COVER SHEET FOR PROPOSAL TO THE NATIONAL SCIENCE FOUNDATION PROGRAM ANNOUNCEMENT/SOLICITATION NO./CLOSING DATE/If not in response to a program announcement/solicitation enter NSF 00-2 FOR NSF USE ONLY NSF PROPOSAL NUMBER NSF DIGG FOR CONSIDERATION BY NSF ORGANIZATIONAL UNIT(S) (Indicate the most specific unit known, i.e., program, division, etc.) DATE RECEIVED NUMBER OF COPIES DIVISION ASSIGNED EMPLOYER IDENTIFICATION NUMBER (EIN) OR TAXPAYER IDENTIFICATION NUMBER (TIN) FUND CODE DUNS # (Data Universal Numbering System) SHOW PREVIOUS AWARD NO. IF THIS IS FILE LOCATION IS THIS PROPOSAL BEING SUBMITTED TO ANOTHER FEDERAL A RENEWAL AGENCY? YES NO IF YES, LIST ACRONYM(S) AN ACCOMPLISHMENT-BASED RENEWAL NAME OF ORGANIZATION TO WHICH AWARD SHOULD BE MADE ADDRESS OF AWARDEE ORGANIZATION, INCLUDING 9 DIGIT ZIP CODE University of Florida University of Florida Department of Zoology Gainesville, FL 32608 AWARDEE ORGANIZATION CODE (IF KNOWN) NAME OF PERFORMING ORGANIZATION, IF DIFFERENT FROM ABOVE ADDRESS OF PERFORMING ORGANIZATION, IF DIFFERENT, INCLUDING 9 DIGIT ZIP CODE PERFORMING ORGANIZATION CODE (IF KNOWN) IS AWARDEE ORGANIZATION (Check All That Apply) (See GPG II.D.1 For Definitions) FOR-PROFIT ORGANIZATION SMALL BUSINESS MINORITY BUSINESS WOMAN-OWNED BUSINESS TITLE OF PROPOSED PROJECT Understanding Woody Plant Dynamics in African Savannas: Integrating Herbivory, Precipitation, Soil Nutrients and Plant De- fense REQUESTED AMOUNT PROPOSED DURATION (1-60 MONTHS) REQUESTED STARTING DATE 36 August 2008 $ SHOW RELATED PREPROPOSAL NO., IF APPLICABLE months CHECK APPROPRIATE BOX(ES) IF THIS PROPOSAL INCLUDES ANY OF THE ITEMS LISTED BELOW BEGINNING INVESTIGATOR (GPG I.A.3) VERTEBRATE ANIMALS (GPG II.D.12) IACUC App. Date DISCLOSURE OF LOBBYING ACTIVITIES (GPG II.D.1) PROPRIETARY & PRIVILEGED INFORMATION (GPG I.B, II.D.7) HUMAN SUBJECTS (GPG II.D.12) Exemption Subsection or IRB App. Date NATIONAL ENVIRONMENTAL POLICY ACT (GPG II.D.10) INTERNATIONAL COOPERATIVE ACTIVITIES: COUNTRY/COUNTRIES HISTORIC PLACES (GPG II.D.10) SMALL GRANT FOR EXPLOR. RESEARCH (SGER) (GPG II.D.12) FACILITATION FOR SCIENTISTS/ENGINEERS WITH DISABILITIES (GPG V.G.) RESEARCH OPPORTUNITY AWARD (GPG V.H) PI/PD DEPARTMENT PI/PD POSTAL ADDRESS Zoology PI/PD FAX NUMBER 411 Carr, Department of Zoology, University of Florida Gainesville, FL 32608 USA NAMES (TYPED) High Degree Yr of Degree Telephone Number Electronic Mail Address B.S 2005 843-323-1351 [email protected] PI/PD NAME Megan Gittinger CO-PI/PD CO-PI/PD CO-PI/PD CO-PI/PD NSF Form 1207 (10/99) Page 1 of 2 CERTIFICATION PAGE Certification for Principal Investigators and Co-Principal Investigators I certify to the best of my knowledge that: (1) the statements herein (excluding scientific hypotheses and scientific opinions) are true and complete, and (2) the text and graphics herein as well as any accompanying publications or other documents, unless otherwise indicated, are the original work of the signatories or individuals working under their supervision. I agree to accept responsibility for the scientific conduct of the project and to provide the required project reports if an award is made as a result of this proposal. I understand that the willful provision of false information or concealing a material fact in this proposal or any other communication submitted to NSF is a criminal offense (U.S.Code, Title 18, Section 1001). Name (Typed) PI/PD Signature Social Security No.* Megan Gittinger Date February 18, 2008 Co-PI/PD Co-PI/PD Co-PI/PD Co-PI/PD Certification for Authorized Organizational Representative or Individual Applicant By signing and submitting this proposal, the individual applicant or the authorized official of the applicant institution is: (1) certifying that statements made herein are true and complete to the best of his/her knowledge; and (2) agreeing to accept the obligation to comply with NSF award terms and conditions if an award is made as a result of this application. Further, the applicant is hereby providing certifications regarding Federal debt status, debarment and suspension, drug-free workplace, and lobbying activities (see below), as set forth in the Grant Proposal Guide (GPG), NSF 00-2. Willful provision of false information in this application and its supporting documents or in reports required under an ensuing award is a criminal offense (U.S. Code, Title 18, Section 1001). In addition, if the applicant institution employs more than fifty persons, the authorized official of the applicant institution is certifying that the institution has implemented a written and enforced conflict of interest policy that is consistent with the provisions of Grant Policy Manual Section 510; that to the best of his/her knowledge, all financial disclosures required by that conflict of interest policy have been made; and that all identified conflicts of interest will have been