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of the
FLORIDA STATE MUSEUM
Biological Sciences
Volume 30 1987 Number 3
TERRESTRIAL PLANT AND WILDLIFE
COMMUNITIES ON PHOSPHATE-MINED LANDS
IN CENTRAL FLORIDA
ROGER S. SCHNOES AND STEPHEN R. HUMPHREY
UNIVERSITY OF FLORIDA
GAINESVILLE
Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCI-
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WAYNE R. MARION
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persons the results of researches in the natural sciences, emphasizing the
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Publication date: 12 May 1987
Price: $3.50
TERRESTRIAL PLANT AND WILDLIFE
COMMUNITIES ON PHOSPHATE-MINED LANDS IN
CENTRAL FLORIDA
ROGER S. SCHNOES AND STEPHEN R. HUMPHREY'
ABSTRACT: A study of plant and animal communities documented the recovery of phos-
phate-mined land in central Florida. Hypotheses about community structure were tested
in two sets of treatments. One set consisted of the relatively stable end results of post-
mining land use. These treatments were: consolidated waste clay soil; late successional
forest on unreclaimed overburden spoil piles with and without interspersed lakes; and
grazed and ungrazed pastures on reclaimed overburden soil. The other set of treatments
tested for differences in community structure among several seral stages on unreclaimed
overburden spoil piles with interspersed lakes. The responses of biological communities
were measured as the identity, diversity, and abundance of plants, mammals, birds, rep-
tiles and amphibians, and insects. Major findings include:
1) Succession on clay waste areas was slow, resulting in senescent forest and a depau-
perate animal community.
2) Unreclaimed spoil piles underwent a rapid primary succession culminating in xeric
or mesic oak forest with rich animal communities. The faunas were different on treat-
ments with and without lakes, but both had moderate-to-high wildlife value, high aes-
thetic quality, and much potential for enhancing animal populations.
3) Succession on reclaimed sites was arrested by grazing or mowing. Both treatments
provided relatively poor wildlife habitat, though animal diversity and abundance gener-
ally were higher on ungrazed sites. However, stocking with cows resulted in high live-
stock biomass and slightly more diverse vegetation.
4) Aboveground lenses of hardening clay appear to be properly characterized as waste-
lands, so implementing optimal reclamation procedures for clay wastes should be a top
priority.
5) By contrast, both unreclaimed sites and the type of reclaimed sites examined here
are valuable land with a variety of potential uses. The advantage these have in common is
the presence of overburden soil.
6) The biological factors underlying the value of reclaimed land are the abundant,
balanced nutrients in overburden soil and the free services provided by colonizing biota,
notably creation of topsoil through symbiotic nitrogen-fixation by Frankia and Myrica,
seed dispersal by wind and later by migratory birds, and possibly by introduction of my-
corrhizal fungi spores by rodents. Operation of these processes should be built into plans
for managing reclamation.
7) Succession on overburden soil was to mesic oak forest. Because its soil hardpan is
destroyed during mining, restoration of pine flatwoods is not possible without new engi-
neering designs for soil reclamation.
8) The study supports reclamation to rangeland as a legitimate post-mining land use,
even though its wildlife value is low. However, the current trend of maintaining improved
IRoger S. Schnoes is a Forester with the Bureau of Land Management, Medford District Office, 3040 Biddle Road,
Medford OR 97501. Stephen R. Humphrey is Associate Curator in Ecology, Florida State Museum, and Affiliate Asso-
ciate Professor of Wildlife Ecology, School of Forest Resources and Conservation, University of Florida, Gainesville FL
32611.
Schnoes, R. S., and S. R. Humphrey, 1987. Terrestrial Plant and Wildlife Communi-
ties on Phosphate-mined Lands in Central Florida. Bull. Florida State Mus., Biol. Sci.
30(3):53-116.
BULLETIN FLORIDA STATE MUSEUM
-. j
u -~
'AN
ii-p
FRONTISPIECE: Part of the central Florida mining district photographed from a U-
2 airplane in 1973. Active or recent mines show parallel lines of pits and spoil piles. Dark
water bodies are active clay settling ponds. Floodplain forest of the Alafia River is in upper
left, the town of Mulberry is at the major crossroads in top center, and Hooker's Prairie is
in bottom center. Scale: 1 cm = 1.5 km, 1 inch = 2.9 miles.
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 55
pasture on nearly all reclaimed land will result in a region-wide decline of wildlife re-
sources in comparison to both pre-mining conditions and the present transitional habitats.
The impending losses could be reduced by including modifications in reclamation plans
that will provide wildlife habitat or by directing some reclamation projects specifically at
wildlife values.
9) A second trend, toward the use of sand tailings as the major surface soil in recla-
mation, has the potential to diminish post-mining land quality on a large scale. Because
the nature of reclaimed soil is a key variable, regulations should address soil quality and
depth.
10) Evaluating the productivity of mined lands would be enhanced by closing infor-
mation gaps including succession on sand tailings, wildlife communities in pre-mining
habitats, socioeconomic value of the wildlife resources lost, field trials of game and vege-
tation management techniques, habitats restorable on soils that combine sand tailings
with consolidated clay, and agronomic and forestry potential of reclaimed soils.
RESUMEN: El present studio document la recuperaci6n de tierras para extracci6n de
fosfatos en el centro de Florida. Varias hip6tesis sobre estructura de comunidad fueron
examinadas en dos grupos de tratamientos. Un grupo consisti6 de las consecuencias, rel-
ativemente estables, de usos de la tierra despues de actividades de extracci6n minera.
Estos tratamientos fueron: suelo de residues consolidados de arcilla; bosques sucesionales
tardios en monticulos de suelo removido en Areas mineras sobrecargadas y no recupera-
das, con y sin lagos interdispersos; y pastizales expuestos y no expuestos a ramoneo en
suelos sobrecargados, no recuperados. El otro grupo de tratamientos incluy6 various esta-
dios serales en monticulos de suelo removido y sobrecargado, interdisperso con lagos. Las
respuestas de las comunidades biol6gicas fueron medidas con respect a la identidad,
diversidad y abundancia de plants, mamiferos, aves, reptiles, anfibios e insects.
Los resultados mas importantes incluyen:
1) La sucesi6n en areas de residues de arcilla fue lenta, dando como resultado un
bosque senescente y una comunidad animal empobrecida.
2) Monticulos de suelo removido, no recuperado experimentaron una ripida sucesi6n,
terminando en bosques m6sicos o x6ricos de roble con ricas comunidades animals. La
fauna fue diferente en los tratamientos con y sin lagos, pero ambos tuvieron un moderado
a alto valor de fauna silvestre, gran calidad est6tica y much potential para promover
poblaciones animals.
3) La sucesi6n en tierras no recuperadas fue impedida por pastoreo y corte. Ambos
tratamientos indican habitats relativamente pobres en fauna; si bien la abundancia y div-
ersidad animal fueron, generalmente, mayores en sitios no expuestos al pastoreo. Sin
embargo, pasturas para vacunos resultaron en una gran biomasa de ganado y una diversi-
dad de vegetaci6n ligeramente mayor.
4) Capas superficiales de arcilla endurecida parecen estar caracterizadas como tierras
improductivas. Por ello, se debe dar primera prioridad a procedimientos que permitan la
recuperaci6n adecuada de los deshechos de arcilla.
5) En contrast, tanto los sitios recuperados como los no recuperados examinados en
el present studio, estan tierras valiosas gracias a la variedad de sus usos potenciales. La
ventaja que estos tienen en comdn es la presencia de suelo sobrecargado.
6) Los factors biol6gicos que contribuyen al alto valor de las tierras recuperadas son:
la abundancia de nutrients balanceados en suelo sobrecargados y los servicios gratuitos
proporcionados por la biota colonizadora, especialmente la formaci6n de suelo a trav6s de
la fijaci6n simbi6tica de nitr6geno por Frankia y Myrica; dispersi6n de semillas por el
viento y posteriormente por aves; y posiblemente la introducci6n de esporas de hongos
formadores de micorrizas por roedores. El funcionamiento de estos process deberia ser
incorporado en cualquier plan de manejo para recuperaci6n de tierras.
BULLETIN FLORIDA STATE MUSEUM
7) La sucesi6n de suelos sobrecargados fue hacia bosques m6sicos de roble. Debido a
que la cubierta endurecida de suelo en bosques pianos de pinos es destruida durante las
operaciones mineras, la restauraci6n de estos bosques no es possible sin nuevos disefios de
ingenieria para reclamaci6n de suelos.
8) Este studio respalda la recuperaci6n de tierras como una alternative legitima de
uso despu6s de actividades mineras, aun en cuando su valor para fines de fauna silvestre
sea bajo. Mas aun, la tendencia actual de mantener pasturas mejoradas en casi todas las
tierras recuperadas resultard en una disminuci6n regional de recursos de fauna en com-
paraci6n a las condiciones previas a las actividades mineras y a las condiciones presents
de habitats transicionales. Las p6rdidas inminentes podrian ser reducidas si se modifica
los planes de recuperaci6n, de tal manera que se prove a la fauna silvestre de hAbitat, o
si se orienta algunos de estos proyectos de recuperaci6n de tierras especificamente hacia
la explotaci6n de los valores de la fauna.
9) Una segunda tendencia que propugna el uso de tierras arenosas como la principal
superficie en recuperaci6n, tiene el riesgo potential de disminuir la calidad de la tierra, a
gran escala, despu6s de la extracci6n minera. Debido a que la naturaleza del suelo a ser
recuperado es una variable clave, se deberia dar regulaciones acerca de la calidad y pro-
fundidad del suelo en cuesti6n.
10) La evaluaci6n de la productividad de tierras con aptitud minera se veria benefi-
ciada si se obtiene mas informaci6n detallada acerca de sucesi6n en tierras arenosas, co-
munidades animals en hAbitats aun no expuestos a minerfa, valor socio-econ6mico de los
recursos faunisticos a perder, experiments de campo sobre t6cnicas de manejo de fauna
y flora, habitats recuperables en suelos que combinan arena y arcilla consolidada, y poten-
cial agron6mico y forestal de los suelos recuperados.
TABLE OF CONTENTS
Introduction ............................ ... ............................. 57
Acknowledgments .............. ... ....................................... 58
Methods .............. ................................................ 58
Research Design ................ ...................................... 58
Study Areas ............... ................... ....................... 59
Field Work ............................................................ 60
Sampling Techniques ................................................... 60
D ata Analysis ............. .... .............................. .......... 62
Statistical Analysis ................................... .................. 62
Results.. ............ ........................................ ............ 63
Vegetation ...................................... ...................... 63
Small Mammals ........................................................ 77
Birds ............ ..... ................................... ........... 86
Insects ......................................... ........... 97
Herpetofauna .......................................................... 98
Large Mammals ....................................................... 99
Statistical Analysis ...................................................... 101
Discussion ................ .................................... ............ 104
Synthesis of Results .............. ................................... 104
Comparison with Other Studies .......................................... 107
Biological Factors Affecting Values and Uses of
Mine-created Lands ................. ................................. 108
Future Landscapes of the Mining Region ................................. 111
Information Gaps ............... ..................................... 112
Literature Cited ............... ................... ..................... 114
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 57
INTRODUCTION
About 5180 square kilometers in central Florida have large phosphate
resources accessible by surface mines. Beginning in the 1880s, mining
has concentrated in the northern portion of the district, and about 750
square kilometers have been mined. These northern reserves will be
depleted in the next 10 to 30 years, and mining operations will shift to
the southern part of the district.
Loss of valuable environments and wildlife because of surface mining
for phosphate has become a major conflict in Florida's land use planning.
Mining of a non-renewable resource is a temporary but total use of the
land, and its destructiveness makes mining unpopular among citizens
whose individual benefits from the activity are small. However, demand
for fertilizer to grow human food is increasing, and mining is expected to
occur throughout the phosphate district. Government's challenge is to
permit the extraction of necessary resources while minimizing losses over
the long term. Consequently regulatory agencies need to focus on min-
ing and reclamation practices that return land to attractive and useful
conditions. At present, neither the preferred conditions nor appropriate
reclamation practices have been determined or agreed upon by mining
companies and regulatory agencies. Carefully considered criteria for quality
need to be established, and future land use options (residential/industrial
development, agriculture/silviculture, water storage, wildlife areas) need
to be retained.
Because mineral extraction technology has become more efficient and
because pumping rights for water (a limiting factor for mine operation)
have been regulated as a per-acre water crop, mining companies have
tried to hold their lands as cheaply as possible for eventual re-mining
and to qualify for consumptive water-use permits. Consequently no in-
terest has operated to return the land to any other use, and no intrinsic
incentive for reclamation has existed. Lands that have escaped re-mining
have undergone natural recovery without prescribed manipulation of soil,
topography, or pioneer biota.
Reclamation of newly mined lands became required by a 1975 state
severance-tax law. A revision of the law in 1978 provided a mechanism
to use tax money to reclaim pre-1975 mined lands and required mining
companies to reclaim future mined lands as an internal feature of their
operating economies. During the past decade, county governments have
imposed reclamation requirements with specific regulations designed to
protect the future contribution of mined lands to the property tax base.
County and state regulations often conflict, and they have poorly under-
stood implications for biological recovery.
The shift of new mining southward and the areawide practice of rec-
BULLETIN FLORIDA STATE MUSEUM
lamation are major developments in determining land use. Results to be
expected in the northern part of the district are nearly ubiquitous recla-
mation with unknown impacts, followed by either conversion of the land
to productive non-mining uses by the companies or release of large tracts
for sale. In the southern part of the district, the entire cycle of mining-
related land use will occur, from destruction of existing uses, through
temporary types, to subsequent reclamation for unknown purposes.
Now that the decision to reclaim mined land has been made, the re-
sults of reclamation practices need to be evaluated. The quality of re-
claimed land is reflected by biological responses to site treatments. Plant
growth and wildlife use are meaningful criteria of quality because they
show the ability of the land to support natural communities, agriculture,
or silviculture, and they are major components of the aesthetic character
of the land. The purpose of this study was to document plant succession
and wildlife use on the abandoned and reclaimed sites in the central
Florida phosphate district.
ACKNOWLEDGEMENTS
This work was conducted under contract for the Office of Environmental Services of
the Florida Game and Fresh Water Fish Commission and was funded by the State Uni-
versity System Board of Regents as STAR Grant 78-1122. It was conducted under the
guidance of H. Eugene Wallace of the Florida Game and Fresh Water Fish Commission.
