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Bird communities of isolated cypress wetlands along an urban gradient in hillsborough county, florida
h [electronic resource] /
by Nathaniel Goddard.
[Tampa, Fla] :
b University of South Florida,
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Thesis (M.S.)--University of South Florida, 2010.
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ABSTRACT: Migratory bird communities are sensitive to landscape alteration. Urban development significantly impacts bird communities on breeding grounds, as well as en-route during migration. One current theory is that Neotropical migratory birds are not limited by breeding or wintering habitat constraints but by food and habitat availability along major migration routes. The eastern flyway is the route taken by neotropical land-birds through eastern North America that follows coastal areas denoted by intense urban development. Coastal areas funnel birds to major departure points along the northern coast of the Gulf of Mexico and the western coast of Florida. Birds were monitored for 12 consecutive months along a decadal time gradient of urban development. Cypress domes are present through a broad scale of urban development in Hillsborough County creating ideal natural sampling units for long term monitoring of wetland bird communities in urban areas. Residential non-migratory bird communities were least influenced by development and did not change significantly with urban development. Neotropical and short-distance migratory birds, however, declined significantly for both richness and bird abundance with increased urban land cover. Migratory birds positively correlated with forested area at a spatial scale of 500 meters surrounding sites. Wintering migrants hit a critical point in development between 10 and 20 years of age, after which they disappeared. Neotropical migrants were most sensitive to declines significantly at sites classified as heavily degraded by the UMAM (Uniform Mitigation Assessment Method) a 'wetland integrity index'.
Advisor: Thomas Crisman, Ph.D.
Neotropical migratory birds
x Biology Integrative
t USF Electronic Theses and Dissertations.
Bird Communities of Isolated Cypress Wetlands Along an Urban Gradient in Hillsborough County, Florida by Nathaniel L. Goddard A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science Department of Inte grative Biology College of Arts and Sciences University of South Florida Major Professor: Thomas Crisman, Ph D Earl Mccoy, Ph D Mark Rains, Ph D Date of Approval: March 29, 2010 Keywords: Neotropical migrant, urban impacts, land use Copyright 2010, Nathaniel L. Goddard
i Tab le of Contents List of Ta bles i ii List of Figures i v Abstract v Introduction 1 Methods 6 Study Area 6 Water Quality 7 Vegetation 7 Wetland Habitat Assessment 8 Land use 9 Birds 1 0 Data Analysis 11 Results and Discussion 1 3 Water Quality 13 Wetland Vegetation 16 Land Use 16 Wetland Condition Index 18 Bird Community 20 Community Statistics 21 Feeding Strategies 2 7 Residents 28 Wint ering Migrants 30 Neotro pical Migrants 31
ii Summary and Concl usions 33 References 36 Appendices 42 Appendix A. Bird Di stribution A mong Sites 43
iii List of Tables Table 1 Wetland C haracteristics and Land Use 16 Table 2 Bird Species and Individual Abundance 25 Table 3 Migratory B ird G uild R elation ships to L and C over 28
iv List of Figures Figure 1 Locations of 16 Study Cypress D omes 7 Figure 2 Wetland pH and Conductivity 13 Figure 3 L and Cover with in 500 Meters of Wetlands 17 Figure 4 Wetland UMAM Values and Age of D evelopment 20 Figure 5 Monthly Species Richness Urban versu s Rural 22 Figure 6 Monthly Species Richness by Migratory Guild 24 Figure 7 Bird Diversity of Wetland C l asses 26 Figure 8 R esident Abundance and Richness 29 Figure 9 Wintering Bird Abundance by C lass 31 Figure 10 Neotropical Bird A bundance and Richness 32
v Bird Communities of Isolated Cypress Wetlands Along an Urban Gradient in Hillsbo rough County, Florida Nathaniel L. Goddard ABSTRACT Migratory bird communities are sensitive to landscape alteration. Urban development s ignificantly impact s bird communities on breeding grounds as well as en route during migration. One current theory i s that Neotropical migratory birds are not limited by breeding or wintering habitat constraints but by food and habitat availability along major migration routes. The eastern flyway is the route taken by neotropical land birds through eastern North Americ a that follows coastal areas denoted by intense urban development. Coastal areas funnel birds to major departure points along the northern coast of the Gulf of Mexico and the western coast of Florida. Birds were monitored for 12 consecutive months along a decadal time gradient of urban development Cypress domes are present through a broad scale of urban development in Hillsborough County creating ideal natural sampling units for long term monitoring of wetland bird communities in urban areas. Resident ial non migratory bird communities were least influenced by development and did not change significantly with urban development. Neotropical and short distance migratory birds, however, declined significantly for both richness and bird abundance with incr eased urban land cover. Migratory birds positively correlated with forested area at a spatial scale of 500 meters surrounding sites. Wintering migrants hit a critical point in development between 10 and
vi 20 years of age, after which they disappeared. Neo tropical migrants were most sensitive to declines significantly at sites classified as heavily degraded by the UMAM (Uniform
1 Introduction Freshwater wetlands are common landscape featu res of southwest Florida and have important environmental functions They filter water and sequester contaminants reducing watershed pollution and eutrophication (Gopal 1999). Wetland habitats promote floral and faunal diversity in general ( Ewel and Odum 1984), and m any birds utilize them for nesting, roosting, and feeding. Forested wetlands are important foraging and roosting areas for many residentia l and migratory bird species. En route n eotropical migratory birds rely on these habitats key stopover p oints as resting and foraging areas during spring and winter migrations (Bryce and Hughes 2002). Forested wetlands also serve as wintering grounds for short distance migratory birds who wintering in the southeastern United States (Buler et al. 2007). W etlands declined by 53% i n the continental United States from 1780 to 1980 (Dahl 1990). Florida suffered the greatest loss of any state (3.8 million hectares) (Dahl 1990) in the past two decades Florida lost 44,500 hectares of forested wetlands 59% attr ibuted to urbanization (Kautz et al. 2007). Urban development is a significant ongoing threat to wetlands of the United States especially Florida, because of disproportional human populations (54%) living in coastal areas (Crossett et al. 2004). The T ampa/St. Petersburg metropolitan area in southwest Florida includes parts of Hernando, Hillsborough, Polk, and Pinellas counties and has experienced a 26% population increase in the past two decades contributing to its current population of approximately 4 million (Claritas 2008). I n 1986 23% of land area was covered by freshwater wetlan ds (Haag et al. 2005), 90% of which were considered isolated, lacking