satisfactorily managed, reduced or eliminated prior to the institution’s expenditure of any funds under the award, in accordance with the institution’s conflict of interest policy. Conflicts that cannot be satisfactorily managed, reduced or eliminated must be disclosed to NSF. Debt and Debarment Certifications (If answer “yes” to either, please provide explanation.) Is the organization delinquent on any Federal debt? Is the organization or its principals presently debarred, suspended, proposed for debarment, declared ineligible, or voluntarily excluded from covered transactions by any Federal Department or agency? Yes No Yes No Certification Regarding Lobbying This certification is required for an award of a Federal contract, grant or cooperative agreement exceeding $100,000 and for an award of a Federal loan or a commitment providing for the United States to insure or guarantee a loan exceeding $150,000. Certification for Contracts, Grants, Loans and Cooperative Agreements The undersigned certifies, to the best of his or her knowledge and belief, that: (1) No Federal appropriated funds have been paid or will be paid, by or on behalf of the undersigned, to any person for influencing or attempting to influence an officer or employee of any agency, a Member of Congress, an officer or employee of Congress, or an employee of a Member of Congress in connection with the awarding of any federal contract, the making of any Federal grant, the making of any Federal loan, the entering into of any cooperative agreement, and the extension, continuation, renewal, amendment, or modification of any Federal contract, grant, loan, or cooperative agreement. (2) If any funds other than Federal appropriated funds have been paid or will be paid to any person for influencing or attempting to influence an officer or employee of any agency, a Member of Congress, and officer or employee of Congress, or an employee of a Member of Congress in connection with this Federal contract, grant, loan, or cooperative agreement, the undersigned shall complete and submit Standard Form LLL, “Disclosure of Lobbying Activities,” in accordance with its instructions. (3) The undersigned shall require that the language of this certification be included in the award documents for all subawards at all tiers including subcontracts, subgrants, and contracts under grants, loans, and cooperative agreements and that all subrecipients shall certify and disclose accordingly. This certification is a material representation of fact upon which reliance was placed when this transaction was made or entered into. Submission of this certification is a prerequisite for making or entering into this transaction imposed by Section 1352, Title 31, U.S. Code. Any person who fails to file the required certification shall be subject to a civil penalty of not less than $10,000 and not more than $100,000 for each such failure. AUTHORIZED ORGANIZATIONAL REPRESENTATIVE NAME/TITLE (TYPED) SIGNATURE DATE Megan Gittinger 02/2008 TELEPHONE NUMBER ELECTRONIC MAIL ADDRESS 843-323-1351 [email protected] FAX NUMBER *SUBMISSION OF SOCIAL SECURITY NUMBERS IS VOLUNTARY AND WILL NOT AFFECT THE ORGANIZATION’S ELIGIBILITY FOR AN AWARD. HOWEVER, THEY ARE AN INTEGRAL PART OF THE NSF INFORMATION SYSTEM AND ASSIST IN PROCESSING THE PROPOSAL. SSN SOLICITED UNDER NSF ACT OF 1950, AS AMENDED. Page 2 of 2 PROJECT SUMMARY. Savannas represent one of the world’s largest biomes and are one of the most important and widespread habitats in sub-Saharan Africa. Trees and grasses dominate these ecosystems; however, the dominance fluctuates from heavily woody areas to grassland. Despite a historical research interest and investment in understanding the structure and function of savannas, we are unable to explain why some areas are converting to desert and others to dense bushland. Recent literature suggests that factors (precipitation, soil nutrients, fire and herbivory) regulating tree versus grass dominance, interact and their strength varies within certain thresholds. However, it is not apparent how these factors interact at local scales in concert with species-specific traits such as plant defense to determine plant productivity. This is particularly important given complex feedbacks that can exist between herbivores and plants. These interactive effects have large implications for the conservation of these systems considering increasing human population, climate change and changing herbivore abundances. The proposed research examines how herbivory and soil nutrients interact across a precipitation gradient to influence woody plant cover and productivity, and plant-herbivore interactions. The research will address three specific questions 1) Does simulated browsing stimulate re-growth of plants?, 2) Is re-growth synonymous with increased forage availability, and will regrowth facilitate re-browsing (and therefore habitat use)? , and 3) Does plant defense mediate the interactions between browsers and plants across nutrient and precipitation gradients?. My research takes advantage of a model system (the Laikipia plateau) where two Acacia species dominate on two soil types that differ in soil fertility (A. drepanolobium on nutrient rich “black cotton” soils, A. mellifera on nutrient poor sandy “red” soils, see6) and are distributed across a broad range of precipitation (400 mm/yr – 750 mm/yr). This research will further our understanding of herbivore mediated effects on woody plants across abiotic gradients, which can directly contribute to conservation and land management in African savannas. Project Summary – Page 1 TABLE OF CONTENTS For font-size and page-formatting specifications, see GPG Section II.C. Total No. of Pages in Section Section Page No.* (Optional)* Cover Sheet (NSF Form 1207) (Submit Page 2 with original proposal only) A Project Summary (not to exceed 1 page) 1 B Table of Contents (NSF Form 1359) 1 C Project Description (including Results from Prior NSF Support) (not to exceed 15 pages) (Exceed only if allowed by a specific program announcement/solicitation or if approved in advance by the appropriate NSF Assistant Director or designee) 5 D References Cited 4 E Biographical Sketches (Not to exceed 2 pages each) 1 F Budget (NSF Form 1030, plus up to 3 pages of budget justification) 5 G Current and Pending Support (NSF Form 1239) 1 H Facilities, Equipment and Other Resources (NSF Form 1363) 1 I Special Information/Supplementary Documentation J Appendix (List below) Include only if allowed by a specific program announcement/ solicitation or if approved in advance by the appropriate NSF Assistant Director or designee) Appendix Items: *Proposers may select any numbering mechanism for the proposal. The entire proposal, however, must be paginated. Complete both columns only if the proposal is numbered consecutively. NSF Form 1359 (10/99) 46 PROJECT DESCRIPTION. Introduction. African savannas support much of the world’s livestock and wild herbivore biomass (Scholes & Archer, 1997), are among the ecosystems most likely to be influenced by future climate change (Bond & Woodward, 2003), and are under immediate threat from human impact (Du Toit & Cumming, 1999). A mechanistic understanding of the factors that structure savannas and facilitate the coexistence of woody plants and grasses is a key conservation and management priority. While the broad-scale drivers of woody plant dynamics, such as precipitation, herbivory and soil nutrient levels have been identified (Sankaran et al., 2005), how these factors interact at local scales in concert with species-specific traits such as plant defense to determine woody plant productivity and dynamics remains controversial (reviewed in Scholes & Archer, 1997). Parsing the mechanisms underlying woody plant dynamics is a strong priority for several pressing conservation issues, including bush encroachment (Scholes & Archer, 1997), and the availability of browse for both livestock (e.g. goats) and native ungulates (e.g., elephants, giraffe, and other browsing species) within this threatened ecosystem. I propose to examine how herbivory and soil nutrients interact across a precipitation gradient to influence woody plant cover and productivity, and plantherbivore interactions. My research takes advantage of a model system (the Laikipia plateau, see Georgiadis et al., 2007) where two Acacia species dominate on two soil types that differ in soil fertility (A. drepanolobium on nutrient rich “black cotton” soils, A. mellifera on nutrient poor sandy “red” soils, see (Pringle et al., 2007) and are distributed across a broad range of precipitation (400 mm/yr – 750 mm/yr). Both systems are classified as semi-arid thorn savannas. Fire suppression has been practiced by land managers in this landscape for decades, and accidental fire histories are well-documented, eliminating this important factor as a confound within Todd Palmer my experimental approaches. Background. Role of herbivores / Positive interactions. African savanna ecosystems provide excellent models for the study of plant-herbivore interactions from both basic and applied scientific perspectives. Large ungulates consume a significant amount of the plant production in these systems (Phillipson, 1973; Sinclair, 1975), and co-occur with domestic livestock throughout much of their range. As livestock densities have increased over the past decades, it is becoming increasingly important to understand the direct and interactive effects of livestock and wildlife on these threatened rangeland systems. Project Description – Page 2 Effects of herbivory on plants have been widely documented. Herbivores have been shown to change plant structure (branching or thorns; Rohner & Ward, 1997), nutrient levels in leaves (nitrogen and phosphorus; Augustine & McNaughton, 2006; Anderson et al., 2007), and secondary compounds (Karban & Baldwin, 1997). At a larger scale browsing can affect primary productivity (Augustine & McNaughton, 2006; Manier & Hobbs, 2007), plant biomass (McNaughton, 1984), and species composition (Manier & Hobbs, 2007; Augustine & McNaughton, 1998; Hobbs, 1996). Whether or not these effects of herbivory on plants are all happening at once, it is apparent that large herbivores have the potential to play a strong role in the structuring of savanna systems. Large herbivory diversity in savannas is considered a function of landscape spatial and temporal heterogeneity (Bell, 1986). Species can be broadly categorized into a number of different functional groups: size (megaherbivores, mesoherbivores), forage type (grazer, browser, mixed feeder), movement patterns (migratory, resident) and gut type (ruminant, nonruminant) (Hoffman, 1989). Each of these groups, which are not mutually exclusive (e.g. some populations can be migratory or resident), will exert pressure on certain plant types and may also be in competition for similar food resources. Understanding how these functional groups differ in terms of effects on plant communities is essential. For example, migratory versus resident ungulates will exert different levels of grazing or browsing intensity, which may effect the species composition in terms of herbivore tolerance. Wild herbivores, their effects on plants and potential feedback mechanism in savanna grasslands have been well documented. One species of large herbivore moving through a landscape clips down grasses, which stimulates leaf elongation and enhanced primary productivity; this in turn provides a more nutritious food source for another herbivore species (McNaughton, 1983; Verweij, 2006). Not only are herbivores affecting plant structure and nutrient dynamics, but also they are manipulating their food resources in such a way that facilitates the coexistence of multiple herbivore species in that space. These ‘hotspots’ are referred to as grazing lawns and the classic example is the migration of wildebeest, zebra and gazelle across African savannas (McNaughton, 1984; McNaughton, 1976). A similar pattern of facilitation between herbivore species has recently been suggested in woodland areas (van de Koppel & Prins, 1998; Owen-Smith, 2003; Makhabu et al., 2006; Fornara & Toit, 2007). Makhabu et al. (2006) experimentally showed that elephant herbivory had an effect on impala foraging. Trees that had been browsed by elephants responded through re-growth and impala showed a preference for trees previously browsed by elephants. Facilitation among herbivores through resource manipulation may not always occur for a number of reasons. In the case of migratory species, resources are separated through space and time; if this temporal separation was not present then we may see competition among species or the presence of fewer species because of a less heterogeneous landscape (Augustine & McNaughton, 2006). Abiotic factors, such as precipitation could also effect grazing lawns; in areas of low rainfall plants may not be able to compensate and re-growth may be limited (Bond & Loffell, 2001). Slow re-growth may explain why in some areas we see direct effects on plant structure by elephants (megaherbivore mixed feeder) and indirect facilitation between other mesobrowsers (e.g. impala), while in other cases no effect on plant community structure or positive relationships between other mesoherbivores is seen (Augustine & McNaughton, 2006). These direct effects of herbivores on plants as well as indirect effects between herbivores further increase the complexity of factors acting on a local scale. It also provides an example of why understanding the effects of different herbivore types on plant community warrants further Project Description – Page 3 research. Long-term experimental manipulations examining the impacts of different functional herbivores on plant communities using abiotic gradients are needed. In addition to research directly testing local community interactions, larger scale modeling projects could also contribute significantly. Although there has been some work using spatially explicit models to look at herbivore abundance and plant structure (or biomass or productivity) these models did not to my knowledge attempt to look at different herbivore types. Plant defenses. Herbivory can be highly detrimental to plants if they are not defended, causing local or complete extinction of the species under extreme herbivore pressure (Bond & Loffell. 