The project was suggested and sponsored by Brad J. Hartman and Douglas Bailey of the
Commission's Office of Environmental Services. Other office staff-Terry Gilbert and Tim
King-provided helpful discussions throughout the project. Additional perspective was
contributed by staff of other agencies, including J. William Yon and Gregory L. Daughter
of the Department of Natural Resources and Jorge C. Southworth of the Department of
Community Affairs. Field sampling would have been impossible without the able assist-
ance of Lindsey J. Hord. We are grateful to Robert S. Hearon, Jay N. Allen, Jr., and other
personnel of International Minerals and Chemical Corporation for their valuable help in
selecting field sites, arranging living quarters, and solving logistic problems. Similarly, we
thank John Tallent of ESTECH General Chemical Corporation, Joseph Tessatori of W. R.
Grace, Inc., and Edward Holloway of Sanlan Ranch for enabling us to sample sites on
their property. John M. Schollenberger of the University of Florida Center for Instruction
and Research Computing advised us on the statistical analysis. Don E. Marietta and Nancy
R. Halliday prepared the figures, and typing was done by Rhoda J. Bryant, Gladys M.
Jackson, Angela K. O'Brien, and Dawn D. Zalenka. Pamela R. Johnson and A. Darlene
Novak provided essential administrative help, as did E. Frazier Bingham and John Scott
Dailey of the Board of Regents office. Individuals providing helpful criticisms of report
drafts included members of the Florida Game and Fresh Water Fish Commission and
International Minerals and Chemical Corporation staffs mentioned above, plus Robert
Goodrich and Lee Cawley of the IMC reclamation office, as well as Ronnie Haynes, James
N. Layne, Wayne R. Marion, and I. Jack Stout.
METHODS
Research Design
This research was designed to evaluate the terrestrial habitats that have been created
after mining in central Florida. Included were both reclaimed sites and those that were
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 59
left to recover without manipulation. Successional changes were detected by studying
sites of different ages. Major components of the biological community were documented
in terms of their identity, abundance, and diversity, and these were evaluated as re-
sponses to site treatment. These groups of organisms included plants, insects, reptiles
and amphibians, birds, and mammals. Measurements were done with systematic, stan-
dardized sampling methods that enabled comparison of all habitats and sites. Three sepa-
rate sites were studied for each post-mining treatment, with the replicates selected to
represent the full range of site variation evident from visual examination of soil, topogra-
phy, and vegetation. Plants were measured once during the study, whereas animals were
measured quarterly to enable detection of seasonal changes while producing realistic av-
erage values for the year.
STUDY AREAS
The 24 study areas were located in Polk County, Florida, and comprised three repli-
cates each of eight post-mining treatments (Table 1). The first category, consolidated clay
settling ponds, is the terrestrial habitat resulting from consolidation of clay waste stored
in large ponds. After slime ponds are filled with colloidal clay, they slowly dry out by
evaporation and transpiration. When the surface clay of a site compacts into a thick crust
with deep cracks, it has made the transition from aquatic to terrestrial habitat, and the
dewatered pond is no longer subject to regulations of the Environmental Protection Agency.
Data from these sites can be compared with data from the younger artificial marshes (see
Gilbert et al. 1981). Subsequent successional stages seldom occur because these areas
become subject either to re-mining or to reclamation by capping with sand tailings. Be-
cause the latter practice is a recent development, fully reclaimed sites were unavailable
Table 1. Study sites, showing eight treatments with three replicates of each.
Replicates
Treatments 1 2 3
Consolidated Clay
Settling Ponds Swift N-2 A-3
Unreclaimed Pits
and Spoil Piles
0-5 years
with lakes Big Teeth Shark Tooth Young Tiger
5-15 years
with lakes Orange Grove Pits Tiger Tail Tiger Bay South
15-30 years
with lakes Homeland Cemetery Gator Lake Achan-4
>30 years
with lakes Bartow South Saddle Creek Park Sanlan Ranch
>30 years
without lakes Old Spoil Piles Christina Old Clarke James
Reclaimed Pastures
Ungrazed Parcel B Noralyn Kibler
Grazed 6-D Marina East H-4
BULLETIN FLORIDA STATE MUSEUM
for study. However, current trends indicate that a major area of land will be formed by
this process.
Mining activities leave a series of roughly parallel spoil piles and pits. In flat regions
the pits fill with groundwater, leaving steep emergent hills interspersed with lakes whose
size depends on whether water levels are high enough to connect many pits. Along topo-
graphic ridges and in the upland slopes around freshwater marshes, the spoil piles are
sufficiently drained that no lakes develop. With or without lakes, these spoil areas nor-
mally were abandoned without modification prior to the reclamation required by 1975
Florida law, and such lands are viewed by the public as moonscapess." Spoil piles are
steep and easily eroded but become stabilized by vegetation. The soil of spoil piles con-
sists of the mixed sand and clay of the overburden that is set aside to gain access to the
phosphate ore. The steep underwater slopes do not support littoral vegetation, which is
limited to areas where erosion has created gentler gradients.
Because of the long history of mining in central Florida, unreclaimed spoil piles and
pits have become a dominant landform in the region, with different tracts available in a
continuous age sequence up to about 50 years old. The research design included four
successional stages of this sequence, to determine what communities result from natural
recovery after mining. Successional categories were established as 0-5, 5-15, 15-30, and
>30 years after mining, based on historical records of the mining companies. A fifth treat-
ment included was the most mature category without the aquatic habitat of interspersed
lakes. This more homogeneous site treatment was expected to support a community dis-
tinct from that with lakes, to be more directly comparable to unmined native forests, and
to provide insight to reclamation planners in comparison with the type including lakes.
Reclamation began on a significant scale on these mined lands in 1975. Consequently
reclaimed sites are young and few alternative types of treatment are fully accomplished.
The most widely practiced reclamation treatment has been to fill old mine pits with sand
tailings and cap the area with overburden. The sand and clay soil is contoured to a flat or
gently rolling surface and planted with exotic legumes and grass to stabilize the soil. These
reclaimed pastures are either ungrazed or leased to private individuals who fence the
areas, fertilize as needed, and stock them with cattle. Reclaimed areas that are not grazed
are mowed twice annually to maintain the potential of these areas as pastures. The mow-
ing arrests succession at a stage of non-native pasture. Both ungrazed and grazed pastures
were included in the research design to document wildlife use on reclaimed land and to
indicate cattle stocking levels achieved in practice. Other types of reclamation that are
beginning to be practiced will be considered in the Discussion.
FIELD WORK
Field work was conducted from 17 October 1978 trough 3 July 1979 and was split into
four quarters to measure seasonal variation in wildlife communities and habitat use. The
quarterly field periods were: Autumn, 17 October-3 December 1978; Winter, 8 January-
2 March 1979; Spring, 13 March-6 May 1979; and Summer, 1 June-3 July 1979.
Bird, small mammal, and insect communities were systematically sampled on each
site during each of the four quarters. Plant communities were sampled in May and June.
Evidence and observations of larger mammals and herpetofauna were recorded through-
out the field work. Not all sites were selected in time to be sampled during the autumn
quarter. These sites are indicated in Tables 3, 7, and 11, and missing data are identified
where appropriate in the Results. The full complement of study sites was available for
sampling during the other three quarters.
SAMPLING TECHNIQUES
VEGETATION.-Plant community sampling was conducted along a 100 m transect in
each study site. Percent cover of shrubs, woody vines, and tree seedlings <5 cm dbh was
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 61
measured along the transect with a line-intercept method. Herbaceous vegetation was
sampled using ten 20 cm x 50 cm plots, located at 10 m intervals along the transect. Plants
were identified to species level, and percent cover in each plot was visually estimated.
Plant names agree with those used by Radford et al. (1964) and Long and Lakela (1976).
Tree species were sampled using 20 of 40 cells, each 10 m long and 5 m wide, by
selecting every other cell on each side of the 100 m transect. In each cell, trees (>5 cm
dbh) were identified and counted. Height was measured with an optical clinometer, and
diameter at breast height (dbh) was measured with a tape measure. Basal area (BA) was
computed for each species using the formula
BA = D
4
where D = dbh. Frequency of occurrence for each species was obtained for the 20 cells.
Relative frequency (RF), relative abundance (RA), and relative dominance (RD) for each
species were calculated using the frequency of occurrence, number of stems, and basal
area respectively compared with the totals for each transect. An importance value (I) was
then assigned to each species such that I = RF + RD + RA.
SMALL MAMMALS.-Small mammal communities were sampled once each quarter using
both Sherman live traps and museum special snap traps. A removal technique was em-
ployed and all animals caught were identified, weighed, and measured, and age class and
reproductive status were noted. Each site was trapped for three consecutive nights using
64 stations consisting of one live trap and one snap trap baited with peanut butter and
oatmeal. These stations were arrayed in two 4 x 8 grids, with trap stations 8 m apart. Traps
were checked each morning and rebaited as necessary.
BIRDs.-Avian communities were sampled using a variable-width transect method
(Emlen 1971). Transects were walked through study sites during the first 2.5 hours after
sunrise, and all birds seen or heard were tallied, along with the distance from the transect
when first detected. Birds were excluded if they occurred beyond 100 m of the transect
line. Transect counts were standardized by extrapolating to the equivalent of a 1 km tran-
sect. In all but the first quarter, time permitted transects to be run twice on different days
on each site. During autumn quarter, only one run was conducted.
Following Emlen's method, a coefficient of detectability (CD) was calculated for each
species, each type of habitat, and each quarter. These coefficients were then used to
correct the original count data to provide a more reliable estimate of the number of birds
present within the transect boundaries. A separate set of CDs was computed for each of
the 8 treatment classes, except that the two types of reclaimed pastures were considered
as one group.
Biomass estimates were made from several sources, including the University of Flor-
ida bird collection, Clench and Leberman (1978), Greenewalt (1975), and incidental weight
measurements made on birds during field work.
INSECTS.-Insects were sampled using light traps with 8 watt blacklight bulbs (BioQuip
Inc., 1320 E. Franklin Ave., El Segundo CA 90245). Light traps were set 1-2 m above
the ground in the evening and retrieved early the next morning. The samples were oven-
dried for 48 hours at 700C and then were weighed to yield biomass data. Time permitted
only 1 set of samples (those from winter quarter) to be sorted to Order. The diversity
values used in this report are from that quarter.
Only three light traps were available for use, and technical problems at times reduced
that number to two or even one. This caused the sampling of all sites to be spread out
over at least ten days and at times much longer. This factor added to the high variance in
the data expected from the nature of insect hatches and insect responses to weather vari-
ation.
HERPETOFAUNA. -Original plans called for pitfall-with-drift-fence trap arrays to be
BULLETIN FLORIDA STATE MUSEUM
constructed on each site, but this proved to be impractical due to the time required in
maintenance of the traps, as well as the destruction of traps by humans and cows. Con-
sequently, reptiles and amphibians were simply noted when observed in the course of
other field work. The result was a list of species without quantification of relative abun-
dance.
LARGE MAMMALS.-Much the same situation existed with this class of wildlife as with
the herpetofauna. A systematic, quantified method of counting tracks along dirt roads was
applied but proved unfeasible because of the clay substrate and heavy vehicular travel.
Tracks, scats, and sightings were recorded when encountered during other activities on
the sites.
DATA ANALYSIS
The simplest method of characterizing post-mining treatments was by comparing lists
of the kinds of plants and animals that were present after recovery. However, more rig-
orous comparisons involved indices of abundance and diversity that reflected differences
among the communities supported by the post-mining treatments. The number of indi-
viduals and biomass provided measures of relative abundance. These two variables nor-
mally show similar trends for animal communities, but not for plants. The Shannon-Wie-
ner index (Shannon and Weaver 1949) was used to measure species diversity, defined as
H' = Yp, In pi/p,, where p, is the proportion of individuals in species i. This diversity
index has two major components. The first is the number of species, or species richness,
which is the fourth type of variable used in our comparisons. The second component is
equitability, or the evenness of distribution of the species, and is defined as E = H'/H,,,
(Sheldon 1969), where H._ is the natural log of the number of species.
Equitability was not extensively used in this report. Like the measures of abundance, H'
and the species number usually show similar trends of diversity (for both plants and ani-
mals). Generally all four quantitative evaluations are presented together to provide a thor-
ough view of community structure and to emphasize the differences and similarities be-
tween treatments.
STATISTICAL ANALYSIS
Statistical analyses of these data were performed by version 79.2B of the Statistical
Analysis System, using the computational facilities of the Northeast Regional Data Center
at the University of Florida.
For statistical analysis the results were separated into two data sets-the annual or
seasonal measurements of 12 variables involving plants, herpetofauna, and large mam-
mals, and the seasonal (quarterly) measurements of 9 variables involving small mammals,
birds, and insects. Two sets of hypotheses were posed for each data set. One tested for
differences among the relatively stable end results of post-mining land use, including five
treatments (consolidated clay settling areas, unreclaimed pits and spoil piles) > 30 years
with lakes and without lakes, and both ungrazed and grazed pastures). The other tested
for trends over the years of the successional sequence on unreclaimed pits and spoil piles
with lakes. Rather than evaluating seral sites simply as replicated sites within treatments,
each was considered as an observation with a specific age in years. Throughout the anal-
ysis, some data were transformed to ensure constant variances (Steel and Torrie 1960).
Small value counts and measurements of area were transformed by (y + 0.5)1/2. Large
value counts were transformed by log (y + 1). And weights were transformed by y1'3. F-
tests were made with alpha = 0.05 using multivariate analysis of variance. Because of
sites and variables missed during autumn sampling, the autumn data were deleted from
the seasonal data set. For both site age and final land uses, analysis of variance was begun
by testing for interaction between season and age or treatment. No significant interaction
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 63
occurred, though for most variables there was a strong seasonal effect. Therefore the
analyses of seasonal and seasonal data were identical. Distinctness of final land uses was
first tested by applying Duncan's multiple range test to treatment means. Natural con-
trasts anticipated in the experimental design were confirmed as groups by the Duncan's
test, so the more powerful analysis of variance specifying these contrasts was conducted
(combined over seasons for seasonal data) to look for more subtle distinctions between
contrasted treatments. Contrasts were as follows: mature unreclaimed sites with vs. with-
out lakes, ungrazed vs. grazed pastures, clay vs. mature unreclaimed sites, and clay vs.
reclaimed pastures. Significance of hypothesized age effects was established by pooling
sums of squares of non-significant responses in with the error sum of squares until only
significant effects remained. The effect of site age was modelled by computing equations
of significant variable responses to site age, using linear regression of the transformed
data. Finally, simple relationships between animal and plant variables were sought by
stepwise multiple regression.
