Avian communities in Florida habitats

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Title:
Avian communities in Florida habitats analysis and review
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v, 46 p. : ill. ; 28 cm.
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Engstrom, R. Todd
Florida -- Nongame Wildlife Program
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Bird populations -- Florida   ( lcsh )
Birds -- Florida   ( lcsh )
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bibliography   ( marcgt )
non-fiction   ( marcgt )

Notes

Bibliography:
Bibliography: p. 30-37.
Statement of Responsibility:
R. Todd Engstrom.
General Note:
"August 1993."
General Note:
"Submitted as final report for Nongame Wildlife Program project NG88-022".

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University of Florida
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Table of Contents
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Full Text




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Avian Communities in Florida Habitats:
Analysis and Review






R. Todd Engstrom*

Cornell Laboratory of Ornithology
159 Sapsucker Woods Road
Ithaca, New York 14850


















Submitted as final report for
Nongame Wildlife Program project NG88-022


August 1993

*Current Address: Tall Timbers Research, Inc. Route 1, Box 678
Tallahassee, Florida 32312-9712
(IVEtSITY OF FLORIDA LIBRARIES


























This report is the result of a project supported by the Florida
Game and Fresh Water Fish Commission's Nongame Wildlife
Program. It has been reviewed for clarity, style, and typographical
errors, but has not received peer review. Any opinions or
recommendations in this report are those of the authors and do not
represent policy of the Commission.













Suggested citation:

Engstrom, T.R. 1993. Avian communities in Florida habitats:
analysis and review. Fla. Game and Fresh Water Fish Comm.
Nongame Wildl. Program Final Rep. 46 pp. + v. Tallahassee, FL.


















AVIAN COMMUNITIES IN FLORIDA HABITATS:
ANALYSIS AND REVIEW


R. TODD ENGSTROM
Comell Laboratory of Ornithology, Cornell University, Ithaca, New York 14850



Abstract: Quantitative studies of bird communities in Florida provide valuable
baseline data for comparisons of the relative abundance of birds in different habitats.
The most extensive sources of data for bird communities in North America are the
Breeding Bird Census (BBC) and Winter Bird Population Study (WBPS). Over 50
counts in both seasons have been made in Florida. In order of decreasing number,
bird counts have been made in coniferous and hardwood forests, wetlands, waterfronts,
agricultural areas, urban areas, rangeland, and beaches.

Studies of bird communities from sources other than the BBC and WBPS are
mainly from the academic community, especially masters theses and doctoral
dissertations. A problem with comparing the results of these studies is that various
methods have been used to count birds. The largest number of bird counts in Florida
were designed to determine the effects of timber management on bird populations.
Bird communities have also been studied to examine the effects of phosphate
mining, agriculture, urbanization, forest fragmentation, and fire.

The objectives of this study are to review the literature on Florida bird
communities, to evaluate the coverage of different habitat types, and to obtain
estimates of bird species richness in different habitats of Florida by using a statistical
technique called rarefaction.

Bird species richness (standardized to the number expected on an 8.1-ha plot)
ranged from 3 in salt marshes to 19 in mature beech-magnolia forest during the
breeding season. In winter the number of bird species (standardized to 8.1 ha) ranged
from 9 in south Florida slash pine forest to 29 in oak-palm-hickory hammock and
maple swamp. Comparison of bird species richness between general habitat types
(e.g., hardwood and pine forests) is confounded by the peninsula effect.

Recommendations to improve the ornithological database for Florida include:
(1) conducting a bird habitat survey, (2) encouraging researchers to use standardized
counting methods, and (3) tying issuance of permits to destroy natural habitat to
conducting pre- and post-disturbance studies. Development of alternative timber
management practices and use of native plants in suburban settings would benefit
Florida birdlife.








NONGAME WILDLIFE PROGRAM FINAL REPORT


ACKNOWLEDGMENTS


Many ornithologists in Florida took the time to review my
list of studies. I thank Stephen R. Humphrey, Robert W. Loftin,
Reed F. Noss, William B. Robertson, James A. Rodgers, Jr., Henry
M. Stevenson, and Glen E. Woolfenden for their comments on the
manuscript. Ronald F. Labisky provided a valuable list of theses
and dissertations produced by students at the University of Florida.
I especially thank James A. Cox for helpful suggestions and
help during all phases of this project. James D. Lowe, Diane L.
Tessaglia, Mitch Hartley, and Kelly Cranfield provided
invaluable help with the computerized Breeding Bird Census
data base at the Cornell Laboratory of Ornithology. Susan I.
Cerulean, J. Larry Landers, W. Wilson Baker, David G. Cook, and
Frances C. James also made useful comments on the manuscript.
This research was supported by the Florida Game and Fresh Water
Fish Commission's Nongame Wildlife Program NG 88-022.








AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom






TABLE OF CONTENTS


ABSTR A CT ............................................... ......................... iii
ACKNOWLEDGMENTS ...................................... ........... iv
INTRODUCTION ................................................................. 1
Breeding Bird Census and
Winter Bird Population Study ........................................... 1
M ETH O D S................................................... ......................... 3
Habitat Classification .......................................... .............. 3
R arefaction ................................................ ....................... 3
RESULTS ...................................... .............. ....... ............. 6
BBCs and W BPSs ................................................................. 6
Comparisons of Species Richness ........................................... 9
D ISC U SSIO N .................................................... ...................... 16
Birds, Land Use, and Habitat Management ........................ 16
Forest Management ...................................... .......... .. 16
A agriculture ................................................. ......... ..... 19
Urbanization ............................................... ............. 20
Phosphate M ining ........................................................... 21
Forest Fragm entation....................................................... 21
Fire .................................................. .......................... 22
Coastal Development .................................... ........... 23
Miscellaneous Studies .................................... .......... 23
Evaluation of the BBC and WBPS ...................................... 24
The Peninsula Effect .......................................... .......... ... 25
CO NCLUSIO NS .....................................................................27
RECOMMENDATIONS ..................................... ........... .. 28
LITERATURE CITED.......................................... ............. 30
APPENDICES .................................... .....................................38











AVIAN COMMUNITIES IN FLORIDA HABITATS -Engstrom 1


INTRODUCTION

Studies of bird communities by habitat and season provide
valuable information about the general distribution of birds in relation
to their resources, if made using similar methods. With proper
planning, comparisons among communities can be made that suggest
the responses of birds to ecological plant succession, fire, logging,
agriculture, and other land uses. Carefully collected information also
serves as baseline data, so comparisons can be made in the future.
While use of such data may not permit strong inference of the causes
of change in populations in experimental studies (Temple and Wiens
1989), some environmental effects are not amenable to
experimentation (DeSante and Geupel 1987). This study is a first
attempt to summarize quantitative studies of birds in Florida.

Ornithological studies of Florida bird populations range from
simple species lists for a locality or a region to standardized counts
in specific habitats. The line transect (Emlen 1971), spot map
(Robbins 1970), and point count methods (Reynolds et al. 1980) are
used most frequently to obtain estimates of relative abundance or
density of birds for a specific area or habitat. The relative merits of
these methods are discussed by Ralph and Scott (1981). mparisons
should only e deamong results of counts that have been conducted
with similar methods. Even when count methodology is standardized,
comparison of results can be complicated by variation effort,
observer competence, season, and plot size.

Data for Florida bird populations are included in national
monitoring and research programs, including the Breeding Bird
Survey (BBS), Christmas Bird Count (CBC), Breeding Bird Census
(BBC) and Winter Bird Population Study (WBPS). The BBS and
CBC provide indices of abundance across large geographical areas,
but do not measure bird populations within specific habitat types;
these programs will not be evaluated here.

Breeding Bird Census and Winter Bird Population Study

The Breeding Bird Census (BBC), initiated by the National
Audubon Society in 1937, is the largest source of standardized plot-
based counts of birds conducted in North America. In the last 51
years over 4,100 BBCs have been conducted in the United States and
Canada. The objectives of the BBC are (1) to determine the species








NONGAME WILDLIFE PROGRAM FINAL REPORT


and density of breeding birds found in each habitat type throughout
North America; (2) to measure the effects of various land-use
practices on breeding-bird populations; (3) to quantify the amount of
yearly variation in densities of breeding birds occupying various
habitat types; and (4) to establish the nesting requirements for each
species of bird throughout its range (Stewart 1949). The National
Audubon Society created the Winter Bird Population Study (WBPS),
the winter analogue of the BBC, in 1948.

BBCs and WBPSs are conducted by experienced volunteers
mostly in the United States and Canada. BBC participants use the
"spot mapping method" (Williams 1936) to count the number of
territorial birds on a measured plot of land (Hall 1964, Robbins 1970,
Van Velzen 1972). A modified spot map method is used in the
WBPS, but relative abundance is measured as the number of
individuals detected per trip instead of the number of territorial
individuals (Kolb 1965). Spot mapping is the most precise technique
for estimating numbers of birds in most terrestrial habitats short of
mark-recapture, but it is not appropriate for colonial and wide-
ranging species.

A method of quantitative habitat description was recommended
for forested habitats to accompany the BBC and WBPS bird counts
by James and Shugart (1970). Density and basal area by size class is
estimated for major tree species. Estimates of shrub density, percent
canopy cover, and ground cover are based on a series of 0.04-ha
circular samples. The combined results of random samples within a
study plot provide data for a quantitative description of the count
area. The 'James-Shugart' method has been adopted by BBC and
WBPS participants as a standard technique for describing vegetation
structure. This technique permits quantitative comparisons of the
vegetation of a variety of habitats that can be used to interpret
comparisons of their avifaunas (James and Wamer 1982).

The objectives of this study were to review the literature on
Florida bird communities, identify habitats for which the bird
communities are poorly documented, and to obtain estimates of bird
species richness in the habitats of Florida. The method of obtaining
comparable estimates of species richness involves a statistical
technique called rarefaction (James and Wamer 1982).







AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 3


METHODS

To review the literature I searched Dissertation Abstracts
International for the years 1951 through 1990 using the following
keywords: birds, ecology, Florida, and community. In addition, I
placed a request for information in the newsletter of the Florida
Ornithological Society. Finally, I wrote to ornithologists who have
conducted research in Florida for comments on and additions to my
initial list of references.

Habitat Classification

There are many different habitat classification systems available
for Florida. Cox et al. (1987) gives a cross-reference table for
three such systems. I used the Florida Land-use and Cover
Classification System (Florida Department of Transportation 1985),
because this system includes human-modified environments. For
this study, I classified all BBCs and WBPSs according to habitat
type. As an example, I included 'Suburban School Park' (BBCs 29-
37 in Appendix II) under the category of Residential, subcate
gory City Park. Some habitat categories were slightly modified. I
combined live oak (Quercus virginiana) and cabbage palm (Sabal
palmetto) into one habitat and changed longleafpine (Pinuspalustris)-
xeric oak forest to pine-xeric oak.

Rarefaction

Variation in sampling methods and study plot size often present
difficulties to researchers who wish to compare species richness
among different terrestrial bird communities. The observation that
the number of species varies with area (the species-area effect) is one
of the oldest in ecology (Arrhenius 1921). Because the relationship
between species richness and area is non-linear (Connor and McCoy
1979), it is unreliable to estimate species richness directly from area.

Rarefaction is a statistical technique that can be used to estimate
species richness (Sanders 1968, Hurlbert 1971, Fager 1972,
Simberloff 1972, 1978; James and Rathbun 1981) and variance of
species richness (Heck et al. 1975) for subsamples of a given sample.
For example, with rarefaction one might ask, if you had 25 species
in a sample of 100 individuals, how many species would be expected
if the sample size were only 50 individuals? Given the number of
individuals in each species of a sample, rarefaction draws individuals







NONGAME WILDLIFE PROGRAM FINAL REPORT


randomly without replacement to estimate the species richness [E(S)]
and the variance of a subsample smaller than the original sample. In
this way, species richness of two samples, originally of different size,
can be compared. For example, one sample can be reduced to the
number of individuals that occurs in another sample. Or, in the case
here, samples can be reduced to the number of individuals expected
on study plots of equal size (James and Rathbun 1981; James and
Wamer 1982).

