1. science
2. biology
3. breeding

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Characterization and manipulation of cooperative breeding in the Brown-headed Nuthatch
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Jordan Victoria Smith
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Jordan Smith
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Page 2 of 2
Project Summary.
This project will contribute to a fundamental understanding of cooperative breeding. It
entails long-term social and genetic characterization of Brown-headed Nuthatch (Sitta
pusilla) breeding behavior in two populations along the Florida-Georgia border. Unlike
previous studies, this project will allow for comparison of breeding system dynamics
between populations while controlling for geographic and genetic variation. Studies
which contribute to an understanding of the breadth of both intra and interspecific
variation among cooperative breeders, such as this, are instrumental for resolution of
broad evolutionary patterns in this rare breeding system.
In addition, this project addresses a specific question in a basic aspect of cooperative
breeding, delayed dispersal. Delayed dispersal is considered a precondition for helping by
establishing group living however, this aspect of cooperative breeding remains
controversial. This project will directly test the ecological constraints hypothesis to
provide insight into factors promoting delayed dispersal. Uncovering the evolutionary
origin of cooperative breeding will provide further insight into the paradox of helping
behavior.
A broader impact of this study is aimed at community outreach in a child friendly format.
The project will be featured on the Tall Timbers Research Station webpage, a
collaborating institution in the study. The website will feature project progress including
field and lab notes, photographs, and general life history of the Brown-head Nuthatch.
Also, the project will be represented in the annual open-house for the field station for the
duration of project. The ultimate goal is to: 1) inform the public of the scientific process
and 2) stimulate interest in science and environmental awareness. The Brown-headed
Nuthatch is a threaten species and its conservation status will be featured to stimulate
interest from the public.
This study will also impact the young members of the scientific community through
undergraduate mentoring. Characterization of the breeding system will require intense
field work such as nest searching and mist-netting, and intense lab work such as DNA
extractions and PCR. Both of these stages of research require assistance from multiple
undergraduate researchers. Dedicated students will have the opportunity to conduct
independent projects and will be expected to participate in an undergraduate research
symposium providing either a poster or oral presentation.
TABLE OF CONTENTS
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NSF Form 1359 (10/99)
46
Characterization and manipulation of cooperative breeding in the Brown-headed Nuthatch
Jordan Smith
Background.
Cooperative breeding (CB) is rare in birds and is found in only three percent of the
world’s avifauna1. It is most frequently observed in kin groups when offspring remain on
the natal territory and assist with nestling care1,2. This helping behavior at the nest is an
evolutionary paradox that has fascinated scientists for decades2. Why do individuals
remain on home territory and forgo personal reproduction thereby decreasing their fitness
2
(see 1 for a full review)? Theory on the evolutionary origin and maintenance of this
peculiar breeding system is centered on kin-based cooperative breeders.
There are two aspects of CB which are traditionally considered important
to a fundamental understanding of the breeding system: delayed dispersal
and helping1,2, 3,4. Delayed dispersal is important because it is regarded
as a precondition to helping by establishing group living at the natal
territory3,4,5,6. In a stepwise process, helping behavior is motivated by
incest avoidance and kin selection once group living is established1.
However, recent studies have revealed a large variation in helping
behavior which conflicts with our fundamental understanding of this
breeding system. For instance, recent molecular studies on the western
bluebird revealed that this species tolerates non-kin help. Not only does
this bird disperse from the natal territory, but it is also inclined to help at
the nest by a factor other then kin-selection. As these divergent
cooperative breeders increase in prominence, it raises doubt as to whether
the initial assumptions in the fundamental theory of CB are properly
founded. The purpose of this study is to reevaluate these fundamental
assumptions in a traditional kin-based species, the Brown-headed
Nuthatch.
The primary objective of this study is thorough social and genetic characterization of a
traditional cooperative breeder to evaluate assumptions in helping behavior. The second
objective of the study is to manipulate dispersal behavior and evaluate the effects on CB
frequency.
The Study System.
The Brown-headed Nuthatch (BHNU) provides an ideal study system to investigate
cooperative breeding. They are small, ten-gram cavity-nesting birds found in mature pine
forests of the Southeastern US and Bahamas12. Small territory sizes (~200m)12 make
them particularly easy to study. Characterization of cooperative breeding systems
requires thorough knowledge of family groups, and in the field this requires extensive
nest searching and monitoring. High population densities in small areas are therefore
advantageous; not only is there a manageable area for fieldwork but still a useful sample
size. Many of the most well studied North American cooperative breeders such as the
Red-cockaded Woodpecker 13 and the Acorn Woodpecker 14 have considerably larger
territories, generally at least ten times the size of those of BHNU.
The cooperative breeding behavior of BHNU has only been preliminarily evaluated. The
majority of work has been social characterization via field observations concentrated on a
north Florida population at Tall Timbers Research Station (TTRS; 30.66 N, 84.22 W).
Based on limited field data, the species appears to be a kin-based helping system with
30% of the family groups cooperating12,16. Males remain on the natal territory as
expected in a traditional system and are active in nesting care by provisioning nestlings
with food (although other duties are suspected)12,16. Understanding social helper duties is
important for evaluating the cost and benefits influencing helping behavior; however,
social characterization may not reflect the underlying genetic motivations of CB.
