heavily affected the southern portion of the kites range (Martin 2007; Gawlik et

al. 2007; FLDEP 2007), causing uncharacteristic nesting patterns that could have potentially

biased Toho's relative contribution high. We also censored 2004, the year in which the managed

drawdown and scraping of Toho was completed, as it may have biased Toho's contribution low.

In addition to these trend models, we also estimated Toho's average contribution to the total kite

population nesting effort for the pre- and post-invasion eras, and since droughts and drawdowns

have tangible affects on the kite population, no years were censored in the calculation of these

estimates (i.e., 2004, and 2007 were included).

Measures commonly used to assess the reproductive success of snail kites include 1) the

proportion of successful nests to the total number of initiated nests (i.e., nest success) and 2) the

average number of young fledged per successful nest (i.e., nest productivity) (Sykes 1987b;

Bennetts & Kitchens 1997; Martin 2007). Nest success may be influenced by innumerable

environmental variables (Sykes et al. 1995). Snail kites inhabit a broad and fragmented

landscape, in which spatiotemporal heterogeneity cannot be ignored (Bennetts et al. 1998, Martin

et al. 2006). Several extraneous variables may affect the fate of a nest, and due to our inability to










account for such environmental stochasticity across the range of the snail kite, comparing nest

success among wetland units over time was not warranted. However, accounting for

environmental variation on a local scale was feasible, thus we compared the annual nest success

on Toho before and after the invasion of the exotic snail.

Nest productivity, the number of young produced per successful nest, is less affected by

drastic stochastic events (e.g., droughts, hurricanes, cold fronts), as these events usually result in

complete nest failure, which is not included in the calculation of nest productivity. The number

of young that adults can provision is dictated strongly by the availability of suitable prey

(Beissinger 1990). Thus, when making comparisons among areas or years, more subtle

differences in local foraging conditions can be revealed by comparing measures of nest

productivity rather than nest success (Murray 2000; Kosciuch et al. 2001). If adult kites cannot

sufficiently exploit exotic snails we would expect lower relative productivity in the face of such

conditions. We compared the number of young produced per successful nest on Toho before and

after the invasion of the exotic snail. We also compared the nest productivity of Toho during the

post-invasion era with that of all other wetlands (pooled) during the same time period.

Reliable nesting data for Toho was available for the period 1995 to 2007; however, no

nests of known fate were observed on Toho in 1998, and only one nest of known fate was

observed in 1999. Additionally, no nesting occurred on Toho in 2004. Therefore, estimates of

nest success and nest productivity for 1998 and 2004 were not calculable, and since estimates for

1999 were based on a sample size of one, confidence intervals were not calculable.

All 95% confidence intervals (95% CI) around our derived estimates of nesting effort, nest

success, and nest productivity were approximated as follows: CI = x + toz SE, where x is the

sample estimate (i.e., mean), t is the test statistic (evaluated at a= 0.05 on n-1 degrees of









freedom), and SE is the standard error of the mean. Such estimation is valid for positive sample

estimates of count, measurement, or proportion that generally follow a lognormal distribution

(Burnham & Anderson 2002). All statistical analyses were performed using the software

package R version 2.5.1.

Survival

Mark-resight data and radio telemetry data can each be used to generate reliable estimates

of apparent survival for snail kites, and both can incorporate site-specificity. While inferences

from mark-resight data are limited to between-year estimates, radio-telemetry can be used to

generate between- and within-year estimates of apparent survival (Bennetts et al. 1999, Martin et

al. 2007b).

From 1992 to 2006, 121 juveniles on Toho were marked with unique alphanumeric bands

(an additional 74 juveniles were banded on Toho in 2007 and will be used for future estimates).

Using mark-resight data from 1992 to 2007, we estimated the apparent annual survival of adult

and juvenile snail kites that were hatched on Toho. No juveniles were banded on Toho in 1993

or 1998, so no annual juvenile survival estimates were generated for these years. Additionally,

no juvenile survival estimates were generated for 2004 because the kites did not nest the year

that the drawdown and scraping treatment was completed. In the computation of annual juvenile

survival during the post-invasion era, we excluded 2003 because we could not account for

drawdown effects, which may have biased our estimate. Annual adult survival, on the other

hand, could be generated for all years (1992-2006). No survival estimates were generated for

2007 because this analysis is contingent on the completion of the 2008 population surveys.

Radio telemetry allows for the estimation of apparent monthly survival probabilities for

kites (Bennetts et al. 1999; Bennetts & Kitchens 1999). Ten juveniles on Toho were equipped

with radio transmitters in 1992 and five juveniles were equipped in 1994, so these years were










used to represent the pre-invasion era for juvenile survival. The only telemetry data available for

juveniles during the post-invasion era came from 2005, in which 14 juveniles on Toho were

equipped with radio transmitters (Bennetts and Kitchens 1997, 1999; Martin et al. 2006a).

Although working with limited sample sizes, we used this radio-telemetry data to estimate the

apparent monthly survival of juveniles hatched on Toho. Radio transmitters had an average

lifespan of over two years; therefore, we also generated estimates of apparent monthly survival

for adults that were hatched on Toho. Although kites often disperse to other wetlands after

fledgling (Bennetts & Kitchens 1997), natal area has a greater affect on juvenile survival than

dispersal area (Bennetts et al. 1999; Martin et al. 2007b). Kites also express natal philopatry,

thus adult survival is also linked with natal location (Martin et al. 2007b). We compared

monthly survival estimates for adults and juveniles hatched on Toho between the pre- and post-

invasion eras.

To generate annual and monthly estimates of apparent survival, we used program MARK

V 4.1 (White & Burnham 1999). We created a suite of apriori survival models based on our

predictions (see above) and the existing literature (Bennetts & Kitchens 1997, 1999; Bennetts et

al. 1999; Martin et al. 2006a, 2007a, 2007b). In our model sets, we allowed survival and

detection probability to vary over time or to be held constant. In addition, we tested an age effect

(i.e., adult versus juvenile), and we also tested group effects corresponding to the pre- and post-

invasion eras. After creating our model sets, we selected the most parsimonious model for

apparent annual survival using Akaike's Information Criteria corrected for overdispersion

(QAIC) and for apparent monthly survival using Akaike's Information Criteria corrected for

small sample size (AICc) (Burnham & Anderson 2004). The selected models were used

generate the respective estimates of annual and monthly survival. All associated standard errors









and confidence intervals were also generated using program MARK V 4. 1 (White & Burnham

1999).

Results

Reproduction

Nesting effort

The number and distribution of snail kite nests varies widely over time (Table 3-1; Figure

1-1), and this may be influenced by innumerable of factors (Sykes et al. 1995); therefore, these

results must be interpreted with caution. We found that the average number of nests initiated

annually on Toho was significantly greater during the post-invasion era (excluding 2004, x= 42,

95% CI= 29-55) than it was in the pre-invasion era (x=17, 95% CI= 9-26) (Figure 3-2). This

ran contrary to Prediction 1. Even though there was a high degree of variation in nesting effort

among some years (Figure 3-3), the difference between pre- and post-invasion nesting effort on

Toho was significant.

We found that Toho's proportional contribution to the annual population nesting effort

ranged from 0.01 to 0. 11 during the pre-invasion era and from 0. 17 to 0.81 during the post-

invasion era (or including 2004, the year in which no kites nested on Toho due to the drawdown,

from 0.00 to 0.81) (Table 3-1; Figure 3-4). Using our first trend model (including all years 1995-

2007) to assess the relative contribution of Toho to the overall population nesting effort over

time, we found a slightly significant positive trend (p= 0.06) (Figure 3-5). Our second trend

model did not include the drought years of 2001 and 2007, in which Toho' s relative contribution

was 0.74 and 0.81 respectively, nor did it include the drawdown year of 2004, in which Toho' s

relative contribution was 0.00 (Table 3-1). This second trend model showed a significant

positive trend in Toho's relative contribution (p= 0.04) (Figure 3-6). We also found that Toho's

average contribution to the overall nesting effort of the kite population was significantly higher









during the post-invasion era (x= 0.33, 95% CI= 0.30-0.35) than it was during the pre-invasion

era (x= 0.06, 95% CI= 0.05-0.07) (Figure 3-7). Thus Prediction 1 was false.

Nest success and productivity

While nest success on Toho varied significantly among some years (Figure 3-8), we did

not find any significant difference in the average nest success on Toho between pre- and post-

invasion eras (Figure 3-9), which were 0.25 (95% CI= 0.12-0.38) and 0.39 (95% CI= 0.27-0.51)

respectively. Thus Prediction 2 was false.

Similarly, nest productivity on Toho varied significantly among some years (Figure 3-10),

but again, we found no difference in the average number of young produced per successful nest

between the pre- and post-invasion eras, which were 2. 13 (95% CI= 1.82-2.43) and 1.73 (95%

CI= 1.51-1.95) respectively (Figure 3-11). Thus Prediction 3 was false. We also found that the

average nest productivity during the post-invasion era did not differ significantly between Toho

(see above) and all other wetlands combined (x= 1.50, 95% CI= 1.38-1.61) (Figure 3-12). Thus

Prediction 4 was fal se.

Survival

Apparent annual survival

The most parsimonious model of the apparent annual survival for kites hatched on Toho

between 1992 and 2006 was one that allowed detection probability to vary by year but that held

survival probability constant for all years. This top model did not include an age effect or a pre-

and post- invasion group effect on survival (Table 3-2); therefore, this model did not make

biological sense based on all previous snail kite literature that shows juvenile survival does, in

fact, vary over time and differ from that of adults (Bennetts et al. 1999; Dreitz et al. 2004; Martin

et al. 2007b). The second most parsimonious model also allowed detection probability to vary

over time, but it included an age effect on survival and allowed juvenile survival probability to









vary among the three pre-defined eras: pre-invasion (1992-1998), decline (1999-2002), and post-

invasion (2003-2006). Since this second model differed by less than two QAIC units (Table 3-

2), it was not significantly different from the top model (Burnham & Anderson 2004), and since

this second model made more biological sense (i.e., it did not set adult and juvenile survival

equal and hold them constant), we used it to generate estimates of apparent annual survival.

We found that the annual survival probability for juveniles hatched on Toho during the

pre-invasion era was 0.45 (95% CI= 0.27-0.66) and that during the post-invasion era it fell to

0.34 (95% CI= 0. 11-0.58) (Figure 3-13). Thus, Prediction 5 was not supported by our estimates

of annual juvenile survival. We found that annual adult survival was 0.81 (95% CI= 0.72-0.88),

and our data did not support allowing adult survival to vary among years or eras, which

supported Prediction 6. Although our data did not support the significance of the fourth most

parsimonious model, which allowed survival and detection to vary by year (Table 3-2), we

decided to generate estimates of apparent annual survival for juveniles hatched on Toho using

this model simply to demonstrate the inherent variation within the study system (Figure 3-14).

Apparent monthly survival

The most parsimonious model of apparent monthly survival for kites hatched on Toho was

one that included a pre- and post-invasion group effect on detection probability and survival, as

well as an age effect on survival (Table 3-3). We found that the monthly survival probability of

juveniles hatched on Toho was 0.93 (95% CI = 0.82-0.97) during the pre-invasion era

(represented by 1992 and 1994) and that it fell to 0.81 (95% CI = 0.65-0.90) during the post-

invasion era (represented by 2005) (Figure 3-15). We found that the monthly survival

probability of adults that were hatched on Toho was 0.92 (95% CI = 0.85-0.96) during the pre-

invasion era (represented by 1992-1996) and that it was 0.90 (95% CI = 0.75-0.97) during the

post-invasion era (represented by 2005-2006). Thus, estimates of monthly adult survival for









kites hatched on Toho were nearly identical between the pre- and post-invasion eras, supporting

Prediction 6. While there was considerably more variation between the pre- and post-invasion

era estimates of monthly juvenile survival from, we found no significant difference, as the

confidence intervals from these estimates overlapped. Although estimates for both monthly and

annual survival for juveniles hatched on Toho during the post-invasion era were lower than those

for the pre-invasion era, our data did not fully substantiate Prediction 5.

Discussion

The effects that exotic apple snails have on snail kite foraging behavior and energetic

may, in turn, result in some negative demographic repercussions. Although we found no

evidence of decreased adult survival, nesting effort, nest success, or nest productivity on Toho

during the post-invasion era, we did Eind some evidence suggesting that juvenile survival has

declined; however, our survival estimates for juveniles were based on severely limited datasets

and thus had overlapping confidence intervals.

Contrary to our predictions, snail kite nesting effort on Toho has not decreased since the

invasion of the exotic snail; in fact, our data suggest an increasing trend in kite nesting effort.

The average annual nest abundance on Toho is significantly higher in the post-invasion era than

it was in the pre-invasion era (42 versus 17 nests per year). The relative contribution of Toho to

the range-wide nesting effort of the kite population has also increased significantly (from 0.06 to

0.33). Nest abundance and distribution are affected by a number of environmental variables. It

is possible that the declining habitat quality in WCA3A has influenced snail kite nesting trends

on Toho. Droughts also influence snail kite behavior, and in 2007 south Florida experienced

moderate to severe drought conditions throughout much of the peak kite breeding season.

Events such as these may be underlying the observed nesting trends on Toho, during the post-

invasion era. However, one cannot discount the possibility that the exotic apple snail attracts









kites to Toho. We do not know the behavioral mechanisms by which kites distinguish habitat

quality, but it is likely that apple snail density (or some cue of snail abundance/availability) plays

a role. For example, the exotic snail lays highly conspicuous pink egg masses on emergent

structures, and these bright pink clusters are prevalent throughout Toho. Kites may be attuning

to these egg masses, or kites may make more direct assessments of foraging conditions. Future

research efforts should focus on teasing out the cues used by kites to assess habitat suitability,

and Toho may provide the conditions necessary to test certain hypotheses related to kite

attraction and site selection.

Nest success and nest productivity remain virtually unchanged on Toho during the post-

invasion era relative to the pre-invasion era. Although statistically insignificant, it is interesting

to note that the average number of young produced per successful nest on Toho during the post

invasion-era is above the range-wide average for this time period. We also found that the

survival probabilities of adult kites that were hatched on Toho did not differ between pre- and

post-invasion eras. These findings, strongly suggest that the exotic snail does not constitute an

environmental stressor for adult kites, as their abilities to survive and to successfully provision

and fledge young have not been negatively affected. On the other hand, the ability of juveniles

to sustain themselves after fledging remains tenuous.

Our estimates for apparent annual survival of juveniles hatched on Toho are rather

inconclusive. The drastically fluctuating annual estimates and wide confidence intervals in

Figure 3-14 are likely the result of insufficient data coupled with environmental stochasticity.

The estimates of apparent annual survival from our best, biologically relevant model provide a

slightly clearer picture; this model shows that juveniles hatched on Toho during the post-

invasion era experienced lower annual survival probabilities than did juveniles hatched on Toho









during the pre-invasion era (0.34 and 0.45, respectively), but these estimates are still relatively

uninformative, as their confidence intervals overlap widely (Figure 3-13). After the completion

of the 2008 population survey, we will have the statistical power to generate more precise

estimates of annual survival for juveniles hatched on Toho during the post-invasion era, and

additional years of band-resight data will continue to increase our precision.

Our estimates of apparent monthly survival are somewhat more revealing. Our Eindings

indicate that juveniles hatched on Toho during the post-invasion era experience lower monthly

survival probabilities than did those hatched during the pre-invasion era (0.81 and 0.93,

respectively), but the confidence intervals of these estimates overlap as well. However, unlike

the confidence intervals around our estimates of annual juvenile survival, the confidence

intervals around our estimates of monthly juvenile survival each exclude the mean estimate of

the comparative group (Figure 3-15).

On average, juvenile survival reaches a level comparable to adult survival around 4 months

post-fledging (Bennetts & Kitchens 1999). If the monthly survival estimates for the pre-invasion

(0.93) and post invasion (0.81) eras are raised to the fourth power (simulating survival to Hyve

months of age), they result in drastically different outcomes, 0.75 and 0.43 respectively. If these

same estimates are raised to the eleventh power (simulating survival to one year of age), they

result in survival probabilities of 0.45 and 0. 10 for the pre- and post-invasion eras, respectively.

The simulated annual survival of juveniles on Toho during the pre-invasion era (0.45), which

was calculated with monthly survival estimates from telemetry data, is equal to the estimate of

apparent annual survival (0.45) that was generated directly from band-resight data; therefore, we

are quite confident in these pre-invasion estimates. However, for juveniles hatched on Toho

during the post-invasion era, we do not find the same agreement between simulated annual









survival (0.10) and the direct estimate of apparent annual survival (0.34), but more data will

likely improve these estimates. While our estimates of survival for juvenile kites hatched on

Toho may not be statistically significant, this does not negate the possibility that there may be a

significant biological difference, as lower juvenile survival during the post-invasion era will

likely result in a significant reduction in recruitment from Toho. In spite of this, juvenile

survival varies widely over time, and these suppositions should be viewed with caution, as our

inferences are limited to the years for which we had sufficient data on juvenile survival (1992,

1994, and 2005).

The kite population is critically endangered, and its sustainability is extremely sensitive to

recruitment. However, at this time, we cannot definitively say whether the exotic apple snail

negatively affects juvenile survival. This is regrettable considering that the implementation of

management actions that could potentially combat the threat of the exotic snail are likely

contingent on such definitive answers. Therefore, we recommend expediting the quest for

suitable answers in relation to juvenile survival on Toho during the post-invasion era. Radio

transmitters should be deployed on the next cohort of juveniles, and a fastidious radio-telemetry

protocol (Bennetts & Kitchens 1997) should be enacted. In addition, conducting two robust

population surveys per year (instead of just one) over the course of the next two years would also

expedite our ability to generate more reliable post-invasion survival estimates for juveniles

hatched on Toho. With suitable assistance, these recommendations will generate the data

necessary to answer these pressing questions, which will lead to a better conservation strategy

for the snail kite.

While management actions focused on the exotic snail may be pending further study, there

is no reason for management actions focused on the native apple snail to be delayed or ignored.









CERP mandates that management actions should "restore and maintain a network of snail kite

foraging habitats and promote habitat that supports primary prey (apple snails) recruitment

throughout the South Florida ecosystem" (USFWS 1999; RECOVER 2005). Exotic snail

populations are already established in close proximity to several of the other critical wetlands

utilized by the snail kite in Florida, but the presence of healthy native apple snail populations

may help buffer any negative effects imposed on the snail kite by the exotic snail. More

resources should be invested in monitoring native apple snail populations within the kite's range

in Florida, as the current spatial scope of such data is limited. Estimating snail densities before

and after management actions should also become standard protocol for all proj ects directly

influencing the wetland habitats on which the kites depend. Such data will be critically

important to adaptive management strategies involving the snail kite and other species,

especially with the onset of DECOMP (Decompartmentalization and Sheet Flow Enhancement

Proj ect) in WCA3A.










Table 3-1. The number of nests initiated range-wide and the contribution from Toho, 1995-2007
1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007
BICY 24 8 1 6 5 0 0 0 0 0 0 0 0
ENP 2 5 0 0 0 12 0 0 0 0 0 23 5
ETOHO 1 0 0 0 0 0 4 0 0 0 0 1 1
GW 1 0 2 3 0 3 0 0 2 0 15 1 0
KISS 0 0 1 2 0 0 0 4 10 7 9 1 7
VVCA1 1 0 1 14 0 0 0 0 0 0 0 1 0
OKEE 23 34 3 8 0 0 0 0 5 10 23 18 0
SJM 19 16 25 9 12 6 1 1 8 21 8 19 0
TOHO 14 18 37 2 3 8 14 25 15 0 47 30 79
WCA2A 8 0 0 0 0 0 0 0 0 0 0 0 0
WCA2B 126 3 22 124 0 0 0 11 11 16 0 2 0
WCA3A 50 79 246 221 70 112 0 60 78 40 12 61 2
WCA3B 2 0 0 3 5 26 0 3 2 5 0 17 1
Total 271 163 338 392 95 167 19 104 131 99 114 174 95
Contribution
from Toho 5% 11% 11% 1% 3% 5% 74% 24% 11% 0% 41% 17% 81%





Table 3-2. Model selection table for apparent annual survival of kites hatched on Toho.
Model QAIC delta(QAIC) Parameters Deviance
1 {Phi(Juv(.) Ad(.) P(t) SIN} 444.09 0.00 18 217.12
2 {Phi(Juv( inv grps) Ad(.) P(t) SIN} 446.03 1.94 20 214.17
3 {Phi(Juv(.) Ad(.) P(~ iny Gps) SIN} 454.53 10.44 4 259.04
4 {Phi(Juv(t) Ad(.)) P(.) use SIN} 455.41 11.32 30 197.43
5 {Phi(.) P(t) SIN} 470.13 26.04 17 245.57




Table 3-3. Model selection table for apparent monthly survival of kites hatched on Toho.
Model AICc delta (AICc) Parameters Deviance
{Phi(Pre(12 mon)not=Post(juv(1 2mon))
1 P(Pre(.)not=Post(.)) SIN} 338.77 0.00 6 268.28
{Phi(Pre(4mon)not=Post(juv(4mon))
2 P(Pre(.)not=Post(.)) SIN} 340.11 1.34 6 269.62
{ Phi (Pre (.) no t=Post (.)) P (Pre (.) no t=Post (t))
3 SIN} 372.80 34.02 39 218.57
{Ph i(P re(.)n ot= Post(.)) P(P re(.)= Post(.))
4 SIN} 409.37 70.60 3 345.17
5 {Phi(Pre(.)=Post(.)) P(Pre(.)=Post(.)) SIN} 413.97 75.19 2 351.82
{ Phi (Pre (.) no t=Post (.)) P (Pre (t) no t=Post (.))
6 SIN} 422.60 83.83 45 249.58
{ Phi (Pre (.) no t=Post (.)) P (Pre (t) no t=Post (t))
7 SIN} 483.53 144.76 75 191.62









MM Toho
O All Other Wetlands


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1995 1996 1997 1998 1999 2000 2001 2002 2003


2004 2005 2006 2007


year

Figure 3-1. Annual number of snail kite nests initiated range-wide, 1995-2007.


r


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Figure 3-2. Average number of nests initiated annually on Toho during the pre- and post-
invasion eras. Post-invasion estimate excludes 2004. Error bars represent 95% CIs.































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Figure 3-3. Number of nests initiated annually on Toho, 1995-2007.






































































124


1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007







































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Figure 3-5. Trend model expressing the increasing contribution of Toho to the total population

nesting effort over time (includes all years 1995-2007). Dashed lines represent 95%

CIs.





















































126





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pre-i invasion post-i nvas ion





Figure 3-7. Relative contribution of Toho to the total population nesting effort during the pre-
and post-invasion eras. Error bars represent 95% CIs.





















~LIll
1995 1996 1997 1999


2002 2003 2005 2006 2007


Figure 3-8. Annual nest success on Toho, 1995-2007. Error bars represent 95% CIs.




















129














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pre-i invasion post-i nvas ion


Figure 3-9. Average nest success on Toho during the pre- and post-invasion eras. Error bars
represent 95% CIs.



























LO







1995 1996 1997 1999 2000 2001 2002 2003 2005 2006 2007

year


Figure 3-10. Annual number of young fledged per successful nest on Toho, 1995-2007. Error
bars represent 95% CIs.


































131










































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pre-i nvas ion


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Figure 3-11. Average nest productivity on Toho during the pre- and post-invasion eras. Error
bars represent 95% CIs.

















132















































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Toho


Figure 3-12. Average number of young fledged per successful nest on Toho vs. all other
wetlands during the post-invasion era. Error bars represent 95% CIs.




















133


All Other Wetlands


































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o





a









O


o


pre-i nvas ion


post-i nvas ion


Figure 3-13. Apparent annual survival of juveniles hatched on Toho during the pre- and post-
invasion eras. Error bars represent 95% CIs.





















134




















O





a)
O


m


7 'D
o;-
m


m
E d
g! d
m
n
n
m

hi
d




o
d-


1992 1994 1995


Figure 3-14. Apparent annual survival of juveniles hatched on Toho, 1992-2006. Error bars

represent 95% CIs.



























































135


1996 1997 1999 2000 2001 2002 2003 2005 2006














O





CO
O




CO














O


o


pre-i nvas ion


post-i nvas ion


Figure 3-15. Apparent monthly survival of juveniles hatched on Toho during the pre- and post-
invasion eras. Error bars represent 95% CIs.


















136









CHAPTER 4
IS LAKE TOHOPEKALIGA FUNCTIONING AS AN ECOLOGICAL TRAP FOR THE
SNAIL KITE IN FLORIDA?

Introduction

Ecological Trap Theory

Through geological time, animal populations adapt to environmental conditions via a

number of mechanisms. One form of adaptation manifest in animal behavior is the cue-response

system. Environmental cues eliciting behavioral responses that, on average, result in a positive

net fitness outcome can be selected for and incorporated into decision making processes through

natural selection (Stevens & Krebs 1986, Sih 1987). Ecological traps can be created when 1)

rapid changes to an ecosystem result in the decoupling of habitat selection cues from the positive

behavioral responses they once elicited and 2) an animal continues to make settlement decisions

based on ingrained environmental cues, but due to the abrupt change in habitat, these decisions

now result in negative fitness consequences unbeknownst to the individual (Tinbergen 1963,

Sherman 1988, Robertson & Hutto 2006).

The criteria necessary to demonstrate an ecological trap are as follows: 1) individuals of

the study population must exhibit preference for one habitat (i.e., the potential trap) that is

greater than or equal to preferences for other available habitats, 2) individual fitness (or an

appropriate surrogate measure) must differ among available habitats, and 3) negative fitness

consequences must result from settling in the preferred (or equally preferred) habitat trap

(Schlaepfer et al. 2002; Robertson & Hutto 2006).

A Hypothetical Example

For the snail kite population in Florida, an ecological trap would be constituted by the

following scenario: Adult kites are attracted to Toho due to the abundance and availability of

exotic apple snails or by the plethora of exotic snail egg masses. Adults, which are larger and









have more foraging experience than juvenile kites, are able to attain ample energy for survival

and reproduction in this altered habitat; therefore, they preferentially nest on Toho even though

other habitats, devoid of the exotic snail, are available. However, juvenile kites, once left to

forage on their own, are unable to sufficiently handle and extract the larger exotic snails, and

subsequently, they starve. Nevertheless, adult kites continue to preferentially select Toho for

nesting because of the lake's plethora of seemingly abundant and available exotic snails. In

demographic terms, adult survival is unaffected, nest success is high, but recruitment is

significantly suppressed due to high juvenile mortality. This situation would be catastrophic

because, as stated earlier, recruitment is one of the predominant factors limiting the growth and

viability of the snail kite population in Florida (Martin 2007).

Small populations, especially those inhabiting landscapes comprised of fragments

displaying source-sink dynamics, are particularly vulnerable to extinction (Pulliam 1988, Pulliam

& Danielson 1991). Therefore, from a management perspective, understanding the effects that

the exotic apple snail has on snail kite foraging success and consequent demographic parameters

has precluded our ability to adaptively manage the system in the kite's favor.

Recent Nesting History

From 1996 to 2003, snail kite reproduction occurred predominantly in WCA3A. Smaller

contributing fragments included ENP, Toho, and Kiss (Bennetts & Kitchens 1997; Dreitz et al.