I used rarefaction to provide standardized estimates of species
richness from BBC data. For example, the mature American beech
(Fagus grandifolia)-southern magnolia (Magnolia grandiflora) forest
(Appendix I; number 10) had a total of 30 species and 83.5 territorial
males. Northern parula, the most abundant species on the plot, had
an estimated 17 territories. Twenty species had at least one territory
on the plot and ten species had less than one territory (denoted by '+').
The distribution of the numbers of territories arranged in order of
decreasing abundance was: 17 14.5 9.5 5 5 5 5 3 2.5 2.5 2.5 2 2 1.5
1.511 1 11++++++++++.

Rarefaction can only be used to analyze a community if the
numbers of individuals in each species are integers. In the example
above, ten species were dropped because they had less than one half
territory on the plot ('+'). Note that seven species had half territories
(one territory straddled the plot boundary). For these, the numbers
were rounded upward, which increased the total number of territories
to 87. These procedures (round all half territories upward and
exclude species that had less than a half territory on the study plot)
were used throughout this analysis.

To obtain an estimate of the number of species for a standard
plot size of 8.1 ha, I estimated the number of individuals on 8.1 ha by
linear interpolation. The plot size for the beech-magnolia forest
above was 15.8 ha. The number of territories estimated for an 8.1-ha
plot is 45 (87 territories x 8.1 ha / 15.8 ha). I then used rarefaction
to estimate species richness for 45 territorial individuals. The
assumption that the number of individuals increases linearly with
increasing area is weak if the habitat is heterogeneous, because the
birds would be unlikely to be evenly distributed.

The Breeding Bird Census and Winter Bird Population Study
were the only two sources of data that I used in the rarefaction
analysis. Most other bird counts in Florida were either conducted








AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 5


with a different method than spot-mapping or the plot size used was
judged to be too small. Only plots of 8.1 ha or greater were used in
the comparison of species richness, because density tends to be
overestimated on smaller plots (Engstrom 1981, Engstrom and James
1981, Verner 1985).

WBPS results can be analyzed directly with the rarefaction
program because the average number of individuals seen on each
trip is rounded to the nearest integer. Species that were encountered
on less than half of the trips (designated with a '+' in the
published results) were not included in the analysis.








NONGAME WILDLIFE PROGRAM FINAL REPORT


RESULTS

Until 1984 BBC and WBPS data were published by the National
Audubon Society for more than 50 and 40 years, respectively. Now
the data are published annually in a supplement to the Journal of Field
Ornithology. Many of the BBCs from 1937 to present (and all of
those conducted in Florida) are stored in a computerized database at
the Cornell Laboratory of Ornithology. The WBPS computerized
database contains only the 1989 counts.

All Florida BBCs are listed alphabetically by author in
Appendix A and all WBPSs in Appendix B. The sequence numbers
in Appendix A and B are used to identify counts throughout the text.
Bird species of all Florida BBCs are indexed by count number in
Appendix C. Scientific names of birds not given in the text are listed
in Appendix C.

BBCs and WBPSs

Fifty-one BBCs were made in Florida on 25 different sites (Table
1; Appendix A). Nineteen sites were censured for only 1 year, 2 sites
for 2 years, 3 sites for 3 years, 1 site for 4 years, 1 site for 5 years, and
1 site for 10 years. The earliest BBC in Florida was conducted in
1958. Seventeen BBCs were made between 1966 and 1969 and 26
from 1980-88.

Eighty-two bird species were observed in Florida BBCs (Appendix
C). Northern mockingbird (Mimus polyglottos) and great crested
flycatcher (Myiarchus crinitus) were the most commonly encountered
species on BBCs. This is a reflection of the study sites selected, not
necessarily an indication of statewide abundance or distribution.

Although the BBC was intended to be a program supported by
amateur participants, many of the BBCs in Florida have been
conducted by people associated with academic institutions, research
stations, and the federal government. Florida BBCs were located in
Leon County (Tall Timbers Research Station and Florida State
University), Franklin County (St. Vincent's National Wildlife Refuge),
Alachua and Levy counties (University of Florida), Pasco County
(University of South Florida), Brevard and Indian River
counties (St. John's National Wildlife Refuge and Entomological
Research Center), Highlands County (Archbold Research Station),
and Collier and Dade counties (Big Cypress and Everglades









AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 7






Table 1. Florida habitat types in which Breeding Bird Censuses (BBC) and Winter Bird
Population Studies (WBPS) took place.



BBCb WBPSb
Ref. No. No. No. No.
Num. Habitat type Sites Cts. Sites Cts.


Urban and developed
Residential, low density
City park

Agriculture
Improved pasture
Citrus groves
Fruit orchard

Rangeland
Coastal scrub

Upland forests
Upland coniferous forests
Pine flatwoods
Pine xeric oak
Sand pine scrub
Upland hardwood forests
Live oak-cabbage palm
Beech-magnolia

Water
Streams and waterways
Bays and estuaries

Wetlands
Wetland hardwood forests
Wetland mixed hardwoods
Vegetated non-forested wetlands
Freshwater marshes
Sawgrass
Saltwater marshes
Cordgrass


1 9


S 1
3 2
1


1 1


-1
-1


1 4 1 3


1 1 1


1 12


Barren land
Beaches other
than swimming beaches


1 1


25 51 23 54


a Habitat types are subsets of the classes identified by the Florida land use, cover, and forms
classification system (Florida Department of Transportation 1985).
b Number of counts differ from the number of sites when sites were sampled for more than
one year.


100
110
1851


400
410
411
412
413
420
427.5
431

500
510
540

600
610
617
640
641
6411
642
6421


700
710


TOTAL







NONGAME WILDLIFE PROGRAM FINAL REPORT


l# of BBCs per County

70 70






Figure 1. The distribution of Breeding Bird Censuses (BBCs) in Florida, by county.












# of WBPSs per County

1 0

I 1 to 5

J6 to 10

> 10
Figure 2. The distribution of Winter Bird Population Studies (WBPSs) in Florida, by county.







AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 9


NationalPark) (Figure 1). BBCs have been conducted in only 10 of
67 counties in Florida.

Florida BBCs were conducted most often in coniferous forests
(23 counts) followed by wetlands (13), and hardwood forests (12)
(Table 1). Only 2 counts were made in agricultural habitat (citrus
grove). Two counts were made of the avian communities in early
stages of secondary succession following clearcutting of pine forests.
The effects of fire on bird communities were studied in pinewoods in
Leon (numbers 42 and 43 in Appendix A), Highlands (numbers 45
and 46), and Collier (numbers 4-7) counties.

Although the WBPS is a younger and much smaller program
than the BBC (approximately 2,000 WBPSs have been conducted
since 1947), more WBPSs than BBCs have been conducted in
Florida. Fifty-four counts have been made in 23 different sites
(Table 1). Counts were made at 13 WBPS sites for only 1 year, 4 sites
for 2 years, 3 sites for 3 years, 1 site for 4 years, and two long-term
counts, Suburban School Park, and Barrier Beach and Saltwater
Estuary, were conducted for 9 and 11 years respectively. The
results of the Barrier Beach and Saltwater Estuary, and Sheltered
Beach studies were not analyzed, because these habitats are very
different from the other habitats considered. Except for counts in
Leon and Duval counties, WBPSs have been conducted
only in peninsular Florida (Figure 2). Fewer winter counts have
been done in natural plant communities than in man-modified
environments (Table 1).

Comparisons of Species Richness

Of 51 BBCs in Florida, 46 were conducted on plots of 8.1 ha (20
acres) or greater. I estimated species richness for a sample of
individuals standardized to 8.1 ha [E(S 8.ha)] for each of these plots.
E(Sg.1ha) ranged from 2.3 in Gulf Coast saltmarsh to 22.3 in mature
beech-magnolia forest (Table 2). Of 54 WBPSs in Florida, 30 were
conducted on plots of 8.1 ha or greater. Using rarefaction, E(S8.ha)
ranged from 9.2 in South Florida slash pine to 29.6 in oak-palm-
hickory hammock and maple swamp (Table 3). Within a single plot,
the number of species may vary considerably among years. For
example, species richness in the mature beech-magnolia forest plot
ranged from 16 in 1981 to 22.3 species in 1988 (Table 2). Variation
on this plot was caused in part by gaps created by treefalls during
Hurricane Kate in 1985.










10 NONGAME WILDLIFE PROGRAM FINAL REPORT






Table 2. Species richness estimated for Breeding Bird Censuses in Floridaa.


Habitatf Plot Ref.b Ind per
County Habitat Type Size No. Year 40 ha S N 8.had E(Sglha)e


Alachua Longleaf Pine 412 10.1
-Turkey Oak




Brevard Cordgrass 6421 25.0
Salt Marsh I

Brevard Cordgrass 6421 25.0
Salt Marsh II

Collier Slash Pine- 411 40.0
Cypress Mosaic

Collier Virgin 411 16.6
Subtropical
Slash Pine Forest

Dade Cutover 411 20.2
Subtropical
Slash Pine Forest

Dade Everglades 6411 44.6
Marsh


50 1958
51 1959
1 1960
11 1961
12 1962

22 1981


23 1981


36 1979


25 1980


39 1980
3 1981


21 1976


19 6.9
12 5.5
16 5.6
19 7.1
16 4.0


51 4 10 3.0


67 6 13 4.5


49 16 9 6.9


147 15 29 13.2


7 4.0
6 4.2


39 10 7 4.5


Franklin Gulf of
Mexico


411 51.4


Highlands Slash Pine- 412 8.1
Turkey Oak


8 1983


47 1969


51 21 10 7.5


7 21 7.0


Highlands Scrubby
Flatwoods

Highlands Recently
Burned Scrubby
Flatwoods

HighlandsSand Pine
Scrub


411 8.1 45 1969


411 8.1



413 8.1


Indian Citrus Grove 221 16.2
River


Indian Oak-Palm 617 23.9
River Hickory Hammock
and Maple Swamp


46 1969



44 1969


19 1967
20 1968
40 1969

16 1966
17 1967
18 1968
41 1969


138 11 28 11.0


49 5 10 5.0



84 6 17 6.0


188 12
193 13
198 11


38 10.7
39 11.4
40 9.7

22 7.8
25 9.2
25 9.6
26 8.1









AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom


Table 2. (continued)


Habitatf Plot Ref.b Ind per
County Habitat Type Size No. Year 40 ha Sc N8.1had E(S8.lha)e


Indian Live Oak- 427.5 14.2
River Cabbage Palm
Coastal Hammock


1966
1967
1968


141 13
177 15
155 10


8.6
10.2
7.9


Leon Longleaf Pine 412
Forest

Leon Longleaf Pine 412
Forest April bum


20.0


15.0


Leon Longleaf Pine
Forest 412 15.0
Unburned control


35 1980


76 18 15 10.2


42 1988 131 24 26 16.5


43 1988 115 21 23 15.0


Leon Slash Pine- 412
Turkey Oak Forest

Leon Mature Beech- 431
Magnolia Forest


Levy Gulf Coast
Salt Marshg


Pasco Live Oak-
Cabbage Palm
Hammock


16.0


15.8


6421 15.0
31.0
15.0
15.0
20.0
20.0
20.0
20.0

427.5 13.5


24 1980


1980
1981
1988

1980
1981
1982
1983
1984
1985
1987
1988


40 .11 8 6.7


291 22
220 20
359 26


1967 181 13
1968 184 11


a BBCs 4-7 and 48 were not included in the analysis because plot size was <8.1 ha.
b Reference numbers refer to Appendix A.
c S represents the total number of species found on the plot.
d N.lha is the number of individuals expected on a plot of 8.1 ha.
e E(S.lha) is the number of species estimated for a sample of the number of individuals estimated to
occur on 8.1 ha (N8s.1h) from rarefaction.
f Habitat type numbers refer to Table 1.
g Two counts (ref. nos. 31 and 34) were not included. Count 31 was not entered in to the computer
and Count 34 was based on an abbreviated sample period.