2 direct surface water outlets downstream under no rmal flow conditions (Leibowitz 2003). Of th ese, 79% were less than 2 hectares in size. Such small isolated wetlands mostly consist of either open freshwater marshes or cypress domes. The study area in Hillsborough County was originally covered primarily by pine flatwoods, palmetto prairie, and forested we tlands (Xian et al. 2007 Friesen et al. 1995). As urban development increased within the urban landscape compared to upland forest, and ha ve become increasingly isolated from one another and other natur al surroundings Most of the surrounding upland forest and scrub has been converted to urban development isolating cypress domes and removing natural wildlife corridors. Urban development e ffect s bird community altering species richness, abundance and d iversity within incorporated wetlands (Whited et al. 2000). Sensitivity of birds to urbanization varies greatly, and w hile density and nesting rates can be higher in urban areas, species richness is often greatly reduced (Gravereski 1976, Bessinger and Os born 1982, Chace and Walsh 2006). Urbanization favors both synanthropic non migratory species and exotic species such as house sparro ws, rock pigeons, and European starlings (Garaffa et al. 2009), while excluding many species sensitive to human disturbanc e such as neotropical migratory wood warblers (Parulidae) (Rottenborn 1999, Allen and C onner 2 000, Whited et al. 2000). Bird guilds most tolerant to urban environments are mainly granivores, omnivores, and areal insectivores that can better utilize the urban
3 Urban development and surrounding land use have far greater impacts on migratory bird species opposed to residential communities Neo tropical migratory songbirds are most affected by urba n development while short distance wintering migrants such as American robin ( Turdus migratorius ) and yellow rumped warbler ( Dendroica coronata ) are to a lesser degree (Whitcomb et al. 1981, Flather and Sauer 1996). En route n eotropical migratory birds a re particularly susceptible to changes in land us e and decrease drastically with increased urbanization. On the southeastern Atlantic and G ulf coast of North America this poses a problem because neotropical migratory species utilize coastal areas such as the Tampa/St. Petersburg metropolitan area for rest stops and foraging before open ocean flights. The time spent at stopover sites greatly exceeds flight time and determines the duration of total migration time (Alerstam 1981, Buler et al. 2007). Neotrop ical migrants must build fat reserves during fall migration before embarking on long open ocean flights through the Gulf of Mexico and Caribbean (Moore and Kerlinger 1987). D uring s pring these same wetlands are used to replenish fat reserves depleted dur ing the return flight. Habitat quality and food availability during stopovers determine the rate at which migrants replenish lost fat reserves. Birds that can recover faster and build reserves are able to spend more time on the breeding grounds, increasi ng reproductive potential (Moore and Kerlinger 1987, Kuenzi and Moore 1991). N eotropical migratory birds that breed in northeastern North America and winter in the Caribbean, Central and South America have declined in recent decades (Robbins et al. 1989 Askins et al. 1990, Peterjohn et al. 1995). Both local and national studies using data from the North American Breeding Bird Surveys indicate declines in many
4 neotropical migratory species (Finch and Stangle 1993, Robbins et al. 1989). From 1980 to 199 1, 35 specie s of Class A neotropical land birds declined, excluding s wimmi ng and most wading birds, while o nly 15 species have increased in numbers (Peterjohn et al. 1995). Several species of w ood warblers which comprise the greatest proportion of neotro pical migrants have decline d in recent decades (Moore and Kerlinger 1987) Significant declines in Cerul ean Warbler ( Dendroica cerulea ) and Prairie Warbler ( Dendroica pinus ) populations have been observed but d efinitive reasons for these are not yet esta blished (Robbins et al. 1992). D eclines are speculated to involve habitat destruction of breeding territory in North American and wintering territory in the Caribbean and South America. However, habitat utilization along complete migratory route s is an i mportant factor for bird migrations that is not yet well understood (Moore and Kerlinger 1987 Newton 2006 ) Migration is a time of extremely high bird mortality, in black throated blue ounting for 85% of total annual mortality (Sillett and Holmes 2002) Major reasons for high mortality rates during migration include unfavorable weather conditions, predation, and food availability. Variation in food availability at stopover sites is oft en density dependent leading to longer stopover times as migration intensity increases (Newton 2006). In woodland sites along the Gulf of Mexico, passerine migrants rapidly depleted insect abundance by up to 67% during periods of high migratory intensity (Moore and Young 1991).
5 Urban growth results in conversion of natural habitats into manage d urbanized systems N ative forested wetlands are reduced and fragmented into heterogeneous disjunct patches or islands (Alig and He aly 1987 and Garaffa et al. 20 09). Neotropical migratory birds take visual cues for habitat quality based on various factors including patch size, vegetative structure, adjoining upland tree cover, distance to roads, and urban intensity ( Askins et al. 1990, Mills et al. 1991, Whited e t al. 2000, Marzluff and Ewing 2001 ). Bird species richness has been decreases with increasing urban land cover, while diversity increases with increasing tree development, even in urban settings (Donnelly and Marzluff 2006, Pennington et al. 2008). The p urpose of this study was to determine the effects of urban development on bird communities utilizing isolated cypress domes in Hillsborough County, Florida. Since bird guilds respond differently to disturbance the effects were measured for each migratory b ird guild by computing total abundance and species richness. A gradient of urban intensity was used to explore how land cover and local site features including vegetation and wetland area influence site selection by residential, wintering short distance m igrants, and en route neotropical migratory birds. Finally bird species richness was compared to a wetland integrity index (UMAM) to determine its value as an estimator for bird utilization.