Much of the controversy about the fundamental concepts of CB stems from recent
molecular studies that are beginning to uncover sneaky helping behavior. Of most
importance is revealing helper reproductive contributions that may be hard to detect
socially. In particular, BHNU have some odd behavior that may result in discordant
genetic and social characterization. A high frequency of incest has been reported in the
TTRS population12,16 (although it has not been confirmed genetically) and contradicts a
primary assumption that kin-based helpers are nonbreeding1. Additional data needs to be
collected on paternity to reveal the possibility of helper contributions to extra-pair
fertilization and effective population size. Genetic assessment of helper reproductive
contributions will be essential for reevaluating theory on CB behavior.
For this particular study two populations along the Florida-Georgia border will be
investigated. One is the TTRS population and the second is the Wade Track population
(WT; 30.45 N, 84.00 W) that has not been monitored previously. These populations were
selected because: 1) they control for interspecific life-history characteristics such as a
long life expectancy and limited dispersal capabilities thought to encourage CB; and 2)
they are in close proximity to each other which controls for geographic and genetic
variation. Previous multi-population studies do not control for these factors making it
difficult to draw strong comparative conclucsions2,6,7. Understanding intraspecific
variation is critical for resolving broad evolutionary patterns.
In addition, preliminary research has identified a possible mechanism for delayed
dispersal at TTRS. In this population, predation at the nest during incubation and
brooding is creating a male biased adult sex skew resulting from lower female survival.
This shortage of females limits breeding opportunities and may be constraining the
population such that group living is established. Therefore, mate availability, an
ecological constraint, is the proposed mechanism for delayed dispersal. Interestingly,
preliminary genetic data suggest that there may be a sex skew in offspring production
such that more females are produced then males.
There are three main hypotheses which account for mechanisms delaying dispersal. The
predominant explanation is the ecological constraint hypothesis proposed by Emlen in
19828, that suggests ecological factors such as mate shortage, habitat saturation, or
unstable environments restrict breeding opportunities and lead to delayed dispersal. This
hypothesis is controversial because it neglects to address how species under similar
ecological constraints can be noncooperators. The broad constraints hypothesis,
summarized by Hatchwell9, integrates the ecological constraint and a life-history
hypothesis summarized by Russell 198910. It suggests that in addition to ecological
constraints, factors such as long life expectancy and restricted dispersal capabilities result
in delayed dispersal. It is important to note that not all cooperative breeders exhibit these
characters4. A final hypothesis, the benefits-of-philopatry, incorporates assessment of the
costs and benefits of staying at home rather then just evaluating the cost of leaving in
dispersal decisions11. This study will directly test the ecological constraints hypothesis.
Project Description.
Objective One: Characterization of the Brown-headed Nuthatch breeding system
Social and genetic characterization of both populations
Field Methods:
Nest searching is required to associate adults with particular nest sites and to determine
the breeding status (helper, breeder, nonbreeder). Nests will be monitored to document
social helper duties at the nest and to establish territory size and quality. Adult birds will
be mist-netted prior to the breeding season as well as target netted at nest sites during
active nesting. All captured adults will be fitted with leg bands for identification,
standard morphological characters will be measured, and either feather or blood samples
(depending on the condition and age of the bird) will be taken for DNA extraction.
Nestlings will be sampled for DNA (blood or feather) as soon as possible and fitted with
an aluminum ID band.
Molecular Methods:
All molecular procedures will be conducted in Dr. Rebecca Kimball’s lab at the
University of Florida. To start, genomic DNA will be extracted from all field samples
using the PUREGENE® DNA Purification Kit. Fifteen microsatellite markers will be
PCR amplified from each individual bird. Haas et al. (in review)16 developed five
markers and ten are currently under optimization. Genotyping of the amplified loci will
be conducted on an ABI PrismTM 3100-Avant genetic analyzer. The resulting allelic
data will be analyzed in the program GeneMarker® v.1.5. The program GenePat v4.1
will be used to assess paternity, the central component of the genetic characterization.
Subsequent analyses to estimate the frequency of inbreeding, frequency of extra-pair
fertilization, dispersal distance, effective population size, and sex skews will be analyzed
according to the methods of Seddon et al 18. The sex of each individual is determined
using the molecular sexing method described by Fridolfsson and Ellegren19.
This study will be done in collaboration with James Cox, research biologist at TTRS. He
has provided DNA samples from the 2006 and 2007 breeding season which are currently
being analyzed in the Kimball lab12. Four undergraduate researchers have been involved
in sex identification and assistance with microsatellite optimization. Haas et al. (in
review) 16 collected preliminary data on the genetic structure of the TTRS population.
To test the assumption that this study
controls for large genetic differences
between the two populations, the
degree of gene flow between TTRS
and the WT will be assessed as in
Hass et al16. Earlier study of gene
flow between TTRS and the Pebble
Hill population16 suggests that genetic
differences between the TTRS and the
WT will be minor.
Objective 2: Manipulation of dispersal behavior
The goal of this manipulation is to directly test the ecological constraints hypothesis for
delayed dispersal and to evaluate the subsequent effects of this manipulation on the
frequency of CB.