18.2
16.0
22.3


10.2
9.4









12 NONGAME WILDLIFE PROGRAM FINAL REPORT






Table 3. Species richness estimated for Winter Bird Population Studies in Floridaa.


Habitat Plot Ref.b Ind per
County Habitat Type Size No. Year 40 ha S N8. hac E(Ss.ha)d


Collier


Dade

Dade


Slash Pine- 411
Cypress Mosaic

Everglades Marsh 6411


Pineland, Scrub,
and Residential


Highlands Slash Pine-
Turkey Oak


Highlands Scrubby Flatwoods 411

Highlands Recently Burned 411
Scrubby Flatwoods

Highlands Sand Pine Scrub 413


Highlands


Mature Citrus
Orchard


Indian River Citrus Grove


Indian River Live Oak-Cabbage 427.5
Palm Coastal
Hammock

Indian River Oak-Palm Hickory 617
Hammock
and Maple Swamp


40.0 41 1980


44.6 21 1975


8.1 5
10.1 6
7
8


213 31 43 16.5


151 34 30 16.2


1975
1976
1977
1978


8.1 54 1969


8.1 52 1969

8.1 53 1969
3 1989

8.1 51 1969
2 1989


221 13.8 28 1989


16.2 18
19
20

14.2 12
13
14

23.9 15
16
17


1967
1968
1969

1967
1968
1969

1967
1968
1969


26 183 26.0
25 66 22.6
24 44 21.2
27 71 23.6


348 23 70 22.4


193 13 39 13.0

237 12 48 12.0
281 17 57 17.0

385 16 78 16.0
193 10 39 10.0

301 15 61 12.3


588 32 119 25.5
494 30 100 22.6
785 34 159 28.8

1494 30 302 25.5
412 28 83 18.7
530 29 107 22.4


1040
748
1000


41 210 29.6
40 151 28.1
39 202 29.6


Longleaf Pine Forest 412


Mature Beech-
Magnolia Forest


South Florida 411
Slash Pine

Live Oak-Cabbage 427.5
Palm Hammock


Grazed Pasture
with Ponds


20.0 39 1982

15.8 38 1983
1 1988


40.0 40 1981


13.5 42 1967


60.7 22 1969
23 1970


222 19 44 15.3

535 22 108 18.0
767 29 155 25.5

96 19 19 9.2


421 29 85 24.7


262 25 53 12.2
252 25 51 12.5


a WPBSs 9-11, 29-37, and 44 were not included in the analysis because plot size was <8.1 ha.
b Reference numbers refer to Appendix B.
c Ns.ha is the number of individuals expected on a plot of 8.1 ha.
d E(S8. ha) is the number of species estimated for a sample of N8. ha individuals from rarefaction.


Leon

Leon


Monroe


Pasco


Volusia









AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 13


20



15 Uf

r" pinewoods
10- wetlands
0* U citrus grove
W) ue hardwoods
5 -
*


0 10 20 30 40 50 60

Num. ind. per 8.1 ha
Figure 3. Scatterplot of estimated species richness [E(S)] vs. the estimated number of
individuals for a standard 8.1-ha plot for BBCs. The average E(S) was taken for plots that were
censused for more than one year.

To compare species richness among habitats I took the average
of all E(S8.1ha) for all years that counts were made for a given site
and then rounded the average to the nearest integer.

In both the BBC and WBPS, species richness and the number of
individuals on an 8.1-ha plot are positively correlated (Figures 3 and
4). In the breeding season, mature beech-magnolia forest (the mean
of 3 years) contained the most individuals and species of all habitats.
Saltmarsh and everglades marsh had the fewest species and
individuals. In winter, hardwood forests were more species-rich than
were pinewoods (Figure 4). The citrus grove in Indian River County
(numbers 18-20 in Appendix B) was surprisingly species-rich,
although a second citrus grove study site in Highlands County
(number 28) had considerably fewer species. Among pinewoods,
clearcut areas had few species and individuals, and longleaf pine-
turkey oak in north Florida was the most species-rich (Figure 5).

A simple ordination of BBCs along an axis of E(S8.1ha) clearly
indicates relative species richness (Figure 6) among habitats. Although
mature beech-magnolia forest is the most species-rich, several
pinewood sites and even citrus groves ranked higher than the hardwood
hammocks from central and south Florida. In winter the oak-palm-
hickory hammock and maple swamp held the most species (Figure 7).
As in the breeding season, the citrus grove in Indian River County
(numbers 18-20 in Appendix B) ranked unexpectedly high.











NONGAME WILDLIFE PROGRAM FINAL REPORT


I I I I I 5
0 10 20 30 40 50


pinewoods

wetlands

citrus grove

hardwoods


Num. ind. per 8.1 ha

Figure 4. Scatterplot of estimated species richness [E(S)] vs. the estimated number of
individuals for a standard 8.1-ha plot for WBPSs. The average E(S) was taken for plots that
were censused for more than one year.


flatwoods
pine-oak

sand pine
clearcut


Num. ind. per 8.1 ha


Figure 5. Scatterplot of estimated species richness [E(S)] vs. the estimated number of
individuals for a standard 8.1-ha plot for BBCs in pine woods. The average E(S) was taken
for plots that were censused for more than one year.


D







*
0


[]0

[]













AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 15


E(Ss.lha)


Habitat Type (reference numbers to Appendix A)


u
Gulf Coast Salt Marsh (26-32,37)
Cordgrass Salt Marsh I (22)
Cutover Subtropical Slash Pine Forest (3,39)
-_Cordgrass Salt Marsh II (23)
Everglades Marsh (21)
Recently Burned Scrubby Flatwoods (46)
5 ----Longleaf Pine-Turkey Oak Assoc. (1,11,12,50,51)
Sand Pine Scrub (44)
-- Slash Pine-Cypress Mosaic (36)
Slash Pine-Turkey Oak Forest (47)
Gulf of Mexico Barrier Island (8)
- Oak-Palm-Hickory Hammock and Maple Swamp (16-18,41)
Live Oak-Cabbage Palm Coastal Hammock (13-15)
10 -Live Oak-Cabbage Palm Hammock (38,49)
0 Longleaf Pine Forest (35)
^uCitrus Grove (19,20,40)
Scrubby Flatwoods (45)
Virgin Subtropical Slash Pine Forest (25)


15 Longleaf Pine Forest-Unburned Control (42)


Longleaf Pine Forest-April Burn (43)


Mature Beech-Magnolia Forest (2,9,10)
20


Figure 6. Ordination of habitats along an axis of estimated species richness for a standard 8.1-
ha plot for BBCs.


E(Sgs.ha) Habitat Type (reference numbers to Appendix B)

South Florida Slash Pine (40)
10
Mature Citrus Grove (29)
S Grazed Pasture with Ponds (22,23)
Suburban School Park (29-37)
-Scrubby Flatwoods (52)
Sand Pine Scrub (2,51)

15 -====Recently Burned Scrubby Flatwoods (3,53)
Longleaf Pine Forest (39)
Everglades Marsh (21)
Slash Pine Cypress Mosaic (41)



20
Mature Beech-Magnolia Forest (1,38)
Live Oak-Cabbage Palm Coastal Hammock (12-14)
Slash Pine-Turkey Oak Association (54)
------Pineland, Scrub, and Residential (5-8)

25 Live Oak-Cabbage Palm Hammock (42)

Citrus Grove (18-20)


Oak-Palm-Hickory-Hammock and Maple Swamp (13-15)
30

Figure 7. Ordination of habitats along an axis of estimated species richness for a standard 8.1-
ha plot for WBPSs.








NONGAME WILDLIFE PROGRAM FINAL REPORT


DISCUSSION

Birds, Land Use, and Habitat Management

Studies of bird communities in Florida other than BBCs and
WBPSs have been conducted with a variety of methods for diverse
purposes (Table 4). Below, I have provided a brief review of most of
these studies under the general headings of forest management,
agriculture, urbanization, phosphate mining, forest fragmentation,
fire, and coastal habitats for migrating birds.

ForestManagement.- Changing land use patterns and methods
of forest management are extremely important to wildlife populations
in Florida. Forests occur on approximately 16.5 million acres (43%)
of the surface area of Florida, excluding inland waters (Brown and
Thompson 1988). From 1980 to 1987 the area of timberland in
Florida decreased by over 4%. Over half of this loss can be attributed
to conversion to urban and related land use.

The forests available for commercial harvest are about evenly
divided between softwood and hardwood types; however, almost all
of the research on bird populations in relation to timber management
in Florida has occurred in pinewoods (softwoods).

Timber management practices that most affect bird populations
in Florida are clearcutting, rotation length (time between harvests),
site preparation, and conversion from natural forests to plantations.
Select-tree harvest and natural regeneration are relatively rare. Studies
have been made of bird populations in relation to each of these
management practices.

In general, more species are found in longleaf pine than in slash
pine forests, and the older and more heterogeneous plantations
become, the better they serve as habitat for birds (Repenning 1983,
O'Meara et al. 1985, Repenning and Labisky 1985, Labisky and
Hovis 1987).

A major change to natural forests of Florida is the timber
management practice of conversion from longleaf pine forest to
plantations of loblolly (Pinus taeda), slash (P. elliotii), or sand pine
(P. clausa). Reduction of the longleaf pine ecosystem has been
estimated to be as large as 98% throughout the southeastern coastal
plain since pre-settlement time (Noss 1989). In the last 7 years,









AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 17


Table 4. Summary of published bird counts in Florida


County Citation Count Method

Alachua Caine (1986) spot-mapping


Alachua Land (1986) line transects
Bradford

Bradford Harris and spot-mapping
McElveen (1981)

Bradford Rowse (1980) transect


Bradford O'Meara et al. (1985) transect

DeSoto Cutright (1981) transect


Humphrey (1985)

Maehr (1980)


Harris et al.
(1975)


Engstrom et al.
(1984)

Repenning (1983)


Labisky and Hovis
(1987)

Hirth and Marion
(1979)

FGFWFC (1976)


Monroe Robertson (1955)


point count

point count


mist netting



spot-mapping


transect


line transects


transect


point count






spot-mapping


La.


Habitat Type Year(s)

slash pine flatwoods 1985-86
clearcut

slash pine plantations 1985


cypress dome 1977-78
flatwoods clearcut

pine flatwoods 1977-78
cypress-bay

flatwoods 1978-

pasture 1976
meadow
cropland

coastal forest 1984

flatwoods, settling ponds, 1978-79
restored mines

70-year old longleaf pine 1973-74
slash pine plantation
mature slash pine

successional oldfield 1967-82
pine forest

longleaf pine flatwoods 1981
slash pine plantations

longleaf pine forests 1980-81
slash pine forest

pasture 1978


loblolly pine hammock 1975-76
pine flatwoods
mature scrub
turkey oak
xeric hammock
mixed swamp

abandoned key lime grove 1951-52
tropical hammock
coastal hammock
pine woods
edge habitats


Duval

Hamilton


Hamilton



Leon


Liberty


Liberty


Manatee


Marion









NONGAME WILDLIFE PROGRAM FINAL REPORT


Table 4. (continued)


County Citation Count Method Habitat Type Year(s)


Nassau ESE (1984)


Nassau
Duval


Cox (1988)


Orange Humphrey
et al. (1985)

Osceola O'Meara (1984)
(1984)

Pinellas Rohwer and
Woolfenden (1969)


Polk


Schoes and
Humphrey (1987)


Putnam Humphrey
et al. (1985)

Putnam Maehr et al. (1982)



[North Harris and Wallace
Florida] (1984)


point count

point count


transect


point count


transect


transect


transect


transect


point count


coastal habitats

maritime hammock


abandoned citrus grove


cypress swamp


wet maple woods
oak ridge

8 post-phosphate mining
treatments

pine savanna


flatwoods
sandhill
hammock


mesic hardwood
hammocks


a Breeding Bird Censuses and Winter Bird Population Studies are not included.





longleafpine acreage in Florida is estimated to have declined by 23%
(Brown and Thompson 1988). The rapid disappearance of longleaf
pine forest has been compared to the conversion of natural prairies to
farmland in the midwestern U.S. (Noss 1989).