RESULTS
VEGETATION
CONSOLIDATED CLAY SETTLING PONDS.-The three sites were cho-
sen to exhibit a successional sequence within the treatment-the Swift
site supporting the youngest community (Fig. 1) and A-3 having the
oldest (Fig. 2). This sequence was reflected in the vegetation analysis
(Table 2). The Swift site was dominated by a dense, young stand of willow
(Salix caroliniana) and a few Baccharis, with a well-developed unders-
tory of grasses. This site was wet enough that cattails (Typha sp.) and
rushes (Juncus sp.) also were established in places. Most willows in this
situation were too small to be considered trees (dbh <5 cm) and thus
were measured as shrubs.
The intermediate-age settling area, N-2, also was dominated by wil-
low, but these were much larger than in the Swift site. Willows of tree
size in N-2 had a mean dbh of 6.3 cm and a mean height of 4.4 m. Herb
cover, shrub cover, and shrub diversity were somewhat higher than at
the Swift site. However, willows still heavily dominated the shrub layer.
Tree-size wax myrtle (Myrica cerifera) dominated A-3, the oldest set-
tling area, but willows were still well established. These willow trees
were generally larger than the wax myrtles and occurred in wet depres-
sions. In addition, numerous dead trees all were found to be willows.
Vines dominated the shrub and herbaceous cover, but this may be mis-
leading because the crowns of the abundant, young wax myrtles were
intermixed with the crowns of the mature trees, and it was impossible to
determine what percent of the cover was contributed by the younger
trees. In this case only shrubs less than 3 m in height were measured.
No herbaceous vegetation was present. This late successional community
on clay settling ponds is unlike undisturbed mature vegetation found in
BULLETIN FLORIDA STATE MUSEUM
Figure 1.-Thicket on a Consolidated Clay Settling Pond, the Swift site. Willows are
in the Foreground, light seed heads ofAndropogon are in the center, and dark wax myrtle
and Baccharis shrubs are scattered in the background.
the region. The single black cherry (Prunus serotina) found in A-3 is
considered to be an artifact, because this individual was located near a
spoil pile island, and it may have been rooted in that substrate rather
than in the clay itself.
Combined as a class, the consolidated clay settling ponds showed
moderate abundance values for herbaceous and shrub vegetation, and
diversities were generally quite low (Figs. 3-5). Both abundance and
diversity values for trees were very low, except that the number of indi-
viduals was higher than in any other site category (Table 2).
It appears that clay settling ponds tended to progress toward a mono-
type of wax myrtle, an unnatural successional pattern. However, effects
and interactions of the clay crust, the underlying colloidal clays, distance
from native seed sources, and the allelopathic effects of wax myrtle (Du-
nevitz and Ewel 1981) are not fully understood. Further research is nec-
essary on these factors, but it will require study of older and more iso-
lated settling areas.
UNRECLAIMED PITS AND SPOIL PILES.-Beginning the successional
sequence of unreclaimed treatments, vigorous oldfield succession oc-
curred in the 0-5 year age class (Figs. 6-7). The three measures of her-
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA
,%,rI:
Figure 2.-Interior of the A-3 clay waste site. The ground is devoid of herbaceous
plants, and a dense forest canopy is formed by wax myrtle (left) and willow (leaning from
right).
baceous growth began at low to moderate levels soon after mining, as
ragweed (Ambrosia artemisiifolia), dog fennel (Eupatorium album), and
natal grass (Rynchelytrum repens) began to colonize the bare spoil piles.
Only a few shrubs (Baccharis) were present, and trees were absent.
Herbaceous communities reached peak levels of abundance and di-
versity in the 5-15 year age class (Figs. 8-9), as Caesar weed (Urena
lobata), cogongrass (Imperata cylindrica), and Andropogon sp. became
established and dog fennel disappeared. Shrubs, vines, and saplings in-
creased markedly in this period, the major species being Baccharis, Lan-
tana camera, grape (Vitis rotundifolia), blackberry (Rubus spp.), and
several kinds of saplings. Trees also became established in this interval,
primarily pioneer species like willow, wax myrtle, and Baccharis, along
with the exotic Brazilian pepper (Schinus terebinthifolius).
Shrubs dominated the vegetation in the 15-30 year age class (Figs.
10-11), as herbaceous growth became less prominent and trees contin-
ued their invasion of the sites. Cogongrass was no longer present, and
panic grass (Panicum dichotomiflorum) and shade-tolerant ferns (Polysti-
chum acrostichoides and Thelypteris normalis) joined the association.
Shrub composition was similar to the previous age class, with significant
Table 2. Summary of plant diversity and abundance.
Herbaceous Vegetation Shrubs Trees
Diversity Abundance Diversity Abundance Diversity Abundance
Treatments Number of Number of Number of Number of Total basal
Sites species H' % cover species H' % cover species H' individuals area
Consolidated Clay
Settling Ponds
Swift
N-2
A-3
Mean + standard
deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard
deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard
deviation
4
2
0
2.0
2.0
3
5
7
5.0
2.0
13
11
6
10.0
3.6
0.71 17.7
0.67 28.8
0 0
0.46 15.5
0.39 14.5
1.03 5.0
1.33 3.0
0.84 39.5
1.06 15.8
0.24 20.5
1.49 43.7
1.54 39.3
1.54 51.8
1.52 44.9
0.02 6.3
4
5
9
6.0
+2.6
1
1
3
1.7
1.2
10
12
10
10.7
1.2
0.46 47.9 1
0.79 77.9 1
1.11 33.3 2
0.78 53.0 1.3
0.32 22.7 0.6
0 4.0 0
0 1.7 0
0.82 11.3 0
0.27 5.7 0
0.47 5.0
1.46 71.9
1.88 122.4
1.62 53.4
1.65 82.6
0.21 35.7
4
2
3
3.0
1.0
0 7 191
0 134 4,244
0.66 331 14,879
0.22 157.3 6,438
0.38 163.2 7,586
S 0
-- 0
-- 0 -
-- 0
1.30 17 1,183
0.40 36 1,559
0.76 11 487
0.82 21.3 1,076
0.45 13.0 544
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard
deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard
deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard
deviation
Reclaimed Pastures
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard
deviation
Grazed
6-D
Marina East
H-4
Mean standard
deviation
6
7
6
6.3
0.6
7
6
4
5.7
1.5
4
9
0
4.3
4.5
3
6
5
4.7
1.5
2
6
9
5.7
3.5
0.86 29.7
1.54 33.1
1.21 36.0
1.20 32.9
0.34 3.2
1.61 17.8
1.30 15.0
0.72 11.8
1.21 14.9
0.45 0.30
1.00 23.4
1.55 26.5
0 0
0.85 16.6
0.78 14.5
0.12 51.3
0.72 47.5
0.42 69.6
0.42 56.1
+0.30 +11.8
0.11 43.5
0.61 63.4
1.52 62.3
0.74 56.4
0.71 1.5
12
13
6
10.3
3.8
18
19
15
17.3
2.1
17
18
17
17.3
0.6
1
1
1
1.0
0.0
0
1
3
1.3
0.32
1.76 23.5
1.96 27.1
0.88 78.1
1.53 42.9
0.57 30.5
2.29 63.0
1.78 125.5
2.12 23.5
2.06 70.7
0.20 51.4
2.09 31.7
2.06 35.4
1.56 82.9
1.90 50.0
0.29 28.6
0 0.0
0 0.2
0 0.1
0 0.1
0.1
-- 0
0 0.4
0.46 4.4
0.23 1.6
2.4
8
3
3
4.7
2.9
8
10
10
9.3
1.2
4
4
4
4.0
0.0
0
0
0
0
0
0
0
0
0.82 200
0.33 69
0.41 60
0.52 109.7
+0.26 78.4
1.64 113
1.76 93
1.69 68
1.69 91.3
0.06 22.5
0.76 31
1.23 18
0.81 44
0.93 31.0
0.25 13.0
0
0
-- 0
-- 0
-- 0
-- 0
-- 0
0
19,655
8,364
3,061
10,360
S8,475
18,672
19,084
20,797
19,518
1,127
23,994
21,968
25,671
23,877
1,854
BULLETIN FLORIDA STATE MUSEUM
DIVERSITY
15-
U
U
LLuJ
0C
V) 10-
0
5-
LU
cO
Z
z
TREES
SHRUBS
H/O
-----.- -
LU
u
V)
0 5
U,
5
z
CLAY UNRECLAIMED PITS & SPOILS RECLAIMED
0-5 5-1515-30 )30 >30 UNG. G.
WITH LAKES -- W/O
Figure 3.-Summary of plant diversity (as measured by species number) on Consoli-
dated Clay Settling Ponds, Unreclaimed Pits and Spoil Piles, and Reclaimed Pastures.
Abbreviations refer to whether pastures are grazed or ungrazed and to the age and pres-
ence or absence of lakes on unreclaimed sites.
- FA
I I -, . . .. . . . . .
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 69
DIVERSITY
H' 1-
0-
2-
H' 1
0-
2 -
H' 1-
0-
TREES
SHRUBS
Es -A
CLAY UNRECLAIMED PITS & SPOILS RECLAIMED
0-5 5-15 15-30 >30 >30 UNG. G.
WITH LAKES -0 W/O
Figure 4.-Summary of plant diversity, as measured by the diversity index H'. Treat-
ments and abbreviations are as in Figure 3.
BULLETIN FLORIDA STATE MUSEUM
ABUNDANCE
TREES
'001 SHRUBS
HERBS
CLAY UNRECLAIMED PITS& SPOILS RECLAIMED
0-5 5-15 15-30 >30 >30 UNG. G.
WITH LAKES IW/O
Figure 5.-Summary of plant abundance on Consolidated Clay Settling Ponds, Un-
reclaimed Pits and Spoil Piles, and Reclaimed Pastures. Abbreviations are as in Figure 3.
VOL. 30 NO. 3
Figure 6.-The Shark Tooth site, Unreclaimed Pits and Spoil Piles, 0-5 years with
lakes. Near the beginning of the time range, the mine is being enlarged by a dragline in
the background. The low water level in pit bottoms is maintained by pumping while
mining continues.
Figure 7.-Young Tiger, the oldest of the 0-5 years with lakes, Unreclaimed treat-
ments. Oldfield succession is well under way and shrub invasion has begun, with Lantana
on the left.
Figure 8.-Tiger Tail, 5-15 years old with lakes, Unreclaimed Pits and Spoil Piles,
showing advanced oldfield growth and shrub invasion. Foreground has Baccharis high on
the left, dog fennel downslope, and willow near the lake edge.
Figure 9.-Tiger Bay South, Unreclaimed Pits and Spoil Piles, 5-15 years with lakes,
showing extensive shrub growth.
Figure 10.-Gator Lake, 15-30 years old with lakes, Unreclaimed Pits and Spoil Piles,
showing extensive shrub and tree growth. In the foreground are wax myrtle, and the
background trees include a volunteer slash pine near the right.
Figure 11.--Achan-4, Unreclaimed Pits and Spoil Piles, 15-30 years with lakes. Two
patches of cattail remain along the bank at center and left.
BULLETIN FLORIDA STATE MUSEUM
addition of sapling wax myrtle and Brazilian pepper and reduction of
blackberry. Small wax myrtle trees became very abundant, and many
pioneer and shade-tolerant species entered the community.
Trees were the most abundant plants in the >30 year age class with
lakes (Figs. 12-13), though shrubs remained more diverse than trees.
Herbaceous vegetation continued to decline in prominence, with ferns
and panic grass becoming more important components. Changes in shrubs
included a reduced Baccharis contribution, appearance of new shade-
tolerant species, prominence of Brazilian pepper at one site, and contin-
ued diversification of vines. Trees increased strikingly in diversity and
abundance. Wax myrtle and other pioneer species diminished in impor-
tance as exotic Brazilian pepper and camphor tree (Cinnamomum cam-
phora) and native forest species like sweetgum (Liquidambar styraci-
flua), water oak (Quercus nigra), live oak (Q. virginiana), and American
elm (Ulmus americana) dominated the canopy.
Vegetation on the >30 year age class without lakes (Figs. 14-15) dif-
fered in several respects from that on the sites with lakes, though it was
similar in high tree abundance. Herbaceous associations were similar,
Figure 12.-Bartow South, >30 years with lakes, Unreclaimed Pits and Spoil Piles.
Most of the trees are water oaks. Grass in the foreground is the edge of a road used
frequently by recreational visitors. With spoil piles partly leveled, this habitat was called
"land and lakes" prior to the uniform reclamation regulations implemented in 1975.
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 75
Figure 13.-Saddle Creek Park, >30 years with lakes, Unreclaimed Pits and Spoil
Piles, with an elm and several red maples on the left. The forest canopy completely shades
the water's edge.
characterized by Caesar weed, ferns, and panic grass. One site had little
herbaceous growth but an unusual abundance of grape vines. Shrubs
were similar in identity on the two treatments, but grape vines were
dominant, and young trees and shrubs were few on all of the sites with-
out lakes, giving the shrub layer an open appearance. The abundance of
vines may be underestimated, because much vine growth was high in
the forest canopy where our sampling technique was relatively ineffec-
tive. Unlike the with-lakes treatment, all sites without lakes were domi-
nated by a closed canopy of live and water oaks, with more biomass but
much less diversity. Relictual pioneers from earlier seral stages were scarce.
To summarize the response of vegetation on unreclaimed pits and spoil
piles, we documented a primary successional process, with an oldfield
community developing in 0-5 years and peaking in 5-15 years, a shrub-
dominated community in 15-30 years, and a maturing, closed-canopy
hardwood forest at >30 years. Mean total basal area of trees steadily
increased with the age of the spoil piles (Fig. 5), as biomass accumulated
in the growing trees. Forest composition differed in old sites with and
without lakes. Presumably the more diverse forests on with-lakes sites
resulted from the gradients of groundwater and sunlight along lake edges,
BULLETIN FLORIDA STATE MUSEUM
Figure 14.-Old Spoil Piles, Unreclaimed >30 years without lakes. The forest canopy
is dominated by live oaks and grape vines. Caesar weed and ferns are in the foreground.
Four piles of overburden are visible. The terrain slopes to Hooker's Prairie toward the
rear.
whereas the more uniform live oak/water oak forests resulted from more
uniform physical characteristics of environments lacking lake edges.
Compared with communities maintained on consolidated clay settling
ponds and reclaimed areas (Figs. 3-5), the maturing forests on unre-
claimed pits and spoil piles were far higher in diversity and abundance
of plant life.
RECLAIMED PASTURES.-The reclaimed sites were grassy fields, either
ungrazed (periodically mowed, Fig. 16) or grazed (Fig. 17). Both treat-
ments were dominated by exotic bahia grass (Paspalum notatum), hairy
indigo (Indigofera hirsuta), and native partridge pea (Cassiafasciculata).