2001; Martin et al. 2006b, 2007c). Historically, Toho served as an important refugium for snail

kites, particularly when regional droughts affected the wetlands throughout the southern portion

of their range (Bessinger & Takekawa 1983; Takekawa & Beissinger 1989; Bennetts & Kitchens

1997, Mooij et al. 2002; Martin et al. 2006a), but Toho did not harbor significant numbers of

nesting kites (Bennetts & Kitchens 1997; Martin et al. 2006b). However, the proportion of









nesting activity on Toho during the post-invasion era has been disproportionately high when

compared to traditional nesting areas (Martin et al. 2007c) (Chapter 3).

Lake Tohopekaliga

We have shown that the exotic apple snail affects the foraging behaviors of snail kites

(Chapter 2), with some demographic consequences (Chapter 3), but to provide evidence that

Toho is functioning as an ecological trap for the snail kite in Florida, the following three

hypotheses must be supported:

* Hypothesis 1. Snail kites exhibit a habitat preference for Toho that is greater than or equal to
preferences for other available wetland units.

* Hypothesis 2. Snail kite fitness differs among these wetland units.

* Hypothesis 3. Negative fitness consequences to the snail kite result from selecting Toho.

Methods

Hypothesis 1

Since we do not have a detailed understanding of snail kite cognition, it is hard measure

habitat preference directly; therefore, we used disproportionate habitat use as a proxy for habitat

preference. We compared the average number of nests initiated annually on Toho during the

pre- and post-invasion eras. Then we assessed the trend in Toho's relative contribution to the

range-wide snail kite nesting effort over time (For methods of data collection see Bennetts &

Kitchens 1997; Dreitz et al. 2004; Martin et al. 2007, 2007c) (For methods of analysis see

Chapter 3). To establish whether snail kites show an equal or greater habitat preference for Toho

relative to other wetlands during the post-invasion era, we made among area comparisons of the

relative abundance of nests (and in parallel, of breeding individuals) and the relative length of the

breeding season. When calculating the average number of nests initiated annually on Toho

during the post-invasion era we excluded 2004 because the managed drawdown overlapped with










the kite breeding season (Chapter 1). We tested the following predictions that would provide

supporting evidence for Hypothesis 1:

* Prediction la. Kite nesting effort (as measured by annual nest abundance) on Toho has
increased significantly since the introduction of the exotic apple snail.

Prediction lb. The relative annual contribution of Toho to the total number of nests found
range-wide has increased significantly since the introduction of the exotic apple snail.

Prediction 10. During the post-invasion era, the average number of nests initiated annually
is significantly higher on Toho relative to other wetlands.

Prediction Id. During the post-invasion era, kite nesting effort (as measured by the duration
of the breeding season) on Toho is longer relative to breeding season patterns of other
wetlands within the kite's range.

Hypothesis 2

Using survival, nest success and nest productivity (i.e., the number of young fledged per

successful nest) as surrogate measures of fitness, we used evidence from the literature to

established whether kites on Toho experience different fitness consequences than kites utilizing

other wetlands within their range. The following predictions would provide supporting evidence


foPr H) pthesi 2:.Autsalkt uvvldfessgiiatyaogwtaduis

* Prediction 2a. Aduvnlt snail kite survival differs significantly among wetland units.

* Prediction 2b. Jueniesni kt su rvival differs significantly among wetland units.


* Prediction 2d. The average number of young fledged per successful nest (i.e., nest
productivity) differs significantly among wetland units.

Hypothesis 3

Using the demographic parameters from Hypothesis 2 as surrogate measures of fitness, we

tested the following predictions that would provide evidence for Hypothesis 3:

* Prediction 3a. Adult survival has decreased on Toho since the invasion of the exotic apple
snail, and during the post-invasion era, adult survival is lower on Toho relative to other
wetlands.










* Prediction 3b. Juvenile survival has decreased on Toho since the invasion of the exotic
apple snail, and during the post-invasion era, juvenile survival is lower on Toho relative to
other wetlands.

Prediction 3c. Nest success has decreased on Toho since the invasion of the exotic snail,
and during the post-invasion era, nest success is lower on Toho relative to other wetlands.

Prediction 3d. The average number of young fledged per successful (i.e., nest productivity)
nest has decreased on Toho since the invasion of the exotic apple snail, and during the post-
invasion era, nest productivity is lower on Toho relative to other wetlands.

Results

Hypothesis 1

We found that snail kites do in fact preferentially select Toho (as measured by

disproportionate use) over other available habitats (Chapter 3). The average annual nesting

effort on Toho has increased since the invasion of the exotic apple snail (Figure 3-2, 3-3), which

validates Prediction la. There is also an increasing trend in Toho's relative contribution to the

overall nesting effort of the entire kite population (Figure 3-5, 3-6), and Toho' s average

contribution to the overall nesting effort is significantly greater during the post-invasion era

when compared to the pre-invasion era (Figure 3-4, 3-7), validating Prediction lb. We found

that during the post-invasion era, a significantly greater number of nests were initiated annually

on Toho relative to all other individual wetlands except for WCA3A; however, the estimate for

Toho (x= 42, 95% CI= 29-55) was higher than that for WCA3A (x= 31, 95% CI= 20-42),

lending support to Prediction lc. Additionally, from 2005 to 2007, kites on Toho expanded their

nesting efforts beyond the typical breeding season (i.e., March to June) (Bennetts & Kitchens

1997; Martin et al. 2006b, 2007c). Kites were actively nesting on Toho from early-March

through early-November in 2005, from early-March through late-August in 2006, and from

early-March through mid-October in 2007. During these years, the vast maj ority of the nesting

activity in other wetlands ceased in May or June, and the last documented nesting activity in any









of these other wetlands was in early-July, late-July, and early-June respectively, thus lending

support to Prediction Id. Due to the effects of the managed drawdown on suitable snail kite

nesting and foraging habitat, 2003 and 2004 were not considered, as the drawdown likely cut the

breeding season short in 2003 and likely deterred nesting in 2004 (Chapter 1).

Hypothesis 2

Barring the occurrence of severe region wide-droughts, we found no supporting evidence

in the literature that adult snail kite survival varies significantly over time or that it differs

significantly among wetland units (Bennetts & Kitchens 1997; Bennetts et al. 1999; Martin et

al.2007b); therefore, Prediction 2a was invalid. On the other hand, the literature did support the

supposition that juvenile survival varies among wetlands (Bennetts et al. 1999; Dreitz et al.

2004), thus validating Prediction 2b. We also found evidence in the literature that average nest

success and average nest productivity vary among wetlands (Snyder et al. 1989; Bennetts &

Kitchens 1997; Dreitz et al. 2001), which supports Predictions 2c and 2d.

Hypothesis 3

There is no evidence for decreased nesting success (Figure 3-8, 3-9) or nest productivity

(Figure 3-10, 3-1 1) as a result of the exotic apple snail on Toho, and during the post-invasion era,

nest success and nest productivity are not significantly lower on Toho relative to other wetlands

(Figure 4-2, 4-3), thus discrediting Predictions 3c and 3d respectively. There is also no evidence

suggesting that the exotic apple snail negatively affects adult survival (Chapter 3), and adult kites

foraging on native versus exotic snails maintain comparable energy balances (Figure 2-32).

Additionally, adults on Toho during the post-invasion era acquire sufficient surplus energy to

successfully provision and fledge young (Chapter 3). Therefore, Prediction 3a is null.

While conclusive evidence is lacking for the supposition that apparent survival of juveniles

from Toho is lower in the post-invasion era relative to the pre-invasion era (Figure 3-13, 3-14, 3-










15), there is indirect evidence based on foraging and energetic that suggests juvenile mortality

on Toho during the post-invasion era should be high. Daily gross energy gains are significantly

lower for juveniles on Toho relative to juveniles on other wetlands that are foraging on native

snails (Figure 2-31), and on average, juveniles on Toho do not maintain positive daily energy

balances (Figure 2-32). While our energetic analyses do not provide conclusive evidence for

Prediction 3b, they do give rise to concern.

Discussion

There is sufficient support for H~gpothes~i\ 1 showing that during the post-invasion era,

kites preferentially select (i.e., disproportionately use) Toho relative to the other available

wetlands within their range (Figure 4-1). However, we cannot say that this disproportionate use

is directly attributable to the presence of the exotic snail on Toho because we did not test

alternative hypotheses that may have led to the observed patterns of habitat use (e.g., drought or

habit decline in other wetlands may have influenced kites to move to Toho) (Chapter 3).

Regardless of the underlying factors) leading to the shift in habitat use, the annual contribution

of Toho to the range-wide kite nesting effort is over 500% greater in the post-invasion era

relative to the pre-invasion era, and nests on Toho during the post-invasion era account for over

30% of the range-wide nesting effort (Chapter 3). Given the number of nesting attempts and the

number of young fledged on Toho relative to other wetlands (Figure 4-4, 4-5), variations in

habitat quality on Toho could have tremendous influence the snail kite population.

The extended breeding seasons observed on Toho from 2005-2007 may be more directly

attributable to the exotic snail. Although little is known about the life history strategy of the

exotic apple snail (Ramakrishnan 2007, Youens & Burks 2007), the absence of an annual

population-wide post-reproductive die-off in the exotic snail (Ramakrishnan 20007), such as that









observed in the native snail (Darby et al. 2008), likely allows kites on Toho to continue their

nesting efforts through the late-summer and into the fall.

Since kites disproportionately use Toho during the post-invasion era, we continued our

tests for an ecological trap. H@psthes~i\ 2 is supported by numerous studies showing that nest

success, nest productivity, and, most applicably, juvenile survival vary among wetlands. And,

although tentative, there is some support for Hypothesis 3. If the effects of the exotic apple snail

had significant negative impacts on adult and juvenile foraging success, then Predictions 3a

through 3d may have been true. However, the foraging success of adults and juveniles is

differentially affected, and only juveniles suffer severe energetic repercussions (Chapter 2).

Therefore, the only support for H@psthes~i\ 3 comes from Prediction 3d, but direct evidence of

suppressed juvenile survival is lacking (Chapter 3). While estimates of juvenile survival are

lower for the post-invasion era than the pre-invasion era, they are not significantly different

(Figure 3-13, 3-15). Nonetheless, the discrepancies in DEE (Figure 2-32) suggest that Prediction

3d may be true, especially given that our estimates of juvenile survival were derived from

relatively small sample sizes.

Adult fertility (and implicitly, recruitment via juvenile survival) is the most influential

factor limiting growth in the snail kite population. Given that the appropriate criteria for

Hypothesis 1 and Hypothesis 2 were met, the validation of Prediction 3d would corroborate the

hypothesis that Toho is functioning as an ecological trap for snail kites. But our findings do not

provide unequivocal evidence, and with the data presently available, we cannot say whether or

not Toho is functioning as an ecologic trap. Continuance of the snail kite population monitoring

protocols coupled with specific radio-telemetry studies of juveniles on Toho should soon provide









us with the data necessary to reassess these critical questions and to arrive at more definitive

answers (see "Discussion" in Chapter 2 and Chapter 3).

While our current Eindings should give rise to serious concerns, we believe the drastic

management actions that would be necessary to resolve the problems on Toho (if it is an

ecological trap) warrant more conclusive evidence. Even if future studies suggest that the exotic

apple snail does not negatively affect juvenile survival, we are in no way advocating that the

exotic snail should be allowed to persist in or spread throughout the wetlands of central and

southern Florida for the sole benefit of the snail kite. In fact, quite the opposite is true.

Notwithstanding the fact that the exotic snail may have unknown deleterious affects on

innumerable species and ecosystem functions, allowing the exotic snail to persist/spread may

jeopardize snail kites in ways not dealt with in this study. Therefore, we support the eradication

of the exotic snail but underscore the importance of carefully assessing all eradication methods,

as negative effects on wetland habitats and native snails must be minimized. Anecdotal evidence

suggests that kites are foraging on exotic apple snails (and possibly even attempting to breed)

along canals and irrigation ditches that are disconnected from the maj or wetlands comprising

their historical range. Are kites attracted to these unconventional areas by the exotic snail? We

do not know, but evidence suggests that traveling through the matrix of unsuitable foraging

habitat, which intersperses wetland fragments, may lead to increased mortality in snail kites

(Bennetts & Kitchens 1997, 2000; Martin 2007). Therefore, the exotic snail may have indirect

negative affects on the kite population even if kites can sustain themselves by feeding on the

exotic snail. It is also possible that kites are being driven into peripheral habitats out of

necessity. Furthermore, we strongly advocate ecosystem and water management strategies that

take into account the life history strategy of the native apple snail, as such strategies will likely









provide the greatest benefits for the snail kite population. Maintenance and monitoring of

suitable native snail populations within wetlands utilized by the kite are necessary to sustain the

snail kite population in Florida (USFWS 1999; RECOVER 2005) and will likely help buffer

against any negative influences imposed by the exotic snail.























O
















EN 3A 2 B OE W J IS TH



Figure -.Aeaenme fnssiiitdanal nnn elnsdrn h otivso


er.Errbr epeet9%Cs










































2005

10-


08-


2 06- -


2 04-


02-


00-




Fiur -2.Anua nstsuces



Figur4represealnt 95 sCces.


(DLDOONI-O
m~mO
Z~mnw Bml
NWCr)
W y O
Y y)
O


2006 2007


06-lmI 06-


04-1' 04-


02-11 I 02-


00 1 I I I I I I I 1 0
00-"- 00- -" ""






on nine wetlands during the post-invasion era. Error bars
































































mmw~lroo
r" Wo "I
W r o
V






2005


mw~l"o
,4m Wo "I
W "ro
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t


"mmw~l"
r w to o
o
w 1 to
o o
t


"mmw~l"
r w to o
o
w 1 to
o o
t


Figure 4-3. Annual nest productivity on nine wetlands during the post-invasion era. Error bars


represent 95% CIs.








































































149



















80




60



40




20


00(9NO-(9m~
a5mm; gsmo
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w ~L C1 I
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40




20


nl
"""-"""""
a4mmwgsm
znrrnw mo
w ~L C1(O~LI
O O


0-00FILnm~C
N-----~
a--mm gm
z~rrnwg mo
w I
w CI(O~L
~L O
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Figure 4-4. Total number of nests initiated annually on nine wetlands during the post-invasion

era.










































150


a~alJ
"""""-"-"
""~--~-~"
m g m
a~Nmw gmO
Z FIWCI
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O

















































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2005


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o
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o


~mmwgi"o
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C) I
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Figure 4-5. Total number of young fledged per year (uncorrected counts) on nine wetlands

during the post-invasion era.



















































151


~mmwgi"o
orow
C) I
W
O
O
t


O~a
'"""
""""









APPENDIX
DERIVATION OF DAILY ENERGY EXPENDITURE

The following series of equations comes from Koplin et al. (1980):

DEB = NFA[(EIM,) (1 P)
where. NFA = duration of diurnal nonf~ight activities as a proportion of the
photoperiod (daylength) (NYFA 1 FA),
FA = duration of flight activities as a proportion of the photoperiod
(PA = 1 -- NFA),
EM,. = existence mretabolismn of nonpasserine bir-ds during winter as a
function of average daily air temzperature (T,):
E~W-r = EMo.~c + (TXA) (2)
EM,,oBc EML,
wthere: & =
30
E Afoo c = .455 Weam (3B )
EIMoc 4.235 We **91 (4)
WY = body mass in g
SMa,h = standard metabo~lism of nonpasserine birds during wvinter at
night as a furnction of average night timne air temperature (Tu):
SMif, = S"IO'C (bXT,) (5)
where: SB;o~c: = 1.YI10 we SLw 16
bi = 0.047W oman~B I)
at the lower critica.1 Itmperature (Tech) SLW M
T, = 47.17 W-eIson gs)
BM = basal metabism
BMI = 0.4fi616 Wess 9)
P = photoperiod as a proportion of the 24,h day.
FC = flight coeffcient; the multiple of BMW expressing; energy expendi-
ture of flight.









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BIOGRAPHICAL SKETCH

Christopher Cattau earned a B.S. in ecology and evolutionary biology from the University

of Tennessee, Knoxville. Then he moved to Florida.





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1 EFFECTS OF THE INVASIVE EXOTIC APPLE SNAIL ( Pomacea insularum ) ON THE SNAIL KITE ( Rostrhamus sociabilis plumbeus ) IN FLORIDA, USA By CHRISTOPHER CATTAU A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2008

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2 2008 Christopher Cattau

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3 To Florida

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4 ACKNOWLEDGMENTS I would like to thank m y committee memb ers (Wiley Kitchens, Phil Darby, and Ken Meyer) for their mentorship, advice, and enc ouragement. Although not officially on my committee, Julien Martin deserves the same gratit ude, as his guidance and instruction, like that of my committee, was indispensable to the deve lopment and completion of this work. I would also like to thank all of the graduate students and field tech nicians who assisted with the collection and processing of data, including An drea Bowling, Brian Reichert, Sara Stocco, Christina Rich, Danny Huser, Will deGravelles, J ean Olbert, Derek Piotro wicz, Nate Richardson, Wesley Craine, Michaela Speirs, Melinda Conne rs, Bridget Deemer, Cour tney Hooker, Andrea Ayala, Melissa Desa, Carolyn Enloe, and all of my fellow coworkers at the Florida Fish and Wildlife Cooperative Research Unit. I would lik e to express particular gratitude to Brian Reichert for helping me with survival analyses This project would not have been possible without the unwavering dedicati on that each of the aforementi oned individuals expressed for conservation of the snail kite and the challe nging (and often arduous) work each performed to advance our knowledge of this endangered species. This cooperative effort has allowed us to elucidate threats to the snail k ite population and to increase our understanding of their ecology, which will lead to more reliable conservation stra tegies. I am enormously grateful to everyone involved with this work.

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........8 LIST OF FIGURES.........................................................................................................................9 LIST OF ABBREVIATIONS........................................................................................................ 13 ABSTRACT...................................................................................................................................14 CHAP TER 1 INTRODUCTION................................................................................................................. .16 Study Population............................................................................................................... ......17 Invasion of the Exotic Snail....................................................................................................19 Population Demography......................................................................................................... 21 Study Objectives and Outline................................................................................................. 23 2 EFFECTS OF THE EXOTIC APPL E SNAIL ON SNAIL KITE FORAGING BEHAVIOR ............................................................................................................................27 Background.............................................................................................................................28 Foraging Behavior...........................................................................................................28 A Note on Pomacea Measurem ents................................................................................ 30 The Native Apple Snail................................................................................................... 30 The Exotic Apple Snail................................................................................................... 31 Previous Foraging Studies...............................................................................................32 Prey availability........................................................................................................32 Prey negotiability and drop rate............................................................................... 34 Searching and handling time.................................................................................... 35 Age and sex effects on foraging behavior................................................................ 36 Energy budgets and profitability.............................................................................. 37 Economic models of foraging behavior.......................................................................... 38 Predictions.......................................................................................................................39 Methods..................................................................................................................................40 Time Activity Budgets.................................................................................................... 40 Sampling locations...................................................................................................41 Selection of focal individuals...................................................................................41 Timing and duration of observations....................................................................... 42 Classification of individual kites..............................................................................43 Collection of behavioral data................................................................................... 45 Assumptions of behavioral observations.................................................................. 46

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6 Apple Snails.....................................................................................................................47 Collection and measurement of empty snail shells.................................................. 47 Collection and processing of live snails................................................................... 48 Statistical Analysis a nd Confidence Intervals ................................................................. 49 Analysis of Foraging Behavior........................................................................................49 Searching and handling time.................................................................................... 51 Drop rate................................................................................................................... 52 Capture rate (i.e., consumption rate)........................................................................ 52 Analysis of Energetics..................................................................................................... 53 Regression models and estimation of aver age snail weight and caloric content...... 53 Profitability............................................................................................................... 54 Daily activity times and energy budgets.................................................................. 54 Results.....................................................................................................................................56 Diet..................................................................................................................................56 Nutritional content of apple snails ...........................................................................57 Live exotic snail measurements............................................................................... 57 Empty snail she ll dim ensions................................................................................... 58 Estimates of total snail weight................................................................................. 58 Foraging Behavior...........................................................................................................59 Comparing adults, subadults, and juveniles............................................................. 59 Effects of the exotic snail on average searching times.............................................60 Effects of the exotic snail on average handling times.............................................. 60 Effects of the exotic snail on drop rates...................................................................61 Effects of the exotic snail on capture rates............................................................... 62 Effects of the Exotic Snail on Energetics........................................................................ 63 Estimates of caloric content..................................................................................... 63 Profitability............................................................................................................... 64 Validation of time activity budget extrapolations.................................................... 64 Daily activity patterns..............................................................................................65 Daily energy balances..............................................................................................65 Discussion...............................................................................................................................66 3 DEMOGRAPHIC EFFECTS OF THE E XOTIC APPLE SNAIL ON THE KITE .............104 Introduction................................................................................................................... ........104 Reproduction.................................................................................................................105 Survival..........................................................................................................................105 Predictions.....................................................................................................................106 Methods................................................................................................................................107 Nesting Effort and Reproductive Success..................................................................... 108 Survival..........................................................................................................................111 Results...................................................................................................................................113 Reproduction.................................................................................................................113 Nesting effort..........................................................................................................113 Nest success and productivity................................................................................114

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7 Survival..........................................................................................................................114 Apparent annual survival.......................................................................................114 Apparent monthly survival.....................................................................................115 Discussion.............................................................................................................................116 4 IS LAKE TOHOPEKALIGA FUNCTIONING AS AN ECOLOGICAL TRAP FOR THE SNAIL KITE IN FLORIDA? ...................................................................................... 137 Introduction................................................................................................................... ........137 Ecological Trap Theory................................................................................................. 137 A Hypothetical Example...............................................................................................137 Recent Nesting History.................................................................................................. 138 Lake Tohopekaliga........................................................................................................139 Methods................................................................................................................................139 Hypothesis 1.................................................................................................................. 139 Hypothesis 2.................................................................................................................. 140 Hypothesis 3.................................................................................................................. 140 Results...................................................................................................................................141 Hypothesis 1.................................................................................................................. 141 Hypothesis 2.................................................................................................................. 142 Hypothesis 3.................................................................................................................. 142 Discussion.............................................................................................................................143 APPENDIX DERIVATION OF DAILY ENERGY EXPENDITURE....................................... 152 LIST OF REFERENCES.............................................................................................................153 BIOGRAPHICAL SKETCH.......................................................................................................164

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8 LIST OF TABLES Table page 2-1 Number of TABs conducted by mo nth and location in 2003, 2004, 2005, 2006 (postinvasion) and in 1993, 1994, 1996 (pre-invasion). ............................................................ 71 2-2 Nutritional contents of P. insularum ..................................................................................71 2-3 Nutritional composition of P. insularum versus P. paludosa. ...........................................72 2-4 Average daily activity pattern of the snail kite in Florida. ................................................ 72 3-1 The number of nests initiated range-wid e and the contribution from Toho, 1995-2007. 121 3-2 Model selection table for apparent annual survival of kites hatched on T oho................ 121 3-3 Model selection table for apparent m onthly survival of kites hatched on Toho. ............. 121

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9 LIST OF FIGURES Figure page 1-1 Major habitat fragments within the range of the snail kite in Florida ............................... 24 1-2 Confirmed locations, as of 2007, of t he exotic snail ( P. insularum ) within the range of the snail kite in Florida. Larger ci rcles encompass multiple small populations in close proximity................................................................................................................ ...25 1-3 Movements of approximately 50 radi o-tagged adult sna il kites am ong wetland fragments over a one-year period. April 1992-April 1993................................................ 26 2-1 Relative sizes of e xotic apple snails................................................................................... 73 2-2 Measurement of snail shells............................................................................................... 74 2-3 Location of apple snail traps on Toho................................................................................ 74 2-4 Linear regression models of e xotic snail weight and m orphology.................................... 75 2-5 Number of native and exotic apple snails captured pe r year with funnel traps on Toho, 2005-2007. ...............................................................................................................76 2-6 Proportion of native to exotic apple sn ails captured in five locations on Toho, 20052007....................................................................................................................................77 2-7 Average length of apple snail shells colle cted from feeding perc hes and nests in nine wetlands throughout the range of the snail kite, 2004-2007. ............................................. 78 2-8 Average shell dimensions of native and e xotic apple snails co nsum ed by kites, 20042007....................................................................................................................................79 2-9 Estimated whole weight of native and exotic snails negotiated by foraging kites during the post-invasion era. ..............................................................................................80 2-10 Average handling times for adult, ju venile, and subadult kites on Toho and KissWCA3A during the post-invasion era................................................................................81 2-11 Average drop rates for adult, juveni le, and subadult kites on Toho and Kiss-W CA3A during the post-invasion era...............................................................................................82 2-12 Average capture rates for adult, juvenile, and subadult ki tes on Toho and KissWCA3A during the post-invasion era................................................................................83 2-13 Average searching times for adult, su badult, and juvenile kites on Toho and KissWCA3A during the post-invasion era................................................................................84

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10 2-14 Average searching times for adult male and fem ale kites on Toho and Kiss-WCA3A during the post-invasion era...............................................................................................85 2-15 Average searching times for adult kites on Kiss and W CA3A during the postinvasion era and on Toho during the preand post-invasion eras..................................... 86 2-16 Average searching times for juvenile kites on Toho and Kiss-WCA3A during the post-invasion era. ............................................................................................................. ..87 2-17 Average handling times for adult male and female kites on Toho and Kiss-WCA3A during the post-invasion era. ..............................................................................................88 2-18 Average handling times for adult kite s on Kiss-WCA3A during the post-invasion era and on Toho during the preand post-invasion eras. ......................................................... 89 2-19 Average handling times for juvenile kites on Toho and Kiss-WCA3A during the post-invasion era.. ..............................................................................................................90 2-20 Raw group drop rates for adult and juve nile kites on Toho and Kiss-W CA3A during the post-invasion era..........................................................................................................91 2-21 Average drop rates for adult male and fe male kites on Toho and Kiss-WCA3A during the post-invasion era...............................................................................................92 2-22 Average drop rates for adult and juven ile kites on Toho and Kiss-W CA3A during the post-invasion era.............................................................................................................. ..93 2-23 Average capture rates for adult male and female kites on Toho and Kiss-WCA3A during the post-invasion era.. .............................................................................................94 2-24 Average capture rates for adult kite s on Kiss and WCA3A du ring the post-invasion era and on T oho during the preand post-invasion eras.................................................... 95 2-25 Average capture rates for juvenile kites on Toho and Kiss-WCA3A during the postinvasion era. .................................................................................................................. .....96 2-26 Estimated energetic content of native a nd exotic snails consum ed by kites during the post-invasion era................................................................................................................97 2-27 Average proportion of time spent flyi ng during day-long and hourly observations by kites on Toho, Kiss, and WCA3A in the post-invasion era. .............................................. 98 2-28 Average capture rates achieved by fo raging kites during m orning, afternoon, and evening observations on Toho and Ki ss-WCA3A in the post-invasion era...................... 99 2-29 Average proportion of time spent fl ying by all kites on Kiss, Toho, and WCA3A during the post-invasion era.. ...........................................................................................100