Experimental design at TTRS:
The ecological constraint of this population is sex skew favoring males;12 therefore,
males will be removed from pair bonds. The number of males removed will depend on
the frequency of cooperative breeders but should provide the opportunity for all helpers
to disperse. Removals will be controlled for territory quality and proximity to
cooperative groups taking into consideration average male dispersal distance. Also
removals will be conducted as soon as possible following pair formation to control for
nest site availability. It is important to control for both territory quality and nest site
availability because these have been identified as potential ecological constraints among
other cooperative breeders.
The methodology for removals will be similar to that of Pruett-Jones and Lewis20, a study
on superb fairy-wrens. Translocated males will be held for three weeks in aviaries at the
University of Florida before release into a neighboring population. This should
maximize the time for helpers to fill territory vacancies while minimizing the effects of
captivity on translocated males. There is a possibility that the translocated males will
return to their original territories upon release from captivity however, this will not affect
the initial measure on dispersal behavior. In the fairy-wren study, vacancies were filled
within a day or two of the initial removals20. Female condition will not be affected by
male removals. The results of this manipulation will be statistically analyzed by a t-test
or an ANCOVA analysis
Predictions:
1) If fewer cooperative groups are found as expected from a release on dispersal
restrictions, this study provides support that dispersal behavior does influence CB.
Furthermore, it provides strong support that ecological constraints drive delayed
dispersal. Although this study does not directly test the broad constraints hypothesis, it
cannot be ruled out based on this result. In further experiments, it will be important to
evaluate this hypothesis in BHNU populations influenced by an ecological constraint
other then sex skew.
2) If no change or an increase in the number of cooperators are found and therefore no
change or an increase in the frequency of delayed dispersal is observed then, this study
provides strong evidence against the influence of ecological factors on delayed dispersal.
Some other non-ecological factor may be influencing delayed dispersal but the effects of
delayed dispersal on CB behavior remain unresolved.
Estimated Timeline:
Preliminary Data: Genetic characterization of TTRS- 2006/2007/2008
Year one:
Complete collection at TTRS and begin WT collection
Year two:
Manipulation at TTRS, continue collection at WT
Year three:
Post hoc collection at TTRS, final collection at WT
Conclusion.
This project will contribute to a fundamental understanding of CB in several ways. First,
manipulation of delayed dispersal and examination of the effects on CB levels directly
test assumptions of the evolutionary influence of delayed dispersal on CB behavior.
Second, a thorough social and genetic characterization of the kin-based cooperative
breeder BHNU, allows reevaluation of assumptions regarding the evolutionary origin and
maintenance of helping behavior in this species. Also, comparisons between the two
populations provide initial estimates of intraspecific variation that are essential for
isolating broader evolutionary patterns. Understanding the evolutionary context of
cooperative breeding is important because helping behavior contradicts current
evolutionary theory in which organisms intrinsically behave as to increase their personal
fitness.
Additional Significance.
Many cooperative breeders are characterized by limited dispersal capabilities and habitat
specialization12,16,21 and both qualities exaggerate sensitivity to habitat destruction21. If a
general mechanism for delayed dispersal is identified, it could potentially lend
contributions to management across a wide variety of cooperative breeders. Specifically,
if dispersal can be increased, effective population size can be increased as well.
In addition, this study has important conservation implications for the Brown-headed
Nuthatch. This species is experiencing range contraction and steady population declines
throughout its natural habitat12,16. In fact, a small Bahamian population is nearly extinct12.
Data from this project is intended for application towards informed conservation
strategies. Beside the identification of dispersal mechanisms, breeding characteristics
such as average dispersal distance, territory size, and the frequency of inbreeding and
extra-pair fertilization can be utilized in management schemes of north Florida and south
Georgia populations and at minimum can serve as a reference for management of
populations over the entire range.
Also, the microsatellite markers developed for this project are applicable in the study of
any Brown-headed Nuthatch population (possibly close relatives too) and will be
accessible in GenBank. Aside from the obvious conservation importance, as stated
previously, this study species is uniquely conducive to controlled yet natural experiments
and should be encouraged for further study both manipulative and comparative in
manner.
Broader Impacts.
This study involves intensive field and laboratory work and requires the assistance of
multiple undergraduate researchers. Technicians will learn field skills such as nest
searching, mist-netting, and bird handling. In the lab, technicians will assist with all
stages of genotyping such as DNA extractions, PCR, and subsequent analyses. Dedicated
technicians will be encouraged to assist with publications and poster presentations. As
mentioned earlier, four undergraduates are already involved in the project and have
performed all sex determining PCR analyses and have assisted with microsatellite primer
optimization.
In addition to providing mentoring to young scientist, this project also provides
opportunity for outreach to the general public. TTRS has an annual open-house aimed to
stimulate interest in science and environmental awareness. This is a perfect opportunity
to engage the public about my specific research and will also provide a glimpse into the
scientific process as they can monitor the progress of the project year to year, starting
with the project design through the final analyses. The target audience of my
presentation will be children and teenagers who usually have minor, if any involvement
in the scientific community. Also, a webpage detailing project progress and notes will be
linked to the TTRS website which can reach beyond just the local community. This
webpage will hopefully foster communication and data sharing between other nuthatch
studies and even among other cooperative breeding studies as well.