Both treatments had a few colonizing shrubs, mainly Baccharis.
The most heterogeneous sites (H-4 and Kibler) were inadequately
represented by our samples. In each reclaimed treatment, sites were
chosen to span the range from homogeneous bahia grass pasture (as in
Fig. 17 for the grazed treatment) to patchy habitat that included pasture
and mesic swales, ditches, and ponds that were subject to less grazing or
mowing pressure (as in Fig. 16 for the ungrazed treatment). The full
range of plant diversity among sites was underestimated because of un-
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 77
Figure 15.-Old Clarke James, Unreclaimed >30 years without lakes. This site had
the spoil piles partly leveled, and the forest was more mesic than at Old Spoil Piles, with
more water oaks (background) mixed with the live oaks (foreground and left).
sampled patches of various habitats on H-4 and Kibler. We avoided the
non-pasture patches of habitat in sampling both plants and animals, but
these areas nonetheless affected the data for mobile animals.
Though the two treatments were similar in amount of plant cover, the
grazed sites were consistently more diverse than the ungrazed sites (Figs.
3-5), resulting from selective cropping by grazers and non-selective har-
vest by mowing machines. The herbaceous cover was more abundant
than for any of the other treatments, as is expected in the absence of
competition from shrubs and trees. Herbaceous diversity, however, was
comparable to that on both consolidated clay settling areas and mature
unreclaimed pits and spoil piles.
SMALL MAMMALS
The results of sampling for small mammals are summarized in Tables
3-6. A total of nine species was sampled during the study.
CONSOLIDATED CLAY SETTLING PONDS.-The clay areas supported
populations of cotton mice (Peromyscus gossypinus) and some rice rats
(Oryzomys palustris), cotton rats (Sigmodon hispidus), house mice (Mus
BULLETIN FLORIDA STATE MUSEUM
Figure 16.-Kibler, the most heterogeneous Reclaimed, Ungrazed site. Reclamation
consisted of leveling the overburden piles, leaving scattered ponds (right background) and
marshes. Oldfield succession is arrested by mowing twice a year. Since the implementa-
tion of reclamation regulations in 1975, a combination of mowed grassland with ponds or
lakes is referred to as "land and lakes reclamation."
musculus), and two species of shrews (Blarina carolinensis and Cryptotis
parva). As the willow thicket developed and later the wax myrtle canopy
closed, the semi-arboreal P. gossypinus became very abundant, Sigmo-
don populations remained unchanged, and the Mus and shrews became
less abundant. The result was that diversity values generally decreased
from the youngest stage (Swift) to the oldest (A-3), whereas abundance
measures increased during the sequence. In comparison with other
treatments (Fig. 18), the diversity and abundance of small mammals on
clay wastes were moderate.
UNRECLAIMED PITS AND SPOIL PILES.-Beginning the successional
sequence, moderately high small mammal diversity and abundance val-
ues were found in the 0-5 year old treatment (Fig. 16), indicating a
strong colonization of the young herbaceous vegetation. Colonization of
the unreclaimed land was by Mus, Sigmodon, Oryzomys, and some P.
gossypinus. Single oldfield mice (Peromyscus polionotus) were captured
at two of the youngest sites, Big Teeth and Shark Tooth (Fig. 6), but none
occurred on older sites.
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 79
*.n '.* -'* -. a; . . . -,'. ,
Figure 17.-6-D, the pasture with the least plant cover among Reclaimed, Grazed
sites. Reclamation here involved filling in old mine cuts with fine sand tailings (resulting
fiom recent flotation treatment of ore) and then flattening emerged spoil pile tops as a
shallow overburden cap. Vegetation is bahia grass with some hairy indigo and rattlesnake
weed in the foreground.
Additional species of small mammals established populations in the
5-15 year old treatment (Fig. 18), resulting in very high values. The
population of Mus declined somewhat between the 0-5 and 5-15 year
classes, but the major change was a substantial increase in the abundance
of Sigmodon, P. gossypinus, and Oryzomys. Both events were normal
responses to the increase in plant cover, particularly herbaceous species
and vines. The pattern of increasing abundance was accompanied by similar
increases in diversity, because most populations continued to grow and a
new species (the eastern woodrat, Neotomafloridana) invaded.
In later seral stages, the small mammal community became less prom-
inent, with moderate diversity and abundance levels. Mus became very
uncommon and population levels of Sigmodon and Oryzomys fell. P. gos-
sypinus populations remained fairly constant, whereas Neotoma became
slightly more common. The reduction of Mus populations probably was
caused by changing habitat characteristics, resulting from development
of shrub and tree vegetation, and possibly by increased competition from
other small mammal species. The drop in Oryzomys numbers may have
Table 3. Summary of small mammal diversity and abundance during autumn quarter.
Asterisks denote sites not sampled during this quarter.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (g)
Consolidated Clay
Settling Ponds
*Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
*Big Teeth
*Shark Tooth
*Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
*Homeland Cemetery
*Gator Lake
*Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
*Saddle Creek Park
*Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
*H-4
Mean standard deviation
1
3
2.01.4
5
7
4
5.31.5
1
3
2
2.01.0
1
1
3
1.71.2
0
2
1.01.4
0.00
0.74
0.370.52
1.36
1.69
1.15
1.400.27
0.00
0.86
0.24
0.360.44
0.00
0.00
0.64
0.220.36
0.00
0.69
0.340.48
5
32
18.019.1
48
38
62
49.312.0
6
13
15
11.34.7
4
5
89
32.748.8
0
2
1.01.4
156.0
1374.0
765.0861.2
3305.3
2193.0
3636.0
3044.8 756.0
1712.2
1712.2
188.0
380.0
490.6
352.9153.1
39.5
58.5
5197.0
1765.02972.2
0
62.5
31.244.2
Table 4. Summary of small mammal diversity and abundance during winter quarter.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (g)
Consolidated Clay
Settling Ponds
Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean + standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
4
3
3
3.30.6
4
3
5
4.01.0
4
6
6
5.31.2
3
4
4
3.70.6
1
3
3
2.31.2
1
5
3
3.02.0
1
1
3
1.71.2
0
4
4
2.72.3
1.47
0.71
0.48
0.88 0.51
0.90
0.34
1.11
0.780.40
0.89
1.38
1.52
1.260.33
0.74
1.23
1.12
1.030.26
0
1.06
1.06
0.700.61
0
1.33
0.80
0.710.66
0
0
0.40
0.13 0.23
0
0.25
1.19
0.480.62
22
17
29
22.76.0
14
25
65
34.726.8
67
46
56
56.3 10.5
8
34
11
17.7 14.2
4
5
5
4.70.6
11
14
7
10.73.5
8
1
37
15.3 19.1
0
12
22
11.311.0
473.4
654.0
992.1
706.5263.3
262.5
301.5
1952.0
838.7 964.4
4212.0
2699.4
3808.7
3570.0 788.9
334.0
2840.8
525.0
1233.3 1395.4
457.0
574.0
519.5
516.858.5
296.5
321.0
291.5
303.0 15.8
98.5
16.5
2458.0
857.7 1386.5
0
169.0
1184.0
451.0640.4
Table 5. Summary of small mammal diversity and abundance during spring quarter.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (g)
Consolidated Clay
Settling Ponds
Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean + standard deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
2
3
2
2.30.6
2
3
3
2.70.6
4
5
4
4.30.6
2
3
4
3.01.0
2
2
1
1.70.57
2
3
1
2.01.0
2
0
4
2.02.0
1
0
4
1.62.0
0.66
0.83
0.17
0.550.34
0.50
0.86
0.83
0.730.19
1.21
1.39
1.25
1.280.09
0.47
1.08
1.12
0.890.36
0.68
0.64
0.00
0.440.38
0.24
1.04
0.00
0.420.54
0.69
0.00
1.06
0.590.54
0.00
0.00
0.89
0.290.51
8
16
25
16.38.5
10
11
26
15.79.0
27
36
27
30.05.2
11
15
11
12.32.3
12
3
1
5.33.8
15
4
3
7.36.6
2
0
50
17.328.3
1
0
21
7.311.8
683.3
706.5
761.0
716.939.9
147.0
139.0
710.5
332.2 327.7
1708.0
1817.7
1264.4
1596.7 293.0
355.0
957.5
1288.5
867.0 473.3
1081.5
249.5
22.0
451.0557.8
280.0
198.3
77.5
185.3101.9
61.0
0.0
3830.1
1297.0 2193.8
12.5
0.0
1767.5
593.3 1767.5
Table 6. Summary of small mammal diversity and abundance during summer quarter.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (g)
Consolidated Clay
Settling Ponds
Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
3
3
1
2.41.2
3
2
2
2.30.6
5
3
5
4.31.2
2
4
4
2.91.0
3
3
2
2.70.6
1
3
2
2.01.0
0
1
3
1.31.5
2
1
4
2.31.5
0.99
0.85
0.00
0.610.53
0.79
0.38
0.65
0.600.20
1.14
0.97
0.99
1.03 +0.09
0.66
1.00
1.24
0.96 +0.29
0.74
0.84
0.64
0.640.10
0.00
1.05
0.27
0.440.54
0.00
0.00
0.25
0.080.14
0.68
0.00
0.85
0.510.44
11
13
14
12.71.5
13
8
14
11.73.2
28
42
55
41.713.5
8
20
6
11.37.6
8
9
3
6.73.2
18
5
13
12.06.6
0
1
25
8.714.2
5
1
18
8.08.9
419.0
783.5
376.0
523.8 225.6
397.9
101.1
409.5
302.8+ 174.8
2275.0
3019.0
4829.1
3374.3 1313.6
343.5
1443.8
494.0
760.4596.6
704.0
763.1
487.0
651.3 145.3
471.3
393.0
410.0
424.841.2
0.0
4.5
2129.4
711.3 1228.1
72.0
13.0
1623.7
569.6913.4
BULLETIN FLORIDA STATE MUSEUM
MAMMALS
DIVERSITY
1.25-
S 1.00-
S .75-
I
.50-
- .25
I
ABUN DANCE
UNR.CLAY RECL. UNRECL.
: 0 U.UG. o
'n
0
Figure 18.-Small mammal diversity and abundance, annual mean of quarterly means
(n =4). Abbreviations are as in Figure 3.
UNR.CLAY RECL. UNRECL.
3 O U. G. no n, o
3: o g^
/s
n .... .. .... . ... .. !
l i # ", r f i l *I I I
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 85
resulted from several factors. When pits were first abandoned, broad
areas were available for marsh development because water levels were
kept low by pumping. Over time water rose on the steep sides of the
pits, narrowing the zone for littoral vegetation. When shrubs and trees
grew on the bank, shading of cattails and other interactions may have
become important. Whatever the mechanism, older sites characteristi-
cally had very little emergent marsh vegetation.
Lowest abundance occurred in the >30 year old class without lakes.
However, the difference between the unreclaimed areas with lakes and
those without was more evident in the species composition than in the
diversity and abundance measures shown in Figure 18. The strictly ter-
restrial sites were heavily dominated by P. gossypinus, and other species
were uncommon. The Christina site contributed much to the diversity
of the treatment because both species of shrews (Blarina carolinensis
and Cryptotis parva) and Mus occurred there. But these animals may
have immigrated from adjacent habitats-a residential community and a
grassy field. Woodrats were absent from these sites, except for one im-
mature individual caught at Christina during the summer quarter. This
animal also may have dispersed from neighboring locations.
In contrast, the mature sites with lakes had considerably fewer Pero-
myscus, while moderate populations of Neotoma and Sigmodon were found
on most areas. This caused the difference in biomass between the two
treatments, because these species are much larger than the Peromyscus
that dominated the without-lakes community. Sigmodon probably was
favored by the retention of herbaceous cover (Fig. 12) along recreational
roads. Neotoma may have benefitted from the combination of mature
forest and the structurally diverse vegetation along water edges.
Reclaimed Pastures.--In general the reclaimed pasture sites showed
low to intermediate small mammal community measures compared with
the other treatments (Fig. 18). These summary values are probably
somewhat higher than the "typical" case for reclaimed sites in the region,
because two sites, Kibler and H-4, accounted for 87% of the individuals
and 97% of the biomass on both reclaimed treatments (Tables 3-6). This
high concentration resulted from the dense populations of Sigmodon found
in the tall stands of grass that escaped mowing or intensive grazing. Most
of the modern reclamation treatments in the area more closely resemble
the other sites, but the Kibler and H-4 sites may indicate the potential
for succession if mowing and intensive grazing were to be reduced.
The difference between grazed and ungrazed reclaimed sites was af-
fected by these two sites. The extraordinarily high numbers of Sigmodon
at the Kibler site resulted in very high abundance values, whereas H-4
consistently had four species present in more even proportions, resulting
in higher diversity for the grazed category.
BULLETIN FLORIDA STATE MUSEUM
BIRDS
The results of bird transects are summarized in Tables 7-10. A total of
150 species was observed in the mining region during the study, and 114
were recorded from the 24 discrete study sites.
CONSOLIDATED CLAY SETTLING PONDS.-In this treatment, both
abundance and diversity values were low compared with the other seven
treatments (Fig. 19). Most of the avifauna on these sites consisted of
resident passerines that usually were associated with brushy habitats,
including gray catbird, white-eyed vireo, common yellowthroat, red-
winged blackbird, cardinal, and rufous-sided towhee. Only six non-pas-
serine species were recorded from the treatment, and wading and water
birds generally were absent. Two wetland species, common moorhen
and common snipe, were seen in the Swift site, which was the wettest
and least consolidated of the three. The low bird diversity and abun-
dance of clay waste sites probably was related to the uniform and simple
structure of the vegetation.
Most of the species in this treatment were permanent residents, and
the bird community showed very little seasonal variation in either diver-
sity or abundance (Tables 7-10). The abundance values in particular were
low and remained low through the winter, when other categories of sites
were used by flocks of either warblers (unreclaimed pits) or sparrows
(reclaimed sites). Though non-resident wintering birds heavily used wax
myrtle on unreclaimed sites, wintering birds were scarce on consolidated
clay settling ponds despite abundant wax myrtle. Evidently an important
component was lacking. Low amounts of insect biomass (Table 11) may
have resulted in a short supply of protein for insectivorous or seasonally
omnivorous birds.
UNRECLAIMED PITS AND SPOIL PILES.-In the successional se-
quence, the diversity and abundance measures increased with increasing
age of the site but, unlike the small mammal data, reached their peak at
a much later stage-either the 15-30 year class or the >30 year old class
with lakes (Fig. 19).