6 Methods Study Area This study was conducted on cypress dome wetlands within the Tampa Bay metropolitan area of northern Hillsborough County, Florida, an area of expanding urban development in a landscape rich in small, forested cypress wetlands (Haag et al. 2005). Cypress domes make up a substantial portion of re maining forested (88%) and wetland (52%) areas in urbanized northern Hillsborough County (FGDL 2006). Cypress domes are depressional, shallow forested wetlands with longer hydroperiods than freshwater marshes (Ewel and Odum 198 4 ). They are typically hydro logically isolated from other wetland and riverine features of the landscape and are dominated by pond cypress ( Taxodium ascendens ) and bald cypress ( Taxodium distichum ), with swamp tupelo ( Nyssa sylvatica var. biflora ) often co dominant. Other common can opy and sub canopy tree species include red maple ( Acer rubrum ), dahoon holly ( Ilex cassine ), and swamp bay ( Persea palustris ).
7 Figure 1. Locations of S tudy S ites in Hillsborough County FL. Study areas were analyzed for impacts to bird communiti es related to urban development. The sixteen sites which ranged between 0.5 and 4.0 hectares were selected based on the decade initial development began within a 200 met er radius of the site. Four reference sites were located in the Lower Hillsborough Flood Detention Area, conservation land owned by Southwest Florida Water Management District, which is devoid of major urban development and/or agricultural activity (FGDL 2006).
8 Water Quality Wetland areas were determined using Arc GIS 9.2 and Florida Wetland Inventory Map data. Monitoring stations were established at the deepest point of each wetland where water levels and quality were surveyed monthly during perio ds of inundation. Water quality data (pH, conductivity, and dissolved oxygen) were collected using a YSI 6920 V2 multi meter sonde. Vegetation Analysis Wetland tree community parameters included canopy cover, species composition, basal area. Canopy c omposition and mortality were assessed using cross sectional belt transects radiating from the wetland center in the four primary directions. The wetland center was established using Florida W etland Inventory polygons and a hand held Garmin Etrex GPS was used to find data points in the field. Fallen trees were counted only if the base originated within or intersected the transect line. Belt transects ran from the center to the wetland edge and were 5 meters wide divided into 10 meter long sub plots arran ged linearly along the transect. All trees within transects with diameter greater than 2 cm were identified to species and measured for diameter at breast height. The latter were used to estimate average basal area for each wetland as a measure of forest density. Ten density measurements were made randomly along transect lines for each wetland using a hand held densiometer to calculate average canopy density. Wetland Habitat Assessment Habitat assessments have been widely used to estimate wetland integr ity and value to wildlife (Lonard et al. 1981). Bird species richness correlates with wetland
9 habitat assessment values and has been used as an indicator group in environmental indices such as the Habitat Assessment Technique, HAT (Cable et al 1989). The wetlands of this study were evaluated using the Florida Department of of wetlands li kely to be impacted by development. It evaluates wetlands based on three 0, with 1 considered a pristine wetland and scores near 0 for heavily degraded sites with low functional value. For this study, UMAM scores were used to determine relative status of wetlands and degree of impact to the wetland and surrounding area. Indicators of location and landscape support include: support of surrounding habitat, invasive and exotic plant species presence in proximity to the wetland, wildlife access (presence or absence of barrier), adverse impact of landscape on wildlife, and hydrological connectivity. Wetland water environmental indicators include: water quantity, timing of inundation, frequency, duration, depth, and saturation of soils. Finally, community structure indicators include: wetland plant cover, proportional presence of invasive and exotic species, health of plant community (stress or increased mortality) and recruitment. Land use Digital land use and cover data from the Florida Geographic Data Library (FGDL) and the U. S. Geological Survey were used to analyze surrounding land use.
10 Hi llsborough County Property Appraiser maps and FGDL parcel maps were used to determine the average age of the earliest 25% of urban development within 500 meters of each study wetland. Land use within the 500 meter buffer was determined using the 2006 Flor ida Land Use, Land Cover Classification System (FLUCCS) map layer, and assigned to three categories of land use for analysis: urban, forested and open water/open land. Percent tree coverage was determined using high altitude area photographs projected on to digital orthophoto quarter quadrangle (DOQQ) maps. A 1 km fishnet grid containing 1600 individual pixels, each 25m x 25m, was overlain on top of DOQQ map layers and centered on each study area. Sites were analyzed for percent tree cover and imperviou s surface area (Donnelly and Marzluff 2006, Botsford 2000). Tree cover cover and 20 to 60% impervious structure, and open land ( urban cover and 0% impervious surfaces (Botsford 2000). Birds Avian surveys were conducted monthly from September 2008 to August 2009. Fixed radius point counts with a maximum radius of 40 meters (Whitcomb et al. 1981), established using a Nikon 550 handheld laser range finder. The area surveyed was approximately 0.50 hectares, which approximates the size of the smallest wetlands in the study. A single bird sampling station was selected at a midpoint between wetland edge and center for each wetland. Birds were surveyed using auditory and visual census techniques for species and individual counts via standard point count techniques (Bibby et al. 2000). Only birds utilizing wetlands were counted in surveys, and both flyovers
11 and those detected beyond the wetland boundary were excluded. Each survey was conducted for at least 10 minutes within the first 3 hours of daylight, coinciding with the period of greatest avian activity (Bibby et al 2000). Point counts began following a 5 minute rest period after arrival at each s ite so birds could re acclimate after disturbance. All bird surveys were conducted by a single observer for consistency. Data Analysis Bird data were summarized for each site by individual abundance and species richness and then grouped by feeding stra tegy and migratory guild. Migratory bird guilds were classified as residential non migratory, neotropical migrants, and short distance wintering migrants according to Whitcomb et al. (1981) and the American Species richness was calculated using total species present for each wetland and compared with predicted species richness extrapolated using jackknifing methods described by Zahl (1977) to determine community representation. Birds were also assigned a feeding guild ( inse ctivorous, omnivorous, granivorous, carnivorous, or piscivorous ) for further analysis. Species richness and abundance present w ere selected as dependent variables for data analysis. Diversity was determined using dices for relative comparison among sites. Bird communities were compared cumulatively and monthly between urban and rural sites. Independent variables corresponded to age of urban development, intensity of urban development (percent urban landcover), w etland size, tree cover in the surrounding landscape, and UMAM wetland int egrity score. Forested area spatial distribution within 500 meters of sample sites was determined using the T Square distance method ( Diggle 1976 we re used to evaluate correlations between
12 landscape variables and wetland index parameters. Multiple regression models were constructed to evaluate relationships between bird migratory groups and land cover variables using PA SW 18 (SPSS) statistical softwa re significance was determined using standard 95% confidence interval s
13 Results Water Quality Due to drought conditions during the study, w ater quality data were only collected from August 2008 through October 2008 when most study w etlands were inundated. Both pH and conductivity were significantly greater in urban than reference wetlands. Wetland pH quickly increased with onset of development (reference mean pH 4.8, urban mean pH 6.3), then remained relatively constant between 6.1 and 6.9 among urban wetlands with no significant differences regardless of development age (Figure 2 a ). Conductivity also significantly increased immediately with initiation of urban development from a mean of 70 S/cm 7.6 in reference wetlands to a mean of 222 S/cm 118 for all urban classes combined (Figures 2b). Wetland 2000 2 had greatly elevated conductivity (566 S/cm 387) coinciding with ongoing construction on surrounding property It was the only site near active construction and had significantly higher conductivity than all other urban sites 269 S/cm ) in spite of silt fences designed to reduce runoff and erosion around its perimeter Urbanization leads to multiple non point and point source i mpacts on wetland water quality. I t increases overland flow associated with impervious surfaces leading to deposition of contaminants in wetlands and elevated conductivity (Lee et al. 2009). U se of groundwater for lawn maintenance also can contribute to increased pH in wetland s and lakes ( Martin et al. 197 6 ).