The conversion to plantations of slash or loblolly pine (species
that did not occur originally on the upland sites in Florida) has
boosted the percentage of Florida forests in plantation management
to the highest of any state in the Southeast (Brown and Thompson
1988). The soil is frequently prepared for plantations by disking,
chopping, or bedding, practices that greatly disturb the natural
groundcover. In commercial forests rotation is frequently limited to
30 years or less, while in national forests the rotation is extended to
60 to 70 years.


1987


1984


1983


1964


1978-79


1984


1980
(2 days)


1978







AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 19


Harris et al. (1975) conducted a study of the response of birds to
different intensities of site preparation. Two stands of mature trees,
1 longleaf and 1 slash pine, and 9 0.4-ha plots of 9-year-old planted
slash pine were compared. The 9 plots of planted slash pine were
divided among treatments of 3 plots each: high, medium, and low
intensity site preparation. Data were collected by transect counts and
mistnets. The mistnet data showed the highest numbers of captures
in the intermediate site intensity treatment, but the transect results had
many more species and individuals in the low intensity site prepared
treatment plots. All site prepared treatments had fewer bird species
and individuals than did the two mature forests. Of the two mature
forests, longleaf had significantly more species and individuals than
slash pine. Intensive site preparation enhanced midstory woody
species, reduced grasses and forbs, and increased structural uniformity
(White et al. 1975), which strongly affected bird abundance (Harris
et al. 1975).

Conversion of natural pinelands to plantations also affects birds
by altering the number and quality of snags. Land (1986) conducted
line transects in 20 slash pine plantations from 17 to 38 years old. Of
29 species of birds counted, 7 species were primary cavity nesters and
4 species were secondary cavity nesters. The number of clusters of
snags (caused by beetle infestations) and stand diameter at breast
height (dbh) were significant positive correlates of cavity nesting
bird density. In Alachua County Caine (1986) found that in clearcut
plots that had snags and nest boxes added the number of birds was
more than doubled over the number in control plots. Based on a study
in Bradford and Alachua counties Land (1986) recommended that
pine plantations could be improved for wildlife by (1) increasing
rotation age, (2) burning on a 4- to 5-year cycle, and (3) leaving
lightning struck trees.

Agriculture.- The effects of agricultural practices on bird
populations in Florida are poorly studied. Conversion from natural
to agricultural environments, field size, the quantity and quality of
buffer habitat, cultivation methods, and application of chemicals are
all known to affect bird populations in Great Britain (O'Connor and
Shrubb 1986).

Cutright (1981) documented seasonal bird species abundance in
five major habitats in west-central Florida: pinewoods, unimproved
pasture, improved pasture, wet meadow, and cropland. Seventy-
seven bird species were observed. Species richness was greatest in







NONGAME WILDLIFE PROGRAM FINAL REPORT


the pinewoods and least in cropland. Using transect sampling, Hirth
and Marion (1979) recorded bird populations during four seasons in
native range in sparse pine flatwoods used for cattle grazing in
Manatee County. Out of a total of 49 species, they found 32
permanent residents, 13 winter residents, and 4 summer residents.
Eastern meadowlark (Sturnella magna) and northern bobwhite
(Colinus virginianus) were the most abundant breeding birds and
palm warbler (Dendroica palmarum), eastern meadowlark, and
savannah sparrow (Passerculus sandwichensis) were the most
common wintering species.

One comparative study has been made of bird populations in an
abandoned citrus grove in Orange County and natural longleaf pine/
turkey oak (Quercus laevis) in Putnam County (Humphrey et al.
1985). This study is part of a project to examine the feasibility of
restoring the grove site to pre-settlement vegetation of longleaf
pine and wiregrass (Aristida stricta). Abandonment of citrus and
conversion to other habitat types, especially urban or suburban types,
is likely to increase in central Florida because so many citrus trees
were killed in two severe freezes in the winters of 1985 and 1989.
The authors concluded that a restored longleaf pine savanna would
have fewer bird species than an orange grove, although the orange
grove lacked notable members of the savanna avifauna.

Urbanization.- Urban and suburban environments in Florida
are increasing. The current population is 13 million and it is expected
to reach nearly 16 million by the year 2000 (Bureau of Economic and
Business Research 1989). The only research on bird communities in
suburban settings in Florida was done in the 1960s in Pinellas County
(Woolfenden and Rohwer 1969a, b, c). In 3 study plots of a combined
area of 40.7 ha, Woolfenden and Rohwer (1969a) estimated that a
new suburb held a bird density of approximately 200 pairs per 40 ha,
but that 2 older suburbs with mature oak and pine trees contained 500
to 600 pairs per 40 ha. The density of birds in the new suburb was
comparable to the density found in natural habitats before urbanization.
Four species, house sparrow (Passer domesticus), mourning dove
(Zenaida macroura), blue jay (Cyanocitta cristata), and northern
mockingbird accounted for 90%_of the total number of birds in the 3
study plots. At least 10 species found in natural habitats were
eliminated by the spread of suburbs: red-tailed hawk (Buteo
jamaicensis), red-shouldered hawk (Buteo lineatus), bald eagle
(Haliaeetus leucocephalus), yellow-billed cuckoo (Coccyzus
americanus), chuck-will's-widow (Caprimulgus carolinensis), red-







AVIAN COMMUNITIES IN FLORIDA HABITATS -Engstrom 21


cockaded woodpecker (Picoides borealis), scrub jay (Aphelocoma
coerulescens), brown-headed nuthatch (Sitta pusilla), pine warbler
(Dendroica pinus), and Bachman's sparrow (Aimophila aestivalis).
Densities of 5 species probably changed little with the replacerrnt
of native s rbs- northerflicker (Colaptes auratus),
redbellied woodpecker (Melanerpes arnlin u), downy woodpecker
(Picoides pubescens), great crested flycatcher, common grackle
(Quiscalus quiscalus), and northern cardinal (Cardinalis cardinalis).

Phosphate Mining.- Phosphate mining has altered the landscape
of over 66,800 ha of land in Florida (Marion 1986). It is usually
accomplished by first using draglines remove "overburden" soil and
expose phosphate ore. The ore is then moved to piles. The piles of
ore are liquified by high-pressure water streams and the slurry is
sent to processing plants. After phosphate has been removed, the
waste slurry is dumped into settling ponds called "slime ponds." In
1975, the Florida Legislature passed a law that required reclamation
of newly mined lands. Reclamation of lands mined before 1975 was
required by a revision of the law in 1978. Mined areas in northern and
central Florida are a mixture of spoil mounds, lakes, and "reclaimed"
areas. Early unreclaimed mines were simply abandoned and have
undergone plant succession.

In a study of clay settling ponds, unreclaimed pits and spoil piles
(various ages after mining and with and without lakes), and reclaimed
pastures, Schoes and Humphrey (1987) found the highest number of
bird species and individuals in older unreclaimed pits and spoil piles.
Maehr (1980) and Maehr and Marion (1984) compared seasonal bird
use in five habitat types in Hamilton County in 1979 and 1980. Bird
species richness was greatest in early succession settling ponds,
lowest in reclaimed mines with natural flatwoods, and intermediate
in unreclaimed mines and late successional ponds. Of all habitats,
late successional ponds had the greatest density of birds; reclaimed
mines had the lowest density. Maehr (1984) compared the species
lists and qualitative measures of abundance of 3 studies of birds on
phosphate mines.

Forest Fragmentation.- The size and shape of natural habitats
are factors that may affect species richness and composition of brd
communities (Robbins 1 .979). Harris and Wallace (1984) conducted
point counts during April and May in 12 mesic hardwood hammocks
that ranged in size from 0.4 to 30 ha. Ten of the study sites were less
than 10 ha. Thirty-one species were encountered on the 12 study







NONGAME WILDLIFE PROGRAM FINAL REPORT


sites. Four species, northern cardinal, Carolina wren (Troglodytes
aedon), red-eyed vireo (Vireo erythrocephalus), and northern parula
(Parula americana), were found in nearly every habitat island
regardless of size. Barred owl (Strix varia), great homed owl (Bubo
virginianus), red-shouldered hawk, red-tailed hawk, and common
flicker were only observed on habitat patches greater than or equal to
8 ha.

In a study of the effects of habitat-island size on species
composition, Harris and McElveen (1981) found a strong positive
relationship between species number and area in small cypress
domes (<8 ha), but O'Meara (1984) did not find a similar relationship
among larger islands (7 to 229 ha). O'Meara concluded that habitat
islands 10 to 20 ha are large enough to maintain the bird species that
typically use cypress ponds in Florida.

Fire.- Prescribed fire is an essential management tool used to
maintain upland pine forests in the southeastern United States.
Without it, pinelands are replaced by hardwood forests. In spite of its
importance, relatively little research has been done in Florida on the
direct and indirect effects of fire on bird populations. Mortality and
nest loss would be direct effects of fire. Alteration of vegetation
structure and composition have indirect and possibly long-term
effects on bird populations. Fire frequency, season, uniformity, and
intensity are components that determine the direct and indirect
effects.

W. W. Baker and R. L. Crawford conducted bird counts on a plot
of old-field pinewoods for 15 years following fire exclusion in 1966
(Engstrom et al. 1984). Substantial successional changes in the
vegetation composition and structure were matched by changes
in the bird community. Open woods species, such as Bachman's
sparrow and blue grosbeak (Guiraca caerulea), left the plot within a
few years of fire exclusion. Species associated with mesic habitats
(e.g. wood thrush [Hylocichla mustelina] and hooded warbler
[Wilsonia citrina]) were observed during the last five years of the
study. Canopy nesting species were relatively unaffected by the
changes, but this may change as pines are replaced by hardwoods
over a longer time period.

Vogl (1973) failed to observe any direct mortality of birds during
a fire set in January in wetland vegetation next to a pond in Leon
County. In fact, birds seemed either undisturbed or even attracted to








AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 23


the fire. The results of counts during four months following the fire
resulted in over three times as many observations of birds along the
burned shoreline as along an unburned control.

Coastal Development.- Coastal forests and habitats may be
particularly important for birds during fall and spring migration.
Humphrey (1985) studied fall migration in forest and spoil
island habitats on Fort George Island. Because he concluded that
coastal forests are very important for migrating birds, he offered the
following suggestions to help minimize the effects of suburbanization:
(1) leave as much of the original vegetation as possible, including
buffers around houses and golf courses, and (2) portions of original
habitat that can be protected shouldn't be disrupted by clearing
shrubs or mowing groundcover. In a study of Amelia Island,
Environmental Science and Engineering (1984) found that migrants
used a housing development as much as forest habitats that had not
been cleared; however, the authors didn't give a detailed description
of the study areas. Cox (1988) counted birds during spring migration
in large and small maritime hammocks in Duval and Nassau counties.
He found that certain species, such as northern parula, black-and-
white warbler (Mniotilta varia), ovenbird (Seiurus aurocapillus),
and summer tanager (Piranga rubra) exhibited a preference for large
hammocks; mourning dove, fish crow (Corvus ossifragus), and
brown-headed cowbird (Molothrus ater) were detected more
frequently in small hammocks. Cox concluded that size and quality
of coastal habitat are probably important for migrant birds.