2001). Large herbivores have been present over a long time scale in Africa; therefore we expect plant species to have evolved defense traits that allow them to tolerate or resist herbivory. A broad range of defense mechanisms has evolved and can be broadly categorized as chemical or mechanical. Chemical defenses include the production of substances such as tannins, phenols, and alkaloids, and they deter herbivores by making plant unpalatable. Mechanical defenses include defense structures (spines) as well as lateral branching (density of branches). The type of defense a plant has may be indicative of the previous type (e.g. herbivore type) and historical levels of herbivory (Fornara & Toit, 2007). Plant defenses can also be studied on shorter time scales and have been shown to demonstrate changes in strength with varying levels of herbivory. For example, spine length in Acacia trees varies with levels of herbivory (Young & Okello, 1998; Young et al., 2003; Milewski & Madden, 2006). Over extended time periods of little to no herbivory we may expect that plants would decrease overall defenses. This is based on the resource availability hypothesis within plant defense theory, which states that there is a trade-off between the cost of producing defenses and how much that affects a plant’s ability to grow and reproduce (Grime, 1979; Coley, 1985; Lim & Turner, 1996). Therefore, we would expect that a plant would only produce defensive compounds in cases where the cost of producing compounds increasing protection and that long-lived plants have less of a trade-off. Ultimately, the way in which plants respond to herbivores can affect plant communities. Disentangling regulators of plant communities is inexorably linked to herbivores effects on plants as well as plants responses to herbivory. This aspect of local community dynamics within savannas is usually separated in the literature. I suggest that these responses to herbivory studied along side the role of herbivores on plants. Examining these interactions within the framework of savanna regulators would also provide insight into applied conservation issues such as bush encroachment, which is usually associated with an increase of unpalatable browse. Project Description – Page 4 Research Questions and Experimental Design. This research will focus on three interrelated objectives: 1. To determine if simulated browsing stimulates re-growth. 2. To determine if re-growth is synonymous with an increase in forage availability and if re-growth facilitates re-browsing (and therefore habitat use). 3. To determine if plant defense mediates the interactions between browsers and plants across nutrient and precipitation gradients. Overall Experimental Design: I will evaluate these questions in a factorial design using 2 treatments, soil type and browsing intensity, across a precipitation gradient. A total of 36 plots (1/4 hectare) will be delineated: 3 replicate plots per soil type (black cotton and red) x 3 precipitation levels (low, 400mm/yr; medium, 600mm/yr; high, 750mm/yr) x 2 browsing intensities (low, high). In each plot, 15 trees (540 trees total) between 1.5-2.5 m in height will be marked and GPS locations taken. Acacia density in the study area is relatively uniform and estimated to be ~500 trees (> ! m in height) per hectare (JR Goheen, pers. comm.); treatment trees represent at least 20% of the total number of reproductively mature trees, which I expect will be sufficient to detect a response. Browsing will be simulated on marked branches at two intensities in plots for two soil types across rainfall gradient. Simulated browsing (low damage) will be performed on 10 branches per tree using methods following Gadd & Palmer, 2001; for high damage browsing, two entire branch systems (minimum diameter = 8cm) will be broken. An invisible, non-toxic fluorescent stain Figure 1: Experimental layout of (RiskReactor®; odorless, taste-less, EPA-approved) will be stimulated browsing. Plots will be added to simulated browsing branch tips to assess post- replicated across a rainfall gradient. treatment browsing. Initial wild and domestic herbivore use of areas will be determined through complete dung collection within each plot; dung will be identified to species in both field and lab. My lab has strong background in these techniques, and expert field assistants who are adept at these methods. The lab has experience in clearing plots entirely of dung; with 5 field assistants, a single " ha plot can be cleared of dung and piles identified in approximately one hour. Objective 1: To determine if simulated browsing stimulates re-growth in Acacia drepanalobium. Hypothesis: Simulated browsing may stimulate re-growth of palatable browse, but the magnitude of this response varies with browsing intensity, soil nutrients, and precipitation. Prediction: I predict higher re-growth and nutrient levels in re-growing shoots in nutrient rich soils (red soils) and in plots where rainfall is sufficient for growth compensation (medium and high; see Gadd & Palmer, 2001). Responses will be of lower magnitude where nutrients and rainfall are