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11 2-30 Average proportion of time spent flyi ng by adult and juvenile kites on Toho and Kiss-W CA3A during the post-invasion era..................................................................... 101 2-31 Estimated gross daily energetic gains fo r adult and juvenile kites foraging on Toho, Kiss, and WCA3A during the post-invasion era. .............................................................102 2-32 Estimated daily energy balances for a dult and juvenile k ites on Toho, Kiss, and WCA3A during the post-invasion era..............................................................................103 3-1 Annual number of snail kite ne sts initiated ra nge-wide, 1995-2007. ..............................122 3-2 Average number of nests initiated annually on Toho during the preand postinvasion eras. Post-invasi on estim ate excludes 2004...................................................... 123 3-3 Number of nests initiated annually on Toho, 1995-2007................................................ 124 3-4 Relative annual contribution of Toho to the total nesting effort range-wide, 19952007..................................................................................................................................125 3-5 Trend model expressing the increasing c ontribution of Toho to the total population nesting effort over tim e (i ncludes all years 1995-2007).................................................. 126 3-6 Trend model expressing the increasing c ontribution of Toho to the total population nesting effort over tim e (excludes dr ought years 2001, 2007 and drawdown year 2004)................................................................................................................................127 3-7 Relative contribution of T oho to the total population nest ing effort during the preand post-invasion eras.. ....................................................................................................128 3-8 Annual nest success on Toho, 1995-2007........................................................................ 129 3-9 Average nest success on Toho during the preand post-invasion eras............................ 130 3-10 Annual number of young fledged pe r successful n est on Toho, 1995-2007................... 131 3-11 Average nest productivity on Toho duri ng the preand post-invasion eras. ...................132 3-12 Average number of young fledged per succ essful nest on Toho vs. all other wetlands during the post-invasion era.. ...........................................................................................133 3-13 Apparent annual survival of juvenile s hatched on Toho during the preand postinvasion eras.................................................................................................................. ...134 3-14 Apparent annual survival of juveniles hatched on Toho, 1992-2006. .............................135 3-15 Apparent monthly survival of juveni les hatched on Toho during the preand postinvasion eras.................................................................................................................. ...136

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12 4-1 Average number of nests initiated annually on nine wetlands during the postinvasion era. .................................................................................................................. ...147 4-2 Annual nest success on nine wetla nds during th e post-invasion era............................... 148 4-3 Annual nest productivity on nine we tlands during the post-invasion era. .......................149 4-4 Total number of nests in itiated annually on nine wetlands during the post-invasion era. ....................................................................................................................................150 4-5 Total number of young fledged per year (uncorrected counts) on nine wetlands during the post-invasion era. ............................................................................................151

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13 LIST OF ABBREVIATIONS 95% CI Ninety-five percent confidence interval AOU American Ornithologists Union df Degrees of freedom FFWCC Florida Fish and Wild life Conservation Commission KISS (or Kiss) Lake Kissimmee n Sample size SD Standard Deviation SFWMD South Florida Wate r Management District TAB Time Activity Budget TOHO (or Toho) Lake Tohopekaliga USACE United States Army Corp of Engineers USFWS United States Fish and Wildlife Service WCA3A Water Conservation Area 3A x Sample mean

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14 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science EFFECTS OF THE INVASIVE EXOTIC APPLE SNAIL ( Pomacea insularum ) ON THE SNAIL KITE ( Rostrhamus sociabilis plumbeus ) IN FLORIDA, USA By Christopher Cattau May 2008 Chair: Wiley Kitchens Major: Wildlife Ecology and Conservation The Snail Kite ( Rostrhamus sociabilis plumbeus ) is an endangered raptor in the U.S. that exhibits an extreme form of dietary specializatio n, feeding almost exclusively on one species of freshwater snail, the Florida Apple Snail ( Pomacea paludosa Say). Lake Tohopekaliga, one of the few remaining wetland fragments utilized by th e snail kite in Florida, recently experienced an infestation of the invasive exotic Island Apple Snail ( Pomacea insularum ), which is relatively larger (length, x = 63.5 mm; weight, x = 56.8 g) than the native apple snail (length, x = 37.6 mm; weight, x = 15.9 g). This relative size differen ce raised questions about the ability of kites (especially juveniles) to negotiate exotic snails, and given the sens itivity of the kite population to recruitment, we conducted a comparative observa tional study to elucidat e the effects of the exotic apple snail on snail kite foraging behavior, energetics, nest success, and survival. Relative to native snails, we found that exotic snails requi re longer handling times (for adults, 302 vs. 72 seconds; for juveniles, 496 vs. 97 seconds), lead to increased drop rates (for adults, 0.21 vs. 0.02; for juveniles, 0.33 vs. 0.06), and result in depressed capture rates (for adults, 1.09 vs. 3.30 snails/hour; for juveniles, 0.7 8 vs. 3.46 snails/hour); however, we also found that exotic snails provide more energy than na tives (12.92 vs. 4.84 kcal/snail). Consequently, the effects of the exotic snail on foraging behavior do not have negative ener getic repercussions for

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15 adult kites. In fact, we found that adult kites are attracted to Lake Tohopekaliga and that the relative contribution of the lake to the range-wide nesting effort increased from 6% to 33% after the invasion of the exotic snail. Conversely, the effect s of the exotic snail on juvenile foraging behavior can lead to insuffici ent daily energy balances and ma y suppress juvenile survival. Given the critically endangered status of the snail kite and the propensity of the exotic apple snail to spread, this work suggests that serious management and cons ervation initiatives that address the exotic apple snail may be necessary to preven t further deleterious consequences for the kite population in Florida

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16 CHAPTER 1 INTRODUCTION Species dependent upon specialized niches ar e particularly subject to environm ental perturbations (Brown & Maur er 1989; Owens & Bennett 2000; Purvis et al. 2000): narrow dietary breadth is one such specialization (Real & Caraco 1986; Begon et al. 1996). In fact, niche specialization is positively correlated with extinction risk across a wide range of taxa (Owens & Bennett 2000; Purvis et al. 2000), including birds (Hughes et al. 2000). The snail kite ( Rostrhamus sociabilis plumbeus) is a wetland-dependent raptor that displays an extreme form of dietary specialization, feeding almost exclus ively on a single species of apple snail ( Pomacea paludosa Say) (Howell 1932; Stieglitz & Thompson 1967), the only species of this genus native to Florida (Rawlings et al. 2007). After se veral decades of landscape fragmentation and hydroscape alteration, the kite popula tion is now confined to a patc hwork of freshwater wetlands that remain within its histori cal range, and the viability of the population rests entirely on the conditions and dynamics of these wetland fr agments (Sykes 1979, 1987a; Bennetts & Kitchens 1992, 1997; Martin 2007). However, many of the remaining wetlands are no longer sustained by the natural processes under which they evolved (USFWS 1999; RECOVER 2005), and hence, are not necessarily characteristic of the hist orical ecosystems that once supported the kite population (Bennetts & Kitchens 1992, 1997, 1999; Ma rtin 2007). Snail kites now face another potential threat, the recently es tablished populations of the invasive exotic apple snail ( Pomacea insularum ) in Florida. Evidence suggests that the exotic apple snail negatively affect s the food handling ability of snail kites (Darby et al. 2007), and there is speculation as to what the consequences for the kite population may be (Darby et al. 2007; Rawlings et al. 2007), but many unanswered questions remain. The snail kite already faces a hi gh risk of extinction (Martin 2007), so t he complications

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17 experienced by kites attempting to ex ploit exotic apple snails are of particular concern, especially given that populations of the native appl e snail have been declining throughout the kites range (Darby et al. 2005) while populations of the exot ic snail in Florida have been spreading (Rawlings et al. 2007). Native sna il populations have declined larg ely in the absence of exotic snails; hence, these population trends are li kely coincidental, but they are concerning nonetheless. Both snail species have characteristics that may influen ce their utility to snail kites. The elucidation of these differe nces, and of their respective effects on snail kite foraging behavior, energetics, and population demography, is an essent ial prerequisite to future conservation planning. Study Population Snail kite ( R ostrhamus sociabilis ) populations occur in North, Central, and South America. Based on apparent geographical va riations in body size, distin ctions are drawn among three subspecies (i.e., R. s. major, R .s. sociabilis, and R. s. plumbeus ), which are recognized by the American Ornithologists Union (AOU) (Ama don 1975). However, only a small number of morphometric measurements were used to separate these subspecies, and the tenuous methodology has been called into question by nu merous authors (e.g., Beissinger 1988, Bennetts & Kitchens 1997, Martin 2007). While the AOU states that R. s. plumbeus occurs in both the United States and Cuba (Amadon 1975), there is no documentation of movement between these two geographic locations (Bennetts & Kitchens 1997; Martin 2007). Furthermore, the aforementioned classification scheme of subspecies, which pools the Florida and Cuba populations together, has no ge netic basis (Beissinger 1988; Ma rtin 2007). Although no study has directly addressed the rela tionship between the Florida an d Cuba snail kite populations, multiple studies (e.g., Sykes 1979; Beissinger et al. 1983; Bennetts & Kitchens 1997; Dreitz et al. 2002; Martin et al. 2006a) of movement and population dynamics suggest that the snail kite

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18 population in Florida is isolated. This study addresses the Florida snail kite population only, and for all practical conservation purpos es, we also consider this popul ation geographically isolated. The snail kite ( Rostrhamus sociabilis plumbeus ) is an endangered raptor (Federal Register 1967, 2007) whose range in the U.S. is confined to the freshwater wetlands of central and southern Florida (Sykes 1984; Bennetts & Kitche ns 1992, 1997; Martin 2007). As an extreme dietary specialist, dependent almost entirely on a single species of freshwater apple snail ( Pomacea paludosa from here on referred to as the native snail or native apple snail ) for food, the snail kite is a wetland-dependent species (Howell 1932; Stieglitz & Thompson 1967; Snyder & Snyder 1969; Sykes 1987a). In addition to fora ging, nesting is also tied directly to wetland habitats. Snail kites always build nests in ve getation surrounded by standing water, which aids in the deterrence of terrestrial predators (S tieglitz & Thompson 1967; Sykes 1987b; Beissinger 1988). Over the past several decades, landscape fragmentation and hydroscape alteration have severely jeopardized the quantity and quality of th e historically contiguous wetland habitats that once comprised the range of the snail kite in Florida (Bennetts & Kitchens 1997, 2000; Kitchens et al. 2002; Martin 2007). The major remaining wetland fragment s used by the snail kite are depicted in Figure 1-1 and incl ude the following: Everglades Na tional Park (ENP), Big Cypress National Preserve (BICY), Water Conservati on Areas (WCA) 1A, 2A, 2B, 3A, 3B, Lake Okeechobee (OKEE), Grassy Waters Preserve (GW), Saint Johns Marsh (SJM), Lake Kissimmee (KISS), Lake Tohopekaliga (TOHO) and East Lake Tohopekaliga (ETOHO) (Bennetts & Kitchens 1997; Dreitz 2000; Kitchens et al. 2002; and Martin et al. 2006a). As of 2007, only one of the major wetlands utilized by the snail kite, Lake Tohopekaliga (from here on referred to simply as Toho), has suffered a major invasion of P. insularum (from here on referred to as the exotic snail or exotic apple snail) (Rawlings et al. 2007; Darby et al. 2007); however,

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19 confirmed exotic snail populations exist in close proximity to many of the other primary wetlands used by the kite in Florida (Rawlings et al. 2007) (Figure 1-2). Within Florida, the kite population is de scribed as nomadic, and monthly movement probabilities among wetland fragments may be as high as 0.25 (Bennetts & Kitchens 1997, 2000). More recent analyses, which include multip le levels of spatial and temporal resolution, suggest that the monthly movement probability among contiguous fragments differs significantly from that among isolated fragments (0.29 vs. 0.10, respectivel y) (Martin et al. 2006a, 2007b). However, numerous studies, of both moveme nt (Sykes 1979; Rodgers et al. 1988; Bennetts & Kitchens 1992, 1997; Martin 2007) and genetics (Rodgers & Stangel 1996), confirm that the spatial distribution of kites in Florida shifts te mporally and that suffici ent individual movement among wetlands occurs, thus uniting the entire Florida population (Figure 1-3). Even though the exotic snail has thus far only infe sted one of the major wetlands util ized by the snail kite, the kite population in Florida does not function as a metapopulation (Bennetts & Kitchens 1997, 2000; Martin et al. 2006b, 2007b; Martin 2 007); therefore, the scope of our project covers the entire kite population. Invasion of the Exotic Snail The exotic a pple snail ( P. insularum ), commonly known as the island apple snail, is native to Argentina, Brazil, and Bolivia. The invasion history of the exotic apple snail is somewhat unclear (Rawlings et al. 2007). P. insularum along with other members of the Pomacea genus that have invaded the U.S., have historically been misidentified as P. canaliculata or lumped together with other species in what was calle d the Canaliculata (or, channeled apple snail) complex (Thompson 1997; Howells et al. 2006; Rawlings et al. 2007) Reliable genetic analysis confirming the presence of P. insularum in Florida dates back to only 2002 (Rawlings et al. 2007); however, exotic snails observed on Toho in 2001 were later identified as P. insularum

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20 (Darby, personal communication ). Many new populations of P. insularum were observed in Florida from 2004 to 2006, suggesting that it may be a relatively new invasive species to the state (Rawlings et al. 2007). The exotic apple snail is know n to feed on the eggs of congeneric snails, and some anecdotal evidence suggests that native apple snail populations appear to decline or even disappear after the introduction of the exotic snail (Rawlings et al. 2007); howev er, native snail populations have declined largely in the absence of exotic snails, and the declining native snail populations have been attributed to other sources such as altered hydrologic regimes and shifts in vegetative communities (Kushlan 1975; Turner 1996; Darby et al. 2003). The ecology of Toho has undergone drastic anth ropogenic changes recent ly. As part of a management strategy aimed at improving habitat conditions for fish populations, a drawdown of the lake stage occurred between late-2003 and ea rly-2004, and an intensive scraping treatment of littoral vegetation followed. Afte r this treatment, water levels on Toho remained low until June 2004. This course of action signi ficantly altered the physical and biological conditions of the lakes littoral zone (Welch 2004; Williams et al. 2005). The emergent vegetation in the littoral zone is a critical component of suitable snail kite foraging habitat (Sykes 1 987a; Bennetts & Kitchens 1997), and once it was removed via m echanical scraping, kites temporarily abandoned Toho (Martin et al. 2003; Kitchens et al. 2005), s ubsequently returning in late-2004 (Kitchens et al. 2005; Martin et al. 2006b). In the interim, the distribution and a bundance of exotic apple snails on Toho grew dramatically (Kitchens et al. 2005). Prior to the drawdown and scraping treatment, native snails domina ted the littoral habitats on Toho, while the exotic snails were largely confined to the pelagic habitats on the in terior side of the lake However, the drawdown and scraping treatment, as well as the prol onged dry conditions that followed it, greatly

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21 suppressed the population of native snails on Toho. When the lake stage rose, the exotic snail population spread out and inva ded the littoral zone (Darby, personal communication ). Since 2003, the exotic snail has been the most abundant apple snail in portions of the lake utilized by snail kites (Kitchens et al. 2007). Population Demography Since 1992, the snail kite populat ion has been m onitored annu ally via robust population surveys, which include extensive mark-recapture techniques. Reliable estimates of abundance, survival, and movement are attainable from these data by utilizing multi-state models, such as Cormack-Jolly-Seber models (C JS), which incorporate detection probabilities and spatial variation (Bennetts & Kitchens 1997; 1999; Dreitz et al. 2002; Martin et al. 2006a, 2007a). Estimates of the snail kite population size and growth rate for the years 1992 to 2005 show several alarming trends. During the period 2000 to 2002 the snail kite population essentially halved, falling from around 3500 to around 1700 indi viduals (Martin et al. 2007a). The most recent demographic analyses show no signs of a significant population rebound. In fact, the estimated stochastic population growth rate for the post-population-d ecline period (i.e., 20032006) is less than one (Martin 2007; Martin et al. 2007a). As indicated by a life table response experiment (LTRE), over 80% of the reduction in the stochastic populatio n growth rate after 1998 is attributable to adult fert ility (Martin 2007). Finally, the most recent snail kite population viability analysis (PVA) shows that extinction wi thin the next 60 years is highly probable if current conditions are representative of future conditions. (For a de tailed discussion of snail kite population viability analyses see Martin 2007). Since the snail kite population is at risk of extinction and because adult fert ility plays such an overwhelming role in the population growth rate, it is critical to identify and attempt to remedy all factors that negatively affect snail kite fertility in order to properly manage for the conservation of the species

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22 Adult fertility is simply the product of 1) the number of juveniles produced per adult during the breeding season and 2) the probability that these juve niles survive until the next breeding season (i.e., juvenile survival) (Martin 2007). Using the aforementioned demographic trends, the recent history of the snail kite popul ation in Florida can be divided into two time periods: 1) the pre-1998 period (inc luding 1998), which is representa tive of a stable population in terms of demographic trends and environmen tal conditions, and 2) the post-1998 period (not including 1998), which is representative of th e population decline and subsequent suppression (Martin 2007). The snail kite population, which was in a steady st ate of decline throughout the period 1999 to 2002, has since stabilized. Adult and juvenile survival have, on average, resumed levels comparable to the pre-1998 period, but nes ting efforts and fecundity remain significantly lower. One anomaly is the unprecedented flurry of nesting attempts that have taken place on Toho since the invasion of the exotic apple snail. In spite of these nesting efforts, preliminary analyses show less than expected recruitmen t from kites on Toho during the post-invasion era (Martin et al. 2007c), suggesting that the exotic snail may nega tively affect nest success or juvenile survival. The invasion of Toho by the exotic apple snail has created unique conditions, in terms of available prey species, for kites. Any deleteriou s effects that exotic sn ails have on snail kite foraging behavior may, in turn, have severe demographic reperc ussions. Notwithstanding the very real possibility that the exotic snail will spr ead into other wetlands u tilized by the snail kite, the fact that the kite population in Florida does not func tion as a metapopulation implies that negative effects experienced by kites on Toho may have population-wide consequences, particularly given the sensitivity of the population growth rate to fecundity and recruitment.

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23 Study Objectives and Outline The governm ent agencies responsible for managing natural res ources (e.g., FFWCC, SFWMD, USFWS, USACE) exhibit tremendous in fluence over the remaining wetlands that comprise the range of the snail kite in Fl orida (Bennetts & Kitc hens 1997; USFWS 1999; RECOVER 2005). Water regulation schedules are used to manage the composition and dynamics of wetland ecosystems on a regional scal e. The hydrology is dictated via a complex network of levees, canals, and other water cont rol structures; and smaller-scale management actions targeted at specific flora and fauna ar e often conducted via mechanical and chemical controls (USFWS 1999; Welch 2004). It is the goal of the multi-billion dollar restoration initiative, the Comprehensive Everglades Restor ation Plan (CERP), to create a management paradigm that mimics the historical, natural-fl ow patterns and community compositions that once drove and maintained the integrity and biodive rsity of Floridas natural wetland ecosystems (USFWS 1999). Understanding the effects that the exotic apple snail has on snail kite foraging and demography precludes our ability to strategical ly manage Floridas wetlands in a way that sustains overall ecosystem health and alleviat es any detrimental consequences to the kite population. As stated by the CERP, The desire d restoration condition for the Everglade snail kite is to restore and maintain a network of sna il kite foraging habitats and promote habitat that supports primary prey (apple snails) recruitm ent throughout the South Florida ecosystem (USFWS 1999; RECOVER 2005). To help reach these goals, we conducted a comparative observational study focused on discerning the effects of the exotic apple sn ail on the snail kite population. We hope that this study will have important applications in both species management and conservation biology:

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24 Figure 1-1. Major habitat fragments within th e range of the snail kite in Florida (modified from Dreitz 2000). Southern Range 1. Everglades National Park (ENP) 2. Big Cypress Nationa l Preserve (BICY) 3. Water Conservation Area 1A (WCA1A) 4. Water Conservation Area 2A (WCA2A) 5. Water Conservation Area 2B (WCA2B) 6. Water Conservation Area 3A (WCA3A) 7. Water Conservation Area 3B (WCA3B) 8. Lake Okeechobee (OKEE) 9. Grassy Waters Preserve (GW) Northern Range A. Saint Johns Marsh (SJM) B. Lake Kissimmee (KISS) C. Lake Tohopekaliga (TOHO) D. East Lake Tohopekaliga (ETOHO) C D B 8 A 3 9 4 5 6 6 7 1 2

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25 Figure 1-2. Confirmed locations, as of 2007, of the exotic snail ( P. insularum ) within the range of the snail kite in Florida. Larger ci rcles encompass multiple small populations in close proximity (modified fr om Rawlings et al. 2007).

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26 Figure 1-3. Movements of approximately 50 radio-tagged adult sn ail kites among wetland fragments over a one-year period. April 1992-April 1993 (modified from Bennetts & Kitchens 1997).

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27 CHAPTER 2 EFFECTS OF THE EXOTIC APP LE SNAIL ON SNAIL KI TE FORAGING BEHAVIOR An animals fitness is inextricably linked to it s diet. The ability to meet certain energetic and nutritional thresholds is a requisite for su rvival, growth, and reproduction. The options available to an individual attempting to meet these energetic requirements are dictated by its physical and behavioral traits and by the enviro nment in which it lives. An animals dietary breadth is therefore constraine d by its genetic makeup and by the resources presently available (Fox 1981; Pyke 1984; Stevens & Krebs 1986; Ehlinge r 1990). In ecology, animal species are often broadly classified as gene ralists and specialists. Generalis ts typically incorporate a wide range of prey types into their diets and utilize varying tactic s to attack, subdue, and consume different prey. Dietary specialists, on the ot her hand, have narrow diet ary breadths and often utilize acute skill sets that faci litate the efficient exploitation of specific prey types (McArthur 1972; Pyke 1984; Stevens & Krebs 1986). Adaptations facilitating the abili ty of a species to exploit specific food resources ma y hinder the ability of that species to expl oit alternative prey types. Antagonistic pleiotropy, the positive gene tic feedback loop through which such inhibition may occur, can lead to extreme dietary speci alization in some instances (Futuyma & Moreno 1988; Cooper & Lenski 2000). Thus, an organisms current behavioral tra its and dietary options are restricted, to an extent, by morphological or physiological characteristic s consequent of past adaptations (McArthur 1972; Pyke 1984; Futuyma & Moreno 1988; Fry 1990; Holt 1996). The snail kite ( Rostrhamus sociabilis plumbeus) is one such extreme dietary specialist, and its unique physical traits and foraging behaviors are tied to its nearly exclusive prey the Florida apple snail ( Pomacea paludosa Say) (Cottom & Knappen 1939; Stieglitz & Thompson 1967; Snyder & Snyder 1969; Sykes 1987a). The extreme form of dietary specialization observed in the snail kite has led some authors to question the ability of the kite to efficien tly incorporate a newly

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28 introduced exotic apple snail, Pomacea insularum into its diet (Rawlings et al. 2007; Darby et al. 2007). Background Foraging Behavior Num erous authors (e.g., Holgerson 1967; Snyder & Snyder 1969; Haverschmidt 1970; Collett 1977; Beissinger 1983; Sykes 1987a) have described the details of snail kite foraging behavior, and here we provide a brief synopsis of those details pertinent to our study. Snail kites exercise two methods of fora ging: course-hunting and perchhunting. While course-hunting, kites usually fly between 1.5 to 10.0 meters above the surface of the water with their heads angled downward looking for snails. These flig hts follow a slow flap-glide pattern and are usually directed into the wind, when present. Upon sighting a snail, kites will make an abrupt turn and descend toward the wate r, hovering briefly just above the surface while reaching into the water with the talons to capture the snail. Sn ails are always captured with the talons, but they may be transferred into the bill during flight (Snyder & Snyder 1969; Beissinger 1983; Sykes 1987a). Kites will then return to a feeding perch in order to ma nipulate and consume the snail. Using the talons, kites position the snail shell so th at the aperture is facing away from the perch and the spire is pointed downward. Kites pry op en the hard operculum of the snail using the torque generated by their upper mandible being pressed against the shell wall. The sharp tip of the upper mandible is then used to cut the musc le attachments of the operculum. After the operculum is removed, kites must further negotiate the shell, repositioning it so that the aperture is facing upward then inserting their curved upper mandible into th e spiraling apertu re to cut the columellar muscle that connects the snails body to the shell. The soft body tissue of the snail is removed and consumed either whole or in torn pa rts. The albumen gland of female snails is often discarded, and the viscera of snails is sometimes eaten and sometimes not. The shell is

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29 usually dropped after extraction, but sometimes kites will hold onto the shell until consumption is complete. Shells of consumed snails fr equently accumulate under feeding perches (Snyder & Snyder 1969; Snyder & Kale 1983; Sykes 1987a). While employing the second form of foraging, perch-hunting, kites remain perched wh ile scanning the surrounding water (1 to 12 meters away) for snails (Sykes 1987a). Upon detection, the kite wi ll swoop down from the perch, and just as in course-hunting, hover direc tly above the surface of the water, grabbing the snail with the talons. The kite will then return to the perch of departure in order to consume the snail. After every successful foraging bout, kites perform th e same meticulous process of negotiating, extracting, and ea ting the snail (Sny der & Snyder 1969; Sykes 1987a). It is important to note that snail kites in Fl orida coevolved with the Florida apple snail. Therefore, certain morphological attributes of the kite seem specifically tailored to exploiting native snails that fall within a particular size range (around 42.6 mm or golf ball-sized) (Snyder & Snyder 1969; Sykes 1987a). The bill of the snail kite is especially adapted to extract the soft tissue from these shells. The deeply -hooked upper mandible of the snail kite has a curvature that closely mimics that of the inner spiral of the apple sn ail shell (Snyder & Snyder 1969), and the bill width of adult kites ranges from 26-34 mm, restri cting the depth into the shell that a kite can reach to cut th e columellar muscle (Sykes et al. 1995). Other physical features of snail kites may also affect their ability to handl e alternative prey items. Middle toe plus claw widths for adult kites range from 49 to 68 mm (Sykes et al. 1995). Average weighs of adult females, adult males, and juveniles are 446, 394 (Sykes et al. 1995), and 389 grams (ValentineDarby et al. 1997) respectively. These attributes potentially impose limitations on the sizes, weights, and types of prey items that can be successfully captured, handled, and consumed by kites.

