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22001155 SOUTHEASTERN NATURALIST 1V4o(3l.) :1541,7 N–5o5. 03
The Ichthyofauna of Big Cypress National Preserve, Florida
Marcus Zokan1,*, Greg Ellis2, Shawn E. Clem3, Jerome Lorenz4, and
William F. Loftus5
Abstract - The Big Cypress Swamp (BCS) is a large freshwater wetland system and drainage
basin (640,000 ha) in southwest Florida and an important component of the Greater
Everglades ecosystem. Despite its size and relationship to the Ever glades, the fish fauna of
BCS has received little study. Documentation of its fish fauna is important to better understand
this dynamic natural system and to monitor changes to the fish community, including
the spread of non-indigenous species. To that end, we surveyed the ichthyofauna of freshwater
habitats in Big Cypress National Preserve (BCNP), the largest and most intact wetland
area (295,000 ha) remaining in BCS. Between October 2002 and May 2004, we recorded 63
fish species from freshwater habitats in BCNP, including 9 non-indigenous species. Species
richness was greatest in permanent freshwater habitats and lowest in shallow temporary
wetlands and seasonally fresh coastal marshes. The most speciose families were the native
Centrarchidae (8 spp.) and the non-native Cichlidae (6 spp.), whereas the most abundant and
widely distributed species were members of Cyprinodontidae, Fundulidae, and Poecilidae.
Similar to other coastal drainages of southern Florida, BCNP has a relatively high occurrence
of euryhaline species (28 spp.).
Introduction
The Big Cypress Swamp (BCS) is a large wetland complex and drainage basin
that forms a major component of the Greater Everglades ecosystem and has a
hydrologic boundary encompassing ~640,000 ha of southwestern Florida (Fig. 1;
McPherson and Halley 1996). Its name refers to the extensive Taxodium distichum
(L.) Rich. (Bald Cypress) forests that characterize the landscape, although a variety
of other forested and herbaceous wetlands are present (Davis 1943, Muss et
al. 2003). BCS is unusual among cypress-dominated wetlands in having slightly
alkaline, high-conductivity waters (Carter et al. 1973). The rainfall pattern is highly
seasonal, resulting in a landscape inundated for a portion of the year and dry for
the remainder, with few permanent water bodies (Klein et al. 1970). Due to the flat
topography, BCS is hydrologically continuous with the Everglades to the east, permitting
movement of fishes between these 2 basins, although Kushlan and Lodge
(1974) stated that there may be faunal differences between them. Extensive canal,
ditch, and borrow-pit excavation in the 1900s accelerated drainage in some portions
of BCS and increased connection to other drainage basins in southern Florida.
These alterations have likely changed the fish fauna, but the lack of data has made
1Odum School of Ecology, University of Georgia, Athens, GA 30602. 2College of Marine Science,
University of South Florida, St. Petersburg, FL 33701. 3Audubon Florida, Corkscrew
Swamp Sanctuary, 375 Sanctuary Road West, Naples, FL 34120. 4Everglades Science Center,
Audubon Florida, 115 Indian Mound Trail, Tavernier, FL 33070. 5Aquatic Research and Communication
LLC, Vero Beach, FL 32967. *Corresponding author - zokanm@gmail.com.
Manuscript Editor: Nathan Dorn
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evaluation of these changes difficult. Given the dynamic nature of the natural system
and the anthropogenic changes to it, documentation of BCS’s fish fauna and its
seasonal changes is of importance to provide a baseline for assessing future changes
and their impacts.
Big Cypress Swamp is defined on its northern margin as waters flowing generally
southward off the more elevated Immokalee Rise, on the east by the transition
from Bald Cypress forest to the Everglades marshes, on the west by coastal uplands
near Naples, and to the south by the estuarine wetlands of the Ten Thousand Islands
(Lodge 2004). Some distinct regions within BCS include Fakahatchee Strand,
Picayune Strand, southern parts of Okaloacoochee Slough, and Corkscrew Swamp
(Fig. 1). Corkscrew Swamp is sometimes considered separate from BCS (e.g., Davis
1943, Klein et al. 1970), but is included here due to hydrologic connections and
similar habitats (Lodge 2004, McPherson and Halley 1996). Much of the northern
and western portions of BCS have been altered by drainage, agriculture, and urban
development (Carter et al. 1973, Duever et al. 1986); however, extensive areas of
Figure 1. Map of southern Florida showing the location of Big Cypress Swamp and important
areas within and adjacent to it, including Big Cypress National Preserve (BCNP),
Corkscrew Swamp (CS), Okaloacoochee Slough (OK), Florida Panther National Wildlife
Refuge (FP), Picayune Strand State Forest (PS), Fakahatchee Strand Preserve State Park
(FS), Everglades National Park (ENP), Water Conservation Area 3 (WCA3), Big Cypress
Indian Reservation (SR), and Miccosukee Reservation (MR).