14 Figure 2. Wetland pH and C onductivity. standard deviations Wetland V egetation Pond cypress was the dominant tree in all wetlands, contributing 81% 9 of total basal area; however, the proportionality of recruit size class cy press (dbh < 8 cm) declined with urban development. Both facultative and facultative wet trees species increased proportionately for the recruit size class of the oldest two urban development Pond cypress contributed 90% of t he recruit size class for
15 class trees, while subdominant trees increased including s wamp tupelo ( 35% ) and red maple ( 13 %) These tw o species, along with wetlands, only in lower concentrations. The 1970 wetlands suffered the greatest decline in cypress regeneration to 7% of the recruit size class. Most red maple (62%) Chinese tallow Sapidium sebiferum (20%) and Brazilian pepper Schinus terebenthifolius (11%). The latter two are invasive exotic species and were found in only in the oldest two wetland classes (1970 Exotic invasions have been attributed to urban h abitat isolation, degraded water quality, and altered hydrology in other studies of Tampa area wetlands (Haag et al. 200 5 Rochow 1994). The decline in dominance of pond cypress associated w ith geographic isolation within a mosaic of urban development, combined with increased exotic species, are indicative of losses to cypress dome functionality (Jubinsky and Anderson 1996 ). The shift from cypress dome to mixed wetland hardwoods in developed areas older than 20 years suggest long term alteration to wetland hydrology typically associated with a decrease in annual hydroperiod. This is a problem that commonly takes decades to appear and is not easily corrected (Ewel and Odum 1984). Urban isola tion of forested wetlands often leads to local extirpation of native flora and facilitates invasion by early successional and exotic species (Ehrnfeld 2000).
15 Table 1. Wetland characteristics and land use by age of surrounding urban development. Site Cl ass Time of Initial Develop ment Wetland Area (ha) UMAM Site Quality Index Tree Basal Area (cm) Wetland Canopy Cover (Percent) Surrounding Wetland Area (ha)* Urban Land area* (ha) Forest Tree Cover 70%* (ha) Low Density Forest 70% > Tree Cover > 20 % (ha ) Open Land Tree Cover ) R 3 Ref. 0.95 0.96 47 91 17.25 0 86.19 11.5 2.31 R 4 2.26 0.99 58 95 33.38 0 90.19 8.5 1.31 R 5 5.56 0.96 78 90 33.95 0 87 8.5 4.5 R 6 3.61 0.94 69 88 39.05 0 87.31 11.63 1.06 2000 1 2000's 2000 1.79 0.74 55 91 19.05 62.77 30.25 16.31 53.44 2000 2 2005 4.31 0.82 45 73 21.93 66.62 40 14.19 47.06 2000 3 2000 2.53 0.81 68 80 29.75 38.87 42.5 20.88 36.63 1990 1 1990's 1990 1.15 0.78 51 84 15.81 70.60 19.44 19.13 61.44 1990 2 1997 2.51 0.76 36 40 29.49 54.88 26.44 15.75 57.81 1990 3 1997 0.56 0.77 41 90 15.84 77.28 20.38 12.44 67.19 1980 1 1980's 1987 1.68 0.76 67 91 30.27 54.22 36.5 21.94 41.56 1980 2 1989 0.5 0.77 72 93 30.28 78.75 44.75 17.06 41.31 1980 3 1984 0.75 0. 59 52 93 15.19 57.48 17.38 41.38 43.75 1970 1 1970's 1974 2.75 0.46 76 82 7.77 83.88 23.94 11.44 36.13 1970 2 1978 1.9 0.35 55 92 9.78 86.33 18.19 32.88 48.94 1970 3 1978 2.58 0.44 72 84 6.56 89.98 16.63 41.56 41.81 Within 500m radius of study we tland
16 Land U se The density of urban development within 500 meters of study wetlands significantly 12, p = 0.03) Total urban area classified as medium to high density urban by the Florida Land Use Cover Classification System (FLUCCS) (FGDL 2004) was used to quantify urban land cover. Urban areas of Tampa Bay are characterized by low le vels of forested level of urban land cover within 500 meters, while recent classes wetlands 80 1 and 80 2 having less open land and more forested area than expected due to their close proximity to the Hillsborough River and its extensive rip arian forest. Landscapes surrounding urban sit es differed from those o f reference sites by the presence of expansive open land among the former wetlands had 84% to 90% open land area (within 500 m), recent and intermediate development classes (2000 1980) showed moderately less open land between ( 39 79% ) while reference sites had < 1% open land within the surrounding landscape.