The youngest class, 0-5 years old, was distinctly different from the
older stages, with lower diversity and abundance measures as well as
different species composition. This undoubtedly resulted from the early
stage of oldfield succession on these sites. In some respects the avifauna
on these young sites resembled that found on reclaimed pastures, with
savannah sparrows, palm warblers, killdeer, and other shorebirds being
common in both categories. In addition the lakes on these sites attracted
the largest diversity of waterfowl of the eight categories. Nine species
were observed there, compared with only three in the 5-15 year old
pits. The reason for the high use by ducks is unclear. One possible expla-
VOL. 30 NO. 3
Table 7. Summary of bird diversity and abundance during autumn quarter based on
one-kilometer transects. Asterisks denote those areas that were not sampled
this quarter.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (kg)
Consolidated Clay
Settling Ponds
*Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
*Big Teeth
*Shark Tooth
*Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
*Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
*Homeland Cemetery
*Gator Lake
*Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
*Saddle Creek Park
*Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean + standard deviation
Reclaimed Pasture
Ungrazed
*Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
*6-D
*Marina East
*H-4
Mean standard deviation
8
4
6.02.8
15
14
14.50.7
22
22
3
4
6
4.31.5
10
8
91.41
1.80
1.04
1.42- 0.53
2.32
2.18
2.250.9
1.45
1.45
0.84
1.19
1.57
1.200.36
1.70
1.20
1.45 1.35
247.7
269.6
258.6 15.5
829.5
1228.9
1029.2 282.4
2071.2
2071.2
62.2
118.1
228.6
136.1 84.4
869.6
181.3
844.0 36.3
6.51
7.59
7.050.76
224.98
91.21
158.09 94.58
114.35
114.35
0.66
7.00
17.02
8.22 _8.24
94.03
22.77
58.40 -50.38
Table 8. Summary of bird diversity and abundance during winter quarter based on
one-kilometer transects.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (kg)
Consolidated Clay
Settling Ponds
Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
8
12
7
9.02.6
9
15
26
16.78.6
18
24
25
22.33.71
24
19
26
23.03.6
30
24
22
25.34.2
8
12
16
12.04.0
7
9
16
10.74.7
4
7
12
7.74.0
1.87
2.23
0.70
1.600.80
1.38
1.87
2.18
1.810.40
2.12
1.46
1.94
1.840.34
1.89
1.34
2.38
1.870.52
1.67
1.44
2.38
1.830.49
1.18
2.12
2.26
1.850.59
0.58
1.56
1.96
1.370.71
0.75
0.43
1.37
0.850.48
108.8
309.2
305.1
241.0 114.5
117.0
47.4
297.9
154.1 129.3
731.5
992.8
1208.1
977.5238.7
1474.6
1672.9
1051.9
1399.8317.2
2280.2
383.5
366.1
1009.9 1100.2
196.5
127.5
389.8
237.9136.0
124.6
130.0
239.6
164.764.9
218.8
97.0
640.1
318.6285.0
3.41
10.50
5.54
6.483.64
12.56
6.28
75.40
31.4138.22
77.32
58.87
76.66
70.95 10.47
45.42
124.97
126.48
98.96 46.37
184.75
22.89
136.43
114.69 83.09
2.53
4.17
11.37
6.024.70
10.75
6.50
17.32
11.525.45
50.06
5.18
44.29
33.1824.42
Table 9. Summary of bird diversity and abundance during spring quarter based on one-
kilometer transects.
Diversity Abundance
Treatments Number Number of
Sites of species H' individuals Biomass (kg)
Consolidated Clay
Settling Ponds
Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits
and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
7
5
8
6.71.5
9
13
18
13.34.5
19
25
25
23.03.5
18
24
21
21.03.0
30
17
13
20.00.89
10
11
14
11.72.08
9
7
15
10.34.2
3
3
7
4.32.3
1.75
1.44
1.82
1.670.20
1.70
2.19
1.83
1.91 0.25
2.33
2.64
2.36
2.440.17
2.34
2.57
2.64
2.52 0.16
2.72
2.19
2.28
2.400.28
2.08
2.17
1.96
2.070.10
2.02
1.38
2.20
1.870.43
0.94
1.10
1.79
1.280.45
78.5
50.9
60.4
63.3 14.0
74.6
44.6
202.5
107.2 83.9
291.9
222.2
453.1
322.4 118.4
238.1
350.3
330.4
306.359.9
880.0
148.8
77.3
368.7444.2
101.0
89.8
287.6
159.5111.1
20.6
55.1
132.3
69.357.2
13.4
3.0
82.0
32.842.9
2.85
1.85
1.72
2.140.62
3.16
4.28
84.64
30.69 46.72
32.15
18.18
37.21
29.189.86
27.10
63.11
248.86
113.02 119.01
108.26
9.69
16.88
44.9454.95
5.24
4.93
8.21
6.131.81
6.31
6.04
11.04
7.802.81
1.30
0.22
2.26
1.261.02
Table 10. Summary of bird diversity and abundance during summer quarter based on
one-kilometer transects.
Diversity Abundance
Treatments Number Numberof
Sites of species H' individuals Biomass (kg)
Consolidated Clay Settling Ponds
Swift 10
N-2 8
A-3 6
Mean standard deviation 8.02.0
Unreclaimed Pits and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard deviation
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
1.74
1.83
1.29
1.620.28
3 0.81
5 1.21
12 1.58
6.74.7 1.200.38
13
21
13
15.74.6
1.50
2.57
1.04
1.700.78
15 2.22
13 2.10
15 2.30
14.31.2 2.210.10
21 2.34
20 2.69
15 2.09
18.73.2 2.370.30
9 1.93
9 1.86
13 2.04
10.32.3 1.940.09
6 1.68
5 1.84
10 1.70
7.02.6 1.740.09
1 0.00
4 1.27
6 1.49
3.72.5 0.920.80
75.5
131.1
55.5
87.4039.2
31.2
22.3
107.9
53.8 47.1
190.5
163.2
365.1
239.6 109.5
139.9
153.2
202.9
165.333.2
238.5
62.0
88.4
129.695.2
42.2
66.7
120.8
76.640.2
25.5
37.2
77.9
46.927.5
1.0
26.7
91.3
29.730.3
4.66
5.19
1.41
3.752.04
3.45
1.99
33.38
12.94 17.72
35.76
17.33
39.85
30.9812.00
29.90
66.37
101.84
66.0435.97
58.80
22.61
21.01
34.1421.37
3.59
8.44
8.37
6.802.78
5.85
4.20
12.13
7.39 4.18
0.10
8.07
23.86
10.68 12.09
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 91
BIRDS
DIVERSITY
20,
y,
uj
U
0.
U-
0 10
uj
z
ABUNDANCE
'
U,
[ g
r o
n-I la . *.
UNR. CLAY REC
SU. G.
^o
L. UNRECL.
I -
A
UNR.CLAY RECL. UNRECL.
S0 u. G. a, o
^ o ^
C.)
no
Figure 19.-Bird diversity and abundance, annual mean of quarterly means (n=4).
Abbreviations are as in Figure 3.
900
S600-
z
u-
0
S 300-
CIO
z
Table 11. Insect biomass (g) in light trap samples through the year. Sample sizes are given in parentheses.
Treatments Mean of
Sites Autumn Winter Spring Summer Quarterly Values
Consolidated Clay Settling Ponds
Swift
N-2
A-3
Mean standard deviation
Unreclaimed Pits and Spoil Piles
0-5 Years With Lakes
Big Teeth
Shark Tooth
Young Tiger
Mean standard deviation
5-15 Years With Lakes
Orange Grove Pits
Tiger Tail
Tiger Bay South
Mean standard deviation
15-30 Years With Lakes
Homeland Cemetery
Gator Lake
Achan-4
Mean standard deviation
9.950.44(2)
0.80(1)
5.386.47(2)
36.44(1)
5.48(1)
26.02(1)
22.65 15.76(3)
1.54(1)
0.64(1)
0.48(1)
0.89 0.57(3)
0.00(1)
0.42(1)
0.15(1)
0.190.21(3)
0.94(1)
0.48(1)
2.59(1)
1.341.11(3)
1.60(1)
0.17(1)
0.30(1)
0.69(3)
1.80(1)
8.34(1)
3.58(1)
4.57 3.38(3)
2.00 1.80(2)
2.99 0.85(2)
18.8117.60(2)
7.9311.57(6)
17.07 11.64(2)
1.50 1.87(2)
7.84 4.14(2)
8.80 8.96(6)
3.711.77(2)
7.31 2.31(2)
6.316.29(2)
5.78 3.52(6)
3.410.24(2)
2.490.01(2)
4.11 0.70(2)
3.340.80(6)
5.89 0.17(2)
13.09 7.07(2)
16.01 5.89(2)
11.666.21(6)
6.76 4.60(2)
4.63 3.13(2)
25.58 3.11(2)
12.32 10.70(6)
17.99 0.77(2)
2.490.98(2)
10.284.64(2)
10.257.26(6)
z
cT
C
tr
2.25(3)
5.36(4)
2.24(4)
3.54(4)
2.63(3)
5.50(3)
11.66(3)
6.59(3)
15.30(4)
3.02(4)
15.91(4)
11.28(4)
7.77(3)
3.32(3)
5.63(3)
5.57(3)
>30 Years With Lakes
Bartow South
Saddle Creek Park
Sanlan Ranch
Mean standard deviation
>30 Years Without Lakes
Old Spoil Piles
Christina
Old Clarke James
Mean standard deviation
Reclaimed Pasture
Ungrazed
Parcel B
Noralyn
Kibler
Mean standard deviation
Grazed
6-D
Marina East
H-4
Mean standard deviation
0.48(1)
0.48(1)
2.80(1)
7.68(1)
6.21(1)
5.56 2.50(3)
46.17(1)
12.92(1)
5.72(1)
21.6021.58(3)
2.39(1)
1.19(1)
1.79 0.85(2)
0.18(1)
5.68(1)
0.29(1)
2.05 3.14(3)
0.04(1)
0.76(1)
0.19(1)
0.330.38(3)
0.01(1)
0.00(1)
0.64(1)
0.22 0.36(3)
0.00(1)
0.02(1)
0.26(1)
0.09 0.14(3)
7.56 6.96(2)
2.99 3.45(2)
9.58 13.26(2)
6.71 7.51(6)
68.11 75.75(2)
7.93 8.98(2)
13.28 11.16(2)
29.77 45.57(6)
18.8311.82(2)
15.66 14.19(2)
65.51 (1)
33.3327.91(3)
10.05 11.47(2)
9.73 6.57(2)
4.82 5.04(2)
8.20 6.85(6)
14.690.04(2)
8.640.77(2)
8.252.23(2)
10.53 3.40(6)
14.873.25(2)
14.60 0.24(2)
15.483.82(2)
14.98 2.28(6)
21.125.81(2)
2.242.04(2)
8.384.06(2)
10.58 9.22(6)
10.20 2.27(2)
12.66 9.98(2)
25.77 30.88(2)
16.21 16.36(6)
5.73(4)
5.77(3)
6.04(3)
4.94(4)
21.46(4)
7.74(4)
8.79(4)
12.66(4)
21.53(4)
7.70(4)
20.06(4)
16.43(4)
5.66(4)
5.90(4)
10.28(3)
6.57(4)
BULLETIN FLORIDA STATE MUSEUM
nation may be that these young sites usually were adjacent to active min-
ing operations. Consequently the areas were not open to the public and
were completely closed to waterfowl hunting, resulting in little human
disturbance to the birds. Another possible reason may involve a more
dense or higher quality food source than in older pits.
In a parallel with the distribution of oldfield mice, there appeared to
be a distinct bird community on Young Tiger compared with birds on the
two younger sites in this class, Shark Tooth and Big Teeth. Young Tiger
had 42 species present during the year, compared with 27 and 24 respec-
tively for the other two. In addition, the other diversity and abundance
figures showed comparable differences (Tables 7-10). The higher bird
community measures may be explained by the more advanced stage of
plant succession supported by Young Tiger. Indeed, certain passerine
species, such as palm warblers, savannah sparrows, boat-tailed grackles,
and red-winged blackbirds, were consistently more abundant on the Young
Tiger site than on the other two. However, much of the difference in
species composition resulted from the high number of wading and water
birds found at Young Tiger. This is more likely tied to some difference in
food supply, water quality, or water levels than to terrestrial plant succes-
sion on the exposed spoil piles. We lack the data to evaluate the impor-
tance of these or other factors. In any case, it appears that in terms of
bird communities the arbitrary age category of 0-5 years may be mis-
leading in that the first major increase of bird diversity and abundance
may occur in 3 years after mining. This statement is based on the data
from only one site; however, qualitative observations on other young pits
in the area tended to support that view.
The 5-15 year old treatment was similar in species composition to
Young Tiger, but with the increase in vine and shrub cover we found
additional species of resident passerines, including white-eyed vireos,
cardinals, and blue jays. In addition, several species of winter residents,
such as yellow-rumped warblers and house wrens, were rare in the 0-5
year class but abundant in the 5-15 year sites. Undoubtedly the shrubs
and vines provided good cover for both wintering and nesting species,
but perhaps even more important was the use of wax myrtle berries for
food during the winter months. This became more apparent in the older
sites but was readily observed here.
Wading birds were still quite common in this class, and increased
shoreline and littoral vegetation provided habitat for king rails and soras.
However the only ducks using these sites consistently were blue-winged
teal.
The 15-30 year old pits and those >30 years old with lakes showed
quite similar species composition and diversity measures (Fig. 19). Per-
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 95
haps the most significant difference was with the woodpeckers. Four spe-
cies (red-bellied and pileated woodpeckers, yellow-bellied sapsucker, and
common flicker) were found in the older category, whereas only the com-
mon flicker was found in the 15-30 year class. This reflects the increased
availability of larger trees and dead snags for both nesting and foraging.
Another interesting difference was that wood ducks were found in all
three of the older sites, but in none of the 15-30 year old sites. Again,
this could be due to the presence of trees large enough to contain cavi-
ties. The hypothesis that nesting sites limit wood ducks to use of the
forested sites could be tested by erecting artificial nesting boxes in younger
areas and observing subsequent use of the sites.
Palm warblers, typically occurring in open ground and brushy habi-
tats, declined in importance with increasing age of the sites after reach-
ing their peak numbers in the 5-15 or 15-30 year old category. The fairly
large number of palm warblers on the older Bartow South site may have
been in response to the surrounding areas that were dominated by old-
fields.
Unlike the diversity measures, the abundance figures for the 15-30
year class and the >30 year class with lakes were quite different (Fig.