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BCS have been preserved as public conservation lands, with the largest portion comprising
the 295,000-ha Big Cypress National Preserve (BCNP; Muss et al. 2003).
BCNP encompasses most of the eastern half of BCS and covers eastern Collier
County, part of northern Monroe County, and a narrow strip of far-western Miami–
Dade County. Big Cypress National Preserve is bordered by multiple public
conservation lands including Water Conservation Area 3 and Everglades National
Park to the east and south, and by Fakahatchee Strand Preserve State Park and
Florida Panther National Wildlife Refuge to the west (Fig. 1). BCNP also borders
Big Cypress Indian Reservation to the north and the Miccosukee Reservation to
the northeast, both of which contain some conservation land; the remainder of the
northern border abuts private land (Fig. 1). Although many studies have examined
the distribution, species composition, and seasonal dynamics of fishes within other
components of the Everglades system (e.g., DeAngelis et al. 2010, Loftus 2000,
Parkos et al. 2011, Trexler et al. 2002), ichthyological studies in BCS have been
rare. We attempted to fill that gap by providing the first comprehensive survey of
the ichthyofauna of freshwater habitats in BCNP, which is the largest, mostly intact
wetland habitat remaining within BCS.
The earliest published collections of freshwater fishes within BCS were by
Fowler (1915, 1926) from the northern fringes of BCS. In 1938 and 1941, Fowler
made additional collections in the vicinity of Ochopee, FL (Fowler 1945). Kushlan
(1972) recorded fish species from a single pond in the eastern portion of BCS,
Carter et al. (1973) collected fish from 5 sites within Fakahatchee Strand, and
Kushlan and Lodge (1974) summarized information on freshwater fish species occurring
throughout southern Florida, including BCS. Following the establishment
of BCNP in 1974, the most extensive collections within the region were by Loftus
and Kushlan (1987), who sampled 39 sites within the southern portion of BCNP.
More recently, Dunker (2003) collected fishes within Big Cypress Seminole Reservation
to the north of BCNP and Addison et al. (2006) and Ceilley (2008) collected
in the Picayune Strand to the west. In addition, Kahl (1964) and Carlson and Duever
(1977) provided information on seasonal population dynamics of fishes within
Corkscrew Swamp Sanctuary in the northwestern region of BCS. Subsequent to the
present study, a long-term monitoring program was implemented at 3 sites within
BCNP (Liston and Lorenz 2008). Apart from Loftus and Kushlan (1987), previous
surveys and sampling studies were spatially restricted and the numbers of samples
were generally low. In addition, few studies attempted to sample throughout a full
seasonal cycle. The present study arose from the National Park Service’s need for
data on the distribution and abundance of fishes in BCNP, particularly the vast
inland areas that remained unsurveyed, and on the expanding populations of nonindigenous
fishes in and near BCNP (Kline et al. 2013).
The primary objectives of our study were to document all fish species present
within BCNP and to compare differences between species assemblages by habitat
and season. We also combined data from our study and other studies to determine
the fish fauna of BCS as a whole. Given the lack of prior information on the species
present within BCS, we provide a comparison of BCS with the fish faunas of other
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major drainages of southern Florida to examine whether BCS represents a distinct
zoogeographic unit as proposed by Kushlan and Lodge (1974). Finally, we compare
the fish fauna of BCS with other major swamp ecosystems in the southeastern US
to examine whether there are any parallels among the fish species present in large
forested wetlands because they are characterized by a unique set of environmental
conditions not present in other aquatic habitats.
Field-site Description
BCNP experiences a subtropical climate with an average yearly precipitation
of 135 cm; it is subject to a strong wet/dry seasonal cycle, with nearly 80% of the
annual rainfall occurring between May and October (Klein et al. 1970). During
the wet season, water covers much of the land surface, whereas during the dry season
aquatic habitats are reduced to isolated wetlands. Water levels rise rapidly at
the beginning of the wet season (early summer) and remain high before declining
in the fall; water levels are lowest in spring, when most wetlands dry completely
(Duever et al. 1986). Natural, permanent freshwater is relatively scarce and occurs
primarily as small ponds interspersed within more extensive wetlands. In addition,
a few coastal freshwater streams that drain the southern portion of BCNP flow all
year (Duever et al. 1986). Construction of canals, ditches, and borrow ponds have
increased the extent of permanent freshwater habitat in BCNP (Duever et al. 1986).