Finally, this project will facilitate interactions between the University of Florida and Tall
Timbers Research Station. TTRS has a rich history of fire ecology, game management,
and forestry research and manages over 100,000 acres of land. This institution may prove
a valuable resource to the university as a potential field station and data supply.
Literature Cited
1. Dickinson JL and Hatchwell BJ (2004) Fitness consequences of helping.
In Koenig WD, Dickinson JL [eds.], Ecology and evolution of cooperative
breeding birds. Cambridge University Press.
2. Hatchwell BJ (2007) Avian Reproduction: Role of Ecology in the Evolution of
Cooperative Breeding. Current Biology, 17 (19): 845-847
3.Ligon JD and Burt DB (2004) Evolutionary Origins. In Koenig WD, Dickinson JL
[eds.], Ecology and evolution of cooperative breeding birds. Cambridge
University Press.
4. Ekman J, Hatchwell BJ, Dickinson JL, Griesser M (2004) Delayed Dispersal.
In Koenig WD, Dickinson JL [eds.], Ecology and evolution of cooperative
breeding birds. Cambridge University Press.
5. Koenig WD, Pitelka FA, Carmne WJ, Mumme RL, Stanback MT (1992) The
Evolution of Delayed Dispersal in Cooperative Breeders. The Quarterly Review
of Biology. 67 (2): 111-150.
6. Baglione, V, Marcos JM, Canesrari D, Griesser M, Andreotti G, Bardini C, and
Boglani G (2005) Does year-round territoriality rather then habitat saturation
explain delayed natal dispersal and cooperative breeding in the carrion crow?
Journal of Animal Ecology 75: 842-851
7. Doerr ED and Doerr VJ (2006) Compartive demography of treecreepers: evaluating
hypotheses for the evolution and maintenance of cooperative breeding. Animal
Behavior 72: 147-159
8. Emlen ST (1982) The evolution of helping. I. An ecological constraints model.
American Naturalist, 119; 29-39.
9. Russell EM (1989) Cooperative breeding: a Gondwanan perspective. Emu, 89: 61-62.
10. Hatchwell BJ, Komdeur J (2000) Ecological constraints, life history traits and the
evolution of cooperative breeding. Animal Behaviour, 59:1079-1086.
11. Stacey PB, Ligon JD (1991) The benefits-of-philopatry hypothesis for the evolution
of cooperative breeding: variation in territory quality and group size effects.
American Naturalist, 137, 831-846.
12. Cox JA, Slater GL (2007) Cooperative breeding in the Brown-headed Nuthatch. The
Wilson Journal of Ornithology, 119:1-8.
13.Walters JR, Copeyon CK, and Carter JH (1992) Test of the ecological basis of
cooperative breeding in red-cockaded woodpeckers. Auk, 109, 90–97.
14. MacRoberts MH and MacRoberts BR (1976) Social organization and behavior of the
acorn woodpecker in central coastal California. Ornithological Monographs,21
15. Komdeur J (1992) Importance of habitat saturation and territory quality for evolution
of cooperative breeding in Seychelles warbler. Nature, 358: 493–495.
16. Haas SE, Cox JA, Kimball RT, and Smith JV. Spatial genetic structure in the
cooperative-breeding Brown-headed Nuthatch (Sitta pusilla). In review
17. West DC, Doyle TW, Tharp ML, Beauchamp JJ, Platt WJ, and Downing, DJ (1993)
Recent growth increases in old-growth longleaf pine. Canadian Journal of Forest
Research 23: 846-853.
18. Seddon N, Amos W, Adcock G, Johnson K, Krasijeveld F, Kraddijeveld-Smit JL,
Senapathi GD, Mulder RA, Tobias JA (2005) Mating system, philopatry, and
patterns of kinship in the cooperatively breeding subdesert mesit Monias benschi.
Molecular Ecology, 14:11 3573-3583
19. Fridolfsson AK, Ellegren H (1999) A simple and universal method for
molecularsexing of non-ratite birds. Journal of Avian Biology, 30, 116-121.
20. Pruett-Jones SG and Lewis MJ (1990) Sex ratio and habitat limitation promote
delayed dispersal in superb fairy-wrens. Nature, 348: 541-542
21. Walters JR, Cooper CB, Daniels SJ, Pasinelli G, Schiegg K(1994) Conservation
Biology. In Koenig WD, Dickinson JL [eds.], Ecology and evolution of
cooperative breeding birds. Cambridge University Press.
Biographical Sketch.
Jordan Victoria Smith
Educational background
University of Florida
University of Florida
Zoology/Microbiology
Zoology
B.S., 2006
Master’s degree, in progress
Appointments
2004-08:
Molecular Lab Technician, Dr. Kimball and Dr.. Braun, Univ of Florida
Spring 2008: Teaching Assistant, General Biology Laboratory One, Univ of Florida
Fall 2007:
Teaching Assistant, General Biology Laboratory One, Univ of Florida
Spring 2007: Teaching Assistant, General Biology Laboratory One, Univ of Florida
Summer 2007: Avian Field Technician (Venezuela), Dr. Thomas Martin Univ of Montana
Winter 2006: Avian Surveyor (St. John U.S.V.I.) Dr. David Steadmen, Univ of Florida
Fall 2006:
Avian Surveyor (Elgin Air Force Base) Dr. Richard Fischer, US Army, ERDC
Summer 2006: Avian Surveyor (AZ, NM U.S.A) Univ of North Carolina, Chapel Hill
Summer 2005: Avian Field Technician (AZ U.S.A) Dr. Thomas Martin, Univ of Montana
Summer 2004: Avian Field Technician (AZ U.S.A) Dr. Thomas Martin, Univ of Montana
Synergistic Activities
(i) Undergraduate Mentor: Working in the Kimball and Braun lab involves extensive interaction
will undergraduates. Interactions include teaching of molecular protocols and procedures and
assistance with skill development in touchy lab techniques.