17 Figure 3. Percent Land Cover Within 500 Meters of W etlands Along a Gradient of Increasing A ge o f Urban D evelopment. surrounding urban sites had an reference sites (Table 1). Urban land cover i ncreased logarithmically converting 56% 15 of the landscape to urban within the first 10 years after development (Figure 3 ). U 0.721, n = 16, p = 0.002 ). The remaining tree cover was either clumped into isolated patches, or transformed into low density urban forest (tree co is not suitable habitat for many forest dwelling migratory bird species (Marzluff and Ewing 2001) Spatial landscape analysis using the T square index of spatial pattern (C), within
18 500 meters of wetlands showed that forested was the dominant habitat in rural reference areas and was randomly distributed (C = 0.433, p = 0.20), but fore st area was clumped in urban environments (C = 0.666, p <0.05) into remnant isolated patches. The expansion of u rban development in Hillsborough County in recent decades (Xian et al. 2007) has led to significant losses in forested area ( Figure 3 ) while urban impervious surfaces have increased within the watershed Garcia Fresca (2005) found that, in urban areas, during rain events impervious surface area increased overland flow, erosion, and sediment deposition into local water bodies, and decreased gr ound water recharge. Wetland Condition Index Significant negative correlations were found between UMAM wetland index values and ages of development (R = 0.85, s = 0. 14 p < 0.001) (Figure 4). Reference wetlands had the highest scores ( UMAM = 0. 9 6 0.02 ) indicating little to no impact to wetland condition The moderate disturbance ( UMAM = 0.78 0.03 ) ; the most extensive impacts were decreased water quality and urban isolation. The wetlands h ad moderate disturbance ( UMAM = 0.70 0.10 ) with degraded sites structure, water quality, and moderate habitat isolation. T he the lowest scores ( UMAM = 0.41 0.06 ) showing significant losses to system integrity, associated with habitat isolat ion poor water quality, and abundant invasive and exotic plants. In 1984 rules were revised by state agencies to include general wetland permitting. Wetland r esource permitting was further streamlined by the Florida
19 Departm ent of Environmental Regulat ion in 1992 to jointly permit storm water systems and protect wetlands concurrently in response to the Warren Henderson Wetlands Protection Act (SWFWMD 2008). Cluster analysi s of wetlands UMAM scores grouped wetlands into two urban groups urban prior to 1984 (UMAM = 0.42 0.06, n = 3) and after 1984 (UMAM = 0.75 0.06, n = 6) and one rural group (UMAM = 0.96 0.02) Though these scores did coincided with state regulatory dates for general wetland permitting they also strongly fit a linear regression (R = 0.85), and it is beyond the scope of this project to suggest any impacts of these policies. Urban development can negatively impact cypress dome and their bird communities by altering water chemistry, vegetative structure, hydroperiod and surrounding land use ( Lonard et al. 1981 ) which decreases wetland functional values to their landscapes. Migratory birds, both en route neotropical migrants and wintering short distance migrants, are extremely sensitive to alterations in forested wetland and riparia n structure and decline rapidly with this type of urban development (Rodewald and Matthews 2005).
20 Figure 4 Wetland UMAM Values and Age of Urban D evelopment. Bird C ommunity A total of 2 528 individuals representing 7 9 bird species were recorded over the 12 month sampling period at the 16 study sites (Appendix A) Non migratory permanent residents made up the majority of bird detections (68% abundance 36 species ) wintering migrants were second ( 21% 13 species ) and en route neotropical migrants comp ri se d 11% of observations from 30 species. Six common resident species were found at all sites and made up 53% of total observations In decreasing order of occurrence they were : Carolina w ren (Thryothorus ludovicianus) blue gra y gnatcatcher ( Polioptila caerulea ), northern cardinal ( Cardinalis cardinalis ), tufted titmouse ( Baeolophus bicolor ), blue jay ( Cyanocitta cristata ), and red bellied woodpecker ( Melanerpes carolinus ). Eleven permanent residents were present at 75% of sites Only 3 species of
21 mig ratory birds were present at 50% of urban sites: gray catbird ( Dumetella carolinensis ), black and white warbler ( Mniotilta varia ), and northern parula ( Parula a mericana ). Community Statistics Bird communities declined with conversion of forest to open land and urban low density urban forest Species richness negatively correlated with open land (R = 0.61, p = 0.01) and urban for est area (R = 0.65, p = 0.01) and positively co rrelated with forest area (R = 0.74, p = 0.01). Mean s pecies richness decline d over time in wetlands after initial development from reference wetlands (S = 33 2.2), to recent development to 3.7). Observed species richness for each wetland was ext rapolated using the jackknifing technique ( Zahl 1977 ) to estimate predicted species. The latter did not differ significantly from the observed ( chi square X = 3.57, p = 0.31, df = 3) indicating adequate representation among development groups Monthly s pecies richness (per sampling event) also significant ly declined with onset of urban development, and was also highest at rural sites (S = 10 3.1) intermediate at recent developmen 2.2) and lowest at older development sites ( 1980
22 Figure 5. Monthly Species Richness of Urban and Rural sites Monthly species richness showed distinctly different trends for combined urban versus rural sites (Figure 5). Reference wetlands had significantly greater species ric hness during spring and summer coinciding with migration and breeding seasons of many birds especially migrants that showed stronger preferences for rural sites. Spring and summer are times of intensive calling by territorial males, which increases detec tion potential for reclusive species and upper canopy birds ( Emlen 1971 Farnsworth et al 2002). Common nesting migrants included black and white warbler, northern parula, palm warbler, prothonotary warbler ( Protonotaria citrea) yellow throated vireo ( vir eo flavifrons ), and red eyed vireo ( Vireo olivaceus ) that, with the exception of the northern parula, showed strong selective preferences for reference sites. Neotropical and wintering migrant species richness were significantly lower in urban wetlands d uring peak migration (Figure 6) Neotropical richness increased in rural
23 wetlands from April to June coinciding with spring migration. Neotropical migrant species richness over the entire year for urban wetlands showed a max species richness of 0.83 1. 1 species compared to rural wetlands with 3.7 1.7 species. Urban migrant richness remained low during the entire study opposed to reference migrants. Total abundance follo wed a similar trend as richness. B irds declined as landscapes surrounding wetla nds transitioned from rural to urban, similar to results by Donnelly and Marzluff (2006), and Bryce et al. (2002). Abundance declined steadily before leveling off and ed in community composition.