Natural and artificial permanent water-bodies serve as dry-season refuges for fishes
and other aquatic species.
The natural hydrology of BCNP has numerous connections to other portions of
BCS and to other components of the Greater Everglades ecosystem. The hydrology
of the central and southern regions of BCNP is largely driven by local rainfall,
with water draining south and southwest toward the Gulf of Mexico overland and
through elongated forested wetlands called strands (Klein et al. 1970). The northwestern
corner of BCNP receives water from Okaloacoochee Slough, which gathers
water from agricultural, ranch, and undeveloped land in northern Collier and western
Hendry counties. Okaloacoochee Slough is a major water input to the adjacent
Fakahatchee Strand and also adds water inputs to strands flowing through western
BCNP (Klein et al. 1970, Lodge 2004). The northern and northeastern portions of
BCNP receive water from agricultural, ranch, and undeveloped land in southern
Hendry County, which flow south and southeast. These waters continue southeastward
across northern BCNP and enter Water Conservation Area 3 and eventually
Everglades National Park (Klein et al. 1970). Some of these waters reenter BCNP
in the southeastern corner and flow southwest to the Gulf of Mexico (Lodge 2004).
These natural linkages should allow fishes to move readily between BCNP, the
western BCS, and the Everglades.
Human-caused changes to the hydrology of BCNP include the construction of
several major canals that have increased the drainage of wetland habitats, and serve
as permanent deep-water refuges and movement corridors for fishes (Fig. 2). The
first major canal constructed was the Barron River Canal, which was built parallel
to State Road 29 from Immokalee to Everglades City and forms the western border
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of BCNP. The most important east–west canal is the Tamiami Canal, which bisects
the southern portion of BCNP and directly connects to the Everglades and the canal
system of southeastern Florida. The L-28 canal forms part of the eastern border
between BCNP and water conservation area 3, whereas the L-28 Interceptor canal
cuts across the northeastern corner of BCNP and connects to the canal system that
drains the agricultural lands south of Lake Okeechobee. Other canals include the
borrow canals paralleling Birdon Road, Loop Road, Turner River Road, Wagon
Wheel Road, and Interstate 75. These canals connect BCNP with the larger canal
system of southern Florida, which has allowed movement of fishes between drainage
basins and may facilitate the spread of non-indigenous species (Kline et al.
2013, Shafland et al. 2008).
Habitat types
As its name suggests, BCNP is dominated by Bald Cypress forest; however,
there is a diverse array of aquatic habitats present. Habitat designations used during
this study were modified from Duever et al. (1986) based on hydroperiod (the number
of days per year a wetland is inundated) and dominant vegetation as follows:
Freshwater wet prairie—a short-hydroperiod wetland (inundated 50–150
days/year) on carbonate marl or sand substrate dominated by various
grasses and sedges, especially Cladium jamaicense (Crantz) Kük.
(Jamaica Swamp Sawgrass), Muhlenbergia capillaris (Lam.) Trin.
(Hairawn Muhly), and Rhynchospora spp. (beaksedges).
Freshwater marsh—a medium- to long-hydroperiod wetland (inundated
225–275 days/year) on peat soils usually located within wet prairies
and dominated by various grasses and sedges, particularly Panicum spp.
(panic grasses) and Eleocharis spp. (spikerushes), as well as Pontederia
cordata L. (Pickerelweed), Sagittaria spp. (arrowheads), Thalia geniculata
L. (Bent Alligator-flag), and Salix caroliniana Michx. (Coastal
Plain Willow).
Coastal marsh—a medium-hydroperiod wetland (inundated 150–250 days/
year) that varies seasonally between fresh and brackish water and is
dominated by Spartina spp. (cordgrasses), spikerushes, Juncus roemerianus
Scheele (Needlegrass Rush), Typha domingensis Pers. (Southern
Cattail), and Distichlis spicata (L.) Greene (Saltgrass). Coastal marshes
occur on the southern border of the BCNP and provide a transition into
estuarine habitats.
Cypress savanna—a short-hydroperiod wetland (mean inundation 120 days/
year) similar to wet prairie, but with a sparse canopy of dwarfed Taxodium
ascendens Brongn. (Pond Cypress). It covers large expanses of
eastern BCNP, but also occurs at the ecotone of wet prairie and cypress
forest.