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30 On rare occasions (e.g., during drought conditions or other instances of food scarcity), kites in Florida have been observed attempting to feed on alternative (i.e ., non-snail) prey items (e.g., small turtles, crawfish, a dead bird, a snake) (Sykes & Kale 1974; Woodin & Woodin 1981; Sykes 1987a; Beissinger 1990; Bennetts et al. 1994); however, they are rarely successful, and certainly not efficient, at extracting ed ible body parts (Sykes & Kale 1974; Beissinger 1988, 1990). Many reported cases of non-snail foraging have involved juvenile birds (Sykes 1987a). In another isolated incident, th ree to four snail kites were obs erved feeding on another exotic snail, Pomacea bridgesii in a flooded agricultural fiel d during a drought (Takekawa & Beissinger 1983). Small, is olated populations of P. bridgesii have been present in south Florida since at least 1966. However, unlike the newly introduced P. insularum P. Bridgesii have not spread to any of the major wetla nds utilized by the snail kite a nd show no evidence of being an invasive threat (Rawlings et al. 2007). A Note on Pomacea Measurements The field of m alacology has yet to adopt standardized measurement protocols for Pomacea (Youens & Burks 2007); therefore, throughout much of the literature, identical dimensional measurements have been described using inconsistent terminology. We adopted the measurement protocols for linear dimensions of length and width as pres ented in Darby et al. (2007). In this study, we standardized the use of these terms, and for all of the results from previous Pomacea studies described within, we relabeled the dimensional measurements in terms of length and width by assessing th e measurement methods reported by the respective authors. The Native Apple Snail Of the m any morphological and physiological factors that can influence prey selection and suitability, prey size may be the most pertinent when dealing with extreme dietary specialists (Stevens & Krebs 1986; Sih 1987). The Florida apple snail ( Pomacea paludosa Say), which is

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31 the only apple snail native to Florida, effectively constitutes the entirety of the snail kites diet and fulfills all of the necessary energetic a nd nutritional requirements for kites (Cottom & Knappen 1939; Stieglitz & Thompson 1967; Snyde r & Snyder 1969; Sykes 1987a). Adult snail shells have a normal range of 40 to 70 mm in length (Thompson 1984). Sykes (1987a) collected empty shells (n=697) from beneath feeding perche s used by snail kites. He found that kites selected snails ranging from 25.2 to 71.3 mm in width (x=42.8, SD=4.9) and from 27.4 to 82.4 mm in length (x=45.8, SD=5.1), with 98.5% of the sn ails falling in the 30 to 60 mm range for both measures and 70% falling in the 40 to 50 mm range. Severa l other authors (e.g., Beissinger 1983; Bourne 1983, 1985a; Tanaka et al. 2006) have also noted that kites rare ly eat snails greater than 60 mm in length. Tanaka et al. (2006) recorded the largest native sn ail (86 mm in length) to have been consumed by a kite. Sykes (1987a) fo und that the average wet weight (whole snail with shell) of live native apple snails ranged from 12.7 to 38.1 grams (x=22.3, SD=6.1, n=24). Darby et al. (2007) reported that the wet weight of an average-size native apple snail collected on Lake Kissimmee during the fall of 2004 was 35 grams (n=1). The Exotic Apple Snail From 2003 to 2007, exotic apple snails were dominant throughout Toho in the areas of the lake utilized most heavily by k ites (Kitchens et al. 2007; Darby, personal communication ). Exotic apple snails commonly exceed 90 mm and can reach 150 mm in length (Benson 2007; Darby et al. 2007), possibly creating a conflict with the life history strategy of the snail kite. Darby et al. (2007) sampled exotic apple snails in Goblets Cove on Toho via throw traps and dip nets. Shell widths of these snails ranged from 67 to 97 mm (x=81, SD=6, n=64) with estimated lengths (calculated using a 1.15 wi dth to length ratio) ranging from 77 to 112 mm (x=95, SD=7, n=64). The wet weight (whole snai l with shell) of an exotic snail with average width and length dimensions was 175 grams (n=1).

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32 On average, the exotic snail is large relative to the native snail, and some size classes of the exotic apple snail may not constitute suitable prey types for snail kites (F igure 2-1). The largest exotic snails may be rendered unavailable if kite s cannot capture them or if kites cannot properly negotiate them while attempting to feed. Relative to the native snails, exotic snails may also require longer handling times, which translate into increased energy expenditures for kites. Useful information alluding to how the size dispar ity between the native and exotic apple snail may affect snail kite foraging behavior recently emerged (Darby et al. 2007), but the issue has yet to be fully addressed, especially in the context of energetic repercussions. Previous Foraging Studies Most snail kite foraging studi es in Florida pre-date the invasion of Toho by the exotic apple snail, and given the snail kites extrem e nature of dietary specialization, previous authors were largely unconcerned with prey type or snail size as influential factors of foraging success. Nevertheless, several aspects of snail kite fora ging behavior have been quantified (e.g., average searching times, handling times, and capture ra tes) (Cary 1985; Sykes 1987a; Beissinger 1990; Bennetts & Kitchens 1997, 2000). Most early studies of snail kite foraging in Florida focus on modeling capture rates (as measured by the number of snails captured divided by the time spent hunting) as a function of environmental variab les (e.g., water depth, temp erature, vegetative community) and attempt to make inferences about prey availability and re lative habitat quality (Darby et al. 2006; Karunaratne et al. 2006; Bennetts et al. 2006). Prey availability Foraging snail kites are lim ited by the availability of apple snails (Sykes et al. 1995). Some authors have used prey dens ity as a proxy to prey availability in avian foraging studies, but this may result in biased inferences (Kushlan 1989; Gawlik 2002). In the case of the snail kite, snail densities below 0.14 snails/m2 do not support foraging kites (D arby et al. 2006), but snail

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33 density is only one of the fact ors that influences prey availa bility (Beissinger 1983; Bourne 1985a, 1985b; Sykes 1987a; Bennetts & Kitchens 1997, Darby et al. 2006, Bennetts et al. 2006). To be available, snails must not only be present; they must also be visible and accessible to foraging kites. For example, snail densitie s may be suitably high in certain vegetative communities (e.g., sawgrass, cattai l) but inaccessible due to visual or physical obstruction (Beissinger 1983; Sykes 1987a; Benne tts et al. 2006). In other cas es, snails may be present and visible (e.g., open water habitats ), but without sufficient emergent vegetation, snails cannot surface (Hanning 1979; Turner 1996) and may remain too deep (>16 cm) for kites to capture (Sykes 1987a). A number of authors have iden tified other factors that may influence the visibility and accessibility of snails to kites. Apple snails are poikilothermic and become less active as water temperature decreases. As temperatures fall, the metabolism of apple snails slows down, which lowers their oxygen requirement, and at the sa me time, dissolved oxygen in the water column increases. Thus, snails visit the surface much less frequently when water temperatures are low (Hanning 1979; Stevens et al. 2002 ), and temperature is negative ly correlated w ith the capture rates achieved by foraging kites (Cary 1985; Benne tts & Kitchens 1997). In extreme cases (<10 oC), snails may be present but completely inactiv e, and thus unavailable (Cary 1985; Stevens et al. 2002). Snails also become inactiv e during low water events (<10 cm) (Darby et al. 2002), often burying themselves in the substrate, presumably in preparation for aestivation (Snyder and Snyder 1969; Kushlan 1975, 1989; Dar by & Percival 2000; Darby et al. 2002), again creating a situation where snails are pres ent but undetectable or inaccessible for kites (Stevens et al. 2002 ; Darby et al. 2003, 2005 ). Rain and heavy wind have also been implicated as factors that reduce kites ability to detect sn ails (Cary 1985; Bennetts & Kitchens 1997). All of

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34 t hese studies were conducted be fore the invasion of Toho by the exotic snail, and none truly address the final component of apple snail availability: negotiability Prey negotiability and drop rate Apple snails can be present, de tectab le, and accessible, yet stil l unavailable if they are not within the size range that can be captured and successfully negotiated by kites to the point of consumption. If a snail is droppe d (and not recovered) during a ny phase of the handling process before it is consumed, that snail effectively becomes unavailable. Cary (1985) observed 1% and 9% drop rates for kites feeding on native sn ails while course-hun ting and perch-hunting respectively. However, the author noted that shells from previously consumed snails may remain suspended near the surface of the water in close proximity to feeding perches and that accidental captures of these empty shells may ha ve contributed to the higher drop rates observed for kites employing the perch-hun ting strategy. Nonetheless, Ca ry (1985) did not distinguish between live snails that were dropped due to mish andling and empty shells that were accidentally captured and subsequently rejected, nor did he implicate size as the causal factor of snail dropping. In a recent study, Darby et al (2007) reported a 44% drop ra te for kites foraging on the exotic snail on Toho and did, in fact, verify that no empty shells were counted as dropped live snails. The authors also noted that an average-si ze exotic apple snail may weigh 37% to 45% as much as a snail kite, and due to their relative sizes, many exotic snails may be unavailable even though they exist in high densities (Darby et al. 2007). It is this latter component of availability involving present and accessible, yet nonnegotiable, snails that raises questions about the suitability of the exotic apple snail as prey for the snail kite in Florida.

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35 Searching and handling time Flight is one of the m ost energetically expe nsive activities in whic h birds engage (Masman & Klaassen 1987; Krebs & Davies 1997; Harrison & Roberts SP 2000). Given that snail kites in Florida rely heavily on a course -hunting strategy (94% of all foraging bouts) (Cary 1985), the time spent searching for snails can significantl y affect the net energe tic gains of foraging. As described in Sykes (1987a), who studied the foraging behavior of kites intermittently from 1967 to 1980, the average duration of a coursehunting foraging bout was 72 seconds for adult males (SD=1.2, range=30-180, n=18) and 150 seconds for adult females (SD=2.9, range=30-720, n=34), which the author notes is a significant difference. This conclusion should be viewed with caution considering the relatively small sample sizes used and the fact that spatiotemporal heterogeneity was ignored. No significant difference between the sexes was found for the mean interval between successive captures, 1338 seconds (range=120 to 5400, n=109), or for the average captures per hour, 2.5 snails/h (range=1 .7 to 3.4, n=109) (Sykes 1987a). Taking temperature effects into consideration, Cary (198 5) reported average capture rates of 5.28 snails per hour above 30oC, 2.68 between 21-30oC, 0.72 between 11-20oC, and 0.0 below 10oC. The primary prey of an extreme dietary speci alist must be profitable enough to provide sufficient net energy for survival, growth, and re production. Handling time al so directly affects the profitability of selected prey items and is one of the central determinants of diet breadth (Werner & Hall 1974; Stevens and Krebs 1986). S ykes (1987a) reported the mean combined handling and eating time for a dult kites to be 162 seconds (SD=84, range 60-420, n=16 males and 53 females), with no signifi cant difference between the sexes. Another study of adult kites consuming native apple snails, Beissinger (1990 ) found the mean combined handling and eating time to be 95.7 seconds (SD=37.3, n=197).

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36 More recently, kites foraging on exotic snails on Toho were compared with those foraging on native snails on Kiss, and snail size was imp lied to affect handling time. On Toho, the average extraction time for the exotic snail was 333 seconds (SD=178, n=10), but the handling times for snails captured on Kiss were not record ed (nor were the searching times for either location). However, as mentioned above, k ites on Toho were observed capturing 25 exotic snails, of which 11 were dropped while handli ng. Kites on Kiss were observed capturing 136 native snails, of which none were dropped (Darby et al. 2007). If one calculates the cumulative handling time (and likewise, searching time) by incl uding all of the failed handling attempts that occur between successful foraging bouts, then these instances of sna il dropping could greatly increase the time and energy spent by kites foragi ng for exotic apple snails. Darby et al. (2007) did not test for age or sex effects on handling tim es or drop rates but did identify experience and relative body weights as likely influe ntial factors of prey handling. Age and sex effects on foraging behavior Learning and experience have been recogni zed as im portant factors contributing to foraging behavior. Animals develop search imag es and improve handling techniques for familiar prey types (Krebs & Davies 1997). Several studies have confirmed that adult snail kite survival remains relatively constant from year to year, while juvenile survival varies widely (Bennetts & Kitchens 1997; Dreitz et al. 2004; Martin et al. 2006a). Experience, and in turn, foraging efficiency may play a large role in this disparit y. Juveniles of most vert ebrate species lack the experience accumulated by adults and are thus more sensitive to environmental variation (Stephens & Krebs 1986; Stearns 1986; Martin et al. 2007a). Sykes et al. (1995) notes that compared to adults, recently-fle dged kites do not demonstrate th e same adept negotiation skills when extracting native apple snails and observations of frequent snail dropping by juvenile kites on Toho during the post-invasion er a suggest the same (personal observation). In addition to

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37 lacking experience, juvenile kite s also weigh roughly 10% less th an adults (males and females combined) (Sykes et al. 1995; Valentine-Darby et al 1997); therefore, the ab ility of juveniles to negotiate exotic snails may be further lim ited by their relative physical proportions. Adult kites may also experience handling compli cations due to the decreased predator to prey weight ratios associated w ith exotic snails. Snail kites are sexually dimorphic, with males ranging from 360 to 440 grams (x=394, SD=22.8, n=28) and females ranging from 350-570 (x=446, SD=47.8, n=29) (Sykes et al 1995); therefore, sex may play a role in the ability of adult snail kites to efficiently exploit exotic apple snails. Energy budgets and profitability Many of the com ponents necessary for the computation of energy budgets for snail kites are present in the litera ture, including daily activity budgets (Beissinger 1984; Cary 1985; Sykes 1987a), energetic costs of behavior (Bei ssinger 1984, 1986, 1987; Beissinger & Snyder 1987), and energetic contents of na tive apple snails (Beissinger 1984; Sykes 1987a), but none have holistically addressed energetic ba lances for kites in Florida in the context of alternative prey types. Snail kites in South America (i.e., R. s. sociabilis ) have been the focus of many prey selection and energetics studi es (Beissinger 1983, 1990; Beissi nger et al. 1994; Bourne 1983, 1985a, 1985b, 1993; Tanaka et al. 2006) Of note, the ranges of P. insularum and R. s. sociabilis overlap, but no documentati on of kites feeding on P. insularum in South America exists (Sykes et al. 1995; Darby et al 2007). The diet of R. s. sociabilis commonly includes one species of apple snail ( P. dolioides) and one species of crab ( Dilocarcinus dentatus ), each with varying size classes, creating an ideal envir onment in which to test certain predictions of optimal foraging theory (Bourne 1983; Beissinger et al. 1994). Handling time was found to be positively related to snail size for kites in Venezuela (Beissinger et al. 1994). Furthe r study of negotiability as a

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38 limiting factor of prey availability and suitabl e prey size is necessary in order to draw meaningful conclusions for the conservation of the snail kite if Florida. Research methods aimed at elucidating the effects of different prey type s (as used in these South American studies) are now applicable to Florida snail kites fora ging on native and exotic apple snails. The calculation of profitability values will dem onstrate the utility that each prey species has to snail kites. Searching time, handling time, and energetic content may vary among prey types, and profitability is a meaningful quantita tive scale on which the costs and benefits of foraging on alternative prey items can be compared (MacArthur & Pianka 1966; Stevens & Krebs 1986). Such an approach was used to comp are dietary choices and foraging site selection of snail kites ( R. s. sociabilis ) in Guyana (Beissinger 1983, 1990; Bourne 1985a, 1985b). However, kites in South America remain idle dur ing long periods of the day (periods that could be spent foraging), and profitability may not be the sole determinant of foraging decisions in snail kites (Beissinger 1 983; Bourne 1985a, 1985b). Therefore, a more holistic measure of the costs and benefits associated with alternative foraging conditi ons may be more useful. The calculation of daily energy balances for kites foragi ng on alternative prey types will shed light on the potential effects the exotic snail may have on the kite population. As stated above, negotiability has been largely ignored until recen tly, and unfortunately, data collected in the recent study (i.e., Darby et al. 2007 ) is insufficient to calculate profitability values and daily energy balances for kites foraging on native versus exotic snails. Economic models of foraging behavior In the study of foraging behavior, econom ic mode ls that compare the costs and benefits of alternative food choices are comm onplace (Krebs & Davies 1997). A general assumption of all studies that use this cost-benefit approach to compare alternative decisions is that it pays for an animal to be efficient. That is, natural select ion will act in favor of individuals that maximize

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39 their benefits while minimizing their cost s (MacArthur 1972; Staddon 2003). Using economic models in the study of foraging be havior requires investigators to define a set of costs and benefits that can be quantifie d and compared using a common currency. The ideal currency with which to weigh the costs and benefits of decisi ons would be fitness (Krebs & Davies 1997). Glimcher (2002) points out that Darwinian fitness provides a natural and genuinely onedimensional scale. Unfortunately, directly quantifying the effects that certain foraging behaviors have on fitness can be difficult; ther efore, more easily quantifiable currencies (e.g., time and energy) are often employed by these ec onomic models. An underlying assumption of these surrogate currencies is that they correlate directly with fitne ss (Krebs & Kacelnik 1991; Brown et al ., 1993). Time and energy were the first tw o currencies used in the development of foraging and diet selection models (MacArthur & Pianka 1966; Emlen 1966), and they have remained the dominant measures in these behavi oral studies ever since (Cuthill & Houston 1997; Sih & Christensen 2001). Predictions Based on our knowledge of snail kite ecology, we expect the exotic apple snail to have deleterious effects on kite foragi ng behavior, and we expect these effects to vary in magnitude relative to foraging experience and relative body weight; therefore, we make the following predictions: Prediction 1. If the average drop rate for exotic sn ails is s ignificantly greater than the average drop rate for natives (i.e., if Prediction 3 is true), then the average searching time for exotic snails will be significantly longer than the average searching time for native snails Prediction 2. The average handling time for exotic snai ls will be significantly longer than the average handling time for native snails. Prediction 3. The average drop rate for exotic snails will be significantly greater than the average drop rate for native snails.

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40 Prediction 4. When feeding on exotic snails, juvenile kites will have significantly greater a) handling times and b) dr op rates than adults. Prediction 5. When feeding on exotic snails, adult ma les will have significantly greater a) handling times and b) drop ra tes than adult females. Prediction 6. The average capture rate for exotic snai ls will be significantly lower than the average capture rate for native snails. Prediction 7. The average profitability of the exotic sn ail will be significantly lower than the average profitability of the native snail. Prediction 8. Daily energy balances will be significantly lower for kites feeding on the exotic snail relative to kites feeding on native snails. Methods To address the above predictions, we conducte d a com parative observational study. First, we quantified the necessary set of measures that pertain to fora ging behavior and prey items. Then we used these data to a ddress the anticipated behavioral differences between/among groups of kites. After teasing out the effects of the exotic apple snail on foraging behavior, we assessed the energetic repercussions for kites by calculati ng prey profitabilities, deriving activity budgets, and projecting daily energy balances. Time Activity Budgets In order to obtain quantitative behavioral data related to foraging activities, we conducted tim e-activity budget (TAB) observations on wild snail kites from 2003-2007 using a modification of focal animal sampling (Altman 1974). Such a method was used to collect TAB data on kites in 1993, 1994, and 1996 (Bennetts & Kitchens 1997, 2000), and so that we could make meaningful comparisons between datasets, we followed similar protocols. During each TAB we observed a focal kite continuously for an allotted period of time and recorded the frequency and duration of all per tinent events and behaviors (see Collection of Behavioral Data ).

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41 Sampling locations Most of our TAB sa mpling effort was focu sed in the following wetlands: Toho, Kiss, WCA3A (Table 2-1; Figure 1-1) Together, these fragments re present much of the remaining core of snail kite habitat in Florida. In recen t years, the snail kite population has relied heavily on Toho, Kiss, and WCA3A during the breeding season. During our study, these were the only wetlands that produced significant numbers of obs ervable juveniles. Kites also utilize Toho, Kiss, and WCA3A during the non-breeding season ; therefore, numerous kites can be found foraging in these wetlands throughout most of the year (Bennetts & K itchens 1997; Martin 2007; Martin et al. 2007c). We also collected data, at least opportuni stically, throughout the remainder of the kites range, but much less effort wa s expended conducting TABs in the following nonfocal wetlands: WCA3B, WCA2B, ENP, OKEE, GW, and SJM (Table 2-1; Figure 1-1). Selection of focal individuals Given the extensive range of th e snail kite and the high expense of field operations, on any given day the wetlands in which we m ade obser vations were determined systematically to optimize travel and funding efficiency. However, we randomly selected th e individual birds that were observed within each wetland fragment. To do so, a block grid with numbered cells was overlaid on a map of the wetland area, and we used a random number generator to select the cell in which to start our observations. Using an airboat and GPS unit for transportation and navigation, we traveled to the se lected cell and observed the firs t snail kite detected upon our arrival. Observations started as soon as the boat was parked a nd necessary equipment set up. If no snail kite was detected in the given cell, we randomly moved to one of the eight surrounding cells by blindly spinning a compass and driving to the adjacent cell at wh ich it pointed. This process was repeated until a snail kite was loca ted for observation. The latitude and longitude (measured in UTMs, Datum NAD83) of all observation locations were recorded with a hand-

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42 held GPS unit. After completing a TAB, the rando m number generator was used to select a cell for the next kite observation, a nd the process started again. Timing and duration of observations TABs were conducted during all m onths of the year (Table 2-1). Months were assigned to one of three seasons as follows: Spring included January, February, Marc h and April; Summer included May, June, July and August; and Fall included September, October, November and December (Bennetts & Kitchens 2000). Spring coinci des loosely with the peak of nest initiation during the breeding season and w ith the annual dry season (Benne tts & Kitchens 1997), although both of these measures show some overlap with Summer (Sykes et al. 1995, Martin 2007). Summer connects the post-breedi ng period (i.e., when kites tend to disperse) with the onset of the wet season (Bennetts & Kitchens 1997; Martin et al. 2006a), while Fall encompasses a period of occasional late-season breedi ng and the onset of the dry season (Bennetts & Kitchens 2000). Depending on the sex and reproductive stage of a breeding snail kite, a high degree of variability may be observed in its behavior (Sykes et al. 1995). Since we could not collect enough TAB data on actively breeding individuals dur ing different reproduc tive stages to treat such data differently, such variability would have only confounded our analyses; therefore, we did not conduct TABs on individu als that were actively engaged in any stage of the reproductive process, and if any breeding behavior was observe d (e.g., courtship, snail delivery to a mate, nest building, nest defense, provisioning of young) then the TAB was discontinued and the data censored from our analyses. Thus, our inferences are limited to snail kites that are foraging independently and cannot necessarily be extrapolated to kites that are provisioning nestlings or a mate. Observations were made during all hours of th e day, as kites forage throughout the day in Florida (Cary 1985; Sykes 1987a); however, in following with previous studies (e.g., Cary 1985;

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43 Sykes 1987a; Bennetts & Kitchens 2000), we reco rded the exact time of day during which observations took place. We assigned observations to one of th ree daytime categories: morning (sunrise plus three hours), evening (sunset minus three hours), and afte rnoon (everything in between). To avoid potential confounding effects of inclement weather conditions on foraging behaviors, observations were not made during times of rain or heavy wind (Cary 1985; Bennetts & Kitchens 1997). We conducted a combination of one hour (n=713), two hour (n=48), and day-long observations (n=21). The intended duration of the observation was always determined prior to locating a focal individual. Due to unpredicta ble weather events and equipment malfunctions, observations were sometimes stopped prematurely. Observations lasti ng at least eight hours were classified as day-long observations, and th ose less than 30 minutes were censored from our analyses. Classification of individual kites Whenever possible we d etermined the sex and relative age of the focal kite. Snail kites are sexually dichromatic, and mature adults of each sex have very distinct plumage coloration (Sykes et al. 1995). We visually identified the se x of all mature adults by their plumage. However, full adult plumage may not be achieved until the breeding season of the 3rd year (i.e., until 36 months old), and the sex of younger kites, especially those from 6 to 24 months old cannot be reliably distinguished by plumage cues (Bennetts & Kitchens 1999). Fortunately, during annual breeding season popul ation surveys, which were conducted by the Florida Fish and Wildlife Cooperative Research Unit from 1992 to 2007, numerous kites were banded as juveniles and had feather samp les taken for DNA analysis (B ennetts & Kitchens 1997, 1999; Martin 2007). Therefore, the sex of many young bi rds (i.e., all those with specific alphanumeric bands and corresponding DNA results) could be de termined, despite their ambiguous plumage.

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44 To avoid inducing bias by systematically bypassi ng non-banded adults and/or old juveniles of ambiguous sex, we still observed these individuals when they were enc ountered, though birds of unknown sex were censored from all analyses in which sex was a covariate. We also identified the relative age of the focal kite. Kites with mature adult plumage (see above) were classified as adults. The second ag e class, juveniles, also has definitive plumage coloration, although the sexes canno t be distinguished. The distinct buff brown and cinnamon juvenile plumage is retained until about the 5th month of life (Sykes et al. 1995). Juveniles also have dark brownish black eyes, in contrast to the bright red eyes of adults. Eye color begins to change between 4 and 6 months of age, and juve niles become hard to distinguish from second year birds after 6 months of age (personal obser vation). Therefore, birds from 6 to 36 months old are hard to age with visual cues. The equipping of juvenile s with unique alphanumeric bands during the annual population survey s helped us keep track of many birds ages through the period of ambiguous plumage coloration. The ages of all kites that were banded as nestlings could be estimated to the day (+/_ 7 days) (Sykes et al. 1995). We still observed kites of ambiguous age that we encountered during TABs. These kites (along with banded kites of know ag es falling from 6 to 36 months old) were placed into a third category: subadults. Even though th ere is no evidence for a subadult age class in snail kites, as juvenile survival parallels that of adults by the fifth or sixth month of age (Bennetts & Kitchens 1999; Bennetts et al. 1999) and ki tes can begin breeding as young as 9 months of age (Synder et al. 1989), it was useful to define a third age class. Birds w hose age could not be positively identified in the field (i.e., non-banded individuals from 6 to 36 months old) were, at first, categorized as subadults, but upon comparing measures of foraging behavior among the three

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45 age classes we found evidence that justified combining adults and subadults into a single adult age class (see Comparison of Adults, Subadults, and Juveniles ). Collection of behavioral data TAB observations were m ade from an airboa t with the use of 10x binoculars and a 15x60x spotting scope. Two observers were always present so that observation and data recording could occur simultaneously. Using a twenty-four hour digital clock, we recorded the time of day to the second that the focal kite engaged in each of the following behaviors: started flying, captured a snail, stopped flying (i.e., returned to a perch), extracted a snail, finished eating a snail, or dropped a snail. Each flight was categorized as either a fo raging bout or a non-foraging flight. Each foraging bout was categorized as either a course-hunt or perchhunt. These distinctions were determined by telltale behavioral cues (Snyde r & Snyder 1969; Beissinger 1983; Sykes 1987a). Foraging bouts were marked successful if the kite returned to the perch with a snail and consumed it, and they were marked unsuccessful if the kite returned to a perch without a snail or if the snail was dropped at the perch before cons umption. In cases where a snail was dropped in flight but a kite made multiple attempts during a single foraging flight, the bout was considered successful if the kite eventually returned to a perch with a snail and consumed it, and it was marked unsuccessful if it did not. Even if a snail was dropped, it was still tallied as a capture, but the associated foraging bout was classified as unsuccessful unless a successful capture followed by successful consumption occurred. We distinguished between dropped snails and empty shells that were picked up and rejected by visual cues (e.g., the sp lash on the water; the lag time before sinking) (Darby et al. 2007). While the kites were perched, we distinguished between handling time and idle perch time. Handling time was defined as the time that elapse d between arriving at a perch with a snail and

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46 finishing the consumption of a snail. Upon a rriving at a feeding perch, kites usually began negotiating the captured snail immediately and continued through the extraction and consumption process without interruption (Syke s 1987a). Handling time included negotiating, extracting, and consuming a snail. The remaini ng time spent perched (i.e., all but the handling time) was classified as idle perch time. Although kites occasionally preened or engaged in intraand inter-species interactions while perched, the times of these behaviors were not recorded during our observations unless such a behavior interrupted the hand ling time of the focal kite or induced the focal kite to take flight. Missing data is one of the inescapable shortcom ings associated with field observation as a mode of scientific inquiry (Morri son 2001). In all circumstances we did our best to record every behavior to the second, but oftentimes, due to co nditions beyond our control, particular actions were unobservable, resulting in unknown star ting or ending times for given events. Assumptions of behavioral observations A m ajor assumption using data collected during TABs is that our observation actions did not affect the observed individual s behavior. Snail kites are rela tively docile birds and are not easily disturbed by the presence of humans or airboats from dist ances greater than 49 meters (Rodgers & Schwikert 2003; Benne tts et al. 2006). To minimi ze potential disruption to the natural behavior of the birds, we always stayed at least 70 meters away when making observations. It is highly unlikel y that our presence at such a distance altered the focal birds normal behavior, unless an active nest was presen t. In spite of the care taken by observers, on occasion human activities did affect the behavior of the snail kite under observation, resulting in the bird flushing evasively. There were two comm on scenarios in which this occurred. First, when we temporarily lost sight of a bird and were forced to move the boat in order to locate it so that the observation could continue we sometimes approached too cl osely before getting a visual

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47 on the bird. The second scenario occurred when a third party (e.g., fishing boats, tour boats) passed through our observation area unknowingly close to the sn ail kite under observation. Instances of human intrusion, observer induc ed or otherwise, were recorded. Apple Snails Collection and measurement of empty snail shells Af ter the completion of a TAB, empty snail shel ls were collected from the feeding perches that were used by the focal kite. Snail shells often accumulate beneath frequently used feeding perches, and snails consumed by kites can be distinguished from those consumed by other predators (Snyder & Snyder 1969; Collett 1977; Sykes 1987a). Snail shells may also accumulate on and around snail kite ne sts (personal observation). Shells we re also collected from nests as part of the nest monitoring protocol under the systematic annual breeding season population surveys (Bennetts & Kitchens 1997; Martin et al. 2007c). To ensu re that collection dates were representative of consumption dates, only fresh shells, as determined by coloration and smell (Bourne 1993), were collected from feeding perches and nests. A ll collected shells were bagged and labeled with the date, GPS coordinates, wetland, and specif ic place of collection (i.e., feeding perch or nest). We also recorded the TAB during which the shells were collected; however, it is common for some feeding perche s to be used by several individual kites intermittently or with close temporal proximity (S ykes 1987a), so we could not be certain that all of the shells we collected during TABs were from the focal individual. As in Sykes (1987a), the linear widths and lengths (dimensions renamed) of these shells were measured to the nearest millimeter using vernier calipers (Figure 2-2a, 2-2b respectively), and these dimensions were used to estimate the whole wei ght and caloric content of the snails that once occupied the collected shells (see Regression models and estimation of average snail weight and caloric content ).