2 4 Figu re 6 Monthly Species R ic hness by Migratory G uild
25 Old development had proportionately greater resident bird populations while new had better winte r migrant representation. New development was more representative of reference sites while old developme nt resembled urban bird communities described in both the Mid Atlantic and Pacific Northwest (Cam et al 2000, Donne l ly and Marzluff 2004, and Dowd 1992). The significant differences in bird abundance between urban and rural areas observed here have been reputed in fewer than half of greater than 100 studies published on the topic (Marzluff e t al. 2001 ) Most reputed increased ab undance or only slight declines; however, they indicate substantial declines in species richness with urbanization. Table 2 Mea n Species Richness and Individual Abundance per Development G roup Site Class Totals Residents Migrants Wintering Neotropical Species Richness 1970's 19.3 15.7 1 2.7 1980's 19.3 13.3 1.7 4.3 1990's 22.7 14 5 3.7 2000's 26.3 14.7 5.3 6.3 Referen ce 32.5 12.7 7.7 12 Individuals Abundance 1970's 106.7 97.3 2 7.3 1980's 97 79 7.3 10.7 1990's 138.7 91.7 41.7 5.3 2000's 144 99.3 34.7 10 Reference 268.25 153.75 69 45.5
26 S displayed a similar pattern as species richne ss (Figure 7 ). Rural wetlands had the highest diversity and it was the lowest in s since d evelopment (R = 0.536, p = 0.001) and increased with forest area within the landscape (R = 0.57, p = 0.001) Diversity was significantly lower in cypress dome wetlands than in similar wetland deciduous forests of n 0.16) ( Tramer 1969), possibly due to fewer migratory species breeding in southwestern Florida. Simpsons index was used to assess dominance among species observed and did not significantly vary among wetland classes Figure 7 Bird D iversity of Wetland C lasses
27 Feeding Strategy F eeding strategies within bird communities changed markedly for residential and migratory bird populations along the rural to urban gradient Residential communities w ere composed primarily of omnivores (56% richness, 54% abundance ) followed by insectivores ( 14% richness, 42% abundance ) piscivorous wading birds (19% richness, 2% abundance ) and carnivores ( 11% richness, 2% abundance ) Both e n route n eotropical and s hort distance migrants wintering in Florida were primarily insectivorous passerines (92% and 83% respectively) and accounted for 84% of total insectivores. They spend the majority of the year on wintering grounds and en route ( Robbins et al. 1989 ) Insec tivorous p asserine m igrants included tyrant flycatchers, vireos, warblers and their allies that migrate to breeding grounds in northern latitudes during spring ( Sherry and Holmes 1995 ) coincid ing with insect emergences and longer day s for improved foraging potentials (Alerstam 2001 and Marra et al. 2005). I nsectivorous migrants travel south in autumn and w inter to avoid freezing temperatures and in sufficient insect availability (Alerstam 2001 Keunzi et al. 1991)
28 Table 3 M igratory Bird Guild R e lationships to Land C over. Migratory Guild Area Forest Urban Forest Open Land adj. R Species Richness en route migrants 0.18 0.90 ** 0.68 ** 0.80 ** 0.80 ** Wintering migrants 0.24 0.70 ** 0.86 ** 0.45 0.75 ** Residents 0.17 0.39 0.37 0.30 0. 07 Total Species 0.30 0.74 ** 0.65 ** 0.61 ** 0.60 ** Abundance en route migrants 0.04 0.91 ** 0.53 0.88 ** 0.78 ** Wintering migrants 0.16 0.72 ** 0.79 ** 0.51 0.58 ** Residents 0.17 0.80 ** 0.45 0.77 ** 0.54 Total Abundance 0.16 0.90 ** 0.65 0.77 ** 0.76 ** *. Significant at p = 0.05 level. **Significant at p = 0.01 level Resident Birds The abundance of resident birds positively correlated with forest area ( R = 0.80, p = 0.01 ) and negative ly with urban development ( R = 0.77, p = 0.05) within 500 m of study wetlands S pecies richness however, was not correlat ed with any land use parameter ( R = 0.07) There were no significant difference s in residential bird populations among urban classes suggesting they are more adept at living in urban environments (Garaffa et al. 2009, Pennington et al. 2008) R esidential bird species richness did not change significantly with onset of urban development ( Table 3 ) but their representation within the communities changed considerably with changes in urb an land use Both species richness and abundance were proportionally greatest in the two oldest development classes (Individuals 88%, Sp ecies 76%) (Figure 8 ) attributed to drastic declines in migrants and increas ed abundance of synanthitropic species ass ociated with urban areas as noted by (Lancaster and Rees 1979, Stratford and Robinson 2005 ) Wetlands in recent development were intermediate (Ind ividuals 68%, Sp ecies 58%) and reference wetlands had the lowest
29 proportional representa tion of resident birds (Ind ividuals 58%, Sp ecies 41%) though number of individuals did not decline Figure 8 Proportional species richness and abundance of resident bird Urban areas were characterized by expansive open land that differ ed from upla nd forests surrounding reference wetlands Such o pen area s are preferred habitats for ma ny residential omnivores including European starling ( Sturnus vulgaris ) (Whitehead et al. 1995) northern mocking bird ( Mimus polyglottos ) red wing ed blackbi rd ( Agela ius phoeniceus ) common grackle ( Quiscalus quiscula ) and boat tailed grackle ( Quiscalus major ) ( Melles et al. 2003 ) D ense edge habitats along forested wetland s in urban areas are preferred habitat of brown thrashers ( Toxostoma rufum ) a common edge speci es not found in interior forest s (Aldefer 2006) Wading birds were represented by 6 resident species including one federally endangered species the wood s tork ( Mycteria americana ) : lim pkin ( Aramus guarauna ) little blue heron
30 ( Egretta caerulea ) and white ibis ( Eudocimus albus ). Wading birds were found only in wetlands with water levels > 10 cm depth, which occ urred durin g only 29% of sampling events, and they showed no response to any other parameter. Wintering Migrants Wintering migrants ( short distance migrants ) include species wintering exclusively in the southeastern United States and n eotropical migra nts with substantial population s wintering southern Florida as well as the Ca ribbean, Central and South America They were predominantly insectiv ores (S = 11, 84% abundance) but did include two omnivor es, gra y catbird ( Dumetella carolinensis ) an e dge species that does very well in urban wetland habitats and American robin ( Turdu s migratorius ) whose preferred foraging habi tats include residential and open forested lands (Aldefer 2006) With these exception s the remaining 11 species (86% abundance) of wintering migrants were sensitive to urban alteration, including 5 species of warblers that accounted for 80% of wintering migrant observation s : prairie warbler ( Dendroica discolor ) palm warbler ( Dendroica palmarum ) pine warbler ( Dendroica pinus ) yellow throated warbler ( Dendroica dominica ) and yellow rumped warbler ( Dendroica c oronata ) Wintering migrant s showed sensitivity to landscape alteration by urban development declining in species richness with increasing low density urban forest ( R = 0.86 p = 0.01 ) consistent with previous studies (Pennington et al. 2008, Stratfo rd and Robinson 2005 ). They were positively correlated with forested area (R= 0. 70 p = 0.05 ) preferring reference sites (n = 276, sites = 4) and recent development ) (n = 230, sites = 6) over old development ( n = 29, sites = 6 ) The gray catbird accounted for the majority of old urban observations (79%) I t prefe r red urban sites to rural possibly