Miscellaneous Studies.- Basic quantitative studies of birds
have been made of several regional vegetation types. Rowse (1980)
documented the seasonal avifauna of naturally regenerated pine
flatwoods with extensions of cypress-bay. She found more species
used the cypress-bay habitat, but relative abundance of birds in the
two habitats was not significantly different. Transients were not an
important part of the community.

Robertson (1955) conducted a major study of the birds of many
habitats of south Florida. He conducted 22 spot map censuses
of three pinewoods, five forest-edge areas, two tropical hammock
forests, a coastal mangrove island, two sawgrass prairies, and one
coastal prairie.

Weiser (1973) obtained lists of the avifauna of polluted and
unpolluted marshes in north Florida. S.A. Nesbitt, D.T. Gilbert, and








NONGAME WILDLIFE PROGRAM FINAL REPORT


D.B. Barbour collected relative abundance data from point counts at
8 permanent study sites in 6 major habitats (mature scrub, loblolly
pine hammock, pine flatwood, turkey oak, xeric hammock, and
mixed swamp) in relation to the cross-Florida barge canal (Florida
Game and Fresh Water Fish Commission 1976). Noss (1988) studied
populations of birds in edge and interior habitats in 5.0-ha plots in
hardwood hammock in Alachua County.

Evaluation of the BBC and WBPS

The main programs in North America to monitor changes in
numbers of birds over time and space are the Breeding Bird Survey
(BBS) and the Christmas Bird Count (CBC) (Robbins et al. 1986,
Butcher 1990). Two strengths of the BBS and CBC are that they are
geographically extensive and the number of samples of the
environment is large. Both of these programs provide data on the
numbers and distributions of birds in Florida, but they don't have
information on numbers of birds in specific habitats.

The BBC (not BBS) has been used to monitor local populations.
Occasionally, local changes are viewed within the context of larger-
scale phenomena. For example, the decline of neotropical migrant
species on some long-term BBC study plots has opened a controversy
over whether local declines are part of widespread trends and what
are the possible causes (Robbins 1979, Hall 1984, Johnston and
Winings 1987, Terborgh 1989). In some cases, population trends
from intensive and extensive studies can be compared (Holmes and
Sherry 1988). This could be done more often with the BBC and BBS
or with the CBC and WBPS.

A strength of the BBC and WBPS is that estimates of density, or
at least relative abundances of bird species, are provided for
specific habitats. This is extremely valuable, because availability
and quality of habitat are essential for the long-term preservation of
birds, as well as all other animals. Data provided by these plot-based
counts can be used with a habitat inventory in Florida. If enough
counts are made in a variety of habitats, it might be possible to
construct models of habitat preference for a large number of bird
species in the state.

A number of aspects of the BBC have been criticized. One
criticism is that interpretation of clusters of registrations used to
indicate the location of territorial individuals is too variable among







AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 25


observers (Enemar et al. 1978). In Great Britain, the field maps for
the Common Bird Count are interpreted by experienced staff at the
British Trust for Ornithology. This would be very difficult to do for
the BBC, but the instructions for map interpretation could be
strengthened. The procedure for interpretation of partial territories
should be applied consistently. This would have a bearing on
estimates of species richness. BBC methods also can be strengthened
by emphasizing the need to record simultaneous registrations
(Tomialojc 1980), by making plot size fairly large (Engstrom 1981),
and by maintaining consistent standards of effort over time and
among plots (Engstrom and James 1984).

Research on bird communities in Florida has not been well-
distributed geographically (Figures 1 and 2) nor have all habitats
been studied. For example, mangrove, bay, cypress, and gum
swamps and forests dominated by exotic tree species need further
study. It is also important to note relatively few sites have been
studied over many years during the breeding season and winter
(Table 1).

The strengths and value of the BBC and WBPS far outweigh the
weaknesses. To date, these databases represent the most extensive
collection of information on the bird populations in specific habitats
in Florida. Independent study reports, including theses, dissertations,
and published papers, are highly diverse, and direct comparisons of
results among these studies would be difficult and risky. Expanding
and improving the BBC and WBPS would be an inexpensive way to
provide good data on the status and trends of bird communities of
Florida.

The Peninsula Effect

A factor that complicates interpretation of patterns of bird species
richness in Florida is the peninsula effect: the gradual diminution of
the number of species from north to south on the peninsula. Robertson
(1955), Robertson and Kushlan (1974), Rohwer and Woolfenden
(1969) and Emlen (1978) have documented decreasing numbers of
bird species within habitats in peninsular Florida progressively
farther south from northern Florida. Wamer (1977) conducted a
study of the Florida peninsula effect using Breeding Bird Survey data
and habitat data that he collected. He concluded that most of the
peninsula effect was caused by replacement of habitats that naturally
supported relatively more bird species with species-poor habitats.







NONGAME WILDLIFE PROGRAM FINAL REPORT


Emlen (1978) conducted transects in pinewoods habitats along
the Florida peninsula. He found that the density of birds decreased
in apparently similar forests as latitude decreased. He hypothesized
that the more southern populations were less well adapted to their
environments because of centrifugal gene flow from mainland
populations. Emlen (1980) compared winter bird communities in
pine forests in southern Florida and the northern Bahama Islands.
More than half of the species that used Florida pinewoods in winter
were migrants. He concluded that the niches used by migrants in the
winter were left unfilled in other seasons.








AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 27


CONCLUSIONS

Generally, species richness among Florida BBCs is lowest in
structurally simple habitats saltwater marshes and highest in
forests. Patterns of species richness among forests are more Tdilicult-
to discern (Figures 3 and 6). Comparing species richness among
BBCs at a large geographical scale, James and Wamer (1982) found
coniferous forests to have fewer species than deciduous forests.
Wiens (1975) also found relatively low species richness in
coniferous forests. A detailed examination of patterns of species
richness in relation to structural aspects of forest habitats in Florida
is beyond the scope of this study. However, in this study, I found that
species richness of coniferous forests in Florida was not lower than
that of deciduous forests.

Species richness can be used as a criterion for evaluating a habitat
or parcel of land for conservation purposes (Fuller 1980, Usher
1986). If the number of species in an area is of interest, rarefaction
has many advantages over diversity indices (James and Rathbun
1981). Although species richness may be of interest, the presence of
particular species is also very important. This is especially true for
species that are sensitive to habitat disturbance or are very habitat
specific.

Given the rapid alteration of many natural habitats in Florida,
conducting an inventory and describing the avian community
quantitatively is a high priority for conservation purposes. The BBC
and WBPS provide a substantial foundation for such an endeavor.
Models of habitat use for individual species can be developed if
quantitative vegetation samples are collected with the bird counts.








NONGAME WILDLIFE PROGRAM FINAL REPORT


RECOMMENDATIONS

(1) Work with the Florida Natural Areas Inventory, Department
of Natural Resources, Game and Fresh Water Fish Commission,
and Division of Forestry to locate the best representations of habitat
in the state. Nature Conservancy preserves, state parks, national
forests, and university research areas are logical sites for long-
term bird studies. Encourage particular bird clubs or individuals to
take on responsibility for research on a study site and possibly even
provide small grants. Perhaps the Florida Ornithological Society
could take on the responsibility to organize an "Ornithological
Survey of the Habitats of Florida" as a follow-up to the Breeding
Bird Atlas.

(2) Most of the research on bird populations in Florida has been
done in pine forests. Research on forestry management practices
indicates that the practices of clearcutting, short rotation, and site
preparation are detrimental to bird populations, especially some of
the species endemic to the southeastern U.S. Alternatives to current
commercial timber management practices should be developed for
national and state-owned forests and encouraged on private lands.

(3) Researchers and environmental consultants should be
encouraged to publish their studies of Florida bird communities.
Contributions to the BBC or WBPS if the spot-mapping method is
used in their studies would be especially valuable. This would add to
a widely accessible computerized data base.

(4) Some studies indicate that urbanization or suburbanization
may increase numbers of some already common birds, but deplete
populations of more highly specialized species. It might be possible
to enhance the survival of some Florida birds that have narrow habitat
requirements by encouraging developers and landowners to plant
native plants in new housing developments (see Mills et al. 1989).
However, fire-dependent plant communities would be difficult to
maintain in this manner.

(5) Issuance of permits to destroy large or rare areas of natural
habitat should be tied to conducting pre- and post-disturbance studies
of birds and other vertebrate wildlife.

(6) Bird studies in relation to Development of Regional Impact
(DRI) reviews should be done according to standard ornithological







AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 29




techniques. Guidelines could be established by the state and required
for use by consulting companies to which the state issues contracts.
Data collected in this way would add significantly to the database of
quantitative studies of birds in Florida and would allow comparisons
among studies.







NONGAME WILDLIFE PROGRAM FINAL REPORT


LITERATURE CITED

Arrhenius, 0. 1921. Species and area. J. Ecology 9:95-99.

Brown, M.J., and M.T. Thompson. 1988. Forest statistics for Florida,
1987. U.S. For. Serv., Res. Bull. SE-101. 61 pp.

Bureau of Economic and Business Research. 1989. Florida Statistical
Abstract. Univ. Presses Fla., Gainesville. 718 pp.

Butcher, G.S. 1990. Audubon Christmas Bird Counts. Pages 5-13 in
J.R. Sauer, and S.Droege, eds. Survey designs and statistical
methods for the estimation of avian population trends. U.S. Fish
Wildl. Serv., Biol. Rep. 90(1). 166 pp.

Caine, L.A. 1986. Augmentation of slash pine plantations with snags
to enhance breeding bird habitat. M.S. Thesis, Univ. Florida,
Gainesville. 82 pp.

Connor, E.F., and E.D. McCoy. 1979. The statistics and biology of
the species-area relationship. Am. Nat. 113:791-833.

Cox, J. 1988. The influence of forest size on transient and resident
bird species occupying maritime hammocks of northeast Florida.
Fla. Field Nat. 16:25-34.

D. Inkley, and R. Kautz. 1987. Ecology and habitat
protection needs of gopher tortoise (Gopherus polyphemus)
populations found on lands slated for large-scale development in
Florida. Fla. Game and Fresh Water Fish Comm., Nongame
Wild. Program Tech. Rep. No. 4, Tallahassee, Fla. 75 pp.

Cutright, N.J. 1981. Bird populations in five major west-central
Florida vegetation types. Fla. Sci. 44:1-13.

DeSante, D.F., and G.R. Geupel. 1987. Landbird productivity in
central coastal California: the relationship to annual rainfall and
a reproductive failure in 1986. Condor 89:636-653.

Emlen, J.T. 1971. Population densities derived from transect counts.
Auk 88:565-576.








AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 31


1978. Density anomalies and regulatory mechanisms
in land bird populations on the Florida peninsula. Am. Nat.
112:265-286.

1980. Interactions of migrant and resident land birds in
Florida and Bahama pinelands. Pages 133-143 in A. Keast and
E.S. Morton, eds.Migratory birds in the Neotropics: ecology,
behavior, distribution, and conservation. Smithsonian Inst. Press,
Washington, D.C.

Enemar, A., B. Sjostrand, and S. Svensson. 1978. The effect of
observer variability on bird census results obtained by a territory
mapping method. Omit. Scand. 9:31-39.

Engstrom, R.T. 1981. The species-area relationship in spot-map
censusing. Stud. Avian Biol. No. 6:421-425.

R.L. Crawford and W.W. Baker. 1984. Breeding bird
populations in relation to changing forest structure following fire
exclusion: a 15-year study. Wilson Bull. 96:437-450.

and F.C. James. 1981. Plot size as a factor in Winter
Bird Population Studies. Condor 83:34-41.

_. 1984. An evaluation of methods used in the Breeding
Bird Census. Am. Birds 38(1):19-23.

Environmental Science and Engineering. 1984. Amelia Island bird
survey. Unpublished.

Fager, E.W. 1972. Diversity: a sampling study. Am. Nat. 106:293-
310.