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48 Collection and processing of live snails Funnel traps were used to collect apple snails on Toho (For details on and a discussion of this collection m ethod see Darby et al. 2001). We sampled from 18 traps that were deployed by an ongoing study (Kitchens et al. 2005, 2007) and from 6 additional traps, which we deployed in an area used heavily by kites (i.e., Grassy Island) that was not covered in the scope of the overlapping study (Figure 2-3). Traps were checked every one to two weeks. All snails caught in the traps located on the east, south, and west shores were identified to species (Rawlings et al. 2007) and tallied. Then, for each trap, individual snails were pooled together by species and the cumulative species weights were recorded to the nearest gram using a spring scale. The snails were subsequently released at the location of capture. Snails caugh t in traps located in Goblets Cove and Grassy Island (Figure 2-3), the two areas of the lake us ed heavily by snail kites (Martin et al. 2007c), were removed, placed in a plastic bags labeled with the date and trap location, and immediately frozen. In the lab, we identified each individual to species, took the dimensional measurements of width and length (see Collection and measurement of empty snail shells ), and measured the whole wet weight (i.e., whole snail wi th shell) to the nearest tenth of a gram using a digital scale. A subset of these frozen snails was randomly selected and the snails removed from their shells. The wet weight of each individuals in ternal body parts (i.e., no shell and no operculum) was recorded. Female snails have an albumen gland, which is the only internal tissue commonly discarded by kites prior to consumption (Snyde r & Snyder 1969; Sykes 1987a). The internal bodies of female snails were weighted with and without their albumen glands, and then the gland was discarded before nutritional analyses were conducted. Three pooled samples (consisting of 20, 30, and 30 snails respectively) were packed on dry ice and shipped overnight to Silliker Laboratory in Chicago, IL for biochemical anal ysis. The samples were homogenized with a

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49 laboratory blender, and then the caloric content was determined as well as the percent moisture, ash, protein, lipid, and ca rbohydrate (Table 2-2). Statistical Analysis and Confidence Intervals All confidence intervals (95% CI) around our derived estim ates were approximated as follows: CI = x +/t /2 SE where x is the sample estimate (i.e., mean), t is the test statistic (evaluated at = 0.05 on n-1 degrees of freedom), and SE is the standard error of the mean. Such estimation is valid for positive sample estimates of count, measurement, or proportion that generally follow a lognormal distribution (Burnha m & Anderson 2002). We also report standard deviations (SD) of some measures for comparison with previous studies. All statistical analyses were performed using the software package R version 2.5.1. Analysis of Foraging Behavior Using the ob servational data that we colle cted, we quantified the following foraging behaviors for each TAB: average searching time, average handling time, drop rate, and capture rate (i.e., consumption rate). These measures we re used to compare the behavioral differences between kites foraging on exotic versus native snails. Most of our conclusions were drawn from observations made on Toho (in which exotic sn ails are prevalent) and on Kiss and WCA3A (neither of which harbors the exotic apple snail) In order to account for small sample sizes, we pooled data from Kiss and WCA3 A for many of our analyses; however, if the measure of comparison differed significantly between Kiss and WCA3A or if small sample sizes were not an issue, then Kiss and WCA3A were treated inde pendently. Data collected in the non-focal wetlands (i.e., WCA3B, WCA2B, ENP, OKEE, GW SJM) were, in many cases, not evenly distributed (e.g., observations were only made during certain seasons or years; unbalanced numbers of observations were made on a particular age group or a particul ar sex), and to avoid

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50 biasing our results via nonrandom effects, these data were excluded from our analyses (unless otherwise noted). Exotic snails were first discovered on Toho in 2001 and were likely predominant on by Toho 2003 (Darby, personal communication ), yet native snails were still relatively abundant in the littoral zone of the lake prior to the comm encement of the drawdown and scraping treatment in late-2003 (Kitchens et al., unpublished data; Darby, personal communication ). Reliable data on the selection of snails by kites during this transitional period (2001-2003) is lacking, but anecdotal evidence suggests that kites were foraging heavily on exotics by 2003: kites were commonly seen course-hunting in deeper regions of the lake (i.e., habitats that do not harbor native snails) and piles of exotic snail shells were observed below kite feeding perches (Darby & Welch, personal communication ). During the pilot year of our study (i.e., 2003), only 23 TABs were conducted, and all of these observations were of juvenile kites, none of which were on Toho. By 2004 kites on Toho were feeding almost exclusively on exotic snai ls (see Results), and although our inferences related to foraging on exotic snails were drawn from TABs made during the period 2004 to 2007, we defined 2003-2007 as th e post-invasion era based on the confirmed presence and likely abundance of the exotic sn ail on Toho during 2003. And in order to increase the sample size of observations made on juvenile kites, we included th e 2003 data for juveniles on Kiss and WCA3A in our analyses concerning the post-invasion era. We also incorporated data from Bennetts and Kitchens (1997, 2000) into some of our analyses. These data were collected in 1993, 1994, and 1996. We used these observations to represent the pre-invasion era. None of the aforementioned foraging behaviors differed significantly among Toho, Kiss, a nd ETOHO during the pre-invasion era, so to increase the sample size, we pooled these data. We then co mpared measures of handling time and drop rate

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51 for adults foraging on Toho during the preand pos t-invasion eras. Data on juveniles from the pre-invasion era was lacking; therefore, we could not compare juvenile foraging behavior before and after the invasion of the exotic snail. Searching and handling time The tim e that elapsed during a foraging bout between takeoff and capture was measured as searching time; however, the calculation of the searching time per foraging bout was contingent on the foraging bout being successful (i.e., it resulte d in the consumption of a snail). Since not all foraging bouts were successful yet kites still expended time and energy searching during these bouts, we calculated the searching time for each successful foraging bout as the cumulative time spent searching between successful bouts. Therefore, for a given successful foraging bout, we took the search time of that bout and added the total flight time of all the previous unsuccessful bouts (i.e., foraging bouts in which no snail wa s captured or in which the snail was dropped before consumption) that had occurred since th e last successful one. Non-foraging flights that occurred between successful foraging bouts were not included in the calculation of searching times. Searching time ( ts) was calculated as follows: ts = tf tr + (tu), where tf is the total flight time of the successful foraging bout, tr is the returning flight time after the capture was made, and (tu) is the sum of the total flight times for all preceding unsuccessful foraging bouts that occurred since the last successful bout. Handling time was measured as elapsed time fr om landing on a perch w ith a snail to the end of consumption. In unusual cases where th e focal kite was interrupted (e.g., by a boat or another bird), we recorded the duration of the alternative behavi or and subtracted it from the actual handling time. Handling time ( th) was calculated as follows: th = TeTp ti, where Te is the time at which the kite finishes eating, Tp is the time at which the kite perches with the snail, and ti is the total time of any interruptions that may have occurred during the handling process.

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52 We only analyzed foraging bouts that had co mplete time documentation. Foraging bouts with missing capture times or handling times were censored from respective analyses. In order to avoid problems associated with pseudoreplica tion that may have resulted from treating every foraging bout independently, we calculated the average searching and handling times for each TAB (i.e., each individual bird) and used th ese averages for comparative analyses. Drop rate We calculated the drop rate for each TAB as th e num ber of snails dropped divided by the total number of snails captured by the focal kite. For this anal ysis, we calculated the total number of snails captured as the sum of droppe d snails and successfully consumed snails. Empty shells that were picked up and rejected we re not included in this calculation. Drop rate ( D ) was calculated as follows: D = sd / (sc + sd), where sd is the number of snails dropped during a TAB and sc is the number of snails consumed during a TAB We also calculated raw drop rates for adults and juveniles on Toho and Kiss-WCA3A so that we could compare our results with those repo rted by Darby et al. (2007). The raw drop rate ( Di) was calculated as follows: Di = Sdi / (Sci + Sdi), where Sdi is the total number of snails dropped during TABs by all individuals in group i and Sci is the total number of snails consumed during TABs by all individuals in group i. Capture rate (i.e., consumption rate) Capture rates were m easured as the number of snails captured and successfully consumed per elapsed time. While snails that were capture d and subsequently dropped were included in the total number of captures used in our calculation of drop rates (see Drop rate ), these mishandled snails were censored from the calculation of captu re rates. We calculated the capture rate for each TAB as the total number of snails successf ully consumed during a TAB divided by the total length of the TAB in seconds (the rate was later converted to snails per hour). If the focal

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53 individual was lost from sight for greater than one minute during the TAB, we subtracted the time it was lost from the total observation time befo re calculating the capture rate. Capture rate ( C ) was calculated as follows: C = sc / to, where sc is the number of snails consumed during a TAB and to is the total length of the TAB in seconds Analysis of Energetics Regression models and estimation of av erage s nail weight and caloric content Using the measurements from the live exotic snai ls that were collected in funnel traps, we ran a linear regression of the tota l snail weight (i.e., shell, oper culum, all soft body parts) as a function of shell length (as done on native snails in Sykes 1987a), which yielded the following equation: total snail weight = (-0.08495465 + 0.01279404 shell length) ^ 2 (p<0.001, adjusted R-squared= 0.78, df= 887) (Figure 2-4a). We used this regression model to derive the original weights of the empty exotic sna il shells that were collected fr om feeding perches after being handled by kites on Toho. We also used the live exotic snails from T oho to run a linear regression of the wet weight of edible soft body parts (i.e., no shell, no operc ulum, no albumen) as a function of shell length (as done on native snails in Sykes 1987a), which yiel ded the following equation: wet weight of soft body parts = (0.8143635 shell length 18.08201) (p<0.001, adjusted R-squared= 0.76, df=362) (Figure 2-4b). Then we converted the wet weights from this model to dry weights by multiplying the output by 0.1215 (i.e., the proportion of dry weight to wet wei ght of edible soft body parts) (Table 2-3). Finally, in order to derive the gross ca loric value of exotic snails consumed by kites on Toho, we multiplied the de rived dry weights by the caloric value of the exotic snail, 3.30 kcal/g (Table 2-2 and 2-3), a nd took the average. Our models and respective coefficients of determination were comp arable to those in Sykes (1987a).

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54 We estimated weights and caloric values fo r the empty native snail shells that were collected during TABs using the followi ng linear models from Sykes (1987a): total snail weight = ( 0.6769 shell length 20.3448) / 0.48, where 0.48 is the proportional weight of wet edible soft body parts to total snail weight; caloric content = (0.6769 shell length 20.3448) 0.145 4.60, where 0.145 is the proportion of dry weight to wet weight of edible soft body parts of the native snail and 4.60 kcal/g is the caloric value of the native a pple (for both models, adjusted R^2= 0.75) (Table 2-3). Profitability Prof itability (i.e., the energy gained from a prey item per unit time invested in obtaining and consuming the prey item) is a common meas ure used to quantify the benefit that an individual receives from a foraging bout. The costs associated with a foraging bout include the time and energy spent searching for and handling a pr ey item. Search time is taken as a function of encounter rate (which itself depends of prey density and dete ction probability), while handling time is a function of prey size a nd shape (i.e., prey type). We derived estimates of profitability for the following four groups: adults feeding on native snails, adults f eeding on exotic snails, juveniles f eeding on native sna ils, and juveniles feeding on exotic snails. We calculated profitability (Pij) as follows: Pij = cj (0.9) / (tsij + thij) where cj is the average caloric content of snail species j, 0.9 is the approximated digestion coefficient for kites (Beissinger 1984), and tsij is the average searching time and thij the average handling time for individuals in age class i feeding on snail species j Daily activity times and energy budgets Daily tim e-activity budgets can be constructed with incomplete observations by using the proportional activity times observe d during TABs to extrapolate the actual time spent conducting each behavior on a daily basis (Pearson 1954; Wo lf et al. 1975; Ashkenazie & Safriel 1979;

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55 Goldstein 1988). We employed this method to estimate average daily time-activity budgets for groups of kites. An obvious assumption is that the activity patterns observed during TABs are representative of normal daily behavior. Previous studies do not report signi ficant differences in behavior patterns among kites obs erved during different times of the day (Beissinger 1983; Cary 1985; Sykes 1987a). We validated this assumpti on with our own data by comparing the relative proportion of time spent flying between day-long and hourly observations. Moreover, capture rates are known to be affected by temperature (Cary 1985), and it has been suggested by some authors (e.g., Cary 1885; Sykes 1987a) that there is a lull in foraging activity during mid-day, although these same authors have also shown that there is no statistically significant difference in capture rates throughout the day (Cary 1885; Sykes 1987a ). Nonetheless, we also validated that the average capture rates achieved by kites did not differ among observations made during the morning, afternoon, and evening (see Validation of time activ ity budget extrapolation ). After validating our assumptions, we calcula ted the relative proportion of time that the following five groups of kites spent flying and pe rched: adults on Toho, adults on Kiss, adults on WCA3A, juveniles on Toho, and juveniles on Kiss-WCA3A. Using these estimates, we derived the average daily energy expenditure ( DEEi) for each of the five groups by employing a series of equations from Koplin et al. (1980) that were ad apted for the snail kite by Beissinger (1984) (see Appendix). Then we estimated the average daily energy gain ( DEGi) for each of the five groups as follows: DEGi = Ci cj (0.9) dl, where Ci is the average capture rate for individuals in group i cj is the average caloric content of snail species j, 0.9 is the approximated digestion coefficient for kites (Beissinger 1984), and dl is the day length. Since we had insufficient data to subdivide each of the aforementioned group by season, we used an overall average day length of 12 hours for our calculations. The annual average day length for WCA3A was 12.16 hours

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56 (SD= 1.25) and for Kiss and Toho was 12.15 hours (SD= 1.12). Finally, we calculated the average daily energy balance ( DEBi) for each of these five groups by subtracting DEEi from DEGi. This is the most common approach in such foraging studies (Goldstein 1988), and variations of this approach have been applied to the snail kite previ ously (Beissinger 1983, 1984; Bourne 1985b; Sykes 1987a). Results Diet We collected 336 piles of e mpty snail shells (r anging from 1 to 121 shells each) from snail kite feeding perches and nests from 2004 to 2007 in the following areas: WCA3A (n=4108), WCA3B (n=532), WCA2B (n=27), ENP (n=379) Okee (n=154), GW (n=61), SJM (n=161), Kiss (n=999), and Toho (n=1486). Toho was the only wetland in which we found shells from the exotic, P. insularum ; however, both native and exotic snails were found on Toho throughout our study. The annual frequencies of each species as they appeared in our funnel traps on Toho appear in Figure 2-5, and the proportions of native to exotic snails captured in each of the five major sampling locations on Toho throughout the study appear in Figure 2-6; however, these years and trap locations do not represent equal sampling efforts, and neither relative species proportions nor trends should be in terpreted from these graphs. During the nearly 790 hours spent observing k ite behavior, we observed kites consuming 2697 snails (360 exotics and 2367 nativ es). We also observed 15 cases of kites eating turtles, all of which involved adult kites on Toho during Ja nuary, February, and November of 2006. We observed one female kite feeding on Marisa sp. shells in WCA3A duri ng late summer of 2004, and several Marisa shells were found mixed in with nati ve apple snail shells below two wellused feeding perches in the same location. Snails of the genus Pomacea accounted for 99.5% of the snail kites diet during our observations.

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57 During our TABs, we also observed five occasions of juvenile kites on Toho consuming vegetation. Although anecdotal, all five instances involved juvenile s that had not been observed eating for over an hour and that may have b een starving. We observed these individuals rummaging around in vegetation near the surface of the water and picking up empty shells and pieces of water lily and hydrilla. There were tw o instances in which the juveniles picked apart and swallowed entire water lily s eed heads. The other three obse rvations involved the juveniles ingesting large chunks of plant material, including hydrilla, water lily stem s, and an unidentified woody twig. After searching the liter ature, we believe that this is the first documentation of kites feeding on vegetation. Nutritional content of apple snails The nutritional content of exo tic snails as reported by Sillik er Laboratories, appears in Figure 2-2. We found that exotic sn ails store significantly fewer ca lories per gram of dry weight than do native snails (3.30 versus 4.52 kcal/g) but that other nutritional m easures for the exotic fall within the range of reported values for the native snail (Figure 2-3). Live exotic snail measurements Live exotic snails (n=890) collected in f unnel traps from Toho from 2005 to 2007 ranged from 11.0 to 73.1 mm in width (x=37.7, SD=12.5) and from 16 to 81 mm in length (x=45.5, SD=10.9). Weights of whole snails (with shell) ranged from 1.1 to 117.9 grams (x=27.1, SD=16.2). However, this sample cannot be consid ered an accurate representation of the pool of available snails for several reasons. Firstly, the physical design of the funnel traps used by Kitchens et al. (2005, 2007) system atically restricted snails ov er 80 mm from entering. In addition to this drawback, snails of different si ze classes also have different probabilities of being captured, escaping, and being eaten by other species while in the trap (Darby et al. 2001; Kitchens et al. 2005, 2007). Noneth eless, 70% of the empty shells collected from kite feeding

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58 perches on Toho fell within the 45.5 to 81.0 mm range (see Empty snail shell dimensions ), and the funnel traps that we used to capture live snails covered this range. Empty snail shell dimensions We found that kites on T oho are eating exo tic snails (n=1486) ranging from 17.0 to 92.1 mm (x=58.2, SD=13.6) in width and from 19.0 to 103.5 mm in length (x=63.5, SD=15.5), with the lengths of 98.5% of the snails falling in the 26.3 to 92.9 mm range and 70% falling in the 45.5 to 81.0 mm range. Empty native snails shells (n=6421) discarded by kites after feeding ranged from 8.50 to 65.8 mm (x=37.6, SD=5.6) in width and from 12.0 to 67.1 mm in length (x=40.2 mm, SD=6.3), with the lengths of 98.5% of the snails falling in the 17.5 to 54.1 mm range, and 70% falling in the 35.5 to 45.7 mm range. We found that the average shell dimensions of native snails did not significan tly differ among wetlands from which adequately sized samples were collected (Figure 2-7), but a significant difference was found between the dimensions of the native shells found in these ar eas and the exotic snail shells collected from Toho (Figure 2-8). Estimates of total snail weight W ith the length measurements from the empt y exotic shells, we predicted the weights (whole snail with shell and operculum) of exotic snails captured by kites on Toho using the following linear model: total snail weight = (-0.0 8465614 + 0.01279172 shell length) ^ 2 Our estimated weights ranged from 2.5 to 153.6 grams, and we found that, on average, kites on Toho are negotiating exotic snails that weigh 56.8 grams (95% CI= 53.9.7). With the length measurements from the empt y native shells, we predicted the weights (whole snail with shell and operculum) of native snails captured by kites using the following linear model from Sykes (1987a): total snail weight = (0.6769 shell length 20.3448) / 0.48 Our predicted native snail weight s ranged from 0.1 to 52.2 grams, and we found that the average

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59 weight of native apple snails negotiated by kites was 15.9 grams (95% CI=15.1.7). Thus, the exotic snails consumed by kites are significan tly heavier (over three times heavier) than the natives (Figure 2-9). Foraging Behavior Comparing adults, subadults, and juveniles We com pared measures of foraging behavior among the three age classes defined by our TAB protocol (i.e., adults, subadul ts, juveniles), and consistent with previous literature, we found no supporting evidence for a subadult age class. Averag e handling times on Kiss-WCA3A were similar for adults (x=72 seconds, 95% CI= 68, n= 259) and subadults (x= 72 seconds, 95% CI= 65, n =65) and their confidence intervals barely in cluded the average handling time for juveniles, 96 seconds (95% CI= 71, n= 22). On Toho, we found that the average handling times for adults (x= 303 seconds, 95% CI= 245, n= 78) and subadults (x= 205 seconds, 95% CI= 109, n= 7) were similar and that they differe d significantly from juveniles (x= 496 seconds, 95% CI= 376, n= 44) (Figure 2-10). Likewise, the average drop rates for adults (on Toho, x= 0.17; on Kiss-WCA3A, x=0.02) were comparable to those for subadults (on Toho, x=0.18; on Ki ss-WCA3A, x= 0.02) and differed significantly from the average drop rates of juveniles (on Toho, x= 0.33; on Kiss-WCA3A, x= 0.06) (Figure 2-11). Average capture rates for adults (on Toho, x= 1.09, 95% CI= 1.05.14, n= 171; on KissWCA3A, x= 3.14, 95% CI= 2.85.43, n= 313) were co mparable to those for subadults (on Toho, x= 1.05, 95% CI= 0.92.17, n= 15; on Ki ss-WCA3A, x= 3.35, 95% CI= 2.66.05, n= 65) and differed significantly from the average capture rates of juven iles (on Toho, x= 0.77, 95% CI= 0.69.85, n= 110; on Kiss-WCA3A, x= 3.4 6 snails/hour, 95% CI= 2.49.44, n= 37) (Figure 2-12). Average searching times were similar among all age classes (Figure 2-13). As a

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60 result of these findings, we recla ssified all subadults as adults a nd used only two age classes (i.e., adults, juveniles) in all of the following analyses. Effects of the exotic snail on average searching times We found no evidence that average searching tim es differ between adult males and adult females (Figure 2-14). After pooling the sexes, we found that during the post-invasion era, the average searching time for adults on T oho (x= 67 seconds, 95% CI= 49, n= 117) was significantly less than the average search times on Kiss (x= 111 seconds, 95% CI= 95, n= 118), and WCA3A (x= 76 seconds, 95% CI= 68, n= 270). The average searching time for adults on Toho during the pre-invasion era (x= 32 seconds, 95% CI= 18, n= 79) was significantly less than th e average searching time on Toho duri ng the post-invasion era (Figure 215). For juveniles, we found that the average sear ching time during the post-invasion era was 61 seconds (95% CI= 43, n= 50) on Toho a nd was 75 seconds (95% CI= 58, n= 46) on Kiss-WCA3A (Figure 2-16). Barring the pre-invasion average from Toho, our results showed a trend opposite from that in Prediction 1 and considering all of the data we found no evidence that the exotic snail aff ects average searching times. Effects of the exotic snail on average handling times We com pared handling times between the adul ts of each sex and found that the average handling times for adult males (on Toho, x=261 seconds, 95% CI= 198, n= 27; on KissWCA3A, x= 71, 95% CI= 65, n= 153) and females (on Toho, x= 319 seconds, 95% CI= 245, n= 56; on Kiss-WCA3A, x= 75, 95% CI = 70, n= 132) were not significantly different (Figure 2-17). T hus, we found no evidence for Prediction 5a ; however, these results did support Prediction 2 After combining males and females we found th at the average handling time for adults on Toho during the post-invasion era was 302 seconds (95% CI= 255, n= 85), while it was 78

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61 seconds (95% CI= 75, n= 130) on Toho during the pre-invasion era and 72 seconds (95% CI= 68, n= 324) on Kiss-WCA3A during the post-invasion era and (Figure 2-18). We found that the average hand ling time for juveniles eati ng exotic snails on Toho was 496 seconds (95% CI= 376, n=44). In contrast we found that the av erage handling time for juveniles eating native snails on Kiss-WCA3A was 96 seconds (95% CI= 71, n=22) (Figure 2-19). Thus, we found that the handling times for e xotic snails were significantly higher for each age class relative to the ha ndling times for native sna ils, which also supported Prediction 2. Additionally, juveniles spent sign ificantly longer than adults ha ndling exotic (but not native) snails, which supported Prediction 4a Effects of the exotic snail on drop rates Under normal foraging conditions (e.g., no heavy wind or rain) on Toho, we observed adults capturing 309 snails and juveniles capturing 199 snails, with raw group drop rates of 20.3% and 42.1% respectively. Under norm al foraging conditions in Kiss-WCA3A, we observed adults capturing 1796 snails and juve niles capturing 386 snails with raw group drop rates of 2.7% and 5.3% respectivel y (Figure 2-20). The raw drop rate we found for juveniles on Toho (i.e., 42.1%) was comparable to the raw dr op rate reported for uni dentified kites on Toho (i.e., 44%) by Darby et al. (2007). These raw group drop rates did not account for individual variation and were inappropriate for our comparisons between/among groups; therefore we used the drop rates from all of the individuals within each group to calculate average group drop rates. Doing so, we found a significant difference between the average drop ra tes of adult males and adult females on Toho. Adult males (x= 0.14, 95% CI= 0.11.17, n= 70) ha d a lower average drop rate than adult females (x= 0.19, 95% CI= 0.15.22, n=109), and this trend was opposite from that which we expected; therefore, we found no support for Prediction 5b On Kiss-WCA3A, we found that the

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62 average drop rates for adult males (x= 0.02, 95% CI= 0.01.03, n= 186) and adult females (x= 0.03, 95% CI= 0.01.04, n= 175) were similar (Figur e 2-21), which, when compared to Toho, lent support to Prediction 3 We found that the average drop rate for juveniles on Toho was 0.33 (95% CI= 0.28.39, n= 110) and that on Kiss-WCA3A it was 0.06 ( 95% CI= 0.03.09, n= 39). After pooling adult males and females, we found the average drop rates for adults were 0.17 (95% CI= 0.15.19, n= 187) on Toho and 0.02 (95% CI= 0.01.03, n= 405) on Kiss-WCA3A (Figure 2-22). Hence, we found that drop rates for adults and juvenile s increased significantly when handling exotic snails, which supported Prediction 3 We also found support for Prediction 4b in that the average drop rate of adults was lower than that of juveniles when foraging on the exotic snail. Effects of the exotic snail on capture rates We found that capture rates did not differ significantly between adult m ales (x= 3.13 snails/hour, 95% CI= 2.75.51, n= 179) and adu lt females (x= 3.18 snails/hour, 95% CI= 2.78 3.58, n= 169) when foraging on native snails from Kiss-WCA3A, but when foraging on exotic snails from Toho their capture rates did di ffer (males, x= 1.27 sn ails/hour, 95% CI= 1.13.41, n= 70; females, x= 0.98. snails/hour, 95% CI= 0.95.00, n= 109) (Figure 2-23). However, we did not feel that the difference between the av erage capture rates of adult males and adult females on Toho justified separating the adult age class by sex before making comparisons with juveniles. First of all, aver age adult capture rates did not di ffer between sexes in Kiss-WCA3A, from which we had a larger sample size. Additionally, capture rates are more sensitive to proximate conditions (e.g., temper ature, season specific behavioral patterns) than are handling times and drop rates, so the variation we observe d could have resulted from such conditions. We found that during the post-invasion era the average capture rate of adult kites was significantly lower on Toho (x = 1.09 snails/hour, 95% CI = 1.04.13, n= 187) than it was on

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63 Kiss (x=3.22 snails/hour, 95% CI= 2.76.69, n= 129) or on WCA3A (x= 3.38 snails/hour, 95% CI= 3.03.73, n= 259). We also found that adult cap ture rates on Toho were lower in the postinvasion era (see above) rela tive to the pre-invasion era (1.95 snails/hour, 95% CI= 1.61.29, n= 65) (Figure 2-24). We found similar trends when comparing the average capture rates of juvenile kites foraging on Toho (x= 0.77 snails/hour, 95% CI= 0.69.85, n= 110) with those on Kiss-WCA3A (x= 3.46 snails/hour, 95% CI= 2.49.44, n= 37) (Figure 2-25). These results provided strong support for Prediction 6 Effects of the Exotic Snail on Energetics Estimates of caloric content W ith the length measurements from the empty exotic shells collected during TABs, we predicted the weights of the edible soft body parts (no shell, no operculum, and no albumen gland) of exotic snails captured by kites on Toho using the following linear model: wet weight of soft body parts = (0.8143635 shell length 18.08201) We converted wet weight to dry weight by multiplying by 0.1215 (the proportion of dr y weight to wet weight of soft body parts), and then we took the product of dry weight and caloric content, 3.30 kcal/g dry weight (Table 23). We found that kites obtai n an average of 12.92 kcal (95% CI= 11.62.27) per exotic apple snail. We predicted the energetic content of the na tive shells collected during TABs using the following linear model from Sykes 1987(a): caloric content = ( 0.6769 shell length 20.3448) 0.145 4.60. We found that kites are obtaining an average of 4.84 kcal (95% CI= 4.60.07) per native snail. Thus, kites obtain significantly more gross energy from e xotic apple snails than from native snails (Figure 2-26).