limited. Project Description – Page 5 Measurements: Browsing simulations will be performed at the beginning of each rainy season (March and October). Branches will be re-surveyed annually for re-growth, including total branch length from marked points and the number of branch apices. Re-browsing is easily detectable and will be noted during surveys. I will collect approximately 50g of leaves from regrowth for analysis of leaf nutrient levels (total N and P using Kjeldahl digestion). Objective 2: To determine if re-growth is synonymous with an increase in forage availability and if re-growth facilitate re-browsing (and therefore habitat use). Hypothesis: If simulated browsing increases re-growth and palatability, then herbivore damage will facilitate re-browsing, consistent with the recent “browsing lawns” hypothesis (e.g. an increase in habitat use; Fornara et al., 2007). Prediction: I predict that re-browsing and increased habitat usage will be more pronounced in areas of high precipitation and nutrient rich soils if resource availability and defense do not interact with re-growth palatability. Measurements: Wild and domestic herbivore use of areas will be re-surveyed every 6 months through complete dung collection within each plot and identification in both field and lab. Prior research in my lab has shown a slow decay rate (~10 months) of dung piles over time, even in the presence of dung beetles. Objective 3: To determine if plant defense mediates the interactions between browsers and plants across nutrient and precipitation gradients. Hypothesis: Plant compensatory growth and inducible defenses are resource limited, and will have interacting effects with precipitation and soil richness on re-growth. Prediction: In resource poor areas, plants will have greater structural and chemical defense. I predict compensatory growth will increase with resource availability (Coley et al., 1985). Measurements: Thorn number and length (mechanical defense) will be recorded prior to herbivore treatments and then re-recorded annually on new growth. I will use previously collected leaves from re-growth for analysis of condensed tannins (methods in Ward, 2002), the primary chemical defense within Acacias in this system (Ward, 2002). Significance. Broader Impacts. Laikipia is a model system for studying drivers of local woody plant dynamics due to its ecology and historical records of landscape use. This research will further our understanding of herbivore mediated effects on woody plants across abiotic gradients, which can directly contribute to conservation and land management in African savannas. Funding for this work will support the education and training of local Kenyan research assistants; there would also be an opportunity for independent research related to this work. Project Description – Page 6 1. Anderson, T.M. et. al. (2007) Forage Nutritive Quality in the Serengeti Ecosystem: The roles of fire and herbivory. American Naturalist. 170, 343-357. 2. Augustine, D.J. and McNaughton, S.J. (1998) Ungulate effects on the functional species composition of plant communities: herbivore selectivity and plant tolerance. Journal of Wildlife Management. 62, 1165-1183. 3. Augustine, D.J. and McNaughton, S.J. 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(2007) Savanna herbivore dynamics in a livestock-dominated landscape. II: Ecological, conservation, and management implications of predator restoration. Biological Conservation. 137, 473-483. 16. Grime, J.P. (1979) Plant Strategies and Vegetation Processes. John Wiley and Sons, New York. 17. Henkin, Z., et. al. (1999) Secondary succession after fire in a Mediterranean dwarf-scrub community. Journal of Vegetation Science. 10, 503-514. 18. Higgins, S.I. et. al. (2000) Fire, resprouting and variability: a recipe for grass-tree coexistence in savanna. Journal of Ecology. 88, 213–229. 19. Hobbs, N.T. (1996) Modification of ecosystems by ungulates. Journal of Wildlife Management. 60, 695-713. 20. Hofmann, R. (1989) Evolutionary steps of ecophysiological adaptation and diversification of ruminants - a comparative view of their digestive system. Oecologia 78: 443-457. 21. Hughs, L. (2003) Climate change and Australia: trends, projections and impacts. Austral Ecology. 28, 423-443. 22. Karban, R. and Baldwin, I.T. (1997) Induced responses to herbivory. University of Chicago Press, Chicago, Illinois, USA. 23. Lim, W. H. L. and Turner, I. M. (1996) Resource availability and growth responses to defoliation in seedlings of three early-successional, tropical, woody species. Ecological Research. 11, 321-324. 24. Makhabu, S.W. et. al. (2006) Elephant impact on shoot distribution on trees and on rebrowsing by smaller browsers. Acta Oecologica. 30, 136-146 25. Manier, D.J. and Hobbs, N.T. (2007) Large herbivores in sagebrush steppe ecosystems: livestock and wild ungulates influence structure and function. Oecologia. 