Cypress forest—medium- to long-hydroperiod wetland (mean inundation
250 days/year) with a dense canopy of medium to tall Pond Cypress and
Bald Cypress; the understory usually consists of ferns and herbaceous
vegetation.
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Mixed hardwood swamp—a long-hydroperiod wetland (mean inundation
290 days/year) similar to cypress forest, but with an extensive subcanopy
of Acer rubrum L. (Red Maple), Annona glabra L. (Pond Apple)
and Fraxinus caroliniana Mill. (Carolina Ash).
Rivers and creeks—permanent flowing water bodies bordered by forested
wetland habitats in their upper reaches and coastal marsh and Rhizophora
spp. (mangrove) forests in their lower reaches; these waterways
constitute a direct link between estuarine waters and inland freshwater
wetlands. Turner River is the most notable example, and was the only
one sampled.
Freshwater canals—artificial permanent habitats upstream from salinity barriers
and frequently connected to freshwater wetlands.
Coastal canals—artificial permanent habitats downstream of salinity barriers;
they are connected to estuarine habitats and vary seasonally in
salinity between fresh and brackish water.
Freshwater ponds are permanent water bodies usually located within long-hydroperiod
wetlands, and also present in BCNP. We did not sample the open portions
of ponds, but occasionally sampled pond edges. We did not treat ponds as a separate
habitat type, but categorized them by the surrounding habitat; it should be noted
that there can be quantifiable differences in fish-community structure between
ponds and surrounding wetlands (Parkos et al. 2011)
Methods
Gear types
To sample fishes, we used a variety of gear types similar to those used by Loftus
and Kushlan (1987) in their survey of southern BCNP. These included several types
of fish traps, gillnets, dip nets, cast nets, baited hoop-nets, angling, and electrofishing;
however, we used traps most extensively because of their portability and ease
of use in shallow, vegetated habitats. Traps used included Gee-type minnow traps,
metal box-traps, collapsible nylon-mesh minnow traps, and Breder traps (Breder
1960). We deployed unbaited traps for 24-h periods, except for a series of samples
from freshwater canals that we fished for 1-h intervals. Each trap type may have its
own selection bias; thus, we sampled each site using a combination of trap types to
reduce potential sampling-bias issues.
Electrofishing equipment included a boat-mounted Smith-Root Type-6A electrofisher
(Smith-Root, Vancouver, WA) for canals and a barge-mounted Type-2.5
GPP electrofisher (Smith-Root) for use in wetlands. We used the boat-mounted unit
(1008 v DC max at 120 pulses per second) on 100-m transects along canal margins.
The barge-mounted unit was capable of a maximum current of 1000 v at either 120
pulses per second DC, or at 60 hertz AC; sampling was standardized by 300 seconds
of shock time.
We used dip nets (1-mm mesh) alone or in addition to other gear types to collect
small species. Whether used alone or with other gear, we dip-netted at a site
until a 10-min interval passed without adding a new species. To target catfishes, we
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deployed cheese-baited hoop nets in canals for 24-h intervals. We used cast nets,
gillnets, and angling opportunistically to collect species missed by other gear types
and as a supplement to the primary gear. We recorded visual observations when
specimens could not be captured. We deposited voucher specimens in the Florida
Museum of Natural History Ichthyology Collection in Gainesville, FL.
Sampling and Analysis
We conducted sampling between October 2002 and May 2004. Most sampling
sites were located within 250 m of roads (Fig. 2) due to the impracticality of
carrying sampling gear long distances. We visited more distant sites by hiking,
boat, or ATV when practical, and by airboat and helicopter when these modes of
transport were available. We stratified sample sites by habitat type and selected
them non-randomly based on water availability. We designated freshwater wet
prairie, freshwater marsh, cypress savanna, cypress forest, and hardwood swamp
as freshwater-wetland habitats to distinguish them from the permanent, artificial,
or seasonally brackish environments of Turner River, canals, and coastal marsh. We
separated samples from wetland habitats into wet season (15 May–30 November)
Figure 2. Map of
Big Cypress National
Preserve (BCNP)
showing the locations
of sample
sites (filled circles)
and important geographical
features
(1 = Barron River
Canal, 2 = Wagonwheel
Road canal,
3 = Birdon Road
canal, 4 = Turner
River Road canal,
5 = Turner River,
6 = Tamiami Canal,
7 = Loop Road canal,
8 = L-28 canal,
9 = L-28 Interceptor
canal, 10 = I-75
canal, and star =
Deep Lake). Canals
and Turner River
are marked by thick
black lines.