19). This pattern is misleading because the younger treatment was not
sampled during the fall quarter and, among the older sites, only Bartow
South was sampled. A great many small birds (2071) were recorded; over
half of these were wintering yellow-rumped warblers in large flocks. This
extraordinarily high count affected the four-quarter mean for this treat-
ment. It may have been representative of the site (the winter count was
even higher), but it was not balanced by the other two sites, which gen-
erally were lower. In fact, using the three quarters for which complete
data are available, the mean number of individuals is higher for the 15-
30 year class than for the >30 year class (623 vs. 502). Domination of the
younger site by wax myrtle created a very concentrated, high energy
food source for wintering birds. Because the large flocks of wintering
warblers have such a major effect on the total autumn and winter quarter
counts, their habitat use is directly related to the total year counts. This
is supported by the fact that Bartow South had far more yellow-rumped
and palm warblers than Saddle Creek Park and Sanlan Ranch, where the
importance of wax myrtle was much less.
The larger differences between old sites with lakes and those without
(Fig. 19) resulted from the absence of water and the small wintertime
berry crop in the latter category. Obviously all wading and water birds
were absent from the bird community, which greatly reduced the diver-
sity and biomass of birds found on these sites. Though the passerine
species composition was very similar, abundance values were much lower
BULLETIN FLORIDA STATE MUSEUM
in the without-lakes category, because large flocks of wintering warblers
did not use these habitats. This again suggests the importance of wax
myrtle to the winter ecology of these birds. The most common species
were cardinals, rufous-sided towhees, and Carolina wrens-typical resi-
dent forest species.
As expected, maturing forest without lakes was heavily occupied by
woodpeckers, with six species recorded for the category. The wealth of
large trees and a good supply of dead stubs provided excellent foraging
and nesting habitat.
Reclaimed Pasture.--Fairly low diversity and abundance figures were
obtained for both categories of reclaimed pastures (Fig. 19). These values
were most comparable to those for the consolidated clay settling ponds
and the >30 year class without lakes, although because of the vastly dif-
fering plant structure the avian community was composed of different
species. Typical species found on reclaimed sites included cattle egrets,
red-winged blackbirds, and American kestrels. As on the 0-5 year un-
reclaimed pits and spoil piles treatment, some shorebirds and wintering
savannah sparrows occurred on reclaimed pastures.
The two reclaimed sites with the most advanced and heterogeneous
plant communities, Kibler and H-4, also supported the most diverse
avian communities. In addition to the species mentioned, swamp spar-
rows, common yellowthroats, northern harriers, and a few cardinals were
observed on these two sites and not on the other four. As with the small
mammal data, Kibler and H-4 contributed heavily to the mean diversity
and abundance values for their respective classes of reclaimed pastures.
The diversity and abundance values for ungrazed sites usually were
much higher than those for grazed sites. This was the case during spring
and summer, but the situation was reversed during winter because large
flocks of birds occurred on grazed sites (American robins on H-4 and
boat-tailed grackles on 6-D). The higher values on ungrazed sites did not
result directly as a response to vegetation, which was somewhat more
diverse and abundant on grazed sites (Figs. 3-5). They may have been
related to the very large insect biomass found on the ungrazed pastures
(Table 11), because many of the bird species are at least partially insecti-
vorous.
From a regional perspective, the open grassland provided habitat for
some species that were not found in other types of sites. Eight species
were found only on reclaimed sites: cattle egrets, black vultures, bald
eagles, short-billed marsh wrens, starlings, bobolinks, eastern meadow-
larks (with one exception), and song sparrows. While some of these may
be sampling artifacts (i.e. bald eagle), it is clear that the presence of these
pastures serves to increase the avian diversity of the region as a whole.
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 97
INSECTS
Insect responses to post-mining treatments were presented as an
overview of abundance (Fig. 20), because detailed sample analysis was
completed only for winter quarter. Seasonal levels of abundance are
given in Table 11. The lowest insect abundance occurred on consolidated
clay settling ponds, with consistently low values in all quarters. During
winter, the insect fauna on these sites was dominated by flies and bee-
tles. In the successional range of sites, insects were most abundant in
the young willow seral stage (Swift) and least abundant in the mature
wax myrtle forest (A-3).
In the successional sequence of unreclaimed pits and spoil piles with
lakes, insect abundance peaked in the 5-15 year age class and declined
201 INSECTS r
0
< 10-
0
5-
0-
I
/~J
X /
7il
7< Jf
/i5
I
e1111
/
7O,
7
I I I 1______ -- rI -I -I IIII I -.--
UNREC. CLAY RECL.
>301 >30
U. G.
UNRECLAIMED
0-5 5-15 15-30 >30
W I/O --WITH LAKES--
Figure 20.-Nightly insect biomass in light trap samples through the year, annual
mean of quarterly means (n= 4). Abbreviations are as in Figure 3.
BULLETIN FLORIDA STATE MUSEUM
thereafter. All these communities were characterized by high levels dur-
ing the summer, and the prominence of the 5-15 year treatment was a
result of high levels in autumn. However, the >30 year treatments
lacked this large contribution in the autumn. The two mature forest in-
sect communities resembled each other in having high summer and low
autumn levels, but the without-lakes community was distinctive in hav-
ing very high numbers of large beetles and moths during the spring
quarter. As a result, this treatment had the highest insect abundance of
any unreclaimed treatment.
Both ungrazed and grazed reclaimed treatments had insect faunas
dominated by caddisflies (Trichoptera) and small beetles. Ungrazed sites
had the most abundant insects of any treatment, with high numbers in
spring, summer, and autumn. Seasonal abundance was much different
on grazed sites, which had moderate numbers in spring, high numbers
in summer and low numbers the rest of the year.
HERPETOFAUNA
Observations of the 9 species of amphibians and 23 species of reptiles
found on study sites are presented in Table 12. Perhaps the most striking
figure was the low number of species (total = 5, mean = 2) observed in
the consolidated clay settling ponds. Only one frog species, the green-
house frog (Eleutherodactylus planirostris), was observed in this cate-
gory even though the moist substrate, deep cracks in the clay crust, and
shady aspect of the sites seemed to provide suitable habitat for more
species.
Only six species were observed on each of the two early unreclaimed
treatments. In contrast, high numbers of species were found in older
stages of unreclaimed pits and spoil piles, because these sites provided
both aquatic habitat for frogs, turtles, and water snakes and shaded ter-
restrial habitat for snakes and skinks. The 15-30 year old class had the
highest number of species (16). The highest mean number of species
(7.7) occurred in the >30 year old pits without lakes, despite the lack of
aquatic habitat on these sites. The well-developed forest there supported
a consistently large number of snake, lizard, and arboreal frog species.
Additionally, all three sites contained active and inactive burrows of the
gopher tortoise (Gopherus polyphemus), which provide cover and nest
sites for numerous vertebrates (Auffenberg 1978).
As expected, few reptiles (3 species of snakes) and no amphibians were
found on reclaimed sites because the very open, sunny aspect created
severe conditions that most species find intolerable. In addition, no bur-
rrows were available to provide protection from the sun.
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 99
LARGE MAMMALS
Observations of large mammals and their signs recorded throughout
the study are presented in Table 12. Thirteen species were found, but
no relative abundance values could be attached to the presence/absence
data. The most ubiquitous of the species was the bobcat (Lynx rufus),
which was found in 14 sites-in all 8 treatments. Other species were
nearly as widespread, including raccoons (Procyon lotor, 13 sites), arma-
dillos (Dasypus novemcinctus, 11 sites), and opossums (Didelphis virgi-
niana, 10 sites). River otters (Lutra canadensis) were observed in all age
classes of unreclaimed pits and spoil piles where lakes were present. In
addition, otters were commonly seen in ditches, pools, and active clay
settling ponds in the area. Gray squirrels (Sciurus carolinensis) were
confined to both categories of old unreclaimed pits where oaks were
abundant. No sign of white-tailed deer was noted, however, and their
absence from the mined region was confirmed by mining company em-
ployees.
The highest mean numbers of species (Table 12) occurred in the 5-15
and 15-30 year classes of unreclaimed pits, with 6.3 and 6.0 species
respectively. The high figures probably resulted from two factors: the
habitats had developed sufficiently to allow many species to use the sites,
and most of the sites had little human disturbance because they were
partially closed to the public. The sites in the >30 year old pits-with-
lakes category were generally more accessible to the public and were
heavily used for fishing, picnicking, and other activities that may have
reduced the mean number of large mammals (3.7) using those sites. Old
sites without lakes had a mean of 5.3 species per site, and these sites
were seldom used by the public.
Reclaimed sites showed consistently low numbers of species, although
some of the wide-ranging animals such as bobcats, raccoons, and red
foxes (Vulpes vulpes) were found to either use or travel through some of
the sites.
The most conspicuous large mammal on the grazed treatment was
domestic cattle. Data provided by the landowner, International Minerals
and Chemical Corporation, showed the sites to be stocked at an average
of 0.90 head per ha (0.68 on 6-D, 1.25 on Marina East, and 0.77 on H-
4). With biomass approximated as 205 kg per head, an average of 184 kg
of cattle biomass per ha is indicated. Higher stocking rates may be pos-
sible, but company policy of assuring that grassland cover is maintained
on reclaimed pastures is implemented by lease agreements that prohibit
overglazing.
BULLETIN FLORIDA STATE MUSEUM
Table 12. Number of species of amphibians, reptiles, and large mammals observed
during all four quarters of field work.
Number of Species
Treatments
Sites Amphibians Reptiles Large mammals
Consolidated Clay Settling Ponds
Swift 0 1 3
N-2 1 1 4
A-3 1 1 1
Unreclaimed Pits and Spoil Piles
0-5 Years With Lakes
Big Teeth 2 0 4
Shark Tooth 2 0 3
Young Tiger 3 1 7
5-15 Years With Lakes
Orange Grove Pits 0 1 6
Tiger Tail 1 1 7
Tiger Bay South 0 2 6
15-30 Years With Lakes
Homeland Cemetery 2 5 9
Gator Lake 2 5 6
Achan-4 3 2 3
>30 Years With Lakes
Bartow South 3 4 4
Saddle Creek Park 5 0 3
Sanlan Ranch 1 4 4
>30 Years Without Lakes
Old Spoil Piles 3 5 4
Christina 1 5 3
Old Clarke James 2 4 9
Reclaimed Pasture
Ungrazed
Parcel B 0 0 1
Noralyn 0 1 3
Kibler 0 2 1
Grazed
6-D 0 0 0
Marina East 0 0 3
H-4 0 2 0
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 101
STATISTICAL ANALYSIS
The results of tests of hypotheses on contrasted final land uses are
presented in Table 13. Considering annual data, the shrub and tree var-
iables showed significant differences among end results of treatments,
whereas herb, herpetofaunal, and large mammal variables did not. Dun-
can's multiple range tests of end results of reclamation showed two sig-
nificantly different groups->30 year old unreclaimed sites with and
without lakes had high mean values, and ungrazed and grazed reclaimed
pastures had low values. The clay waste treatment fell between these
groups with intermediate values but was not clearly assignable to either.
The two reclaimed treatments were never significantly different. In cases
where the clay treatment moved up in mean value to second or first
place, it was never significantly different from the third place treatment.
When the two unreclaimed treatments were significantly different, the
treatment with lakes had the higher mean value. For variables with this
relation reversed, the differences were not significant. The contrasts re-
inforced these observations without showing additional distinctions.
Considering seasonal data (Table 13), only bird variables showed sig-
nificant differences among final land uses; insect and small mammal var-
iables did not. Response of seasonal variables to treatment end results
was weaker than their successional responses, for distinct reasons in each
group of animals. For small mammal data, no seasonal effect appeared in
the analysis of variance, but the effect of sites within treatments was
stronger than the overall treatment effect. Hence high site variance pre-
vented distinction of post-mining treatments (Table 13) with small mam-
mal data. By comparison, the strongest effect on bird variables was the
season effect. The Duncan's grouping of final land uses and the contrasts
showed patterns similar to those for annual data, with two different fea-
tures. The clay waste treatment was never significantly different from
the reclaimed pastures. Ungrazed pastures were significantly higher
than grazed pastures in bird H'.
The results of hypotheses on response to site age also are presented
in Table 13. Considering annual data, the shrub and tree variables
showed significant responses to age, whereas herpetofaunal and large
mammal variables did not. Considering seasonal data, small mammal and
bird variables showed significant responses to age over the interval 0-36
years, and insect variables did not. Tests showed no simple pattern of X2,
X3, or X4 functions for all variables.
Models of plant and animal response to age, based on statistically sig-
nificant age effects (Table 13), yield predictions of community succession
on unreclaimed overburden soil (Fig. 21). Shrub species colonized for
the first 9 years after mining (Fig. 21A); then the number of species
BULLETIN FLORIDA STATE MUSEUM
Table 13. F-tests of hypotheses showing significance levels of contrasts between end
results of post-mining treatments and of response to site age. Because of the
large number of tests, an elevated alpha is appropriate, and significance is
judged at P<0.01.
withvs. ungrazed clay vs. clay vs. response to
Variables without vs. grazed with/without pastures site age
Annual data
T. herb species ns ns ns ns ns
Herb H' ns ns ns ns ns
T. herb cover ns ns ns ns ns
T. shrub species ns ns *** *** **
Shrub H' ns ns *** ** **
T. shrub cover ns ns ns *** **
T. tree species *** ns *** *** ***
Tree H' ** ns *** ns **
T. tree individuals ns ns ns ** ***
T. tree basal area ns ns *** **
T. herpetofauna species ns ns ns ns
T. large mammal species ns ns ns ns ns
Seasonal data
T. small mammal species ns ns ns ns *
Small mammal H' ns ns ns ns ***
T. small mammal individuals ns ns ns ns **
T. small mammal biomass ns ns ns ns ***
T. bird species ns ** ns *
Bird H' ns ns ns *
T. bird individuals ns ns ns ns ***
T. bird biomass ** ns ns **
T. insect biomass ns ns ns ns
T = transformed
ns = not significant
* = P<0.05
** = P<0.01
*** = P<0.001
lized as shrub cover and H' continued to increase (the latter because of
increased evenness of species' abundances). Beginning in year 17, a cycle
of colonization and decline occurred, with a peak at 30-31 years. Con-
currently, shrub cover and H' decline as trees became more dominant.
Tree species, H', and individuals (Fig. 21B) all showed steady increases
during succession. However, tree basal area reaches its peak in year 26-
27 and decreased thereafter. Transformed values for small mammal spe-
cies and H' (Fig. 21D) changed very little over time. However, the pat-
tern for small mammal individuals and biomass roughly corresponded
VOL. 30 NO. 3
102
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 103
60 A. SHRUBS 12 B. TREES
55 11
5o"" =-46 10-" 2 .
405 "
> 35 ------------- r" 0996 0> 7
0 30 0 6.