152, 739-750. 26. McNaughton, S.J. (1976) Serengeti migratory wildebeest: facilitation of energy flow by grazing. Science. 191, 92-94. 27. McNaughton, S.J. (1983) Compensatory growth as a response to herbivory. Oikos. 40, 329336. 28. McNaughton, S.J. (1984) Grazing Lawns: Animals in Herds, Plant Form, and Coevolution. American Naturalist. 124, 863-886. Literature Cited – Page 8 29. Milewski, A.V. and Madden, D. (2006) Interactions between large African browsers and thorny Acacia on a wildlife ranch in Kenya. Afr. J. Ecol. 44, 515–522. 30. Owen-Smith, N. (2003) Elephants and ecosystems. In: Vandewalle, M. (Ed.), Effects of Fire, Elephants and Other Herbivores on the Chobe Riverfront Ecosystem. Proceedings of a Conference Organized by The Botswana-Norway Institutional Co-Operation and Capacity Building Project (BONIC). Botswana Government Printer, Gaborone, 17–23. 31. Phillipson, J. (1973) The biological efficiency of protein production by grazing and other land-based systems. The Biological Efficiency of Protien Production (Ed. by J.G.W. Jones) 217-235. Cambridge University Press, London. 32. Pringle, R. M., Young, T. P., Rubenstein, D. I. and McCauley, D. J. (2007) Herbivoreinitiated interaction cascades and their modulation by productivity in an African savanna. Proceedings of the National Academy of Sciences of the USA. 104, 193-197. 33. Rohner, C. and Ward, D.M. (1997) Chemical and Mechanical Defense against Herbivory in Two Sympatric Species of Desert Acacia. Journal of Vegetation Science. 8, 717-726. 34. Sankaran, M. et. al. (2004) Tree-grass coexistence in savannas revisited: insights from an examination of assumptions and mechanisms invoked in existing models. Ecology Letters. 7, 480-490. 35. Sankaran, M. et al. (2007) Determinants of woody cover in African savannas. Nature. 438, 846-849. 36. Scholes, R.J. and Archer, S.R. (1997) Tree-grass interactions in savannas. Annual Review of Ecology and Systematics. 28, 517-544. 37. Sinclair, A.R.E. (1975) The Resource Limitation of Trophic Levels in Tropical Grassland Ecosystems. The Journal of Animal Ecology. 44, 497-520. 38. van de Koppel, J. and Prins, H.T. (1998) The importance of herbivore interactions for the dynamics of African savanna woodlands: a hypothesis. Journal of Tropical Ecology. 14, 565576. 39. van Langevelde, F. et. al. (2003) Effects of fire and herbivory on the stability of savanna ecosystems. Ecology. 84, 337-350. 40. Verweij, J.T. et. al. (2006) Grazing lawns contribute to the subsistence of mesoherbivores on dystrophic savannas. Oikos. 114, 108-116. 41. Ward, D., and Young, T.P. (2002) Effects of large mammalian herbivores and ant symbionts on condensed tannins of Acacia drepanolobium in Kenya. Journal of Chemical Ecology. 28, 921-93. Literature Cited – Page 9 42. Young, T.P. and Okello, B. (1998) Relaxation of an induced defense after exclusion of herbivores: spine length in Acacia drepanolobium. Oecologia. 115, 508-513. 43. Young, T.P. et. al. (2003) Effects of natural and simulated herbivory on spine lengths of Acacia drepanolobium. Kenya. Oikos. 101, 171-179. Literature Cited – Page 10 Megan Cathleen Gittinger Department of Zoology, University of Florida 411 Carr, Gainesville FL, 32611-8525 [email protected] Professional Preparation: Virginia Polytechnic Institute University of Florida Employment: • 08/2007 - present • 08/2007 - present • 11/2005 - 05/2007 • 04/2005 - 08/2005 • • • 02/2005 - 04/2005 09/2005 - 11/2005 08/2003 - 12/2003 08/2004 - 12/2004 08/2002 - 12/2004 Wildlife Science Zoology B.S., 2005 Ph.D, in progress Ph.D. student, University of Florida Teaching Assistant, Biological Sciences, University of Florida GIS and Data Information Manager, WildAid Foundation Thailand Research Intern, Smithsonian Institution, Conservation and Research Center Research Assistant, Dept. of Plant, Soil and Insect Science, University of Massachusetts-Amherst Lab and Field Assistant, Biology Dept., Virginia Tech Field Assistant, Biology Dept., University of Virginia, Mountain Lake Biological Station Synergistic Activities: • During my time at WildAid Foundation Thailand, I was involved in a broad range of outreach and training activities including reporting and interviewing local stakeholders, instructing GIS and GPS application to a range of field and non-government officials, and providing input on national protected area management. I also co-authored an article on tiger conservation that was published in The Bangkok Post. • At Smithsonian Institute, I instructed the usage and application of Remote Sensing technology to local high school teachers in order for them to incorporate novel science technology in their classes. • I trained and mentored several undergraduates while working on my undergraduate research project. • While working as a camp counselor, I designed and instructed classes on Botany and Ecology to students ages 10-17. Adviser: Todd Palmer, University of Florida, Florida Biosketch – Page 11 FOR NSF USE ONLY 5 4 SUMMARY PROPOSAL BUDGET ORGANIZATION PROPOSAL NO. DURATION (MONTHS) University of Florida Proposed PRINCIPAL INVESTIGATOR/PROJECT DIRECTOR Granted AWARD NO. Megan Gittinger 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. Megan Gittinger 36 2. 