Florida Department of Transportation. 1985. Florida land use, cover
and forms classification system.

Florida Game and Fresh Water Fish Commission. 1976. Cross
Florida barge canal restudy report. Wildlife Study. Vol. III. Bird
study. 250 pp.

Fuller, R.J. 1980. A method for assessing the ornithological interest
of sites for conservation. Biol. Cons. 15:229-239.








NONGAME WILDLIFE PROGRAM FINAL REPORT


Hall, G.A. 1964. Breeding Bird Censuses-why and how. Aud. Field
Notes 18:413-416.

1984. Population decline of Neotropical migrants in an
Appalachian forest. Am. Birds 38:14-18.

Harris, L.D., and J.D. McElveen. 1981. Effect of forest edges on
north Florida breeding Birds. Intensive Manage. Practices
Assessment Rep. 6:1-24.

and R.D. Wallace. 1984. Breeding bird species in
Florida forest fragments. Proc. Annu. Conf. Southeast Assoc.
Fish and Wildl. Agencies 38:87-96.

L.D. White, J.E. Johnston, D.G. Milchunas. 1975.
Impact of forest plantations on north Florida wildlife and habitat.
Proc. Ann. Conf. Southeast Assoc. Fish and Wildl. Agencies
28:659-667.

Heck, K.L., Jr., G. van Belle, and D.S. Simberloff. 1975. Explicit
calculation of the rarefaction diversity measurement and the
determination of sufficient sample size. Ecology 56:1459-1461.

Hirth, D.H., and W.R. Marion. 1979. Bird communities of a south
Florida flatwoods. Fl. Sci. 42:142-151.

Holmes, R.T., and T.W. Sherry. 1988. Assessing population trends of
New Hampshire forest birds: local vs. regional patterns. Auk
105:756-768.

Humphrey, S.R. 1985. Fort George Island: fall migrant forest birds.
Unpublished.

J.F.Eisenberg, and R.Franz. 1985. Possibilities for
restoring wildlife of a longleaf pine savanna in an abandoned
citrus grove. Wildl. Soc. Bull. 13:487-496.

Hurlbert, S.H. 1971. The nonconcept of species diversity: a critique
and alternative parameters. Ecology 52:577-586.

and S. Rathbun. 1981. Rarefaction, relative abundance,
and diversity of avian communities. Auk 98:785-800.







AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 33


and H.H. Shugart. 1970. A quantitative method of
habitat description. Aud. Field Notes 24:727-736.

and N.O. Wamer. 1982. Relationships between
temperate forest bird communities and vegetation structure.
Ecology 63:159-171.

Johnston, D.W., and D.I. Winings. 1987. Natural history of
Plummer's Island, Maryland. XXVII. The decline of forest
breeding birds on Plummer's Island, Maryland and vicinity.
Proc. Biol. Soc. Washington 100:762-68.

Kolb, H. 1965. The Audubon Winter Bird-Population Study. Aud.
Field Notes 19:432-434.

Labisky, R.F., and J.A. Hovis. 1987. Comparison of vertebrate
wildlife communities in longleaf pine and slash pine habitats in
north Florida. Pages 201-228 in H. A. Pearson, F. E. Smeins, and
R. E. Thill, eds. Ecological, physical, and socio-economic
relationships within southern national forests. U.S. Dep. Agric.,
For. Serv., South. For. Exp. Stn. Gen. Tech. Rep. SO-68. 293 pp.

Land, E.D. 1986. Snag characteristics and relationships to cavity-
nesting birds in slash pine plantations of north-central Florida.
M.S. Thesis, Univ. Florida, Gainesville. 57 pp.

Maehr, D.S. 1980. Avian abundance and habitat preferences on new
habitats created by phosphate mining. M.S. Thesis, Univ. Florida,
Gainesville. 122 pp.

_. 1984. Status of birds using phosphate-mined lands in
Florida. Am. Birds 38:28-31.

M.C. Conner, andJ. Stenberg. 1982. Bird diversity and
abundance in three plant communities in Putnam Co., Florida.
Fla. Field Nat. 10:69-73.

and W.R. Marion. 1984. Bird abundance and distribution
in a north Florida phosphate mine. Proc. Ann. Conf. Southeast
Assoc. Fish and Wildl. Agencies 38:111-120.

Marion, W. R. 1986. Phosphate mining: regulations, reclamation
and revegetation. Final report, Fla. Inst. Phosphate Res., Bartow,
Fla. 72 pp.







NONGAME WILDLIFE PROGRAM FINAL REPORT


Mills, G.C., J.B. Dunning, Jr., and J.M. Bates. 1989. Effects of
urbanization on breeding bird community structure in southwestern
desert habitats. Condor 91:416-428.

Noss, R.F. 1988. Effects of edge and internal patchiness on habitat
use by birds in a hardwood forest. Ph.D. Thesis, Univ. Florida,
Gainesville. 109 pp.

1989. Longleaf pine and wiregrass and keystone
components of an endangered ecosystem. Natur. Areas Journ.
9:211-213.

O'Connor, R. J., and M. Shrubb. 1986. Farming and birds. Cambridge
University Press, Cambridge.

O'Meara, T. 1984. Habitat-island effects on the avian community in
cypress ponds. Proc. Ann. Conf. Southeast Assoc. Fish and
Wildl. Agencies 38:97-110.

L.A. Rowse, W.R. Marion, and L.D. Harris. 1985.
Numerical responses of flatwoods avifauna to clearcutting. Fl.
Sci. 48:208-219.

Ralph, C.J., and J.M. Scott, eds. 1981. Estimating numbers of
terrestrial birds. Studies in Avian Biology No. 6.

Repenning, R.W. 1983. Effects of even-age timber management on
bird communities of the longleaf pine forest of north Florida.
M.S. Thesis, Univ. Florida, Gainesville. 41 pp.

and R.F. Labisky. 1985. Effects of even-age timber
management on bird communities of the longleaf pine forest in
northern Florida. J. Wildl. Manage. 49:1088-1098.

Reynolds, R.T., J.M. Scott, and R.A. Nussbaum. 1980. A variable
circular-plot method for estimating bird numbers. Condor
82:309-313.

Robbins, C.S. 1970. Recommendations for an international standard
for a mapping method in bird census work. Am. Birds 24:3-6.

1979. Effect of forest fragmentation on bird populations.
Pages 198-212 in R.M. DeGraaf and K.E. Evans, tech. coords.
Management of north central and northeastern forests for nongame
birds. U.S. For. Serv., Gen. Tech. Rep. NC-51.








AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 35


D. Bystrak, and P.H. Geissler. 1986. The Breeding Bird
Survey: its first fifteen years, 1965-1979. U.S. Dept. Interior, Fish
and Wildl. Serv., Res. Publ. 157, Washington, D.C.

Robertson, W.B., Jr. 1955. An analysis of the breeding-bird
populations of tropical Florida in relation to the vegetation. Ph.D.
Thesis, Univ. Illinois, Urbana.

and J.A. Kushlan. 1974. The southern Florida avifauna.
In Memoir 2: Environments of south Florida: present and past.
Miami Geol. Soc.

Rohwer, S.A., and G.E. Woolfenden. 1969. Breeding birds of two
Florida woodlands: comparisons with areas north of Florida.
Condor 71:38-48.

Rowse, L.A. 1980. Avian community composition of a north Florida
flatwoods. M.S. Thesis, Univ. Florida, Gainesville. 66 pp.

Sanders, H.L. 1968. Marine benthic diversity: a comparative study.
Am. Nat. 102:243-282.

Schoes, R.S., and S.R. Humphrey. 1987. Terrestrial plant and wildlife
communities on phosphate-mined lands in central Florida. Florida
State Mus. Spec. Sci. Rep. No. 3, Gainesville, FL. 192 pp.

Simberloff, D.S. 1972. Properties of the rarefaction diversity
measurement. Am. Nat. 106:414-418.

1978. Use of rarefaction and related methods in
ecology. Pages 150-165 in K.L. Dickson, J. Cairns, Jr., and R.J.
Livingson, eds. Biological data in water pollution assessment:
quantitative and statistical analyses. Am. Soc. for Testing and
Materials STP 652.

Stewart, R.E. 1949. Thirteenth Breeding Bird Census. Aud. Field
Notes. 3:255.

Temple, S. A., and J. A. Wiens. 1989. Bird populations and
environmental changes: can birds be bio-indicators? Am. Birds
43:260-270.

Terborgh, J. 1989. Where have all the birds gone? Princeton Univ.
Press, Princeton, New Jersey.








NONGAME WILDLIFE PROGRAM FINAL REPORT


Tomialojc, L. 1980. The combined version of the mapping method.
Pages 92-106 in H. Oelke, ed. Bird census work and nature
conservation. Proc. VI Int. Conf. Bird Census Work / IV Meeting
Eur. Ornithol. Atlas Committee (Gottingen 1979). Dachverband
Deutsche Avifaunisten, Gottingen, W. Germany.

Usher, M.B. 1986. Wildlife conservation evaluation: attitudes, criteria,
and values. Pages 3-44 in M.B. Usher, ed. Wildlife conservation
evaluation. Chapman and Hall, London.

Van Velzen, W.T. 1972. Breeding Bird Census instructions. Am.
Birds 26:927-931.

Verner, J. 1985. Assessment of counting techniques. Pages 247-302
in R.J. Johnson, ed. Current Ornithology. Vol. 2. Plenum Press,
New York.

Vogl, R.J. 1973. Effects of fire on the plants and animals of a Florida
wetland. Am. Midl. Nat. 89:334-347.

Wamer, N.O. 1977. Avian diversity and habitat in Florida: an
analysis of a peninsular diversity gradient. M.S. Thesis, Florida
State Univ. 54 pp.

Weiser, C.E. 1973. A comparison of the avifauna between an
unpolluted marsh and a polluted marsh on the Gulf coast of
Florida. MS. Thesis, Florida State Univ., Tallahassee. 39 pp.

White, L.D., L.D. Harris, J.E. Johnston, and D.G. Milchunas. 1975.
Impact of site preparation on flatwoods wildlife habitat. In Proc.
Ann. Conf. Southeast Assoc. Fish and Wildl. Agencies 28:347-
353.

Wiens, J.A. 1975. Avian communities, energetic, and functions in
coniferous forest habitats. In The symposium on management of
forest and range habitats for nongame birds. U.S. For. Ser., Gen.
Tech. Rep. WO-1, 343 pp.

Williams, A.B. 1936. The composition and dynamics of a beech-
maple climax community. Ecol. Monogr. 6:317-408.

Woolfenden, G.E., and S.A. Rohwer. 1969a. Breeding birds in a
Florida suburb. Bull. Florida State Mus. 13:1-83.







AVIAN COMMUNITIES IN FLORIDA HABITATS -Engstrom 37




1969b. Bird populations in the suburbs and two woodland
habitats in Pinellas County, Florida. Fla. State Board of Health
Monogr. No. 12:101-109.

1969c. Relative abundance of bird populations in
residential suburbs in the Tampa Bay area of Florida. Fla. State
Board of Health Monogr. No. 12:110-117.









NONGAME WILDLIFE PROGRAM FINAL REPORT


Appendix A. List of Breeding Bird Censuses (BBCs) conducted in Florida.

1. Allen, T. T., D. E. Birkenholz, and D. A. Jenni. 1960. BBC 18. Longleaf Pine-Turkey Oak
Association. Aud. Field Notes 14:497-498.

2. Baker, W.W. 1989. BBC 25. Mature Beech-Magnolia Forest. J. Field Omith. 60:39.

3. Caprio, A., and D.L. Taylor. 1982. BBC 87. Cutover Subtropical Slash Pine Forest. Am.
Birds 36:74.

4. 1982. BBC 88. Virgin Subtropical Slash Pine Forest-February Bur. Am. Birds
36:74.