(ii) Citizen Scientist Outreach: While working on a project in Venezuela, I participated in
community outreach by encouraging local college students to visit our study site in Yacambu to
learn about science and field biology in particular. In addition, I was involved in teaching mistnetting technique and bird identification to a local birding club. Also, children often visited the
park on school field trips and the field crew was involved in short introductions to who scientist
are, and what a scientist does as part of the curriculum of their visit.
(iii) National Audubon’s Society: I am a volunteer participant in seasonal bird counts
Collaborators
James Cox, Tall Timbers Research Station
Sarah Haas, University of Florida
Master’s Advisor
Edward Braun, University of Florida
FOR NSF USE ONLY
5
4
SUMMARY PROPOSAL BUDGET
ORGANIZATION
University of Florida
PROPOSAL NO.
DURATION (MONTHS)
(Year 1)
Proposed
PRINCIPAL INVESTIGATOR/PROJECT DIRECTOR
Granted
AWARD NO.
Rebecca Kimball
A. SENIOR PERSONNEL: PI/PD, Co-PIs, Faculty and Other Senior Associates
NSF-Funded
List each separately with name and title. (A.7. Show number in brackets)
1.
2.
3.
Person-months
CAL ACAD SUMR
Jordan Smith
James Cox
Funds
Funds
Requested By
Granted by NSF
Proposer
(If Different)
$
$
$
$
4.
5.
6. (
) OTHERS (LIST INDIVIDUALLY ON BUDGET EXPLANATION PAGE)
7. ( 3 ) TOTAL SENIOR PERSONNEL (1-6)
B. OTHER PERSONNEL (SHOW NUMBERS IN BRACKETS)
1. (
) POSTDOCTORAL ASSOCIATES
2. (
) OTHER PROFESSIONALS (TECHNICIAN, PROGRAMMER, ETC.)
3. (
) GRADUATE STUDENTS
4. ( 4 ) UNDERGRADUATE STUDENTS
5. (
) SECRETARIAL - CLERICAL (IF CHARGED DIRECTLY)
6. (
) OTHER
TOTAL SALARIES AND WAGES (A + B)
C. FRINGE BENEFITS (IF CHARGED AS DIRECT COSTS)
TOTAL SALARIES, WAGES AND FRINGE BENEFITS (A + B + C)
D. EQUIPMENT (LIST ITEM AND DOLLAR AMOUNT FOR EACH ITEM EXCEEDING $5,000.)
TOTAL EQUIPMENT $ 5190
E. TRAVEL
1. DOMESTIC (INCL. CANADA, MEXICO AND U.S. POSSESSIONS)
2. FOREIGN
F. PARTICIPANT SUPPORT
1. STIPENDS
$
2. TRAVEL
3. SUBSISTENCE
4. OTHER
TOTAL NUMBER OF PARTICIPANTS (
)
COSTS
G. OTHER DIRECT COSTS $ 10,600
1. MATERIALS AND SUPPLIES
2. PUBLICATION/DOCUMENTATION/DISSEMINATION
3. CONSULTANT SERVICES
4. COMPUTER SERVICES
5. SUBAWARDS
TOTAL PARTICIPANT
6. OTHER
TOTAL OTHER DIRECT COSTS
H. TOTAL DIRECT COSTS (A THROUGH G) $15,790
I. INDIRECT COSTS (F&A) (SPECIFY RATE AND BASE)
TOTAL INDIRECT COSTS (F&A)
J. TOTAL DIRECT AND INDIRECT COSTS (H + I) $15,790
K. RESIDUAL FUNDS (IF FOR FURTHER SUPPORT OF CURRENT PROJECT SEE GPG II.D.7.j.)
L. AMOUNT OF THIS REQUEST (J) OR (J MINUS K)
M. COST SHARING: PROPOSED LEVEL $
PI/PD TYPED NAME AND SIGNATURE*
AGREED LEVEL IF DIFFERENT: $
DATE
FOR NSF USE ONLY
ORG. REP. TYPED NAME & SIGNATURE*
DATE
NSF Form 1030 (10/99) Supersedes All Previous Editions
*SIGNATURES REQUIRED ONLY FOR REVISED BUDGET (GPG III.C)
INDIRECT COST RATE VERIFICATION
Date Checked
Date of Rate Sheet
Initials-ORG
FOR NSF USE ONLY
5
4
SUMMARY PROPOSAL BUDGET
ORGANIZATION
University of Florida
PROPOSAL NO.
DURATION (MONTHS)
(Year 2)
Proposed
PRINCIPAL INVESTIGATOR/PROJECT DIRECTOR
Granted
AWARD NO.