> 25 r2= 0912
|-----------1
sor' oas
-05 -1
-10 -2, -- -- -
0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 2 4 6 10 12 14 16 18 20 22 24 26 2 30 32 3 36
YEARS YEARS
ro C. BIRDS 14 --'". D. MAMMALS
65 r2=0365 13
60- 12 / =0.669
60- 11
935 5
0 2 4 6 10 214161830222426 28 33 024 10 I2 .14.16.18 ....2422. .32. 3
YEARS YEARS
S5show r a -9 a 30-31 23
peak was higher, and abundance levels were very low by year 36. Bird
decreased through year 36 (Fig. 21C). Bird H' increased slowly through----
Stepwise multiple regression generated a large number of significant
relation s r 0403
10 -2
-2
05 -3
0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 3X 3 34 36 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36
YEARS YEARS
Figure 21.-oModels ofplant and animal response to serat age of unreclaimed overbur-
den piles, based on the age effects shown in Table 13.
with that for shrub species, with two cycles of colonization and decline
showing peaks at 8-9 and 30-31 years and a low at 23 years. The first
peak was higher, and abundance levels were very low by year 36. Bird
species, individuals, and biomass increased until 22-25 years and then
decreased through year 36 (Fig. 21C). Bird H' increased slowly through-
out the sere.
Stepwise multiple regression generated a large number of significant
relationships between bird and plant variables. The most consistent re-
lationships can be used to estimate wildlife potential of field sites. Equa-
tions selected for this purpose were limited to those that were based on
annual (not quarterly) data, were mathematically simple, had very high
probability values, and had their dependent variables already shown to
change predictably during succession (Fig. 21A, B). Two equations met
these criteria. For end treatments,
TBS = 2.0777 + 0.7411 TTS (F = 22.1,) P<0.001,r2 = 0.630),
and for successful sites,
TBS = 2.7161 + 0.4489 TSC (F = 19.4, P<0.001, r2 = 0.661),
where TBS is transformed bird species, TTS is transformed tree species,
and TSC is transformed shrub cover.
BULLETIN FLORIDA STATE MUSEUM
DISCUSSION
SYNTHESIS OF RESULTS
The approach of this study solved a commonly recognized problem in
evaluating ecosystems: "Comparisons of the quality of biomass before
and after mining are . difficult because the wildlife composition is of
different species" (Phosphate Land Reclamation Study Commission 1978).
Measuring both diversity and abundance of several components of the
biota on each study site enabled use of features common to all biological
communities in evaluating the quality of post-mining habitats. A wildlife
community normally is considered valuable if it contains a great diversity
of species, a high abundance of one desirable species, or a balance of
moderate levels of diversity and abundance. For example, valuable com-
munities may include a tropical forest (with many species but low abun-
dance), hoofed animals of the Great Plains (with many bison but low
species diversity), and hoofed animals of East African grassland (with
substantial numbers of both species and individuals). Using this rationale
we have compared the quality of very different communities that de-
velop on post-mining treatments.
Table 14 summarizes the relative diversity and abundance of the five
categories of wildlife over the range of treatments. The values are pre-
sented as a percentage of the maximum value obtained for each variable.
No attempt was made to derive a numerical rating of the treatments from
this table, because we chose not to equate the importance of different
types of animals. Instead we compared wildlife values by dividing the
relative figures into three arbitrary ranges and counting the number of
scores in each range for each site class.
This summary shows that the clay settling ponds consistently had low
to moderate values, with only one measure in the high range (Table 14).
It was apparent from general observations in the area that active and
recently deactivated settling ponds provided excellent wetland habitats
with much higher wildlife values, but once a crust formed and willows
dominated the site much of the attractiveness to wildlife was lost. Values
continued to decline as the clay dried further.
The series of unreclaimed treatments showed a primary succession
that included high-quality wildlife communities (Table 14). Colonization
of mined areas began soon after the disturbance and, without manage-
ment, fairly diverse and abundant communities became established within
5-7 years. The 5-15 and 15-30 year old age classes in particular sup-
ported large communities. The two mature classes had slightly lower
values, especially in abundance, but diversities often remained high.
Natural succession of unreclaimed overburden soil led to maturing
VOL. 30 NO. 3
Table 14. Summary of relative importance for mean diversity and abundance. Values are expressed as percentage of the maximum value
obtained for each parameter, and placed in arbitrary ranges (low = 0-33 percent, medium = 34-66 percent, high = 67-100
percent). Values considered high are in italics. Low values are in parentheses.
Large
Small Mammals Birds Insects Herpetofauna Mammals z
Treatments Number Number of Biomass Number Number of Biomass Biomass Number Number
of Species H' Individuals of Species H' Individuals of Species of Species
Consolidated Clay
Settling Ponds 52 48 39 (23) 34 72 (18) (5) (22) (30) 43
Unreclaimed Pits and
Spoil Piles
0-5 Years
With Lakes 62 56 47 (17) 57 74 (12) (27) 40 35 75
5-15 Years
With Lakes 100 100 100 100 88 94 72 78 69 (30) 100
15-30 Years 0
Without Lakes 67 77 (31) (33) 90 100 70 100 34 87 95
>30 Years
Without Lakes 46 42 (21) (29) 100 91 100 83 (30) 78 59
>30 Years Without
Lakes 47 39 (23) (11) 45 80 (17) (7) 77 100 84 Z
Reclaimed Pasture C
Ungrazed 35 (21) 42 40 43 73 (31) (23) 100 (17) (27)
Grazed 40 (32) (14) (14) (24) 46 (14) (16) 40 (9) (16)
BULLETIN FLORIDA STATE MUSEUM
forest, shown by models (Fig. 21B) to be at a late successional stage at
36 years after mining. The late successional stage had an enormous amount
of biomass stored as living wood and had more diverse and more abun-
dant wildlife than endpoints of other treatments-clay waste areas and
reclaimed pastures. In addition, these old sites had considerable aes-
thetic value that is difficult to quantify. As anticipated, community struc-
ture on late successional sites was strongly affected by the presence of
lakes. The kinds of wildlife occupying these two treatments were very
different, and the numbers of tree and bird species, tree H', and amount
of bird biomass were significantly different (Table 13).
Reclaimed pastures had consistently low wildlife values (Table 14). In
almost all categories the animal community on ungrazed sites was more
diverse and abundant than on grazed sites, with a significant difference
apparent for bird H' (Table 13). However, the selective harvest of grazing
resulted in a somewhat more diverse pasture vegetation. Stocking of pas-
tures with cattle, by adding a large amount of mammal biomass, showed
that reclaimed sites have integrity as good life-support systems. Hence,
reclaimed pastures are valuable because they support an abundant food
source for humans. Though wildlife value of this treatment is low, the
viability of reclaimed pastures suggests that high wildlife potential could
be realized on these sites if a wildlife-oriented land use decision was
made.
Four distinct seral stages are apparent (Fig. 21). First is the oldfield
stage of grasses and forbs, reaching its fullest development at 5-7 years.
Second is the pioneer shrub stage, characterized by Lantana, Baccharis,
vines, and forbs, between 8 and 14 years after mining. Third is the wax
myrtle stage, dominated by wax myrtle and invading forest trees from
about 15 to 30 years. Fourth is the oak forest stage, dominated by water
oak, live oak, and several other species. Later forest types that might
occur were not available for study. The decline of herbs occurred as woody
plants increasingly intercepted the sunlight on which the herbs de-
pended for energy. Minimum values for shrub species and tree H' coin-
cided in time and correlated with dominance of wax myrtle in the tree
and/or shrub data. Apparently the allelopathic wax myrtle forced the de-
cline of the pioneer shrub community, dominated the next community of
young trees, and declined as the forest community emerged into the full
sunlight. Two distinctive small mammal communities occurred with peak
abundances in the pioneer shrub and forest stages. Mammal abundance
correlated with the number of shrub species (and no other plant variable)
and may have responded directly to shrubs for food or shelter. However,
shrub H' and cover cannot be eliminated as causes of the mammal pat-
tern, because significance levels did not permit modelling of possible
effects. The reason for low mammal abundance in the wax myrtle stage
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 107
is unknown. Bird species number and abundance stabilized and began
to decrease during the wax myrtle stage, and the decline continued as
the forest developed. The final stabilization of variables to be expected
in a climax community did not appear by year 36.
COMPARISON WITH OTHER STUDIES
Based on a compilation of pre-existing knowledge of the fish and wild-
life of the central Florida phosphate district, Layne et al. (1977) pre-
dicted general faunal changes likely to result from mining and reclama-
tion. They concluded that impacts of mining on important "species are
likely to be adverse . the overall effect will be a serious reduction in
both diversity and abundance of wildlife resources of the seven-county
area. Some of the deleterious impacts of mining could be reduced by
modification of current reclamation practices and priorities; however, the
net loss of fish and wildlife habitat and populations will still be of major
proportion." Our study suggests the same conclusions, but our interpre-
tation differs in one respect. How large a net loss of habitats and popu-
lations occurs depends strongly upon decisions about both reclamation
practices and subsequent land use. Our data indicate that the adverse
effects of mining can be temporary and that at least some fish and wildlife
resources can be restored after mining. Consequently our conclusions
emphasize the potential for restoration as determined by the new soils
and landforms and the actual restoration accomplished as determined by
post-mining land use decisions.
Recent work by Frohlich (1981) balances our results and those of Layne
et al. (1977). He showed that although small mammals were more abun-
dant in unreclaimed mines than in unmined flatwoods, some species were
lost in the conversion. As in our study, small mammals were more abun-
dant on unreclaimed sites than on reclaimed ones, and reclaimed and
sand tailings treatments were similar in supporting few small mammals.
In a baseline study on endangered species that might be impacted by
the proposed mining of the Osceola National Forest, the National Fish
and Wildlife Laboratory (1978: 272,274) concluded that "Phosphate ex-
traction by surface mining techniques will effectively extirpate all of the
local flora and fauna in the mined area. This impact will be total but, for
some species, need not be permanent if wildlife habitats are considered
in reclamation plans. Osceola National Forest could be mined for phos-
phate and returned to a system of wildlife habitats that, for many species,
would be equal to or improved over habitats now available." In the final
supplement to the final environmental statement on whether Osceola
mining leases should be approved, the basic scenario gave the reclama-
tion goal as "the restoration of the existing resource and reestablishment
BULLETIN FLORIDA STATE MUSEUM
of its multiple use, sustained-yield capabilities including sustained tim-
ber and forage production, wildlife and watershed management, and rec-
reation. Reclamation would .. be aimed at reestablishing .. the same
type of plant and aquatic communities with the same interspersion of
community types, i.e. the pine flatwoods, cypress swamps, creek swamps
and lakes .. (U.S. Bureau of Land Management 1979: 1-6). Though
the original statement emphasized as workable many specific reclama-
tion practices, the supplement stated (p. I-8) that "the present primitive
state-of-the-art of phosphate mine reclamation precludes the evaluation
of the potential impact of future mining and reclamation of the Osceola
by merely extrapolating present technology to future conditions. Re-
search is needed to provide the technological base required to restore to
[sic] the 'natural' system of the Osceola." Our study shows that establish-
ing some kind of forested wildlife habitat is certainly possible, but achieving
specifically targeted habitats that have never before been recreated will
require experimental engineering of soil structure and quality and of hy-
drologic regime. Some habitats, such as pine flatwoods over soil hard-
pan, may not be restorable in a reasonable framework of time or money.
BIOLOGICAL FACTORS AFFECTING VALUES AND USES OF MINE-
CREATED LANDS
Of the three distinct groups of treatments in this study (clay waste
areas, unreclaimed overburden, and reclamation with an overburden cap),
the latter two support valuable biological communities and have high
potential for natural or self-restoration. The clay waste areas appear to
form ecosystems with limited value for wildlife and for future land uses.
We have documented the successful, unsubsidized development of
hardwood forest on overburden spoil piles. The primary attribute of this
treatment is the soil itself. Overburden consists of about 80 percent sand
and 20 percent clay; it exceeds native soil in values for CaO, MgO, and
PO, but is deficient in K20 (Hawkins 1979). Importantly, very little ni-
trogen is present. The invading biota invests free services that enhance
succession, with plants rebuilding topsoil and wildlife dispersing seeds
to reintroduce and diversify the plants. The earliest plant colonization is
mainly of species with seeds strongly dispersed by autumn winds-dog
fennel, grasses, and Baccharis. Significantly, most invading woody spe-
cies have animal-dispersed seeds, whereas few species with water-dis-
persed or weakly wind-dispersed seeds become reestablished. An early
invader, wax myrtle, plays a special role by hosting a symbiotic root bac-
terium (Frankia) responsible for nitrogen fixation. This actinomycete cat-
alyzes the fixation of dinitrogen from the atmosphere and converts it to
ammonia (Torrey 1978). The nitrogen gives a great advantage to the pio-
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 109
neering wax myrtle, because the nitrogen fixed in root nodules is rapidly
transported for use in the aboveground portion of the plant. An annual
increment of nitrogen is returned to the soil as leaf litter, but nitrogen
stored in the wood is unavailable to other plants until the pioneer dies.
Aside from having its own supply of a scarce nutrient, wax myrtle inhibits
the success of competing plants through an allelopathic effect, shown
against the exotic Brazilian pepper and likely against other species as
well (Dunevitz and Ewel 1981). Wide and rapid dispersal is assured be-
cause wax myrtle provides an abundant supply of berries to migrant and
resident birds in autumn and winter. Perhaps the most spectacular ex-
ample is the tree swallow, with millions of individuals wintering in Flor-
ida. Tree swallows are regular users of unreclaimed sites with lakes, where
flocks of hundreds or thousands alternate their feeding between wax myrtle
berries and flying insects (over both water and land). Use of actinomy-
cete-nodulated plants in land reclamation has been recommended by
Silvester (1976). Another process possibly enhancing succession is the
introduction of spores of mycorrhizal fungi by colonizing rodents that
include fungi in their diets (Maser et al. 1978).
The forest community resulting on unreclaimed overburden is aes-
thetically attractive and supports a number of game species. The soil is
likely to be useful for many forms of profitable land use, including agri-
culture and silviculture. The number of uses of this mined land would
be increased by rounding off the spoil piles to produce gently rolling
terrain, and also succession would be speeded by reduced erosion. Sur-
facing the land with topsoil set aside at the beginning of mining probably
would speed the recovery process by supplying nitrogen and seeds, but
providing topsoil appears to be unnecessary if ample time is allotted for
natural recovery on overburden soil. The current regulatory criterion of
80 percent plant cover was exceeded on unreclaimed sites in about 10
years, as compared with the regulatory limit of 5 1/2 years after the ces-
sation of mining. If the piles were rounded to reduce slope erosion,
probably this level of plant cover could be achieved in 6-8 years without
investing in improved pasture.