31 because of thick edge habitats surrounding urban wetlands lacking at rural wetlands W intering migrants show ed a critical change in site classes were except for gray catbird, they disappeared at older sites (Figure 9 ) Wintering migrant species richness did not respond to land use but did positively correlate with UMAM habitat index scores using 810, n = 16, p = 0.01 ) Figure 9 Winte r ing Bird Abundance Neotropical Migrants Neotropical migrants strongly correlated with land use within 500 meters of wetlands. Positive relation ships were found for b oth species richness ( R = 0.91, p = 0.001 ) and abundance ( R = 0.90, p = 0.005 ) versus forest area and negatively correlated with increase d urban area ( R = 0.88 p = 0.005 and R = 0.80 p = 0.005 respectively ) were noted Neotrop ical migrant abundance decline d from a mean of 43 12 birds per
32 site in rural wetlands to 73 in the wetlands (Figure 10 ). Species richness followed a similar pattern, declining from a mean of 12 1 species in rural wetlands to s. Figure s 10 Neotropical M igrant A bundance and R ichness by Wetland C lass
33 Migratory birds did not significantly respond to wetland size (abundance R = 0.04 p = 0.89 species richness R = 0.18 p = 0.49 ) which is consistent with a study of wood lot patch size in urban and rural area s of 4 to 20 hectares (Freisen et al. 1995). Neotropical species richness ) and abundance 16, p = 0.01 ) did positively correlate with wetland integrity index (UMAM) scores for individual wetlands Wood warblers (Parulidae) account ed for 70% of neotropical bird observations and 15 of 30 neotropical migrant species observed. Only one warbler northern parula, appeared insensitive to urban development and was observed at all urban sites of older classes It was the most abundant comprising 30% of neo tropical migrant observations and 50% from class wetlands. The remaining warbler species declined from 20 observations per site at reference sites to just 2 at old urban sites Wood warblers are a group of concern because they are highly migratory and sensitive to habit at alteration not only on their breeding and wintering grounds but along migratory routes ( Minor and Urban 2010 ) Some warbler s have declined significant ly in recent decades including cerulean and pine warblers (Peterjohn et al. 1995) along with the enda Dendroica kirtlandii ) and the presumed extinct Vermivora bachmanii ) Summary and Conclusions Many neotropical migratory birds use Tampa B ay as a staging area before embarking on open water fall migrations to the Caribbean C entral and South America It is also an important recovery area upon returning from wintering grounds during early
34 spring en route to breeding grounds in the northern temperate zone ( Moore et al. 1995 ). These stops just before and right af ter open ocean flights are vital to bird survival and are also important for recovery of lost fat reserves before contin uing to their migratory endpoints (Kuenzi et al. 1991). During migration birds expend great amounts of energy traveling through unfami liar terrain with variable food availability. S topover habitats are vital during migration a time of extremely high mortality for birds accounting up to 80% of annual mortality (Sillett and Holmes 2002 ) Habitats along the migration route can act as nut rient bottlenecks, limiting available resources for late migrants (Alerstam and Hedenstrom 1998) Low habitat quality at stopover sites along migratory route greatly extend stopover time and decrease possible time spent on wintering and breeding ranges (M oore and Kerlinger 1987). Delay ed arrival on migratory grounds can have additional negative consequences for migratory birds including lower body weight and poor nest site selectio n. Reduced nestin g conditions due to late arrival further result in reduce d broods per year increas ed nest predation and nest parasitism, and reduced overall nesting success (Rodewald and Shustack 2008) In recent years it has been proposed that en route migration may actually be the factor limiting migratory bird population s ( Alerstam and Hedenstrom 1998, Newton 2004) Many migratory birds migrate along the eastern coast of North American ( Moore and Kerlinger 1987 ) in coastal areas that have had seen the greatest national population growth in the United States in recent dec ades (Dahl 1990) For this reason preservation of forested wetlands for conservation of migratory bird populations is very important for maintaining current populations and is an issue of concern for migrant species with significant declines in recent de cades like the pine warbler that utilizes forested areas
35 near coasts (Robbins et al. 1989) For cypress domes in Tampa, forest area and habitat quality are important for maintaining migratory bird population. Land managers need to preserve sufficient amo unts of wetland and upland forested area in new development and maintain connection to rural or exurban systems in order to provide adequate wildlife corridors to support native bird populations and other biota
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43 Appendix A. Bird distrib ution among sites Common Name Scientific Name Migratory Status Feeding Strategy Sample Site Species Presence 19 70 1 19 70 2 19 70 3 19 80 1 19 80 2 19 80 3 19 90 1 19 90 2 19 90 3 20 00 1 20 00 2 20 00 3 Ref 3 Ref 4 Ref 5 Ref 6 Acadian Flycatcher Empidonax virescens M I X X American Crow Corvus brachyrhynchos R O X X X X American Goldfinch Carduelis tristis M G X X X X American Robin Turdus migratorius W O X X X American Redstart Setophaga ruticilla M I X Anhinga Anhinga anhinga R P X Barred Owl Strix varia R C X X X Blackburnian Warbler Dendroica fusca M I X X Blue grey Gnatcatcher P olioptila caerulea R I X X X X X X X X X x X X X X X X Brown headed Nuthatch Sitta pusilla R I X Blue headed Vireo Vireo solitarius M I X X Blue Jay Cyanocitta cristata R O X X X X X X X X X X X X X X X X Black Vulture Coragyps atratus R C X X Blackpoll Warbler Dendroica striatta M I X Brown Creeper Certhia americana R I X Brown Thrasher Toxostoma rufum R O X X X X X Boat tailed Grackle Quiscalus major R O X X X X X X X Black throated Green Warbler Dendroica virens M I X Black and White Warbler Mniotila varia M I X X X X X X X X X X X R = residents, W = wintering migrants, M = Neotropical migrants. I = insectivore, O = omnivore, G = granivore, C = carnivore, P = piscivore.