Rebecca Kimball
A. SENIOR PERSONNEL: PI/PD, Co-PIs, Faculty and Other Senior Associates
NSF-Funded
List each separately with name and title. (A.7. Show number in brackets)
1.
2.
3.
Person-months
CAL ACAD SUMR
Jordan Smith
James Cox
Funds
Funds
Requested By
Granted by NSF
Proposer
(If Different)
$
$
$
$
4.
5.
6. (
) OTHERS (LIST INDIVIDUALLY ON BUDGET EXPLANATION PAGE)
7. ( 3 ) TOTAL SENIOR PERSONNEL (1-6)
B. OTHER PERSONNEL (SHOW NUMBERS IN BRACKETS)
1. (
) POSTDOCTORAL ASSOCIATES
2. (
) OTHER PROFESSIONALS (TECHNICIAN, PROGRAMMER, ETC.)
3. (
) GRADUATE STUDENTS
4. ( 4 ) UNDERGRADUATE STUDENTS
5. (
) SECRETARIAL - CLERICAL (IF CHARGED DIRECTLY)
6. (
) OTHER
TOTAL SALARIES AND WAGES (A + B)
C. FRINGE BENEFITS (IF CHARGED AS DIRECT COSTS)
TOTAL SALARIES, WAGES AND FRINGE BENEFITS (A + B + C)
D. EQUIPMENT (LIST ITEM AND DOLLAR AMOUNT FOR EACH ITEM EXCEEDING $5,000.)
TOTAL EQUIPMENT
E. TRAVEL
1. DOMESTIC (INCL. CANADA, MEXICO AND U.S. POSSESSIONS)
2. FOREIGN
F. PARTICIPANT SUPPORT
1. STIPENDS
$
2. TRAVEL
3. SUBSISTENCE
4. OTHER
TOTAL NUMBER OF PARTICIPANTS (
)
COSTS
G. OTHER DIRECT COSTS $ 6590
1. MATERIALS AND SUPPLIES
2. PUBLICATION/DOCUMENTATION/DISSEMINATION
3. CONSULTANT SERVICES
4. COMPUTER SERVICES
5. SUBAWARDS
TOTAL PARTICIPANT
6. OTHER
TOTAL OTHER DIRECT COSTS
H. TOTAL DIRECT COSTS (A THROUGH G) $6590
I. INDIRECT COSTS (F&A) (SPECIFY RATE AND BASE)
TOTAL INDIRECT COSTS (F&A)
J. TOTAL DIRECT AND INDIRECT COSTS (H + I) $6590
K. RESIDUAL FUNDS (IF FOR FURTHER SUPPORT OF CURRENT PROJECT SEE GPG II.D.7.j.)
L. AMOUNT OF THIS REQUEST (J) OR (J MINUS K)
M. COST SHARING: PROPOSED LEVEL $
PI/PD TYPED NAME AND SIGNATURE*
AGREED LEVEL IF DIFFERENT: $
DATE
FOR NSF USE ONLY
ORG. REP. TYPED NAME & SIGNATURE*
DATE
NSF Form 1030 (10/99) Supersedes All Previous Editions
*SIGNATURES REQUIRED ONLY FOR REVISED BUDGET (GPG III.C)
INDIRECT COST RATE VERIFICATION
Date Checked
Date of Rate Sheet
Initials-ORG
FOR NSF USE ONLY
5
4
SUMMARY PROPOSAL BUDGET
ORGANIZATION
University of Florida
PROPOSAL NO.
DURATION (MONTHS)
(Year 3)
Proposed
PRINCIPAL INVESTIGATOR/PROJECT DIRECTOR
Granted
AWARD NO.
Rebecca Kimball
A. SENIOR PERSONNEL: PI/PD, Co-PIs, Faculty and Other Senior Associates
NSF-Funded
List each separately with name and title. (A.7. Show number in brackets)
1.
Person-months
CAL ACAD SUMR
Jordan Smith
2. James
3.
Funds
Funds
Requested By
Granted by NSF
Proposer
(If Different)
$
$
$
$
Cox
4.
5.
6. (
) OTHERS (LIST INDIVIDUALLY ON BUDGET EXPLANATION PAGE)
7. ( 3 ) TOTAL SENIOR PERSONNEL (1-6)
B. OTHER PERSONNEL (SHOW NUMBERS IN BRACKETS)
1. (
) POSTDOCTORAL ASSOCIATES
2. (
) OTHER PROFESSIONALS (TECHNICIAN, PROGRAMMER, ETC.)