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64 Profitability Using average searching times and handling tim es for each group of kites and the average caloric content of each snail species, we found th e following profitabilities: adults feeding on the exotic snail, 2.10 kcal/min; adults feeding on th e native, 1.75 kcal/min; ju veniles feeding on the exotic snail, 1.39 kcal/min; juven iles feeding on the native, 1.47 k cal/min. We found that exotic snails are more profitable than natives for adults but that native snails ar e more profitable than exotics for juveniles. Hence, Prediction 7 holds true for juveniles but not for adults. Validation of time activi ty budget extrapolations We found e vidence validating our assumption th at day-long time activity budgets could be extrapolated from shorter obser vations throughout the day. On Toho, the average proportion of time spent flying during day-long (x= 0.108, 95% CI= 0.086.131) and hourly (x= 0.99, 95% CI= 0.077.121) TABs was almost identical. We found a similar level of agreement between day-long (x= 0.148, 95% CI= 0.112.174) and hourly (x= 0.158, 95% CI= 0.131.184) TABs on WCA3A. On Kiss, the average proportion of time spent flying during day-long TABs was 0.158 (95% CI= 0.113.185), while during hourly TABs it was 0.192 (95% CI= 0.164.221); however, the confidence intervals still overlapped (Figure 2-27). We found additional evidence that behavioral patterns observed during hourly observations were representative day-long ac tivity patterns when we compar ed the average capture rates among morning, afternoon, and evening TABs. For this analysis, we combined Kiss and WCA3A because of the similar average capture rates observed for each area (Figure 2-24). On Toho, average capture rates in the morning and evening were higher than in the afternoon, but there was no significant difference between mo rning and evening or between afternoon and evening. On Kiss-WCA3A, we found that averag e capture rates decrea sed throughout the day from morning to afternoon to ev ening, but all confidence interval s overlapped so this trend was

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65 not significant (Figure 2-28). Si nce we found no clear pattern of average capture rates varying significantly among different times of the day, which agreed with the findings presented in the literature (Cary 1985; Sykes 1987a), we felt comfortable with th e assumption that hourly TABs were representative of day-long TABs. Daily activity patterns On average, we found that kites spent 85.6% of their diurnal activity tim e perched and 14.4% flying (SD=14.4), which agrees with the patterns recorded by previous behavioral studies (e.g., Cary 1985; Bennetts and Kitchens 1997) (Tab le 2-4). However, we found that theses proportions differed signif icantly between Toho (flying, x= 0.104, 95% CI= 0.0761.133) compared to Kiss (flying, x= 0.157, 95% CI = 0.124.190) and WCA3A (f lying, x= 0.142, 95% CI= 0.111.174) (Figure 2-29). This made biologi cal sense given that we found no difference in the average search times for Toho, Kiss, and WCA3A but that we did find that capture rates on Toho were much lower that on Kiss-WCA3A (see Effects of the exotic snail on average handling times and Effects of the exotic snail on drop rates ). In addition, we found a trend across areas that juveniles spend more time flying than a dults, but this trend was not significant (Figure 2-30). Daily energy balances Assuming a 12 hour photoperiod, we used the av erage capture rate from each group of kites (see Effects of the exotic snail on capture rates ) to estimate the number of snails captured per day and to derive the gross daily energetic gain from such consumption. We estimated that adults on Toho consumed between 18 snails/d ay (i.e., 157 kcal/da y, 95% CI= 150) and that they consumed 41 snails/day on Kiss (i.e., 174 kcal/day, 95% CI= 168) and between 44 snails/day on WCA3A (i.e., 166 kcal/day, 95% CI= 158). We estimated that juveniles consumed be tween 15 snails/day on Toho (i.e., 110 kcal/day, 95% CI= 105

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66 116) and between 37 snails/day on Kiss-WC A3A (i.e., 178 kcal/day, 95% CI = 170) (Figure 2-31). We found that kites on Toho expended approxi mately 133 kcal/day while those on KissWCA3A expended approximately 138 kcal/day. Ther efore, the net daily en ergy gain for adults on Toho was 24 kcal/day and, while it was 36 on Kiss and 28 on WCA3A. The daily energy balance for juveniles was 40 kcal/day on Kiss-WC A3A, while, on average, there was a net loss of 23 kcal/day for juvenile s on Toho (Figure 2-32). Discussion It is eviden t that the exotic apple snail affects several as pects of snail kite foraging behavior. When compared to the average hand ling times for native snails, handling times for exotic snails are extremely infl ated, with adults and juveniles respectively taking three and five times longer to negotiate, extract, and consume e xotic snails. Kites al so experience elevated drop rates when foraging for exotic snails. On average, adults drop over eight times more exotic snails than native snails (17% versus 2%). Juveniles, already dropping around 6% of the native snails that they handle, experience an average drop rate of 33% when foraging for exotic snails. We found that, on average, kites are consuming na tive apple snails that fall between 35 to 45 mm in length and weigh around 16 gram s, while the exotic snails that they typically consume range from 45 to 81 mm and weigh between 50 to 60 gram s. It is likely that the larger relative proportions of the exotic snail ar e underlying, at least in part, the increased handling times and drop rates. However, we believe that experien ce may affect handling times and drop rates more so than does the relative size/w eight ratio of predat or to prey. While handling times and drop rates increase significantly for a ll groups of kites when foraging on exotic apple snails, there is an obvious discrepancy in the magnitude of the e ffect between adults a nd juveniles, but this discrepancy cannot be attributed to relative size/weight differenc es alone. Adult kites display

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67 reverse sexual dimorphism, with males weighing up to 10% less than females, yet while foraging on exotic snails, adult males maintain significantl y lower drop rates than adult females. This contradicts our expectations that were based on relative weight ra tios alone and suggests that the lack of experience in juvenile s exacerbates the problems they ha ve with negotiating exotic apple snails. While the negative effects of the exotic apple snail on the aforementioned foraging behaviors of the snail kite may seem debilitati ng, they cannot be viewed, substantively, outside the context of energetic ramifications. By dry weight, the edible body tissu e of the exotic snail contains 3.30 kcal/g, while that of the native snail contains 4.60 kca l/g. Interestingly, our results for the exotic snail (i.e., P. insularum ) are similar to those reported for P. canaliculata 3.37 kcal/g (Catalma et al. 1991; PhilRice 2001). Although the edible tissues of exotic apple snails contain less energy per gram, whole exotic snails still contain more total energy than whole native snails, and this is due to the relative diffe rence in their average sizes. The average exotic snail contains 12.92 kcal, while the average native snail contains 4.84 kcal. In light of these findings, other effects of the exotic snail on snail kite foraging be havior seem to make biological sense. Adult kites on Toho achieve an av erage capture rate of 1.09 sna ils/h, which is significantly lower than their average capture rates on Kiss (3 .22 snails/h) and WCA3A (3.38 snails/h). The average capture rates of juveniles display a si milar trend, 0.77 snails/h on Toho and 3.46 snails/h on Kiss-WCA3A. While the difference in capture rates between adults and juveniles on Toho may be a consequence of juveniles increased ne gotiability problems with the exotic snail, the overall trend in capture rates between Toho a nd Kiss-WCA3A is likely due to the fact that similar energetic benefits can be obtained by consum ing fewer exotic snails. The exotic snail has

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68 no direct measurable affect on searching time. Even though the elevated drop rates associated with exotic snails result in kites making mo re attempts before successfully capturing and consuming exotic snails, both adults and juveni les have shorter average searching times per success on Toho than they do on Kiss and WCA3A. Therefore, the trend we observe in capture rates is not a product of increased searching times for exotic snails. Searching time, however, is strongly affected by the abundan ce and availability of apple snails. Even though some of the exotic snails on Toho may be unavailable due to the inability of kites to negotiate them, kites may still maintain short searching times if exotic snails exist in a high enough abundance. Estimates of exotic snail abundance specific to the vegetative communities exploited by foraging kites on Toho preclude our ability to disentangle the relationship between exotic snails searching times, and capture ra tes. If the short searching times on Toho are being maintained by a superab undance of exotic apple snails, then under different conditions (i.e., lower snail densities) the drop rates and presumably longer searching times associated with the exotic snail could translate into much greater energetic expenditures for the snail kite, potentially causing th e profitability of exotic snails to fall below levels that are energetically sustainable for snail kites. El ucidating the affects of exotic snail density on average snail kite searching times and capture rates should receive imme diate attention given that populations of the native apple snail have been declining throughout the kites range (Darby et al. 2005) while populations of th e exotic snail have been spr eading (Rawlings et al. 2007) and that exotic snail densities will likely vary among wetlands. In S outh America, the range of the snail kite ( R. s. sociabilis ) overlaps the natural range of P. insularum (i.e., what we call the exotic snail) but there is no documentation that kites feed on these snails (Sykes et al. 1995; Darby et al. 2007), which pr ovides reason for further concern if one considers the possibility that

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69 in its natural habitat, P. insularum may exist in densities too low to provide a sustainable wforaging base for kites. Our estimates of DEE are comparable to t hose reported in Beissinger (1984). Assuming that the conditions experienced by each group of kites on Toho, Kiss, and WCA3A during the post-invasion era are representative of average conditions and that the se arching times, handling times, capture rates, and activity patterns of each respective group are representative of average responses to such conditions, then adults kites can maintain co mparable daily energy balances whether feeding on the exotic or the native snai l; however, juveniles, while maintaining energy balances similar to those of adults on Ki ss-WCA3A, are on average experiencing net energy losses of around 20 kcal/day when foraging for e xotic snails on Toho. This is concerning given that multiple authors (e.g., Kirkwood 1981; Newt on 1991) have shown that raptors successively failing to meet daily energy requirements may face a high risk of mortality within several days to a week. Our analyses of energy balances are crude and we present the findings here to demonstrate the possible energetic consequences of the effects that the exotic apple snail has on snail kite foraging behavior. We utilized some parameters (e.g., rates of energetic expenditure for flying and food handling; the caloric contents of native apple snails) that were reported in previous, possibly outdated, literature (e.g., Beissinger 1984; Sykes 1987a ). In order to conclude unequivocally that juvenile kites on Toho fail, on average, to meet their daily energetic requirements, we recommend that more advanced methods of energetic analysis be employed. We provide here a valuable star ting point and suggest that serious efforts should be made to apply recent work by Porter et al. (1 994, 2000, 2002, 2003) to the snail kite. Using spatiotemporally explicit models that can account for variations in the local microclimate as well

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70 as in the biophysical aspects of individual snail kites and their pr ey will provide more robust and reliable estimates of daily energy balances.

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71 Table 2-1. Number of TABs conducted by month and location in 2003, 2004, 2005, 2006 (postinvasion) and in 1993, 1994, 1996 (pre-invasion). Spring Summer Fall Era Location JAN FEB MAR APRMAYJUNJULAUGSEP OCT NOVDEC TOHO 4 15 0 2135535153 36 2925 KISS 17 26 8 27231245 2 122 WCA3A 36 53 8 2342412311 38 4654 WCA2B 0 2 0 000219 0 00 WCA3B 0 0 8 000000 0 00 ENP 0 0 0 000000 0 40 OKEE 0 0 0 000000 0 03 SJM 0 0 5 042200 0 03 Pre-invasion Era ETOHO 0 0 0 000000 0 00 TOHO 4 9 0 2942484947 33 2920 KISS 14 22 5 26211144 18 1219 WCA3A 32 36 7 2252033218 32 3841 WCA2B 0 2 0 000119 0 00 WCA3B 0 0 4 000000 0 00 ENP 0 0 0 000000 0 30 OKEE 0 0 0 000000 0 02 SJM 0 0 0 040200 0 03 Post-invasion Era ETOHO 2 2 2 002301 2 07 Table 2-2. Nutritional contents of P. insularum. Nutritional Measurement S1S2S3Average SD Calories 49.536.843.3 43.2 6.35 Calories from Fat 4.233.2 3.47 0.64 Total Fat (g) 0.470.330.35 0.38 0.08 Saturated Fat (g) 0.190.120.13 0.15 0.04 Total Monounsaturated Fat (g) 0.080.090.08 0.08 0.01 Total Polyunsaturated Fat (g) 0.160.10.11 0.12 0.03 Trans Fat (g) 0.010.010.01 0.01 0 Cholesterol (mg) 94.266.855.772.23 19.82 Sodium (mg) 71.697.289.786.17 13.16 Total Carbohydrate (g) 1.50.72.3 1.5 0.8 Protein (g) 9.827.767.74 8.44 1.2 Calcium (mg) 464535681 560 110.64 Iron (mg) 5943.643.348.63 8.98 Moisture (g) 84.218986.786.64 2.4 Ash (g) 3.992.252.89 3.04 0.88 Note: Nutritional content is measured per 100g of wet weight of edible soft body tissues.

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72 Table 2-3. Nutritional composition of P. insularum versus P. paludosa. P. insularumP. paludosaP. paludosa Cattau et al. 2007Beissinger 1984 Sykes 1987a Kcal/g 3.25 +/0.11 4. 52 +/0.25 4.60 +/0.18 % Fat 2.87 +/0.21 3.11 +/0.49 3.4 % Carbohydrate 11.05 +/5.63 NA 27.4 % Protein 63.63 +/6.28 61.5 +/5.2 36.3 % Moisture 13.37 +/2.40 NA 14.5 % Ash 22.48 +/2.50 9.59 +/1.16 32.1 Note: Nutritional Contents of apple snails reported as % dry weight of edible soft body parts, mean +/standard deviation. Table 2-4. Average daily activity patte rn of the snail kite in Florida. Investigator Years Covered by Study Time Perched Time Flying Standard Deviation Cary 1985 1977, 1978, 197985.30%14.70% 15.60% Bennetts and Kitchens 1997 1993, 1994, 199686.10%13.90% 14.00% Cattau et al. 2007 2004, 2005, 2006, 200785.60%14.40% 14.40%

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73 A B Figure 2-1. Relative sizes of exo tic apple snails. A) 50, 63, and 80 mm in length. B) 50 and 80 mm in length.

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74 Figure 2-2. Measurement of snail shells A) Shell width. B) Shell length. Figure 2-3. Location of a pple snail traps on Toho.

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75 A B Figure 2-4. Linear regression models of exotic snail weight and morphology. A) Whole snail weight as a function of shell length. B) We t weight of edible soft body parts as a function of shell length. 20 30 40 50 60 70 80 246810 length (mm)weight^0.5 (g) 20 30 40 50 60 70 80 01020304050 length (mm)weight (g)

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76 2005 2006 2007 Native Apple Snail Exotic Apple Snail yearnumber of snails 0 500 1000 1500 Figure 2-5. Number of native and exotic apple snails captured per year with funnel traps on Toho, 2005-2007.

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77 E.shoreGob.CoveL.grassyS.shoreW.shore native exotic Lake Locationproportioin of snails trapped 0.00.20.40.60.81.0 Figure 2-6. Proportion of native to exotic apple snails captured in five locations on Toho, 20052007.

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78 ENPWCA3AWCA3BWCA2BOKEEGWSJMKISSTOHO 20 40 60 80100LOCATAIONlength (mm) Figure 2-7. Average length of apple snail shells collected from f eeding perches and nests in nine wetlands throughout the range of the sna il kite, 2004-2007. Shaded boxes represent 95% CIs. Lower and upper whiskers repres ent first and third quartiles respectively. Open circles represent extreme outliers.

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79 native shell widthnative shell lengthexotic shell widthexotic shell length 020406080100 linear dimension (mm) Figure 2-8. Average shell dimensions of native a nd exotic apple snails consumed by kites, 20042007. Shaded boxes represent 95% CIs. Lo wer and upper whiskers represent first and third quartiles respectively. Open circles represent extreme outliers.

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80 Native Snails Exotic Snailsweight (g) 010203040506070 Figure 2-9. Estimated whole weight of native an d exotic snails negotiated by foraging kites during the post-invasion era. Er ror bars represent 95% CIs.

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81 TOHO.adultsKISS-WCA3A.adultsTOHO.subadultsKISS-WCA3A.subadultsTOHO.juvenilesKISS-WCA3A.juveniles 0 20040060080010001200LOCATION.agehandling time (seconds) Figure 2-10. Average handling times for adult, juvenile, and subadu lt kites on Toho and KissWCA3A during the post-invasion era. Shad ed boxes represent 95% CIs. Lower and upper whiskers represent first a nd third quartiles respectively.

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82 TOHO.adTOHO.subadTOHO.juvKISS-WCA.adKISS-WCA.subadKISS-WCA.juv LOCATION.ageproportion dropped 0.00.10.20.30.40.5 Figure 2-11. Average drop rates for adult, j uvenile, and subadult kites on Toho and KissWCA3A during the post-invasion era.

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83 TOHO.adultTOHO.subadTOHO.juvKISS-WCA.adultKISS-WCA.subadKISS-WCA.juv LOCATION.agesnails / hour 012345 Figure 2-12. Average capture rates for adult, juvenile, and subadu lt kites on Toho and KissWCA3A during the post-invasion era. Error bars represent 95% CIs.

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84 TOHO.adultsKISS-WCA3A.adultsTOHO.subadultsKISS-WCA3A.subadultsTOHO.juvenilesKISS-WCA3A.juveniles 050100150200250300350LOCATION.agesearchling time (seconds) Figure 2-13. Average searching times for adult, subadult, and juvenile kites on Toho and KissWCA3A during the post-invasion era. Shad ed boxes represent 95% CIs. Lower and upper whiskers represent first a nd third quartiles respectively.

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85 KISS-WCA3A.femaleTOHO.femaleKISS-WCA3A.maleTOHO.male 050100150200250300350LOCATION.sexsearching time (seconds) Figure 2-14. Average searching times for adult male and female kites on Toho and Kiss-WCA3A during the post-invasion era. Shaded boxes represent 95% CIs. Lower and upper whiskers represent first and third quartiles respectively.

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86 Toho (post-invasion)Toho (pre-invasion)WCA3A (post-invasion)Kiss (post-invasion) 050100150200250300350LOCATION (era)searchling time (seconds) Figure 2-15. Average searching times for a dult kites on Kiss and WCA3A during the postinvasion era and on Toho during the preand post-invasion eras. Shaded boxes represent 95% CIs. Lower and upper whiske rs represent first and third quartiles respectively.

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87 TOHO KISS-WCA3A 050100150200250300350LOCATIONsearching time (seconds) Figure 2-16. Average searching times for juve nile kites on Toho and Kiss-WCA3A during the post-invasion era. Shaded boxes represent 95% CIs. Lower and upper whiskers represent first and third quartiles respectively.

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88 KISS-WCA3A.femaleTOHO.femaleKISS-WCA3A.male TOHO.male 020040060080010001200LOCATION.sexhandling time (seconds) Figure 2-17. Average handling times for adult male and female kites on Toho and Kiss-WCA3A during the post-invasion era. Shaded boxes represent 95% CIs. Lower and upper whiskers represent first and third quartiles respectively.

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89 Toho (post-invasion)Toho (pre-invasion)WCA3A-KISS (post-invasion) 0 200 400 600 800LOCATION (era)handling time (seconds) Figure 2-18. Average handling times for adu lt kites on Kiss-WCA3A du ring the post-invasion era and on Toho during the preand post-in vasion eras. Shaded boxes represent 95% CIs. Lower and upper whiskers represent fi rst and third quart iles respectively.

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90 TOHO KISS-WCA3A 020040060080010001200LOCATIONhandling time (seconds) Figure 2-19. Average handling times for juve nile kites on Toho and Kiss-WCA3A during the post-invasion era. Shaded boxes represent 95% CIs. Lower and upper whiskers represent first and third quartiles respectively.

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91 TOHO.adultsTOHO.juvenilesKISS-WCA.adultsKISS-WCA.juveniles) LOCATION.ageproportion dropped 0.00.10.20.30.40.5 Figure 2-20. Raw group drop rates for adult and juvenile kites on Toho and Kiss-WCA3A during the post-invasion era

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92 TOHO.femalesTOHO.malesKISS-WCA.femalesKISS-WCA.males LOCATION.sexproportion dropped 0.00.10.20.30.40.5 Figure 2-21. Average drop rates for adult ma le and female kites on Toho and Kiss-WCA3A during the post-invasion era. Er ror bars represent 95% CIs.

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93 TOHO.juvenilesTOHO.adultsKISS-WCA.juvenilesKISS-WCA.adults LOCATION.ageproportion dropped 0.00.10.20.30.40.5 Figure 2-22. Average drop rates for adult and juvenile kites on Toho and Kiss-WCA3A during the post-invasion era. Error bars represent 95% CIs.

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94 TOHO.malesTOHO.femalesKiss-WCA3A.malesKiss-WCA3A.females LOCATION.sexsnails / hour 012345 Figure 2-23. Average capture rates for adult male and female kites on Toho and Kiss-WCA3A during the post-invasion era. Er ror bars represent 95% CIs.

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95 KISS (post-invasion)TOHO (post-invasion)WCA3A (post-invasion)TOHO (pre-invasion) LOCATION (era)snails / hour 012345 Figure 2-24. Average capture rates for adult ki tes on Kiss and WCA3A du ring the post-invasion era and on Toho during the preand post-invasion eras. Error bars represent 95% CIs.

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96 KISS-WCA3A TOHO LOCATIONsnails / hour 012345 Figure 2-25. Average capture rates for juvenile kites on Toho and Kiss-WCA3A during the postinvasion era. Error bars represent 95% CIs. `

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97 Native Snails Exotic Snailskcal 02468101214 Figure 2-26. Estimated energetic content of nati ve and exotic snails consumed by kites during the post-invasion era. Error bars represent 95% CIs.

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98 TOHO (daylong)TOHO (hourly)KISS (daylong)KISS (hourly)WCA3A (daylong)WCA3A (hourly) LOCATION (length of observation)proportion of time spent flying 0.000.050.100.150.200.250.30 Figure 2-27. Average proportion of time spent fl ying during day-long and hourly observations by kites on Toho, Kiss, and WCA3A in the postinvasion era. Error bars represent 95% CIs.

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99 Toho.morningToho.afternoonToho.eveningKiss-WCA.morningKiss-WCA.afternoonKiss-WCA.eveningLocation.da y time.snails / hour 012345 Figure 2-28. Average capture rates achieved by foraging kites during morning, afternoon, and evening observations on Toho and Kiss-WCA3 A in the post-invasi on era. Error bars represent 95% CIs.

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100 KISS TOHO WCA3A LOCATIONproportion of time spent flying 0.000.050.100.150.200.250.30 Figure 2-29. Average proportion of time spen t flying by all kites on Kiss, Toho, and WCA3A during the post-invasion era. Er ror bars represent 95% CIs.

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101 TOHO.adultKISS.adultWCA3A.adultTOHO.juvKISS-WCA3A.juv LOCATION.ageproportion of time spent flying 0.000.050.100.150.200.250.30 Figure 2-30. Average proportion of time spent flying by adult and juvenile kites on Toho and Kiss-WCA3A during the post-invasion era. Error bars represent 95% CIs.

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102 TOHO.adultsWCA3A.adultsKISS.adultsTOHO.juvenilesKISS-WCA3A.juveniles LOCATION.agekcal 0 50 100 150 200 Figure 2-31. Estimated gross daily energetic gain s for adult and juvenile kites foraging on Toho, Kiss, and WCA3A during the post-invasion era. Error bars represent 95% CIs.

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103 TOHO.adultsWCA3A.adultsKISS.adultsTOHO.juvenilesKISS-WCA3A.juveniles LOCATION.agekcal -40-20 0 20 40 60 Figure 2-32. Estimated daily energy balances fo r adult and juvenile kites on Toho, Kiss, and WCA3A during the post-invasion era. Error bars represent 95% CIs.