5. 1982. BBC 89. Virgin Subtropical Slash Pine Forest-December Bur. Am.
Birds 36:74-75.

6. 1982. BBC 90. Virgin Subtropical Slash Pine Forest-Summer Bum. Am. Birds
36:75.

7. 1982. BBC 91. Virgin Subtropical Slash Pine Forest-Unburned Control. Am.
Birds 36:75.

8. Christman, S.P. 1984. BBC 171. Gulf of Mexico Barrier Island. Am. Birds 38:119-120.

9. Engstrom, R. Todd. 1981. BBC 32. Mature beech-magnolia forest. Am. Birds. 35:58.

10. 1982. BBC 31. Mature beech-magnolia forest. Am. Birds. 36:61.

11. Jenni, D. A., T. T. Allen, and C. H. Trost. 1961. BBC 43. Longleaf Pine-Turkey Oak
Association. Aud. Field Notes 15:524.

12. D. E. Birkenholz, and J. D. Ligon. 1962. BBC 19. Longleaf Pine-Turkey Oak
Association. Aud. Field Notes 16:527-528.

13. Kale, H. W., II, and L. A. Webber. 1968. BBC 29 (1966). Live Oak-Cabbage Palm Coastal
Hammock. Aud. Field Notes 22:676-680.

14. 1968. BBC 29 (1967). Live Oak-Cabbage Palm Coastal Hammock. Aud.
Field Notes 22:676-680.

15. 1968. BBC 29 (1968). Live Oak-Cabbage Palm Coastal Hammock. Aud.
Field Notes 22:676-680.

16. 1968. BBC 30 (1966). Oak-Palm-Hickory Hammock and Maple Swamp.
Aud. Field Notes 22:680-684.

17. 1968. BBC 30 (1967). Oak-Palm-Hickory Hammock and Maple Swamp.
Aud. Field Notes 22:680-684.

18. 1968. BBC 30 (1968). Oak-Palm-Hickory Hammock and Maple Swamp.
Aud. Field Notes 22:680-684.

19. 1968. BBC 58 (1967). Citrus Grove. Aud. Field Notes 22:708-710.

20. 1968. BBC 58 (1968). Citrus Grove. Aud. Field Notes 22:708-710.

21. Kushlan, J. A., and M. S. Kushlan. 1977. BBC 144. Everglades Marsh. Am. Birds
31:83.










AVIAN COMMUNITIES IN FLORIDA HABITATS -Engstrom 39


22. Leenhouts, W.P. 1982. BBC 186. Cordgrass Salt Marsh I. Am. Birds 36:99

23. 1982. BBC 187. Cordgrass Salt Marsh II. Am. Birds 36:99-100.

24. Little, C.D. 1981. BBC 68. Slash Pine-Turkey Oak Forest. Am. Birds 35:66.

25. Maynard, W.R., D.L. Taylor, and R. Rochefort. 1981. BBC 83. Virgin Subtropical Slash
Pine Forest. Am. Birds 35:70.

26. McDonald, M.V. 1982. BBC 188. Gulf Coast Salt Marsh. Am. Birds 36:100.

27. 1983. BBC 176. Gulf Coast Salt Marsh. Am. Birds 37:100.

28. 1984. BBC 170. Gulf Coast Salt Marsh. Am. Birds 38:119.

29. 1985. Unpublished BBC 113. Gulf Coast Salt Marsh. Am. Birds 39:114.
(Data stored at Cornell Laboratory of Ornithology)

30. 1986. Unpublished BBC 92. Gulf Coast Salt Marsh. Am. Birds 40:71.
(Data stored at Cornell Laboratory of Ornithology)

31. 1987. Unpublished BBC. Gulf Coast Salt Marsh. Cornell Laboratory of
Ornithology.

32. 1988. Unpublished BBC 72. Gulf Coast Salt Marsh. Am. Birds 42:148.
(Data stored at Cornell Laboratory of Ornithology)

33. 1989. BBC 81. Gulf Coast Salt Marsh. J.Field Ornith. 60:68.

34. 1990. Gulf coast salt marsh. J.Field Orith. 61:77.

35. Niemi, G.J. 1981. BBC 81. Longleaf Pine Forest. Am. Birds 35:69.

36. Patterson, G.A., W.B. Robertson, and D.E. Minsky. 1980. BBC 65. Slash Pine-
Cypress Mosaic. Am. Birds 34:61-62

37. Post, W. 1981. BBC 200. Salt Marsh. Am. Birds 35:99.

38. Robertson, H. M., C.F. Sumner, H.W. Werner, and G.E. Woolfenden. 1968. BBC 31.
Live Oak-Cabbage Palm Hammock. Aud. Field Notes 22:684.

39. Vemick, K., and D.L. Taylor. 1981. BBC 82. Cutover Subtropical Slash Pine Forest. Am.
Birds 35:69-70.

40. Webber, L. A., and H. W. Kale, II. 1969. BBC 63. Citrus Grove. Aud. Field Notes
23:744-745.

41. 1969. BBC 73. Oak-Palm-Hickory Hammock and Maple Swamp. Aud. Field
Notes 23:751.

42. West, R.L. 1989. BBC 82. Longleaf Pine Forest-April Burn. J. Field Omith. 60:68-69.

43. 1989. BBC 83. Longleaf Pine Forest-Unburned Control. J. Field Omith.
60:69-70.

44. Woolfenden, G. E. 1969. BBC 50. Sand Pine Scrub. Aud. Field Notes 23:733-734.

45. 1969. BBC 51. Scrubby Flatwoods. Aud. Field Notes 23:734-735.










NONGAME WILDLIFE PROGRAM FINAL REPORT


46. 1969. BBC 52. Recently Burned Scrubby Flatwoods. Aud. Field Notes
23:735-737.

47. 1969. BBC 53. Slash Pine-Turkey Oak Association. Aud. Field Notes 23:737-
738.

48. 1969. BBC 54. Low Flatwoods with Bayhead. Aud. Field Notes 23:738.

49. Woolfenden, G.E., S.A. Rohwer, J.A. Stevenson, C.F. Sumner, and C.E. Winegamer, 1967.
BBC 34. Live Oak-Cabbage Palm Hammock. Aud. Field Notes 21:635-637.

50. and R.G. Allan. 1958. BBC 15. Longleaf Pine-Turkey Oak Association.
Aud. Field Notes 12:450.

51. T. T. Allen, and D. E. Birkenholz. 1959. BBC 14. Longleaf Pine-Turkey Oak
Association. Aud. Field Notes 13:466.









AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 41


Appendix B. List of Winter Bird Population Studies (WBPS) that were conducted
in Florida.

1. Baker, W.W. 1989. WBPS 2. Mature Beech-Magnolia Forest. J. Field Omith. 60:7-8.

2. Cink, C.L., and F.E. Lohrer. 1990. WBPS 16. Sand pine scrub. J. Field Omith. 61:13-
14.

3. 1990. WBPS 23. Recently burned Scrubby Flatwoods. J. Field Omith.
61:17.

4. Connor, H., and R.W. Loftin. 1985. WBPS 54. Barrier Beach and Saltwater Estuary.
Am. Birds 39:119.

5. Fisk, E. J. 1975. WBPS 46. Pineland, Scrub, and Residential. Am. Birds 29:770.

6. 1976. WBPS 38. Pineland, Scrub, and Residential. Am. Birds 30:1055.

7. 1978. WBPS 31. Pineland, Scrub, and Residential. Am. Birds 32:33-34.

8. 1979. WBPS 33. Pineland, Scrub, and Residential. Am. Birds 33:27.

9. Good, J.M. 1982. WBPS 9. Slash Pine. Am. Birds 36:31.

10. 1982. WBPS 51. Canal and Associated Spoil Bank. Am. Birds 36:41.

11. 1982. WBPS 70. Avocado Grove. Am. Birds 36:45.

12. Kale, H. W., II, and L. A. Webber. 1969. WBPS 16 (1967). Live Oak-Cabbage Palm
Coastal Hammock. Aud. Field Notes 23:531-533.

13. 1969. WBPS 16 (1968). Live Oak-Cabbage Palm Coastal Hammock. Aud.
Field Notes 23:531-533.

14. 1969. WBPS 16 (1969). Live Oak-Cabbage Palm Coastal Hammock. Aud.
Field Notes 23:531-533.

15. 1969. WBPS 17 (1967). Oak-Palm-Hickory-Hammock and Maple Swamp.
Aud. Field Notes 23:533-535.

16. 1969. WBPS 17 (1968). Oak-Palm-Hickory-Hammock and Maple Swamp.
Aud. Field Notes 23:533-535.

17. 1969. WBPS 17 (1969). Oak-Palm-Hickory-Hammock and Maple Swamp.
Aud. Field Notes 23:533-535.

18. 1969. WBPS 43 (1967). Citrus Grove. Aud. Field Notes 23:547-549.

19. 1969. WBPS 43 (1968). Citrus Grove. Aud. Field Notes 23:547-549.

20. 1969. WBPS 43 (1969). Citrus Grove. Aud. Field Notes 23:547-549.

21. Kushlan, J. A., and M. S. Kushlan. 1976. WBPS 63. Everglades Marsh. Am. Birds
30:1066-1067.

22. Loftin, R.W. 1970. WBPS 40. Grazed Pasture with Ponds. Aud. Field Notes 24:564-565.

23. 1971. WBPS 36. Grazed Pasture with Ponds. Am. Birds 25:654-655.









NONGAME WILDLIFE PROGRAM FINAL REPORT


24. 1987. WBPS 30. Barrier Beach and Saltwater Estuary. Am. Birds
41:158.

25. 1988. WBPS 35. Barrier Beach and Saltwater Estuary. Am. Birds
42:151.

26. 1989. WBPS 14. Barrier Beach and Saltwater Estuary. J. Field Ornith.
60:13-14.

27. 1990. WBPS 27. Barrier beach and saltwater estuary. J.Field Omith.
61:19-20.

28. Lohrer, F.E. 1990. WBPS 28. Mature citrus orchard. J. Field Ornith. 61:20.

29. Moseley, L. H. 1969. WBPS 49. Suburban School Park. Aud. Field Notes 23:552.

30. 1970. WBPS 46. Suburban School Park. Aud. Field Notes 24:566-567.

31. 1971. WBPS 48. Suburban School Park. Am. Birds 25:660.

32. 1972. WBPS 48. Suburban School Park. Am. Birds 26:683-684.

33. 1973. WBPS 54. Suburban School Park. Am. Birds 27:696.

34. 1974. WBPS 69. Suburban School Park. Am. Birds 28:726-727.

35. 1975. WBPS 80. Suburban School Park. Am. Birds 29:787.

36. 1976. WBPS 78. Suburban School Park. Am. Birds 30:1073.

37. 1978. WBPS 70. Suburban School Park. Am. Birds 32:47.

38. NeSmith, C. 1983. WBPS 14. Mature Beech-Magnolia Forest. Am. Birds 37:34.

39. NeSmith, C., and J. Cox. 1984. WBPS 109. Longleaf Pine Forest. Am. Birds 38:63.

40. Patterson, G.A., and W.B. Robertson, Jr. 1981. WBPS 34. South Florida Slash Pine.
Am. Birds 35:50.

41. 1981. WBPS 35. Slash Pine Cypress Mosaic. Am. Birds 35:30-31.

42. Robertson, H. M., H.L. Shoffner, C.F. Sumner, H. Werner, C.E. Winegarer, and G.E.
Woolfenden. 1968. WBPS 16. Live Oak-Cabbage Palm Hammock. Aud. Field
Notes 22:488.

43. Wass, M. 1955. WBPS 22. Sheltered Beach. Aud. Field Notes 9:301.

44. 1955. WBPS 23. Mixed Palmetto Scrub Grassland. Aud. Field Notes
9:301-302.