3. (
) GRADUATE STUDENTS
4. ( 4 ) UNDERGRADUATE STUDENTS
5. (
) SECRETARIAL - CLERICAL (IF CHARGED DIRECTLY)
6. (
) OTHER
TOTAL SALARIES AND WAGES (A + B)
C. FRINGE BENEFITS (IF CHARGED AS DIRECT COSTS)
TOTAL SALARIES, WAGES AND FRINGE BENEFITS (A + B + C)
D. EQUIPMENT (LIST ITEM AND DOLLAR AMOUNT FOR EACH ITEM EXCEEDING $5,000.)
TOTAL EQUIPMENT
E. TRAVEL
1. DOMESTIC (INCL. CANADA, MEXICO AND U.S. POSSESSIONS)
2. FOREIGN
F. PARTICIPANT SUPPORT
1. STIPENDS
$
2. TRAVEL
3. SUBSISTENCE
4. OTHER
TOTAL NUMBER OF PARTICIPANTS (
)
COSTS
G. OTHER DIRECT COSTS $ 6590
1. MATERIALS AND SUPPLIES
2. PUBLICATION/DOCUMENTATION/DISSEMINATION
3. CONSULTANT SERVICES
4. COMPUTER SERVICES
5. SUBAWARDS
TOTAL PARTICIPANT
6. OTHER
TOTAL OTHER DIRECT COSTS
H. TOTAL DIRECT COSTS (A THROUGH G) $6590
I. INDIRECT COSTS (F&A) (SPECIFY RATE AND BASE)
TOTAL INDIRECT COSTS (F&A)
J. TOTAL DIRECT AND INDIRECT COSTS (H + I) $6590
K. RESIDUAL FUNDS (IF FOR FURTHER SUPPORT OF CURRENT PROJECT SEE GPG II.D.7.j.)
L. AMOUNT OF THIS REQUEST (J) OR (J MINUS K)
M. COST SHARING: PROPOSED LEVEL $
PI/PD TYPED NAME AND SIGNATURE*
AGREED LEVEL IF DIFFERENT: $
DATE
FOR NSF USE ONLY
ORG. REP. TYPED NAME & SIGNATURE*
DATE
NSF Form 1030 (10/99) Supersedes All Previous Editions
*SIGNATURES REQUIRED ONLY FOR REVISED BUDGET (GPG III.C)
INDIRECT COST RATE VERIFICATION
Date Checked
Date of Rate Sheet
Initials-ORG
FOR NSF USE ONLY
5
4
SUMMARY PROPOSAL BUDGET
ORGANIZATION
University of Florida
PROPOSAL NO.
DURATION (MONTHS)
(Total for 3 Years)
Proposed
PRINCIPAL INVESTIGATOR/PROJECT DIRECTOR
Granted
AWARD NO.
Rebecca Kimball
A. SENIOR PERSONNEL: PI/PD, Co-PIs, Faculty and Other Senior Associates
NSF-Funded
List each separately with name and title. (A.7. Show number in brackets)
Person-months
CAL ACAD SUMR
1. Jordan Smith
2.
3.
Funds
Funds
Requested By
Granted by NSF
Proposer
(If Different)
$
$
$
$
James Cox
4.
5.
6. (
) OTHERS (LIST INDIVIDUALLY ON BUDGET EXPLANATION PAGE)
7. ( 3 ) TOTAL SENIOR PERSONNEL (1-6)
B. OTHER PERSONNEL (SHOW NUMBERS IN BRACKETS)
1. (
) POSTDOCTORAL ASSOCIATES
2. (
) OTHER PROFESSIONALS (TECHNICIAN, PROGRAMMER, ETC.)
3. (
) GRADUATE STUDENTS
4. ( 4 ) UNDERGRADUATE STUDENTS
5. (
) SECRETARIAL - CLERICAL (IF CHARGED DIRECTLY)
6. (
) OTHER
TOTAL SALARIES AND WAGES (A + B)
C. FRINGE BENEFITS (IF CHARGED AS DIRECT COSTS)
TOTAL SALARIES, WAGES AND FRINGE BENEFITS (A + B + C)
D. EQUIPMENT (LIST ITEM AND DOLLAR AMOUNT FOR EACH ITEM EXCEEDING $5,000.)
TOTAL EQUIPMENT $5190
E. TRAVEL
1. DOMESTIC (INCL. CANADA, MEXICO AND U.S. POSSESSIONS)
2. FOREIGN
F. PARTICIPANT SUPPORT
1. STIPENDS
$
2. TRAVEL
3. SUBSISTENCE
4. OTHER
TOTAL NUMBER OF PARTICIPANTS (
)
COSTS
G. OTHER DIRECT COSTS $23,780
1. MATERIALS AND SUPPLIES
2. PUBLICATION/DOCUMENTATION/DISSEMINATION
3. CONSULTANT SERVICES
4. COMPUTER SERVICES
5. SUBAWARDS
TOTAL PARTICIPANT
6. OTHER
TOTAL OTHER DIRECT COSTS
H. TOTAL DIRECT COSTS (A THROUGH G) $23,780
I. INDIRECT COSTS (F&A) (SPECIFY RATE AND BASE)
TOTAL INDIRECT COSTS (F&A)
J. TOTAL DIRECT AND INDIRECT COSTS (H + I) $28,970
K. RESIDUAL FUNDS (IF FOR FURTHER SUPPORT OF CURRENT PROJECT SEE GPG II.D.7.j.)
L. AMOUNT OF THIS REQUEST (J) OR (J MINUS K)
M. COST SHARING: PROPOSED LEVEL $
PI/PD TYPED NAME AND SIGNATURE*
AGREED LEVEL IF DIFFERENT: $
DATE
FOR NSF USE ONLY
ORG. REP. TYPED NAME & SIGNATURE*
DATE
NSF Form 1030 (10/99) Supersedes All Previous Editions
*SIGNATURES REQUIRED ONLY FOR REVISED BUDGET (GPG III.C)
INDIRECT COST RATE VERIFICATION
Date Checked
Date of Rate Sheet
Initials-ORG
Budget Justification.