3. 4. 5. 6. ( ) OTHERS (LIST INDIVIDUALLY ON BUDGET EXPLANATION PAGE) 7. ( ) 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. (1) 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.) Funds Funds Requested By Granted by NSF Proposer $64,500 (If Different) $ 10,800 96,800 Field equipment Leaf nutrient analysis Leaf tannin analysis 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 1. MATERIALS AND SUPPLIES 2. PUBLICATION/DOCUMENTATION/DISSEMINATION 3. CONSULTANT SERVICES 4. COMPUTER SERVICES 5. SUBAWARDS 4,000 2,000 12,000 TOTAL PARTICIPANT 5,696.52 15,050 20,746.52 135,546.52 6. OTHER TOTAL OTHER DIRECT COSTS H. TOTAL DIRECT COSTS (A THROUGH G) I. INDIRECT COSTS (F&A) (SPECIFY RATE AND BASE) TOTAL INDIRECT COSTS (F&A) J. TOTAL DIRECT AND INDIRECT COSTS (H + I) 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. Salaries 1. Principal Investigator: I will be the PI throughout the project and spend 4 months total each year in Kenya. The remainder of the year I will spend at University of Florida. To dedicate my time fully to the project, I am requesting a full-time salary (12-months) at $25,000/year. 2. Undergraduate assistant: I will rely on an undergraduate assistant at Univeristy of Florida to process field collections, enter data, and assist in data management and analysis ($7.50 / hr for 1 academic year). Exceptional undergraduates will have the opportunity to visit the field site in Kenya as research assistants. Travel: Two round-trip tickets to Kenya will cost approximately USD 4,000, totally USD 12,000 throughout the project. In addition to airfare, transportation on the ground in Kenya (USD 200 from Nairobi airport to Mpala field station) as well as in Florida (USD 100 from Gainesville to Orlando airport) is necessary. I have also budgeted USD 1000 for domestic travel to one national meeting per year (for two years), at which my results will be presented. Materials and Supplies: Supplies for experiments include but are not limited to large clippers, tape measures, non-toxic fluorescent stain (RiskReactor®; odorless, taste-less, EPA-approved), dung collection bags, analysis of leaf nutrient levels (total N and P using Kjeldahl digestion), and analysis of condensed tannins (methods in Ward, 2002). These total approximately USD 4,000. Other direct costs Station fees (Materials and Supplies): Mpala station fees for students currently amounts to USD 474.71 per month. Total station fees are USD 5,696.52 for 3 years (4 months per year). Kenyan field assistant: The Kenyan participant will be a staff field assistant from the Mpala Research Centre, who will accompany researchers in the field, act as a lookout for dangerous wildlife, and assist in field studies ($200/mo for 3 years). For safety reasons, the MRC requires that all field researchers be accompanied by a station guide. Vehicle license fees, insurance, petrol, and maintenance: I will be able to use the current field station vehicle (Land Cruiser pickup truck. The poor condition of Kenyan roads and the high price of spare Toyota parts makes maintaining a reliable vehicle relatively expensive, but the vehicle is critical to my ability to conduct this project in an efficient and safe manner. Based on my lab’s many years of experience in Kenya, I anticipate that it will cost approximately $200 per month to help pay for license, insurance, maintainence, and fuel the project’s field vehicle. Miscellaneous: Visa fees ($50/ visit/person) and costs of frequent communication between project personnel in Kenya and the U.S. (mail, telephone, and FAX; $400) are included in this category. In addition, I will need to contribute to the renewal of the 2year Kenya Research Clearance at a cost of $100. Budget Justification - Page 12 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: Megan Gittinger Support: Current Pending Submission Planned in Near Future *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. 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: Mpala Research Station The Mpala Research Centre (MRC) is a facility for scientific research, education, and training in central Kenya. Clinical: Animal: Computer: Office: University of Florida, Department of Zoology. Other: MAJOR EQUIPMENT: List the most important items available for this project and, as appropriate, identify the location and pertinent capabilities of each. 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. 5 6 NSF Form 1363 (10/99)
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