45. Wilson, J., and R. W. Loftin. 1979. WBPS 100. Barrier Beach and Saltwater Estuary.
Am. Birds 33:47-48.

46. 1980. WBPS 48. Barrier Beach and Saltwater Estuary. Am. Birds
34:36-37.

47. 1981. WBPS 56. Barrier Beach and Saltwater Estuary. Am. Birds 35:36.










AVIAN COMMUNITIES IN FLORIDA HABITATS --Engstrom 43


48.

49.

50.

51. Woolfenden,

52.

53.
24:559-560.


1982. WBPS 50. Barrier Beach and Saltwater Estuary. Am. Birds 36:41.

1983. WBPS 49. Barrier Beach and Saltwater Estuary. Am. Birds 37:42.

1984. WBPS 69. Barrier Beach and Saltwater Estuary. Am. Birds 38:56.

G. E. 1970. WBPS 29. Sand Pine Scrub. Aud. Field Notes 24:559.

1970. WBPS 30. Scrubby Flatwoods. Aud. Field Notes 24:559.

1970. WBPS 31. Recently Burned Scrubby Flatwoods. Aud. Field Notes


1970. WBPS 32. Slash Pine-Turkey Oak Association. Aud. Field
Notes 24:560.










NONGAME WILDLIFE PROGRAM FINAL REPORT


Appendix C. All bird species encountered on Breeding Bird Censuses in Florida. The
numbers refer to reference numbers in Appendix A.

Agelaius phoeniceus, 4, 8, 19, 20, 21, 22, 23, 40
Aimophila aestivalis, 35, 42, 43
Aix sponsa, 10
Ammodramus maritimus, 26, 27, 28, 29,30, 31,32, 33, 34, 37
Anhinga, see Anhinga anhinga
Anhinga anhinga, 21
Ani, Smooth-billed see Crotophaga ani
Aphelocoma coerulescens, 45, 46, 47
Aramus guarauna, 21
Archilochus colubris, 2, 8, 10, 13, 16, 17, 38, 49
Ardea herodias, 17
Barn-Owl, Common see Tyto alba
Blackbird, Red-winged see Agelaius phoeniceus
Bluebird, Eastern see Sialia sialis
Bobwhite, Northern see Colinus virginianus
Bubo virginianus, 47
Bunting, Indigo see Passerina cyanea
Buteo lineatus, 2, 9, 10, 13, 14, 15, 16, 17, 18, 21, 25, 41, 42, 43
Butorides striatus, 2, 13, 14, 15, 16, 18, 21, 41
Caprimulgus carolinensis, 8, 14, 15, 16, 17, 18, 24, 39, 44, 45, 47, 48
Cardinal, Northern see Cardinalis cardinalis
Cardinalis cardinalis, 2, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 24,
25, 29, 36, 38, 40,41,42,43,44,45,47, 48, 49
Cathartes aura, 2
Chaetura pelagica, 8
Chickadee, Carolina see Parus carolinensis
Chordeiles minor, 1, 11, 19, 20, 22, 39, 40, 42, 43, 46, 50, 51
Chuck-will's-widow, see Caprimulgus carolinensis
Cistothorus palustris, 26, 27, 28, 29, 30, 31, 32, 33, 34, 37
Coccyzus americanus, 2, 9, 10, 13, 14, 15, 16, 17, 18, 19, 20, 38, 40, 41, 49
Colaptes auratus, 2, 3, 13, 14, 15, 16, 17, 18, 25, 36, 41,42, 43, 45, 47
Colinus virginianus, 3, 4, 5, 6, 19, 20, 25, 35, 36, 39, 40, 42, 43, 46, 48
Columbina passerina, 13, 14, 15, 19, 20, 40, 45, 46
Contopus virens, 35, 42, 43
Corvus brachyrhynchos, 2, 4, 9, 10, 25, 36
Corvus ossifragus, 47
Cowbird, Brown-headed see Molothrus ater
Crotophaga ani, 20
Crow, Fish see Corvus ossifragus
Crow, American see Corvus brachyrhynchos
Cuckoo, Yellow-billed see Coccyzus americanus
Cyanocitta cristata, 2, 3, 9, 10, 11, 17, 18, 35, 38, 39, 42, 43, 44, 45, 46, 47, 48, 49
Dendroica dominica, 8
Dendroica pinus, 2, 3, 4, 5, 6, 7, 8, 24, 25, 35, 36, 39, 42, 43, 48
Dove, Mourning see Zenaida macroura
Dryocopus pileatus, 2, 9, 10, 13, 14, 15, 16, 17, 18, 25, 36, 38, 39, 41, 49
Duck, Wood see Aix sponsa
Eagle, Bald see Haliaeetus leucocephalus
Egretta tricolor, 21
Elanoidesforficatus, 25, 36
Empidonax virescens, 2, 9, 10
Falco sparverius, 45, 47
Flicker, Northern see Colaptes auratus
Flycatcher, Acadian see Empidonax virescens
Flycatcher, Great Crested see Myiarchus crinitus









AVIAN COMMUNITIES IN FLORIDA HABITATS Engstrom 45


Gallinule, Purple see Porphyrula martinica
Geothlypis trichas, 8, 21, 23, 25, 46
Gnatcatcher, Blue-gray see Polioptila caerulea
Grackle, Boat-tailed see Quiscalus major
Grebe, Pied-billed see Podilymbus podiceps
Grosbeak, Blue see Guiraca caerulea
Ground-Dove, Common see Columbina passerina
Guiraca caerulea, 8, 42, 43
Haliaeetus leucocephalus, 8
Hawk, Red-shouldered see Buteo lineatus
Heron, Great Blue see Ardea herodias
Heron, Green-backed see Butorides striatus
Heron, Tricolored see Egretta tricolor
Hirundo rustica, 8
Hummingbird, Ruby-throated see Archilochus colubris
Hylocichla mustelina, 2, 9, 10
Jay, Blue see Cyanocitta cristata
Jay, Scrub see Aphelocoma coerulescens
Kestrel, American see Falco sparverius
Kingbird, Eastern see Tyrannus tyrannus
Kite, American Swallow-tailed see Elanoidesforficatus
Lanius ludovicianus, 19, 20, 40
Laterallus jamaicensis, 23
Limpkin, see Aramus guarauna
Meadowlark, Eastern see Sturnella magna
Melanerpes carolinus, 1, 2, 3, 4, 5, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 24, 25, 35,
36, 38, 39, 41,42,43,44,45,46, 47, 48,49, 50
Melanerpes erythrocephalus, 35, 42, 43, 46
Meleagris gallopavo, 2, 8, 10
Mimus polyglottos, 13, 14, 15, 19, 20, 40, 46
Mockingbird, Northern see Mimus polyglottos
Molothrus ater, 2, 6, 8, 10, 35, 42, 43
Myiarchus crinitus, 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 14, 15, 16, 17, 18, 19, 24, 25, 35, 36,
38, 39, 41, 42,43, 44,45, 47, 48, 50, 51
Nighthawk, Common see Chordeiles minor
Nuthatch, Brown-headed see Sitta pusilla
Nuthatch, White-breasted see Sitta carolinensis
Oporornisformosus, 2, 10
Osprey, see Pandion haliaetus
Otus asio, 13, 14, 38, 49
Owl, Barred see Strix varia
Owl, Great Horned see Bubo virginianus
Pandion haliaetus, 8
Parula americana, 2, 8, 9, 10, 38, 49
Parula, Northern see Parula americana
Parus bicolor, 1, 2, 9, 10, 11, 12, 16, 18, 24, 25, 35, 36, 38, 42, 43, 49, 50, 51
Parus carolinensis, 2, 8, 9, 10, 35, 42, 43
Passerina cyanea, 2
Picoides borealis, 25, 35, 36, 42, 43
Picoidespubescens, 2, 3, 4, 5, 6, 8, 9, 10, 14, 15, 16, 17, 18, 25, 35, 36, 39, 41, 42, 43, 47, 49
Picoides villosus, 2, 10, 16, 17, 18, 25, 42, 44
Pipilo erythrophthalmus, 2, 3, 4, 13, 14, 15, 19, 20, 23, 25, 35, 39, 40, 42, 43, 44, 45, 46,
47, 48
Piranga rubra, 1, 2, 8, 9, 10, 11, 12, 24, 25, 35, 42, 43, 50, 51
Podilymbus podiceps, 19
Polioptila caerulea, 1, 2, 4, 5, 6, 7, 8, 9, 10, 11, 12, 24, 25, 35, 36, 38, 42, 43, 49, 50, 51
Porphyrula martinica, 8, 21
Protonotaria citrea, 9, 10









NONGAME WILDLIFE PROGRAM FINAL REPORT


Quiscalus major, 21
Rail, Black see Laterallus jamaicensis
Rail, Clapper see Rallus longirostris
Rail, King see Rallus elegans
Rallus elegans, 21, 22, 23
Rallus longirostris, 26, 27, 28, 29, 30, 31, 32, 33, 34, 37
Screech-Owl, Eastern see Otus asio
Seiurus motacilla, 2, 9, 10
Shrike, Loggerhead see Lanius ludovicianus
Sialia sialis, 4, 5, 6, 7, 25, 36, 42, 43
Sitta carolinensis, 2
Sitta pusilla, 4, 5, 7, 9, 25, 35, 36, 42, 43
Sparrow, Bachman's see Aimophila aestivalis
Sparrow, Seaside see Ammodramus maritimus
Strix varia, 2, 9, 10, 14, 16, 17, 18, 21, 38, 41,49
Sturnella magna, 19, 20, 22, 23, 36, 40, 42, 46
Swallow, Barn see Hirundo rustica
Swift, Chimney see Chaetura pelagica
Tanager, Summer see Piranga rubra
Thrasher, Brown see Toxostoma rufum
Thrush, Wood see Hylocichla mustelina
Thryothorus ludovicianus, 2, 3, 6, 8, 9, 10, 13, 14, 15, 16, 17, 18, 21, 24, 25, 36, 38, 39,
41,43,44,45,47,48,49
Titmouse, Tufted see Parus bicolor
Towhee, Rufous-sided see Pipilo erythrophthalmus
Toxostoma rufum, 13, 14, 17, 18, 20, 24, 41
Troglodytes aedon, 6
Turkey, Wild see Meleagris gallopavo
Tyrannus tyrannus, 8
Tyto alba, 38, 49
Vireo, Red-eyed see Vireo olivaceus
Vireo, White-eyed see Vireo griseus
Vireo, Yellow-throated see Vireoflavifrons
Vireoflavifrons, 1, 2, 9, 10, 11, 12, 35, 42, 43, 50
Vireo griseus, 2, 9, 10, 14, 15, 16, 17, 18, 36, 39, 41, 43, 44, 45, 48
Vireo olivaceus, 2, 9, 10, 11, 18, 24, 38, 41, 49
Vulture, Turkey see Cathartes aura
Warbler, Hooded see Wilsonia citrina
Warbler, Kentucky see Oporornisformosus
Warbler, Pine see Dendroica pinus
Warbler, Prothonotary see Protonotaria citrea
Warbler, Yellow-throated see Dendroica dominica
Waterthrush, Louisiana see Seiurus motacilla
Wilsonia citrina, 2, 9, 10
Wood-Pewee, Eastern see Contopus virens
Woodpecker, Downy see Picoides pubescens
Woodpecker, Hairy see Picoides villosus
Woodpecker, Pileated see Dryocopus pileatus
Woodpecker, Red-bellied see Melanerpes carolinus
Woodpecker, Red-cockaded see Picoides borealis
Woodpecker, Red-headed see Melanerpes erythrocephalus
Wren, Carolina see Thryothorus ludovicianus
Wren, House see Troglodytes aedon
Wren, Marsh see Cistothorus palustris
Yellowthroat, Common see Geothlypis trichas
Zenaida macroura, 12, 13, 14, 15, 18, 19, 20, 40, 41, 42, 43, 45, 46, 47, 48, 50, 51


















































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