Our documentation has shown that the reclaimed pastures are rela-
tively poor wildlife habitat. Nonetheless they are valuable as a prairie-
like monoculture, supporting abundant grass, insects, cotton rats, and
cows. Subsidies involved are leveling of spoil pile tops to form the sur-
face soil, establishing improved pasture, fencing, and animal husbandry,
plus periodic rehabilitative fertilizing and mowing. We have no data to
show whether ranching on this soil is competitive in terms of pasture
maintenance and livestock production on non-mined land, but the over-
burden cap appears to be a justifiable reclamation treatment with a val-
uable post-mining use. Furthermore, the overburden cap, if of sufficient
BULLETIN FLORIDA STATE MUSEUM
depth, should resemble unreclaimed spoils in supporting other agricul-
tural and silvicultural uses and native hardwoods. Hence the Kibler site
(Fig. 16), if not mowed, probably would develop a plant and wildlife
community like that on Bartow South (Fig. 12) or Old Clarke James (Fig.
15), differing only in the amount of lake area. Presumably the depth of
the overburden cap and, if shallow, whether underlain by clay waste or
sand, would be important determinants of whether pine plantations or
particular forest types would thrive. Presumably the overburden should
be about 1 m deep to support mesic-adapted trees with diffuse root sys-
tems, like slash pines and water oaks, and deeper for xeric-adapted trees
with taproots, like longleaf pines and live oaks. A subsoil of clay wastes
probably would result in a perched water table, which would affect the
surficial plant community on a relatively shallow cap.
Our study indicated that dewatered clay waste has only moderate
wildlife value and moderate potential for natural recovery. As active clay
settling ponds, these areas have very high wildlife value as marshes, and
they could be maintained as marshes by proper water level manipula-
tion. However, absorption of water by the clay creates a volume of waste
to be stored that can exceed the volume of mined pits, so at some mines
clay must be stored aboveground, behind dikes. Concern over polluting
clay spills from above-grade ponds prompts efforts to dewater the clay.
Invasion of consolidated clay by willows helps the process, because wil-
lows are phreatophytes with high rates of evapotranspiration. Willows
and the wax myrtles that replace them should enhance soil development
with surface litter and root material, but further succession is not appar-
ent. We found a very old clay waste area on Sanlan Ranch, with native
tree seed sources nearby, that had an open, senescent stand of large red
maple trees scattered in a vine-covered space. Probably the clay waste
areas begin to develop into swamp forests but eventually become too dry
and hard to survive.
The colloid under the clay crust will not support heavy buildings, so
settling areas cannot be used for residential or industrial development.
The soil is fertile, even in potassium (Hawkins 1979), and has proven
capable of growing pasture grass and row crops. The feasibility of culti-
vating consolidated clay has been demonstrated, but low ground-pres-
sure vehicles must be used. More typically, the clay is capped with sand
tailings and improved pasture is established on the cap soil. Because few
options exist for use of these aboveground lenses of hardening clay, top
priority should be placed on identifying and mandating optimal recla-
mation techniques. With a sand or overburden cap, clay lenses may de-
velop perched water tables and probably would favor the growth of swamp-
adapted trees like slash pines. From the standpoint of wildlife habitat, a
preferable way to use the clay and sand wastes might be to replace rela-
VOL. 30 NO. 3
1987 SCHNOES AND HUMPHREY: TERRESTRIAL COMMUNITIES IN FLORIDA 111
tively deep sand around the edge of the central clay-surfaced depression.
This linkage of an aquiclude and a body of water-storing soil should ex-
tend the hydroperiod from winter and summer rains to enhance the cen-
tral wetland while diversifying it with fringing uplands. A major im-
provement of the soil may be achieved if the logistic problems of mixing
sand tailings with clay waste during deposition can be solved.
FUTURE LANDSCAPES OF THE MINING REGION
Past landscapes of the region-dominated by extensive pine flatwoods
and high sand ridges or terraces-have been destroyed by mining and
will not be replaced. The ridges have been leveled and dispersed as
overburden. Low ground has lost the soil hardpan that had maintained
seasonally flooded forests. Though the former habitats and biota (Davis
1967, Layne et al. 1977) will not be replaced, our results show that other
forests and wildlife can be established on mine overburden. Our results
are relevant to the 46,868 ha in Florida mined for phosphate prior to
1975 (Phosphate Land Reclamation Study Commission 1978), but if rec-
lamation practices and subsequent land use decisions change, then there
is no assurance that the natural resource values we have documented
will pertain.
Indeed, new post-mining treatments have been established by a 1978
land reclamation law (Florida Statutes, Chapter 378) and current recla-
mation rules (Florida Administrative Code, Chapter 16C-16). Post-1975
spoil pile areas that do not include settling ponds or recirculating water
systems must be reclaimed, beginning within 18 months after mining
ceases and completed within 4 years thereafter. Consequently, spoil piles
will undergo little more succession than is documented for our Shark
Tooth site (Fig. 6). Clay settling ponds must be reclaimed, beginning
within 10 years after semi-liquids are no longer added to the area and
completed within 4 years thereafter. Consequently, consolidated clay will
be reclaimed at the stage of our Swift or N-2 sites (Fig. 1). Active clay
settling ponds thus will become the dominant transitional mining land
use because of their long periods of filling and dewatering-up to about
20 years, including the 10-year rule for reclamation. If spoil piles are left
in the active settling areas, terrestrial succession will result in islands of
habitat intermediate in maturity between our 5-15 year old (Figs. 8-9)
and 15-30 year old (Figs. 10-11) spoil pile sites, but presumably these
islands will be destroyed when the tops of the overburden piles are lev-
elled during reclamation. Because the trend of post-reclamation land-use
decisions is overwhelmingly toward ranching and agriculture, very little
of the good wildlife habitat lost to mining is being restored. The result of
these trends is that future wildlife values will be sharply truncated, oc-
BULLETIN FLORIDA STATE MUSEUM
curring mainly in the artificial freshwater marsh habitat of active and
inactive clay settling ponds. Wildlife use of this habitat is documented
by Wenner and Marion (1981), Maehr (1980), and Gilbert et al. (1981).
An even greater threat to both productivity and wildlife potential of
post-reclamation soils is suggested by long-term trends in the nature of
surface soils resulting from mining. Table 15 shows that in 20 years most
reclaimed surface may be sand instead of overburden, resulting in a loss
of both wildlife potential and productive agricultural uses. During the
1970s, the desirable overburden has been the dominant surface soil, av-
eraging 72.4 percent (with 1977 data missing). Moreover, the vast major-
ity of this reclaimed overburden has been deep soil. In the 1980's over-
burden surface soil is projected to continue to dominate reclaimed lands,
but most of it will be in the form of a relatively shallow cap over sand
tailings and/or clay. In the following two decades overburden will dimin-
ish to a small minority of the reclaimed surface. Contributing factors to
this pattern are the use of large amounts of overburden to construct set-
tling pond dikes and the lesser depth of overburden in new mines in the
southern part of the district. The change complementary to the decline
of overburden is the projected increase in sand tailings surface soil from
11.5 percent in the 1970s to 68.1 percent during 2000-09. If that soil is
a sand-clay mix, as proposed, the value of the reclaimed soil may be
quite high. However, if the mixing process continues to be unfeasible or
uneconomical, most reclaimed lands will be dewatered clay surfaced with
low-quality sand tailings.
The ability of sand tailings to support wildlife habitat, grazing, agri-
culture, and silviculture needs to be investigated. Small mammal com-
munities on early seral tailings are low in diversity and abundance (Mar-
ion et al. 1981). Fertility of sand tailings is relatively low. This soil consists
of about 99 percent sand and 1 percent clay; it is high in CaO and P0,,
but deficient in MgO and K20 (Hawkins 1979). Like other post-mining
soils, sand tailings also is very low in nitrogen. We have examined both
patchy and complete-cover pastures on sand tailings but do not know
whether pasture is difficult to establish or whether cattle stocking levels
need to be low to prevent overgrazing. A citrus planting on sand tailings
between Bartow and Winter Haven failed to survive (Hawkins 1979).
Successional patterns on sand tailings are unknown but are likely to lead
to a xeric plant association, like native longleaf pine savanna, sand pine
forest, or rosemary (Ceratiola) desert, or else to a scrub oak forest in the
absence of fire (Veno 1976).
INFORMATION GAPS
A substantial number of information gaps prevent a complete evalua-
tion of the capability of mined land to support wildlife populations and
VOL. 30 NO. 3
Table 15. Type of reclamation recently completed or approved for implementation in the central Florida phosphate district (as of
November 1979). The sequence of soil strata is shown from top to bottom. Mixed soil types are indicated by hyphens.
Reclamation sequence Relative area of reclamation types (percent and total hectares)
(top/bottom) 1971-4. 1975. 1976. 1978b 1979bc 1980-89 1990-99" 2000-09
Overburden (area includes some lake surface) 63.8 51.1 53.1 -- 65.9 31.7 8.1
Overburden/sand tailings 1.4 4.0 8.2 28.7
Overburden/sand tailings/clay -
Overburden/sand tailings-clay
Overburden/clay
Overburden-sand tailings
Overburden-sand tailings/natural ground
Overburden-sand tailings/clay
Subtotal
16.7
1.8 16.9
- -- 27.5 9.8
S 4.9 1.2
9.5 5.5 6.4 __ --
74.7 60.6 67.7 28.7 100.1
9.7 17.9 15.8
Clay/natural ground
Clay/sand tailings
Subtotal
Sand tailings
Sand tailings/clay
Sand tailings/clay/natural ground
Sand tailings-clay
Sand tailings-clay/overburden
Sand tailings-clay/natural ground
Subtotal
Peat/sand/tailings/clay
Total hectares
2.8 15.3 -- 5.5
12.5 33.2 15.8 5.5
12.9 6.3 12.6
S 65.8
Cr
n
z
7.5
7.5
4.9 45.6
-2.5
0.0 7.4 45.6
- 0.4
- 11.8
- 1.8
12.9 6.3 12.6 65.8
1565 1037 1742 229
37.9
S 1.5 --
0.0 15.5 37.9
0.7
982 4408 3704
O
z
68.1
68.1
2080
*Phosphate Land Reclamation Study Commission 1978.
hBased on release dates in records of the Bureau of Geology.
'Based on scheduled completion dates in records of the Bureau of Geology.
--
BULLETIN FLORIDA STATE MUSEUM
other productive uses in the future. Research begun on the successional
sequence of sand tailings and to relict native habitats (Marion et al. 1981)
should be continued. This would complete coverage of the array of exist-
ing post-mining treatments and would use natural areas both as controls
for gauging the success of natural reclamation and as indicators of what
wildlife resources have been lost in the mining/reclamation process. An-
other approach to evaluating the fish and wildlife to be lost in a proposed
mining area is to estimate the socioeconomic value of the resource from
a survey of recreational fish and wildlife activities; such a study has been
conducted by the Alberta Oil Sands Environmental Research Program
(1978). Management techniques for enhancing target game populations
on transitional clay settling ponds, unreclaimed land, and reclaimed areas
should be evaluated with field trials. Field trials also could be used to
test the role of fire in altering succession on consolidated clay, as sug-
gested by Breedlove and Adams (1977). Tests should be conducted to
combine consolidated clay settling areas with a complete or fringing sand
cap to create flatwoods or upland/wetland. Continued work is needed to
evaluate the amount and regional diversity of the fish and wildlife re-
source to be restored after mining.
Agronomic studies of reclaimed pastures with both overburden and
sand tailings soils should be conducted to learn what subsidies are needed
to support ranching and whether such ranchland is competitive with non-
mined rangeland. Forestry studies of both native tree communities and
commercial plantations should be done to evaluate the effect of re-
claimed soil composition, depth, and substrate on tree growth. The po-
tential for establishing longleaf pine savanna or sand pine forest on deep
sand tailings should be investigated.
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Contributions to the BULLETIN OF THE FLORIDA STATE MUSEUM,
BIOLOGICAL SCIENCES SERIES, may be in any field of biology. Manuscripts dealing with
natural history of systematic problems involving the southeastern United States or the New World
tropics are solicited especially. Manuscripts should be of medium length-circa 35 to 200 pages
(10,500-60,000 words). Examination for suitability is made by an Editorial Board.
The BULLETIN is distributed worldwide through institutional subscriptions
and exchanges. It is considered the responsibility of the author to distribute his paper to all inter-
ested individuals. To aid in this the authors) receives) 50 copies free, and he(they) may purchase
additional separates at cost if ordered when page proof is returned. The author is also responsible
for any charges incurred for alterations made by him on galley or page proofs. The Museum will
send an invoice to the author for this amount upon completion of publication.
PREPARATION OF MANUSCRIPT
Contributors should consult recent numbers of the BULLETIN for preferred
style and format. Highly recommended as a guide is the CBE Style Manual, 3rd Edition, 1972
(American Institute of Biological Sciences, Washington, D.C.).
Manuscripts must be submitted in duplicate and satisfy the following minimal
requirements. Please submit duplicate copies of manuscripts. They must be double-spaced
throughout, including tables, figure captions, and literature citations. Figure legends and tables
should be typed on separate sheets. Also, please submit a copy of the complete text and tables on
one or more 514" flexible diskettes from (1) IBM or compatible microcomputer with Microsoft
WORD; or (2) Apple microcomputer with Applewriter submit document files. If that is not
possible, the editors will have such diskettes prepared commercially at author's expense upon ac-
ceptance of the manuscript.
All illustrations are referred to as figures. They must comply with the following
standards: Photographs should be sharp, with good contrast, and printed on glossy paper. If the
background of photographs (especially those of specimens) is not desired, amberlith should be cut
out and used to cover the background. Drawings should be made with dense black waterproof ink
on quality paper or illustration board. All illustrations should have a cover sheet. All lettering will
be medium weight, sans-serif type (e.g. Futura Medium, News Gothic) in cutout, dry transfer, or let-
tering guide letters. Make allowance so that after reduction no lower case letter will be less than 1
mm high (2 mm is preferred) nor any capital letter greater than 5 mm high. The maximum size for
illustrations is 9" x 14" (twice BULLETIN typepage size); illustrations should not be less than type-
page width (4Vz"). With soft lead pencil on the back of each illustration, designate the top and iden-
tify each by author's name, manuscript title, and figure number.
All manuscripts not submitted in BULLETIN format will be returned to the
author for retyping.
Manuscripts and all editorial matters should be addressed to:
Managing Editor of the BULLETIN
Florida State Museum
University of Florida
Gainesville FL 32611
U.S.A.
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