44 Appendix A. Bird distribution among sites (Continued) Common Name Scientific Name Migratory Status Feeding Strategy Sample Site Species Presence 19 70 1 19 70 2 19 70 3 19 80 1 19 80 2 19 80 3 19 90 1 19 90 2 19 90 3 20 00 1 20 00 2 20 00 3 Ref 3 Ref 4 Ref 5 Ref 6 Carolina Chickadee Poecile carolinensis R O X X X X X X X X X X Carolina Wren Thryothorus ludovicianus R I X X X X X X X X X X X X X X X X Cedar Waxwing Bombycilla cedrorum R G X X X X Cape May Warbler Dendroica tigrina M I X Common Grackle Quiscalus quiscula R O X X X X Cooper's Hawk Accipiter cooperii R C X Common Nighthawk Chordeiles minor M I X Common Yellowthroat Geothlypis trichas M I X X X X X Chestnut sided Warbler Dendroica pensylvanica M I X X X Downy Woodpecker Picoides pubenscens R I X X X X X X X X X X X X X X Eastern Phoebe Sayornis phoebe M I X X X X X X X X X Eastern Kingbird Tyrannus tyrannus W I X X Eastern Towhee Pipilo erythrophthalmus R I X European Starling Sturnus vulgaris R O X Fish Crow Corvus ossifragus R O X X X X X X X X X Great Blue Heron Ardea herodias R P X X X X Grey Catbird Dumetella carolinensis W O X X X X X X X X X X X X Great Egret Ardea alba R P X X X X X Gray Kingbird Tyrannus dominicensis M I X Hairy Woodpecker Picoides villosus R O X X X X R = residents, W = wintering migrants, M = Neotropical migrants. I = insectivore, O = omnivore, G = granivore, C = carnivore P = piscivore.
45 Appendix A. Bird distribution among sit es (Continued) Common Name Scientific Name Migratory Status Feeding Strategy Sample Site Species Presence 19 70 1 19 70 2 19 70 3 19 80 1 19 80 2 19 80 3 19 90 1 19 90 2 19 90 3 20 00 1 20 00 2 20 00 3 Ref 3 Ref 4 Ref 5 Ref 6 Hermit Thrush Cart harus mimus M O X X X X Little Blue Heron Egretta caerulea R P X Limpkin Aramus guarauna R P X Louisiana Waterthrush Seiurus motacilla M I X X Mourning Dove Zenaida macroura R G X X Magnolia Warbler Dendroica magnolia M I X Northern Cardinal Cardinalis cardinalis R O X X X X X X X X X X X X X X X X Northern Flicker Colaptes auratus R O X X X X X X X X Northern Mockingbird Mimus polyglottos R O X X X X X X X Northern Parula Parula americana M I X X X X X X X X X X X X X X X Northern Waterthrush Seirus noveboracensis M I X Osprey Pandion haliaetus R P X Ovenbird Seiurus aurocapilla M I X X X X X X Palm Warbler Dendroica palmarum W I X X X X X X X X X X Philadelphia Vireo Vireo philadelphicus M I X Pine Warbler Dentroica pinus W I X X X Pileated Woodpecker Dryocopus pileatus R O X X X X X X X X X X X X X X X Prothonotary Warbler Protonotaria citrea M I X X X X X X Prairie Warbler Dendroica discolor W I X X X X X X X X X Red breasted Nuthatch Sitta canadensis W I X R = residents, W = wintering migrants, M = Neotropical migrants. I = insectivore, O = omnivore, G = granivore, C = carnivore, P = piscivore.
46 Appendix A Bird distribution among site (Continued) Common Name Scientific Name Migratory Status Feeding Strategy Sample Site Species Presence 19 70 1 19 70 2 19 70 3 19 80 1 19 80 2 19 80 3 19 90 1 19 90 2 19 90 3 20 00 1 20 00 2 20 00 3 Ref 3 Ref 4 Ref 5 Ref 6 Red bellied Woodpecker Melanerpes carolinus R O X X X X X X X X X X X X X X X X Ruby crowned Kinglet Regulus calendula W I X X X Red eyed Vireo Vireo olivaceus M I X X X X X X X Red shouldered Hawk Buteo lineatus R C X X X X X X X X X X X X X X Red winged Blackbird Agelaius phoeniceus M O X Scarlet Tanager Piranga olivacea M I X X Sora Porzana carolina W O X Swamp Sparrow Melospiza georgiana M I X Tufted Titmouse Baeolophus bicolor W O X X X X X X X X X X X X X X X X White eyed Vireo Vireo griseus W I X X X X X X X X White Ibis B ubulcus ibis R O X X X Wood Duck Aix sponsa R O X Wood Stork Mycteria americana R O X White throated Sparrow Zonotrichia albicollis R O X Yellow breasted Chat Icteria virens M I X X Yellow billed Cuckoo Coccyzus americanus M O X Yellow bellied Sapsucker Sphyrapicus varius W O X X X X Yellow rumped Warbler Dendroica coronata W I X X X X X X X X X Yellow throated Vireo Vireo flavifr ons M I X X X X X X Yellow throated Warbler Dendroica dominica W I X X X X X X X X R = residents, W = wintering migrants, M = Neotropical migrants. I = insectivore, O = omnivore, G = granivore, C = carnivore P = piscivore