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104 CHAPTER 3 DEMOGRAPHIC EFFECTS OF THE EXOTIC APPLE SNAIL ON THE KITE Introduction Com pared to other vertebrate species, birds, especially those that employ active foraging strategies, have high metabolic ra tes and internal temperatures. Physiological conditions of such species can only be maintained through fre quent energy acquisition (Blem 1976; Krebs & Kacelnik 1991; Carey 1996). Many species, such as the snail kite (Beissinger & Snyder 1987; Bennetts & Kitchens 2000), face the risk of starvation daily, as failing to meet minimum energetic thresholds over the course of several days to a week will likely result in mortality (Kirkwood 1981; Newton 1991). Given the vital necessity for kites to maintain proper energy balances, foraging takes precedence over other beha viors. Thus, foraging behavior directly influences the amount of time and energy an indivi dual can afford to allo cate toward growth and reproduction (Krebs & Kacelink 1991; Carey 1996; Newton 1991). As shown, the exotic apple snail has significant effects on the foraging behavior of snail kites, especi ally juveniles (Chapter 2), and thus it is important to determine what demographic consequences the kite population may face as a result. Martin (2007) ur ges that under the current system, adult fertility appears to be crucial [to the viability of the snail kite populat ion], and factors likely to have large effects on adult reproduction and juvenile survival should receive more attention. To determine whether the exotic apple snail constitutes an environmental stressor that has negative demographic consequences for the kite population, we compar e measures of reproduction (including nesting effort, nest success, and nest pr oductivity), along with estimates of survival, from Toho between the preand post-invasion eras.

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105 Reproduction The reprodu ctive activity of snail kites is lim ited, in large part, by the availability of food resources (Nichols et al. 1980; S nyder et al. 1989). The height of the breeding season stretches from March through June, but kites may breed dur ing any month of the year (Beissinger 1988; Snyder et al. 1989; Bennetts & Kitchens 1997). K ites are sensitive to proximate environmental conditions throughout the breeding pr ocess. In most circumstances adult kites attempt to breed at least once a year and, on occasion, have been observed successfully fledging two broods in the same breeding season (Snyder et al. 1989; Benne tts & Kitchens 1992, 1997). On the other hand, kites may forgo nesting completely in years of severe droughts (Sykes 1979b; Nichols et al. 1980; Beissinger 1986; Snyder et al. 1989). Kites may also limit reproductive effort during other times of food scarcity (Beissinger 1986; Bennetts et al. 1988; Snyder et al. 1989). In addition to influencing the decision of whether or not to commence nesting, environmental conditions can also affect kite behavior after nesting ha s begun. Adults may abandon initiated nests or provision fewer young in times of food stress, resu lting in lower nest productivity (Beissinger & Snyder 1987; Sykes 1987b; Bennetts et al. 1988). Nest failures not resulting from parental abandonment are usually the result of either depr edation or structural collapse (Sykes 1979a, 1987b; Bennetts et al. 1988; Snyder et al 1989; Bennetts & Kitchens 1997). Survival Kite survival has been linked to a number of factors, but prey availability plays a critical role, as starvation is thought to be the leading cause of m ort ality (Sykes et al. 1995). After reaching independence, juvenile kites commonly become emaciated, especially when dispersing through unfamiliar territory in which adequate pr ey is not available (Bennetts & Kitchens 1992, 1997; Martin et al. 2007b). Likewise, most a dult mortalities are linked to starvation during severe droughts (Beissinger 1988; Bennetts & Kitchens 1997; Martin et al. 2006a, 2007b).

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106 Demographic evidence suggests that the snail kite populat ion can be divided into two age classes (juveniles and adults) and that after approximately 5 months of age, the survival probability of juveniles effectively parallels th at of adults (Bennetts and Ki tchens 1999; Benne tts et al. 1999; Dreitz et al. 2004; Ma rtin et al. 2006a). Predictions Relativ e to adult kites feeding on native snails, adults feedin g on exotic snails spend more time handling prey, capture fewer snails per day, and achieve lower daily energy balances (Chapter 2). If these behavioral discrepancies translate into significantly less available time or energy for adults on Toho during the post-invasion era, then we would expect to see a decrease in the occurrence and/or success of behaviors requiring additional energy reserves, such as reproduction. It must also be considered that the foraging behaviors of adults and juveniles are differentially affected by the exot ic apple snail (Chapter 2). Gi ven that adult kites maintain positive daily energy balances while feeding on exo tic snails, we assume that the exotic snail does not represent a threat to adu lt survival. The foraging behavior of juvenile kite s, on the other hand, is severely impacted by the exotic snail, and preliminary estimates indicate that juveniles on Toho may not be achieving sustainable daily ener gy balances (Chapter 2). In light of our previous findings, we made the following predictions: Prediction 1. Average annual nesting effort on Toho is significantly lo wer during the postinvasion era than it was duri ng the pre-invasion era. Prediction 2. The average nest success on Toho is significantly lower during the postinvasion than it was during the pre-invasion era. Prediction 3. Nest productivity (i.e., the number of young produced per successful nest) is significantly lower during the post-invasion than it was during the pre-invasion era. Prediction 4. Nest productivity during the post-invasi on era is significantly lower on Toho relative to other wetlands utilized by the snail kite.

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107 Prediction 5. The survival of juveniles hatched on Toho is significantly lower during the post-invasion era relative to the pre-invasion era. Prediction 6. The survival of adult kites that were hatched on Toho does not differ significantly between the preand post-invasion eras. Methods Robust breeding-season population surveys, along with other long-term studies of demography and movement of the snail kite population, began in 1992. Protocols utilized systematic surveys throughout th e range of the kite that c oupled counts with capture-markrecapture methods, including band-resighting and radio-telemetry. A wealth of demographic data (including survival estimates of juvenile s and adults) and moveme nt data have been accumulated. Range-wide snail kite nesting data (including nesting attempts, nest success, and nest productivity) has also been collected in conjunction with the a nnual population monitoring project. However, on Toho, not all data types we re available for all years (see below) (For detailed methods, findings, and discussions se e Bennetts & Kitchens 1992, 1997, 2000; Dreitz et al. 2001, 2004; Martin et al. 2006a, 2006b, 2007a 2007b, 2007c). Yet, comparisons of demographic parameters with spec ific regard to the invasion of Toho by the exotic apple snail have not been made previously. Based on recent demographic trends of the snail kite population and on the invasion history of the of the exotic a pple snail in Florida (Chapter 1), we defined 1992-1998 as the preinvasion era and 2003-2007 as the post-invasion era. These years also corresponded roughly with our preand post-invasion TAB data (C hapter 2). The period 1999 to 2002 corresponds with the severe decline of the snail kite popul ation (Martin et al. 2006a ; Martin 2007) and with the first occurrences of th e exotic snail on Toho (Darby, personal communication ). To avoid confounding effects associated with the populat ion decline and with the unknown relative

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108 proportion of native and exotic sn ails making up the diet of the snail kite on Toho, we defined 1999-2002 as the decline era and treated it independe ntly from the preand post-invasion eras. Nesting Effort and Reproductive Success All nests ob served with at least one egg or one chick dur ing any monitoring visit were classified as initiated (i.e., active). Kites comm only build nests that are never initiated (Chandler & Anderson 1974; Beissinger 1984; Snyder et al 1989; Bennetts & Kitchens 1997); therefore, nests in which no eggs or young were ever observed were censored from our analysis (from here on nests refers to initiated nests only). All nests in which at least one chick was observed to reach potential fledging age, 24 days old (Snyder et al. 1989), were classified as successful, and all chicks that reached at least 24 days old were assumed to fle dge. Nests that failed before fledging young were classified as unsuccessful. Nests of unknown fa te were censored from our analyses. The average fledging age of kites is 28.7 days (Sykes 1987b), but it ranges from 24 to 35 days (Synder 1989). We used this 24-day benchmark to reduce the likelihood of missing fledging events, which would result in the misclassi fication of these nests as failures or as nests of unknown fate. We do not feel that using the 24-day benchmark biased our estimates of nest success or nest productivity, as mortality during th is nesting stage is minimal for kites (Steenhof & Kochert 1982; Bennetts & Kitche ns 1997; Dreitz et al. 2001). Measures commonly used to assess the relative reproductive effort of snail kites in specific wetland units during a given year include 1) the number of breed ing individuals present and 2) the number of nests initiated. The latter is le ss ambiguous and provides a more tangible measure of nesting effort; therefore, we used the average number of nests init iated annually on Toho to compare the preand post-invasion eras (2004 wa s excluded from the post-invasion average). Nest abundance can vary radically between wet and dry year s (Sykes et al. 1995), making it hard to draw definitive conclusions when comparing two eras; therefore, we also looked at the

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109 relative annual abundance of kite nests on Toho in contrast to othe r wetlands. We calculated the relative contribution of Toho to the annual nesting effort of th e entire kite population for the years 1995 to 2007 by dividing the number of nest s on Toho by the total nu mber of nests found range-wide in a given year (no nesting data was available for 1992-1994). Then we developed two trend models to assess whether the contribution of Toho to the overall nesting effort has changed over time. In the firs t trend model we included all ye ars from 1995-2007. In the second trend model we removed anomalous years, in which documented environmental phenomenon may have drastically altered kite behavior. We censored 2001 and 2007, two years in which severe droughts heavily affected the southern portion of the kites range (M artin 2007; Gawlik et al. 2007; FLDEP 2007), causing uncharacteristic nes ting patterns that could have potentially biased Tohos relative contribution high. We also censored 2004, the year in which the managed drawdown and scraping of Toho was completed, as it may have biased Tohos contribution low. In addition to these trend models, we also estimat ed Tohos average contribution to the total kite population nesting effort for the preand post-invasion eras, a nd since droughts and drawdowns have tangible affects on the kite population, no ye ars were censored in th e calculation of these estimates (i.e., 2004, and 2007 were included). Measures commonly used to assess the reprodu ctive success of snail kites include 1) the proportion of successful nests to the total number of initiated nests (i.e., nest success) and 2) the average number of young fledged per successf ul nest (i.e., nest productivity) (Sykes 1987b; Bennetts & Kitchens 1997; Martin 2007). Ne st success may be influenced by innumerable environmental variables (Sykes et al. 1995). Snail kites inhabit a broad and fragmented landscape, in which spatiotemporal heterogeneity cannot be ignored (Benne tts et al. 1998, Martin et al. 2006). Several extraneous variables may affect the fate of a nest, and due to our inability to

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110 account for such environmental st ochasticity across the range of the snail kite, comparing nest success among wetland units over time was not warranted. However, accounting for environmental variation on a local scale was feasible, thus we co mpared the annual nest success on Toho before and after the invasion of the exotic snail. Nest productivity, the number of young produced per successful nest, is less affected by drastic stochastic events (e.g., dr oughts, hurricanes, cold fronts), as these events usually result in complete nest failure, which is not included in the calculation of nest productivity. The number of young that adults can provision is dictated strongly by the av ailability of suitable prey (Beissinger 1990). Thus, when making compar isons among areas or years, more subtle differences in local foraging conditions can be revealed by comparing measures of nest productivity rather than nest su ccess (Murray 2000; Kosciuch et al. 2001). If adult kites cannot sufficiently exploit exotic snails we would expect lower relative pr oductivity in the face of such conditions. We compared the number of young pr oduced per successful nest on Toho before and after the invasion of the exotic sn ail. We also compared the nest productivity of Toho during the post-invasion era with that of all other we tlands (pooled) during the same time period. Reliable nesting data for Toho was availa ble for the period 1995 to 2007; however, no nests of known fate were observed on Toho in 1998, and only one nest of known fate was observed in 1999. Additionally, no nesting occurr ed on Toho in 2004. Therefore, estimates of nest success and nest productivity for 1998 and 2004 were not calculable, and since estimates for 1999 were based on a sample size of one, confidence intervals were not calculable. All 95% confidence intervals (95% CI) around our derived estimates of nesting effort, nest success, and nest productivity were approximated as follows: CI = x +/t /2 SE where x is the sample estimate (i.e., mean), t is the test statistic (evaluated at = 0.05 on n-1 degrees of

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111 freedom), and SE is the standard error of the mean. Such estimation is valid for positive sample estimates of count, measurement, or proportion that generally follow a lognormal distribution (Burnham & Anderson 2002). All st atistical analyses were pe rformed using the software package R version 2.5.1. Survival Mark-resight data and radio telem etry data can each be used to generate reliable estimates of apparent survival for snail k ites, and both can incorporate site -specificity. While inferences from mark-resight data are limited to between-yea r estimates, radio-telemetry can be used to generate betweenand within-year estimates of apparent survival (Bennetts et al. 1999, Martin et al. 2007b). From 1992 to 2006, 121 juveniles on Toho were ma rked with unique alphanumeric bands (an additional 74 juveniles were banded on Toho in 2007 and will be used for future estimates). Using mark-resight data from 1992 to 2007, we estim ated the apparent annua l survival of adult and juvenile snail kites that were hatched on Toho. No juveniles were banded on Toho in 1993 or 1998, so no annual juvenile survival estimates were generated for these years. Additionally, no juvenile survival estimates were generated for 2004 because the kites did not nest the year that the drawdown and scraping treatment was comp leted. In the computation of annual juvenile survival during the post-invasion era, we excluded 2003 because we could not account for drawdown effects, which may have biased our estimate. Annual adult survival, on the other hand, could be generated for all years (1992-2006). No survival estimates were generated for 2007 because this analysis is contingent on the completion of the 2008 population surveys. Radio telemetry allows for the estimation of apparent monthly survival probabilities for kites (Bennetts et al. 1999; Bennetts & Kitchens 1999). Ten juveniles on Toho were equipped with radio transmitters in 1992 and five juveniles were equipped in 1994, so these years were

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112 used to represent the pre-invasion era for juvenile survival. The only tele metry data available for juveniles during the post-invasion era came fr om 2005, in which 14 juveniles on Toho were equipped with radio transmitters (Bennetts and Kitchens 1997, 1999; Martin et al. 2006a). Although working with limited sample sizes, we used this radio-telemetry data to estimate the apparent monthly survival of juveniles hatche d on Toho. Radio transmitters had an average lifespan of over two years; therefore, we also ge nerated estimates of apparent monthly survival for adults that were hatched on Toho. Although kites often disperse to other wetlands after fledgling (Bennetts & Kitchens 1997 ), natal area has a greater aff ect on juvenile survival than dispersal area (Bennetts et al 1999; Martin et al. 2007b). Kite s also express natal philopatry, thus adult survival is also linked with natal loca tion (Martin et al. 2007b). We compared monthly survival estimates for adults and juve niles hatched on Toho between the preand postinvasion eras. To generate annual and monthl y estimates of apparent surv ival, we used program MARK V 4.1 (White & Burnham 1999). We created a suite of a priori survival models based on our predictions (see above) and the existing literature (Bennetts & Kitchens 1997, 1999; Bennetts et al. 1999; Martin et al. 2006a, 2007a, 2007b). In our model sets, we allowed survival and detection probability to vary over time or to be he ld constant. In addition, we tested an age effect (i.e., adult versus juvenile), and we also tested group effects correspondi ng to the preand postinvasion eras. After creating our model sets, we selected the most parsimonious model for apparent annual survival using Akaikes Info rmation Criteria corrected for overdispersion (QAIC) and for apparent monthl y survival using Akaikes Information Criteria corrected for small sample size (AICc) (Burnham & Anderson 2004). The selected models were used generate the respective estimates of annual and monthly survival. All associated standard errors

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113 and confidence intervals were also generate d using program MARK V 4.1 (White & Burnham 1999). Results Reproduction Nesting effort The num ber and distribution of sn ail kite nests varies widely over time (Table 3-1; Figure 1-1), and this may be influenced by innumerable of factors (Sykes et al. 1995); therefore, these results must be interpreted with caution. We found that the av erage number of nests initiated annually on Toho was significantly greater durin g the post-invasion era (excluding 2004, x= 42, 95% CI= 29) than it was in the pre-invasion era (x=17, 95% CI= 9) (Figure 3-2). This ran contrary to Prediction 1 Even though there was a high degree of variation in nesting effort among some years (Figure 3-3), the difference be tween preand post-invasion nesting effort on Toho was significant. We found that Tohos proportional contributi on to the annual population nesting effort ranged from 0.01 to 0.11 during the pre-invasi on era and from 0.17 to 0.81 during the postinvasion era (or including 2004, the year in which no kites nested on Toho due to the drawdown, from 0.00 to 0.81) (Table 3-1; Figure 3-4). Us ing our first trend mode l (including all years 19952007) to assess the relative contribution of T oho to the overall population nesting effort over time, we found a slightly significant positive tr end (p= 0.06) (Figure 3-5). Our second trend model did not include the drought years of 2001 and 2007, in which Tohos relative contribution was 0.74 and 0.81 respectively, nor did it include the drawdown year of 2004, in which Tohos relative contribution was 0.00 (Table 3-1). Th is second trend model showed a significant positive trend in Tohos relative c ontribution (p= 0.04) (Figure 3-6) We also found that Tohos average contribution to the overall nesting effort of the kite popul ation was significantly higher

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114 during the post-invasion era (x= 0.33, 95% CI = 0.30.35) than it was during the pre-invasion era (x= 0.06, 95% CI= 0.05.07) (Figure 3-7). Thus Prediction 1 was false. Nest success and productivity While nest success on T oho varied significantly among some years (Figure 3-8), we did not find any significant difference in the averag e nest success on Toho between preand postinvasion eras (Figure 3-9), which were 0.25 (95% CI= 0.12.38) and 0.39 (95% CI= 0.27.51) respectively. Thus Prediction 2 was false. Similarly, nest productivity on Toho varied significantly among some years (Figure 3-10), but again, we found no difference in the average number of young produced per successful nest between the preand post-invasion eras, wh ich were 2.13 (95% CI= 1.82.43) and 1.73 (95% CI= 1.51.95) respectively (Figure 3-11). Thus Prediction 3 was false. We also found that the average nest productivity during the post-invasi on era did not differ significantly between Toho (see above) and all other wetlands combined (x= 1.50, 95% CI= 1.38.61) (Figure 3-12). Thus Prediction 4 was false. Survival Apparent annual survival The m ost parsimonious model of the apparent annual survival for kites hatched on Toho between 1992 and 2006 was one that allowed detecti on probability to vary by year but that held survival probability constant for all years. This top model did not include an age effect or a preand postinvasion group effect on survival (Tab le 3-2); therefore, th is model did not make biological sense based on all previ ous snail kite literature that shows juvenile survival does, in fact, vary over time and differ from that of adults (Bennetts et al. 1999; Dre itz et al. 2004; Martin et al. 2007b). The second most parsimonious mo del also allowed detect ion probability to vary over time, but it included an age effect on survival and allowed juvenile survival probability to

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115 vary among the three pre-defined eras: preinvasion (1992-1998), decline (1999-2002), and postinvasion (2003-2006). Since this second model differed by less than two QAIC units (Table 32), it was not significantly different from the top model (Burnham & Anderson 2004), and since this second model made more biol ogical sense (i.e., it did not set adult and juvenile survival equal and hold them constant), we used it to ge nerate estimates of apparent annual survival. We found that the annual survival probabil ity for juveniles hatched on Toho during the pre-invasion era was 0.45 (95% CI= 0.27.66) and that during th e post-invasion era it fell to 0.34 (95% CI= 0.11.58) (Figure 3-13). Thus, Prediction 5 was not supported by our estimates of annual juvenile survival. We found that annual adult survival was 0.81 (95% CI= 0.72.88), and our data did not support allowing adult survival to vary among years or eras, which supported Prediction 6 Although our data did not support th e significance of the fourth most parsimonious model, which allowed survival and detection to vary by year (Table 3-2), we decided to generate estimates of apparent annual survival fo r juveniles hatched on Toho using this model simply to demonstrate the inherent va riation within the study system (Figure 3-14). Apparent monthly survival The m ost parsimonious model of apparent m onthly survival for kites hatched on Toho was one that included a preand pos t-invasion group effect on detecti on probability and survival, as well as an age effect on survival (Table 3-3). We found that the monthly survival probability of juveniles hatched on Toho was 0.93 (95% CI = 0.82.97) during the pre-invasion era (represented by 1992 and 1994) and that it fe ll to 0.81 (95% CI = 0.65.90) during the postinvasion era (represented by 2005) (Figure 3-15). We found that the monthly survival probability of adults that were hatched on Toho was 0.92 (95% CI = 0.85.96) during the preinvasion era (represented by 1992-1996) and that it was 0.90 (95% CI = 0.75.97) during the post-invasion era (represented by 2005-2006). T hus, estimates of monthly adult survival for

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116 kites hatched on Toho were nearly identical be tween the preand post-invasion eras, supporting Prediction 6 While there was considerably more va riation between the preand post-invasion era estimates of monthly juvenile survival fr om, we found no significan t difference, as the confidence intervals from these estimates overlapped. Although estimates for both monthly and annual survival for juveniles hatched on Toho duri ng the post-invasion era were lower than those for the pre-invasion era, our da ta did not fully substantiate Prediction 5 Discussion The effects that exotic ap ple snails have on snail kite foraging be havior and energetics may, in turn, result in some negative dem ographic repercussions. Although we found no evidence of decreased adu lt survival, nesting effort, nest su ccess, or nest productivity on Toho during the post-invasion era, we did find some evidence suggesting that juvenile survival has declined; however, our survival estimates for juveniles were based on se verely limited datasets and thus had overlapping confidence intervals. Contrary to our predictions, sn ail kite nesting effort on T oho has not decreased since the invasion of the exotic snail; in fact, our data s uggest an increasing trend in kite nesting effort. The average annual nest abundance on Toho is signi ficantly higher in the post-invasion era than it was in the pre-invasion era (42 versus 17 nests per year). Th e relative contribu tion of Toho to the range-wide nesting effort of the kite popula tion has also increased si gnificantly (from 0.06 to 0.33). Nest abundance and distribu tion are affected by a number of environmental variables. It is possible that the declining ha bitat quality in WCA3A has influe nced snail kite nesting trends on Toho. Droughts also influence snail kite behavior, and in 2007 sout h Florida experienced moderate to severe drought c onditions throughout much of th e peak kite breeding season. Events such as these may be underlying the observed nesting trends on Toho, during the postinvasion era. However, one cannot discount the possibility that the exotic apple snail attracts

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117 kites to Toho. We do not know the behavioral mechanisms by wh ich kites distinguish habitat quality, but it is likely that apple snail density (or some cue of snail a bundance/availability) plays a role. For example, the exotic snail lays highly conspicuous pink egg masses on emergent structures, and these bright pink clusters are prevalent throu ghout Toho. Kites may be attuning to these egg masses, or kites may make more direct assessments of foraging conditions. Future research efforts should focus on teasing out the cues used by kites to assess habitat suitability, and Toho may provide the conditions necessary to test certain hypotheses related to kite attraction and site selection. Nest success and nest productivity remain virtually unchanged on Toho during the postinvasion era relative to th e pre-invasion era. Although statistically insignificant, it is interesting to note that the average number of young produced per successf ul nest on Toho during the post invasion-era is above the range -wide average for this time pe riod. We also found that the survival probabilities of adult kites that were hatched on Toho di d not differ between preand post-invasion eras. These findings, strongly suggest that the exotic snail does not constitute an environmental stressor for adult kites, as their abilities to survive and to successfully provision and fledge young have not been negatively affecte d. On the other hand, the ability of juveniles to sustain themselves after fledging remains tenuous. Our estimates for apparent annual survival of juveniles hatched on Toho are rather inconclusive. The drastically fluctuating annual estim ates and wide confid ence intervals in Figure 3-14 are likely the result of insufficient data coupled with environmental stochasticity. The estimates of apparent annual survival from our best, biologically re levant model provide a slightly clearer picture; this model shows that juveniles hatched on Toho during the postinvasion era experienced lower a nnual survival probabilities than did juveniles hatched on Toho

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118 during the pre-invasion era (0.34 an d 0.45, respectively), but these es timates are still relatively uninformative, as their confidence intervals overlap widely (Figur e 3-13). After the completion of the 2008 population survey, we will have the st atistical power to generate more precise estimates of annual survival for juveniles ha tched on Toho during the post-invasion era, and additional years of band-resight data will continue to increase our precision. Our estimates of apparent monthly survival are somewhat more re vealing. Our findings indicate that juveniles hatche d on Toho during the post-invasion era experience lower monthly survival probabilities than did those ha tched during the pre-invasion era (0.81 and 0.93, respectively), but the confidence intervals of thes e estimates overlap as well. However, unlike the confidence intervals around our estimates of annual juvenile surv ival, the confidence intervals around our estimates of monthly juvenile survival each exclude the mean estimate of the comparative group (Figure 3-15). On average, juvenile survival reaches a leve l comparable to adult survival around 4 months post-fledging (Bennetts & Kitchens 1999). If the monthly survival estimates for the pre-invasion (0.93) and post invasion (0.81) eras are raised to the fourth power (simulating survival to five months of age), they result in drastically diffe rent outcomes, 0.75 and 0.43 respectively. If these same estimates are raised to the eleventh power (simulating survival to one year of age), they result in survival probabilities of 0.45 and 0.10 fo r the preand post-invasion eras, respectively. The simulated annual survival of juveniles on Toho during the pre-invasion era (0.45), which was calculated with monthly survival estimates from telemetry data, is equal to the estimate of apparent annual survival (0.45) th at was generated directly from band-resight data; therefore, we are quite confident in these pr e-invasion estimates. However, for juveniles hatched on Toho during the post-invasion era, we do not find the same agreement between simulated annual

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119 survival (0.10) and the direct estimate of a pparent annual survival (0.34), but more data will likely improve these estimates. While our estima tes of survival for j uvenile kites hatched on Toho may not be statistically significant, this does not negate the possibility that there may be a significant biological difference, as lower juvenile survival du ring the post-invasion era will likely result in a signific ant reduction in recruitment from Toho. In spite of this, juvenile survival varies widely over time, and these s uppositions should be viewed with caution, as our inferences are limited to the years for which we had sufficient data on juvenile survival (1992, 1994, and 2005). The kite population is critically endangered, and its sustainability is extremely sensitive to recruitment. However, at this time, we cannot definitively say whether the exotic apple snail negatively affects juvenile survival. This is regrettable considering that the implementation of management actions that could potentially combat the threat of the exotic snail are likely contingent on such definitive answers. Ther efore, we recommend expediting the quest for suitable answers in relation to juvenile surviv al on Toho during the post-invasion era. Radio transmitters should be deployed on the next cohort of juveniles, and a fastidious radio-telemetry protocol (Bennetts & Kitchens 1997) should be enacted. In addition, conducting two robust population surveys per year (instead of just one) over the course of the next two years would also expedite our ability to generate more reliable post-invasion survival estimates for juveniles hatched on Toho. With suitable assistance, th ese recommendations will generate the data necessary to answer these pressing questions, whic h will lead to a bett er conservation strategy for the snail kite. While management actions focused on the exotic snail may be pending further study, there is no reason for management actions focused on the native apple snail to be delayed or ignored.

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120 CERP mandates that management actions should r estore and maintain a network of snail kite foraging habitats and promote habitat that s upports primary prey (appl e snails) recruitment throughout the South Florida ecosystem (U SFWS 1999; RECOVER 2005). Exotic snail populations are already established in close proximity to several of the other critical wetlands utilized by the snail kite in Fl orida, but the presence of hea lthy native apple snail populations may help buffer any negative effects imposed on the snail kite by the exotic snail. More resources should be invested in monitoring native apple snail populations w ithin the kites range in Florida, as the current spatial scope of such data is limited. Estimating snail densities before and after management actions should also become standard protocol for all projects directly influencing the wetland habitats on which the k ites depend. Such data will be critically important to adaptive management strategies involving the snail kite and other species, especially with the onset of DECOMP (Decomp artmentalization and Sheet Flow Enhancement Project) in WCA3A.