Funding is estimated on calculations to sample forty territories at each field site. The
number of adults to sustain forty active territories should flucuate around 100 (including
helpers), and the average clutch size is 4.5 individuals with a maximum nestling number
of 200.
Genotyping cost: $16. 00 per individual
-PCR cost: $1.00 per individual
(Includes reagents, fluorescently labeled microsatellite primers, sexing primers, and gel
electrophoresis)
-Genotyping: $15.00 per individual
(Includes cost for all fifteen microsatellites)
Year 1
Single Cost Items
3-banding pliers ($20):
Capillary tubes, coolers with ice packs:
Two portable CD players with speakers:
3-Digital caliper ($140 ):
3- Digital and spring scales ($150):
Mist-nets, poles, pole adaptors, rope:
Aviary cost (food, individual cages, cleaning)
PUREGENE® DNA Purification Kit ($500/250 samples):
Sampling Cost
Blood buffer:
Color bands (0.25 each):
Genotyping of remaining 2006, 2007, 2008 individuals:
Genotyping 2009 TTRS:
Genotyping 2009 WT:
Total Year One:
Year 2 Cost
Blood buffer:
Color bands (0.25 each):
Genotyping 2010 WT and TTRS:
$60
$60
$100
$420
$450
$500
$600
$3000
$5,190
$50
$150
$2400 (150 individuals)
$3200 (200 individuals)
$4800 (300 individuals)
$10,600
$15,790
$40
$150
$6400 (400 individuals)
$6,590
Year 3 Cost
Blood buffer:
Color bands (0.25 each):
Genotyping 2011 WT and TTRS:
$40
$150
$6400 (400 individuals)
$6,590
Three Year Total
$28,970
Current and Pending Support
(See GPG Section II.D.8 for guidance on information to include on this form.)
The following information should be provided for each investigator and other senior personnel. Failure to provide this
information may delay consideration of this proposal.
Other agencies (including NSF) to which this proposal has been/will be submitted.
Investigator:
Support:
Current
Pending
Submission Planned in Near Future
*Transfer of Support
Project/Proposal Title:
NOTHING
Source of Support:
Total Award Amount: $
Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project.
Support:
Current
Pending
Cal:
Acad:
Submission Planned in Near Future
Sumr:
*Transfer of Support
Project/Proposal Title:
Source of Support:
Total Award Amount: $
Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project.
Support:
Current
Pending
Cal:
Acad:
Submission Planned in Near Future
Sumr:
*Transfer of Support
Project/Proposal Title:
Source of Support:
Total Award Amount: $
Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project.
Support:
Current
Pending
Cal:
Acad:
Submission Planned in Near Future
Sumr:
*Transfer of Support
Project/Proposal Title:
Source of Support:
Total Award Amount: $
Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project.
Support:
Current
Pending
Cal:
Acad:
Submission Planned in Near Future
Sumr:
*Transfer of Support
Project/Proposal Title:
Source of Support:
Total Award Amount: $
Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project.
Cal:
Acad:
Sumr:
*If this project has previously been funded by another agency, please list and furnish information for immediately preceding funding period.
NSF Form 1239 (10/99)
5
5
USE ADDITIONAL SHEETS AS NECESSARY
FACILITIES, EQUIPMENT & OTHER RESOURCES
FACILITIES: Identify the facilities to be used at each performance site listed and, as appropriate, indicate their capacities, pertinent
capabilities, relative proximity, and extent of availability to the project. Use “Other” to describe the facilities at any other
performance sites listed and at sites for field studies. Use additional pages if necessary.
Laboratory:
Kimball Braun Laboratory, Department of Zoology, University of Florida
All molecular research will be conducted in this lab; it is fully equipped for ALL stages of genotyping including preparatory
work such as DNA extractions through the final analyses.
This lab is located in my home university and is support by two of my committee members. It is accessible as needed.
Clinical:
Animal:
University of Florida Aviary: housing translocated birds
Computer:
Office:
Other:
Field Sites
Tall Timbers Research Station :13093 Henry Beadel Drive Tallahassee, Fl 32312
Both sites are available as needed for sample collection, observation, and manipulation
The sites are located approximately 165 miles Northwest of my home university
MAJOR EQUIPMENT: List the most important items available for this project and, as appropriate, identify the location and
pertinent capabilities of each.
Thermocycler: necessary for PCR of microsatellite loci- Kimball Braun Laboratory
ABI Prism TM 3100-Avant genetic analyzer: necessary for genotyping all individuals- Kimball Braun Laboratory
GeneMarker V.1.5: necessary program for reading genotyping results- Kimball Braun Laboratory
Mist-nets and poles: necessary for capture of birds- Kimball Braun Laboratory, Tall Timbers Research Station
USGS/ color leg bands: critical for individual bird identification-Tall Timbers Research Station
OTHER RESOURCES: Provide any information describing the other resources available for the project. Identify support services
such as consultant, secretarial, machine shop, and electronics shop, and the extent to which they will be available for the project.
Include an explanation of any consortium/contractual/subaward arrangements with other organizations.
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6
NSF Form 1363 (10/99)