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121 Table 3-1. The number of nests initiated rang e-wide and the contribution from Toho, 1995-2007 1995 1996 1997 1998199920002001200220032004 2005 20062007 BICY 24 8 1 6500000 0 00 ENP 2 5 0 00120000 0 235 ETOHO 1 0 0 0004000 0 11 GW 1 0 2 3030020 15 10 KISS 0 0 1 20004107 9 17 WCA1 1 0 1 14000000 0 10 OKEE 23 34 3 80000510 23 180 SJM 19 16 25 912611821 8 190 TOHO 14 18 37 2381425150 47 3079 WCA2A 8 0 0 0000000 0 00 WCA2B 126 3 22 124000111116 0 20 WCA3A 50 79 246 221701120607840 12 612 WCA3B 2 0 0 35260325 0 171 Total 271 163 338 392951671910413199 114 17495Contribution from Toho 5% 11% 11% 1%3%5%74%24%11%0% 41% 17%81% Table 3-2. Model selection table for apparent annual survival of kites hatched on Toho. Model QAICdelta(QAIC)Parameters Deviance 1 {Phi(Juv(.) Ad(.) P(t) SIN} 444.09 0.00 18 217.12 2 {Phi(Juv( inv grps) Ad(.) P(t) SIN} 446.03 1.94 20 214.17 3 {Phi(Juv(.) Ad(.) P( ~ inv Gps) SIN} 454.5310.44 4 259.04 4 {Phi(Juv(t) Ad(.)) P(.) use SIN} 455.4111.32 30 197.43 5 {Phi(.) P(t) SIN} 470.1326.04 17 245.57 Table 3-3. Model selection table for apparent monthly survival of kites hatched on Toho. Model AICcdelta (AICc)Parameters Deviance 1 {Phi(Pre(12 mon)not=Post(juv(12mon)) P(Pre(.)not=Post(.)) SIN} 338.77 0.00 6 268.28 2 {Phi(Pre(4mon)not=Post(juv(4mon)) P(Pre(.)not=Post(.)) SIN} 340.11 1.34 6 269.62 3 {Phi(Pre(.)not=Post(.)) P(Pre(.)not=Post(t)) SIN} 372.8034.02 39 218.57 4 {Phi(Pre(.)not=Post(.)) P(Pre(.)=Post(.)) SIN} 409.3770.60 3 345.17 5 {Phi(Pre(.)=Post(.)) P(Pre(.) =Post(.)) SIN} 413.9775.19 2 351.82 6 {Phi(Pre(.)not=Post(.)) P(Pre(t)not=Post(.)) SIN} 422.6083.83 45 249.58 7 {Phi(Pre(.)not=Post(.)) P(Pre(t)not=Post(t)) SIN} 483.53144.76 75 191.62

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122 1995199619971998199920002001200220032004200520062007 Toho All Other Wetlands yearnumber of nests 0 100200300400 Figure 3-1. Annual number of snail kite nests initiated range-wide, 1995-2007.

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123 pre-invasion post-invasionnumber of nests 0102030405060 Figure 3-2. Average number of nests initiated annually on To ho during the preand postinvasion eras. Post-invasion estimate excl udes 2004. Error bars represent 95% CIs.

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124 1995199619971998199920002001200220032004200520062007 yearnumber of nests 020406080100 Figure 3-3. Number of nests initiated annually on Toho, 1995-2007.

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125 1995199619971998199920002001200220032004200520062007 yearproportion of nests 0.00.20.40.60.81.0 Figure 3-4. Relative annual contri bution of Toho to the total nesting effort range-wide, 19952007.

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126 199619982000200220042006 0.00.20.40.60.81.0 yearproportion of nests Figure 3-5. Trend model expressi ng the increasing contribution of Toho to the total population nesting effort over time (includes all y ears 1995-2007). Dashed lines represent 95% CIs.

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127 1996 1998 2000 2002 2004 2006 0.00.20.40.60.81.0 yearproportion of nests Figure 3-6. Trend model expressi ng the increasing contribution of Toho to the total population nesting effort over time (excludes dr ought years 2001, 2007 and drawdown year 2004). Dashed lines represent 95% CIs.

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128 pre-invasion post-invasionproportion of nests 0.00.10.20.30.40.5 Figure 3-7. Relative contribution of Toho to the total population ne sting effort during the preand post-invasion eras. Error bars represent 95% CIs.

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129 19951996199719992000200120022003200520062007 yearnest success 0.00.20.40.60.81.0 Figure 3-8. Annual nest success on Toho, 19952007. Error bars represent 95% CIs.

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130 pre-invasion post-invasionnest success 0.00.20.40.60.81.0 Figure 3-9. Average nest success on Toho during th e preand post-invasion eras. Error bars represent 95% CIs.

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131 19951996199719992000200120022003200520062007 yearnumber of young 0.00.51.01.52.02.53.0 Figure 3-10. Annual number of young fledged per successful nest on Toho, 1995-2007. Error bars represent 95% CIs.

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132 pre-invasion post-invasionnumber of young 0.00.51.01.52.02.53.0 Figure 3-11. Average nest productivity on Toho during the preand post-invasion eras. Error bars represent 95% CIs.

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133 All Other Wetlands Tohonumber of young 0.00.51.01.52.02.53.0 Figure 3-12. Average number of young fledged per successful nest on Toho vs. all other wetlands during the post-invasion era. Error bars represent 95% CIs.

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134 pre-invasion post-invasionapparent annual survival 0.00.20.40.60.81.0 Figure 3-13. Apparent annual surv ival of juveniles hatched on Toho during the preand postinvasion eras. Error bars represent 95% CIs.

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135 199219941995199619971999200020012002200320052006apparent annual survival 0.00.20.40.60.81.0 Figure 3-14. Apparent annual survival of juveniles hatched on Toho, 1992-2006. Error bars represent 95% CIs.

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136 pre-invasion post-invasionapparent monthly survival 0.00.20.40.60.81.0 Figure 3-15. Apparent monthly survival of j uveniles hatched on Toho during the preand postinvasion eras. Error bars represent 95% CIs.

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137 CHAPTER 4 IS LAKE TOHOPEKALIGA FUNCTIONING AS AN ECOLOGICAL TRAP FOR THE SNAIL KIT E IN FLORIDA? Introduction Ecological Trap Theory Through geological time, anim al populations adapt to environmental conditions via a number of mechanisms. One form of adaptation manifest in anim al behavior is the cue-response system. Environmental cues elic iting behavioral responses that, on average, result in a positive net fitness outcome can be selected for and in corporated into decisi on making processes through natural selection (Stevens & Krebs 1986, Sih 1987) Ecological traps can be created when 1) rapid changes to an ecosystem result in the decoup ling of habitat selection cues from the positive behavioral responses they once elicited and 2) an animal continues to make settlement decisions based on ingrained environmental cues, but due to the abrupt change in habitat, these decisions now result in negative fitne ss consequences unbeknownst to the individual (Tinbergen 1963, Sherman 1988, Robertson & Hutto 2006). The criteria necessary to demonstrate an ecolo gical trap are as follo ws: 1) individuals of the study population must exhibit preference for one habitat (i.e., the potential tr ap) that is greater than or equal to prefer ences for other available habitats 2) individual fitness (or an appropriate surrogate measure) must differ among available hab itats, and 3) negative fitness consequences must result from settling in the preferred (or equally preferred) habitat trap (Schlaepfer et al. 2002; R obertson & Hutto 2006). A Hypothetical Example For the snail kite population in Florida, an ecological trap would be constituted by the following scenario : Adult kites ar e attracted to Toho due to the abundance and availability of exotic apple snails or by the pl ethora of exotic snail egg masses. Adults, which are larger and

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138 have more foraging experience than juvenile kites, are able to attain ample energy for survival and reproduction in this altere d habitat; therefore, they pref erentially nest on Toho even though other habitats, devoid of the exotic snail, are available. However, juvenile kites, once left to forage on their own, are unable to sufficiently handle and extract the larger exotic snails, and subsequently, they starve. Nevertheless, adult ki tes continue to preferen tially select Toho for nesting because of the lakes plethora of seemingly abundant and available exotic snails. In demographic terms, adult survival is unaffect ed, nest success is hi gh, but recruitment is significantly suppressed due to hi gh juvenile mortality. This s ituation would be catastrophic because, as stated earlier, recruitment is one of the predominant factors limiting the growth and viability of the snail kite populat ion in Florida (Martin 2007). Small populations, especially those inhabi ting landscapes comprised of fragments displaying source-sink dynamics, are particularly vulnerable to extincti on (Pulliam 1988, Pulliam & Danielson 1991). Therefore, from a manageme nt perspective, understanding the effects that the exotic apple snail has on snail kite foragi ng success and consequent demographic parameters has precluded our ability to adaptively manage the system in the kites favor. Recent Nesting History From 1996 to 2003, snail kite reproduction occu rred predominantly in WCA3A. Smaller contributing fragments included ENP, Toho, and Ki ss (Bennetts & Kitchens 1997; Dreitz et al. 2001; Martin et al. 2006b, 2007c). Historically, Toho served as an important refugium for snail kites, particularly when regiona l droughts affected the wetlands throughout the southern portion of their range (Bessinger & Takekawa 1983; Take kawa & Beissinger 1989; Bennetts & Kitchens 1997, Mooij et al. 2002; Martin et al. 2006a), but Toho did not ha rbor significant numbers of nesting kites (Bennetts & Kitchens 1997; Martin et al. 2006b). However, the proportion of

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139 nesting activity on Toho during th e post-invasion era has been disproportionately high when compared to traditional nesting area s (Martin et al. 2007c) (Chapter 3). Lake Tohopekaliga We have shown that the exotic apple snail a ffects the foraging behaviors of snail kites (Chapter 2), with som e demographic consequences (Chapter 3), but to provide evidence that Toho is functioning as an ecologi cal trap for the snail kite in Florida, the following three hypotheses must be supported: Hypothesis 1. Snail kites exhibit a habita t preference for Toho that is greater than or equal to preferences for other availabl e wetland units. Hypothesis 2. Snail kite fitness differs among these wetland units. Hypothesis 3. Negative fitness consequences to the sn ail kite result from selecting Toho. Methods Hypothesis 1 Since we do not have a d etailed understanding of snail kite cognition, it is hard measure habitat preference directly; therefore, we used disproportionate habitat use as a proxy for habitat preference. We compared the average number of nests initiated annua lly on Toho during the preand post-invasion eras. Then we assessed th e trend in Tohos relative contribution to the range-wide snail kite ne sting effort over time (For methods of data collection see Bennetts & Kitchens 1997; Dreitz et al. 2004; Martin et al. 2007, 2007c) (For methods of analysis see Chapter 3). To establish whether snail kites show an equal or greater habitat preference for Toho relative to other wetlands duri ng the post-invasion era, we made among area comparisons of the relative abundance of nest s (and in parallel, of breeding indivi duals) and the relative length of the breeding season. When calculating the average number of nests init iated annually on Toho during the post-invasion era we excluded 2004 b ecause the managed drawdown overlapped with

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140 the kite breeding season (Chapter 1). We tested the following predictions that would provide supporting evidence for Hypothesis 1 : Prediction 1a. Kite nesting effort (as measured by annual nest abundance) on Toho has increased significantly since the intr oduction of the exotic apple snail. Prediction 1b. The relative annual cont ribution of Toho to the to tal number of nests found range-wide has increased significantly since the intr oduction of the exotic apple snail. Prediction 1c. During the post-invasion era, the averag e number of nests initiated annually is significantly high er on Toho relative to other wetlands. Prediction 1d. During the post-invasion era, kite nest ing effort (as measured by the duration of the breeding season) on Toho is longer rela tive to breeding season patterns of other wetlands within the kites range. Hypothesis 2 Using survival, nest success and nest produc tivity (i.e., the num ber of young fledged per successful nest) as surrogate measures of fitne ss, we used evidence from the literature to established whether kites on Toho experience different fitness cons equences than kites utilizing other wetlands within their range. The following predictions would provide supporting evidence for Hypothesis 2 : Prediction 2a. Adult snail kite survival diffe rs significantly among wetland units. Prediction 2b. Juvenile snail kite survival di ffers significantly among wetland units. Prediction 2c. Nest success differs si gnificantly among wetland units. Prediction 2d. The average number of young fledged per successful nest (i.e., nest productivity) differs significantly among wetland units. Hypothesis 3 Using the demographic param eters from Hypothesis 2 as surrogate measures of fitness, we tested the following predictions th at would provide evidence for Hypothesis 3 : Prediction 3a. Adult survival has decreased on Toho si nce the invasion of the exotic apple snail, and during the post-invasion era, adult survival is lower on Toho relative to other wetlands.

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141 Prediction 3b. Juvenile survival has decreased on Toho since the invasion of the exotic apple snail, and during the post-i nvasion era, juvenile survival is lower on Toho relative to other wetlands. Prediction 3c. Nest success has decreased on Toho since the invasion of the exotic snail, and during the post-invasion era, nest success is lower on Toho relative to other wetlands. Prediction 3d. The average number of young fledged per successful (i.e., nest productivity) nest has decreased on Toho since the invasion of the exotic apple snail, and during the postinvasion era, nest productiv ity is lower on Toho relative to other wetlands. Results Hypothesis 1 We found that snail kites do in fact pref erentially select Toho (as m easured by disproportionate use) over other available habitats (Chapter 3). The average annual nesting effort on Toho has increased since the invasion of the exotic apple snail (Figure 3-2, 3-3), which validates Prediction 1a There is also an increasing trend in Tohos relative contribution to the overall nesting effort of the entire kite population (Figure 3-5, 3-6), and Tohos average contribution to the overall nesting effort is si gnificantly greater during the post-invasion era when compared to the pre-invasion era (Figure 3-4, 3-7), validating Prediction 1b We found that during the post-invasion era, a significantly greater number of nests were initiated annually on Toho relative to all other individual wetlands except for WCA3 A; however, the estimate for Toho (x= 42, 95% CI= 29) was higher than that for WCA3A (x= 31, 95% CI= 20), lending support to Prediction 1c. Additionally, from 2005 to 2007, kites on Toho expanded their nesting efforts beyond the typical breeding season (i.e., March to June) (Bennetts & Kitchens 1997; Martin et al. 2006b, 2007c). Kites were actively nesting on Toho from early-March through early-November in 2005, from early-M arch through late-August in 2006, and from early-March through mid-October in 2007. During these years, the vast majority of the nesting activity in other wetlands ceased in May or June, and the last documented nesting activity in any

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142 of these other wetlands was in early-July, late -July, and early-June resp ectively, thus lending support to Prediction 1d Due to the effects of the mana ged drawdown on suitable snail kite nesting and foraging habitat, 2003 and 2004 were not considered, as the drawdown likely cut the breeding season short in 2003 and likely de terred nesting in 2004 (Chapter 1). Hypothesis 2 Barring the occurrence of seve re region wide-droughts, we found no supporting evidence in the litera ture that adult snai l kite survival varies significa ntly over time or that it differs significantly among wetland units (Bennetts & Kitchens 1997; Benne tts et al. 1999; Martin et al.2007b); therefore, Prediction 2a was invalid. On the other ha nd, the literature did support the supposition that juvenile survival varies among wetlands (Bennetts et al 1999; Dreitz et al. 2004), thus validating Prediction 2b. We also found evidence in th e literature that average nest success and average nest productivity vary among wetlands (Snyder et al. 1989; Bennetts & Kitchens 1997; Dreitz et al. 2001), which supports Predictions 2c and 2d. Hypothesis 3 There is no evidence for decreased nesting su ccess (Figure 3-8, 3-9) or nest productivity (Figure 3-10, 3-11) as a result of the exotic apple snail on Toho, and during the post-invasion era, nest success and nest productiv ity are not significantly lower on Toho relative to other wetlands (Figure 4-2, 4-3), thus discrediting P redictions 3c and 3d respectively. There is also no evidence suggesting that the exotic apple sn ail negatively affects adult surviv al (Chapter 3), and adult kites foraging on native versus exotic snails maintain comparable energy bala nces (Figure 2-32). Additionally, adults on Toho during the post-inva sion era acquire suffici ent surplus energy to successfully provision and fledge young (Chapter 3). Therefore, Prediction 3a is null. While conclusive evidence is lacking for the s upposition that apparent survival of juveniles from Toho is lower in the post-invasion era rela tive to the pre-invasion era (Figure 3-13, 3-14, 3-

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143 15), there is indirect evidence ba sed on foraging and energetics that suggests juvenile mortality on Toho during the post-invasion er a should be high. Daily gross energy gains are significantly lower for juveniles on Toho relativ e to juveniles on other wetlands that are foraging on native snails (Figure 2-31), and on average, juvenile s on Toho do not maintain positive daily energy balances (Figure 2-32). While our energetic an alyses do not provide conclusive evidence for Prediction 3b they do give rise to concern. Discussion There is sufficient support for Hypothesis 1 show ing that during the post-invasion era, kites preferentially select (i.e ., disproportionately use) Toho re lative to the other available wetlands within their range (Figure 4-1). However, we cannot say that this disproportionate use is directly attributable to th e presence of the exotic snail on Toho because we did not test alternative hypothese s that may have led to the observed patterns of habitat use (e.g., drought or habit decline in other wetlands may have influenced kites to move to Toho) (Chapter 3). Regardless of the underlying factor(s ) leading to the shift in hab itat use, the annual contribution of Toho to the range-wide kite nesting effort is over 500% greater in the post-invasion era relative to the pre-invasion era, and nests on Toho during the pos t-invasion era account for over 30% of the range-wide nesting effort (Chapter 3). Given the numb er of nesting attempts and the number of young fledged on Toho relative to othe r wetlands (Figure 4-4, 4-5), variations in habitat quality on Toho could have tremendous influence the snail kite population. The extended breeding seasons observed on Toho from 2005-2007 may be more directly attributable to the exotic sna il. Although little is known about the life history strategy of the exotic apple snail (Ramakrishnan 2007, Youens & Burks 2007), the absence of an annual population-wide post-reproductive di e-off in the exotic snail (Ram akrishnan 20007), such as that

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144 observed in the native snail (Dar by et al. 2008), likely allows k ites on Toho to continue their nesting efforts through the late-summer and into the fall. Since kites disproportionately use Toho duri ng the post-invasion era, we continued our tests for an ecological trap. Hypothesis 2 is supported by numerous st udies showing that nest success, nest productivity, and, most applicabl y, juvenile survival vary among wetlands. And, although tentative, there is some support for Hypothesis 3 If the effects of the exotic apple snail had significant negative impacts on adult and juvenile foraging success, then Predictions 3a through 3d may have been true. However, the fo raging success of adults and juveniles is differentially affected, and only juveniles suffer severe energetic reperc ussions (Chapter 2). Therefore, the only support for Hypothesis 3 comes from Prediction 3d but direct evidence of suppressed juvenile survival is lacking (Chapter 3). While esti mates of juvenile survival are lower for the post-invasion era th an the pre-invasion era, they are not significantly different (Figure 3-13, 3-15). Nonetheless, the discrepancies in DEE (Figure 2-32) suggest that Prediction 3d may be true, especially give n that our estimates of juven ile survival were derived from relatively small sample sizes. Adult fertility (and implicitly, recruitment via juvenile survival) is the most influential factor limiting growth in the sn ail kite population. Given that the appropriate criteria for Hypothesis 1 and Hypothesis 2 were met, the validation of Prediction 3d would corroborate the hypothesis that Toho is functioning as an ecological trap for snail kites. But our findings do not provide unequivocal evidence, and with the data presently available, we cannot say whether or not Toho is functioning as an eco logic trap. Continuance of th e snail kite population monitoring protocols coupled with specific radio-telemetry studies of juveniles on Toho should soon provide

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145 us with the data necessary to reassess these cri tical questions and to arrive at more definitive answers (see Discussion in Chapter 2 and Chapter 3). While our current findings should give rise to serious concerns, we believe the drastic management actions that would be necessary to resolve the problems on Toho (if it is an ecological trap) warrant more conclu sive evidence. Even if future studies suggest that the exotic apple snail does not negatively affect juvenile survival, we are in no way advocating that the exotic snail should be allowed to persist in or spread throughout the wetlands of central and southern Florida for the sole benefit of the sna il kite. In fact, quite the opposite is true. Notwithstanding the fact that the exotic snail may have unknown deleterious affects on innumerable species and ecosystem functions, al lowing the exotic snail to persist/spread may jeopardize snail kites in ways not dealt with in this study. Ther efore, we support the eradication of the exotic snail but undersco re the importance of carefully a ssessing all eradication methods, as negative effects on wetland habi tats and native snails must be minimized. Anecdotal evidence suggests that kites are foraging on exotic apple snails (and possibly even attempting to breed) along canals and irrigation ditches that are disc onnected from the major wetlands comprising their historical range. Are kite s attracted to these unconventional areas by the exotic snail? We do not know, but evidence suggests that travelin g through the matrix of unsuitable foraging habitat, which intersperses wetla nd fragments, may lead to incr eased mortality in snail kites (Bennetts & Kitchens 1997, 2000; Martin 2007). Therefore, the exot ic snail may have indirect negative affects on the k ite population even if kites can sustain themselves by feeding on the exotic snail. It is also possi ble that kites are being driven into peripheral habitats out of necessity. Furthermore, we st rongly advocate ecosystem and wate r management strategies that take into account the life history strategy of the native apple snail, as such strategies will likely

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146 provide the greatest benefits for the snail k ite population. Maintena nce and monitoring of suitable native snail populations within wetlands utilized by the k ite are necessary to sustain the snail kite population in Florida (USFWS 1999; RECOVER 2005) and will likely help buffer against any negative influences imposed by the exotic snail.

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147 ENP3A2B3BOKEEGWSJMKISSTOHOnumber of nests 01020304050 Figure 4-1. Average number of ne sts initiated annually on nine we tlands during the post-invasion era. Error bars represent 95% CIs.

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148 ENP(0) 3A(43) 2B(6) 3B(2) OKEE(3) GW(1) SJM(6) KISS(8) TOHO(9)2003nest success 0.0 0.2 0.4 0.6 0.8 1.0 ENP(0) 3A(40) 2B(16) 3B(5) OKEE(10) GW(0) SJM(21) KISS(7) TOHO(0)2004 0.0 0.2 0.4 0.6 0.8 1.0 ENP(0) 3A(11) 2B(0) 3B(0) OKEE(23) GW(15) SJM(8) KISS(9) TOHO(47)2005nest success 0.0 0.2 0.4 0.6 0.8 1.0 ENP(23) 3A(60) 2B(2) 3B(17) OKEE(13) GW(1) SJM(19) KISS(1) TOHO(30)2006 0.0 0.2 0.4 0.6 0.8 1.0 ENP(5) 3A(2) 2B(0) 3B(1) OKEE(0) GW(0) SJM(0) KISS(7) TOHO(79)2007 0.0 0.2 0.4 0.6 0.8 1.0 Figure 4-2. Annual nest success on nine wetlands during the post-invasion era. Error bars represent 95% CIs.

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149 ENP(0) 3A(28) 2B(3) 3B(0) OKEE(1) GW(1) SJM(3) KISS(7) TOHO(6)2003number of young 0.0 0.5 1.0 1.5 2.0 2.5 3.0 ENP(0) 3A(19) 2B(13) 3B(5) OKEE(5) GW(0) SJM(5) KISS(4) TOHO(0)2004 0.0 0.5 1.0 1.5 2.0 2.5 3.0 ENP(0) 3A(0) 2B(0) 3B(0) OKEE(3) GW(1) SJM(2) KISS(1) TOHO(12)2005number of young 0.0 0.5 1.0 1.5 2.0 2.5 3.0 ENP(14) 3A(13) 2B(0) 3B(3) OKEE(9) GW(0) SJM(2) KISS(1) TOHO(12)2006 0.0 0.5 1.0 1.5 2.0 2.5 3.0 ENP(0) 3A(0) 2B(0) 3B(0) OKEE(0) GW(0) SJM(0) KISS(2) TOHO(41)2007 0.0 0.5 1.0 1.5 2.0 2.5 3.0 Figure 4-3. Annual nest productivity on nine wetlands during the post-invasion era. Error bars represent 95% CIs.

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150 ENP(0) 3A(43) 2B(6) 3B(2) OKEE(3) GW(1) SJM(6) KISS(8) TOHO(9)2003number of nests 0 20 40 60 80 ENP(0) 3A(40) 2B(16) 3B(5) OKEE(10) GW(0) SJM(21) KISS(7) TOHO(0)2004 0 20 40 60 80 ENP(0) 3A(11) 2B(0) 3B(0) OKEE(23) GW(15) SJM(8) KISS(9) TOHO(47)2005number of nests 0 20 40 60 80 ENP(23) 3A(60) 2B(2) 3B(17) OKEE(13) GW(1) SJM(19) KISS(1) TOHO(30)2006 0 20 40 60 80 ENP(5) 3A(2) 2B(0) 3B(1) OKEE(0) GW(0) SJM(0) KISS(7) TOHO(79)2007 0 20 40 60 80 Figure 4-4. Total number of nest s initiated annually on nine we tlands during the post-invasion era.

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151 ENP 3A 2B 3B OKEE GW SJM KISS TOHO2003number of young (uncorrected counts) 0 20 40 60 80 ENP 3A 2B 3B OKEE GW SJM KISS TOHO2004 0 20 40 60 80 ENP 3A 2B 3B OKEE GW SJM KISS TOHO2005number of young (uncorrected counts) 0 20 40 60 80 ENP 3A 2B 3B OKEE GW SJM KISS TOHO2006 0 20 40 60 80 ENP 3A 2B 3B OKEE GW SJM KISS TOHO2007 0 20 40 60 80 Figure 4-5. Total number of young fledged per year (uncorrected counts) on nine wetlands during the post-invasion era.

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152 APPENDIX DERIVATION OF DAILY ENERGY EXPENDITURE The following series of equations com es from Koplin et al. (1980):

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163 Welch, Z. C. 2004. Littoral vegetation of Lake T ohopekaliga: community description prior to a large-scale fisheries habitat-enhancement pr oject. MS thesis. University of Florida, Gainesville. Werner, E.E. and D.J. Hall. 1974. Optimal foraging and the size selection of prey by bluegill sunfish. Ecology 55: 1042-1052. White, G.C. and K.P. Burnham. 1999. Program MARK: Survival rate estimation from both live and dead encounters. Bird Study 46 (Supplement.):S120-S139. Williams, G.E., J.W. Koebel, D.H. Anderson, S.G. Bousquin, D.J. Colangelo, J.L.Glenn, B.L. Jones, C. Carlson, L. Carnal, and J. Jorg e. 2005. South Florida E nvironmental Report. Chapter 11: Kissimmee River Restoration a nd Upper Basin Initiatives. South Florida Water Management District, West Palm Beach, FL. Wolf, L.L., F.R. Hainsworth, and F.B. Gill. 1975. Foraging efficiencies and time budgets in nectar-feeding birds. Ecology 56(1): 117-128. Woodin, M.C and C.D. Woodin. 1981. Everglade Kite predation on a soft-she lled turtle. Florida Field Naturalist 9: 64. Youens, K.A. and R.L Burks. 2007. Comparing applesna ils with oranges: the need to standardize measuring techniques when studying Pomacea Aquatic Ecology Online: DOI 10.1007/s10452-007-9140-0 (accessed 11/10/07).

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164 BIOGRAPHICAL SKETCH Christopher Cattau earned a B.S. in ecology a nd evolutionary biology from the University of Tennessee, Knoxville. Then he moved to Florida.


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