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and dry season (1 December–14 May) to examine assemblage differences. We did
not include seasonal designations for Turner River, coastal canal, and coastal marsh
due to small sample sizes, or for freshwater canal because most samples were from
the dry season when wetlands lacked water. Our primary goal was to inventory all
fish species present within BCNP. Thus, we did not attempt to create a balanced
sampling scheme regarding habitat type or season, and sample sizes among habitat
types were unequal but somewhat proportional to the area of the habitat type within
BCNP, with the exception of freshwater canal and freshwater marsh. Sample sizes
were also uneven within a habitat type between seasons because water availability
limited when we could sample habitat types. Since a primary goal of the inventory
was to increase sample coverage across BCNP, we rarely sampled the same site
more than once.
In an effort to collect all species present, we used a variety of gear at most
sites. A sampling event usually consisted of deploying a variety of trap types, up
to 2 traps each of the 4 listed types noted above for a 24-h period. We placed each
trap 2–5 m from adjacent traps in patches of vegetation, fallen woody debris, or
gaps within dense emergent vegetation. After processing trap catches, we used
dip nets to sample species not captured with the trap gear. In canals, we also utilized
cast nets and angling to target species not taken with the other gear. Sites at
which we employed trap gear usually were less than 100 m2.
We also established electrofishing and dip-net sample sites. A canal-electrofishing
site consisted of a single 100-m transect, whereas a wetland-electrofishing site
consisted of an area shocked for an interval of 300 seconds. Dip-net sites were sites
where we only used dip nets; these were often remote locations where the transport
of other gear types was impractical.
We defined frequency of occurrence as the proportion of sample sites within
each habitat type (and season for wetland sites) at which a species was collected.
The sample sites used in frequency analysis were distributed by habitat and season
as follows: coastal marsh (n = 10), coastal canal (n = 16), freshwater canal (n =
81), Turner River (n = 11), hardwood-swamp dry season (n = 18), hardwood-swamp
wet season (n = 15), cypress-forest dry season (n = 69), cypress-forest wet season
(n = 17), cypress-savanna dry season (n = 9), cypress-savanna wet season (n = 13),
wet prairie dry season (n = 7), wet prairie wet season (n = 28), freshwater-marsh dry
season (n = 38), and freshwater-marsh wet season (n = 21) (Table 1). We used sites
sampled solely with hoop nets, gill nets, cast nets, or angling strictly to determine
species presence within a habitat. We excluded these sites from frequency calculations
because their use was not standardized and they were often used to target
specific species and not to characterize the whole fish community .
We evaluated sample data with cluster analysis and non-metric multidimensional
scaling (NMDS) to determine spatial (habitat) and temporal (seasonal)
differences in the fish community (Clarke 1993, 2006). We employed Bray-Curtis
similarity to calculate a distance matrix from a frequency-of-occurrence data table
that consisted of the frequency of occurrence for each species by habitat type and
by season within habitat type for wetland habitats. We then used the distance matrix
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to create a group-linkage cluster dendrogram. We selected cut-off values of 50%,
65%, and 75% similarity from the cluster analysis because they encompassed the
most well-defined groupings. We ran NMDS on the similarity matrix to create a
visual plot of the distances between groupings and drew the cut-off values selected
from the cluster analysis on the plot to encompass groups of similar assemblages.
Although we relied largely on passive traps for our fish-community analyses,
they are not ideal for sampling the entire fish community. Studies by Obaza et al.
(2011) in the Everglades found that considerable differences exist in the capture
and retention of different species of fish within the region. The capture of a species
using trap gear depends on encounter rates; therefore, detection was sporadic for
rare species or species that do not readily enter traps.
Throughout this study, we defined freshwater-fish species as primary and secondary
freshwater fishes, whereas we used euryhaline to describe species that are
marine in derivation, but may enter fresh water transiently, or may spend part or all
of their lifecycle in fresh water.
Comparison to regional drainages
We compared the ichthyofauna of BCS to fish communities inhabiting regional
drainages to examine zoogeographic patterns. We combined our data from BCNP
with published and unpublished records from other portions of BCS and compared
the combined BCS fish fauna to those of 8 other drainage areas (Fig. 3) including
the Kissimmee River drainage, Lake Okeechobee and Fisheating Creek, Charlotte
Harbor drainage, the Caloosahatchee River, the northern Everglades (Everglades
Agricultural Area and Water Conservation Areas), the southern Everglades (Everglades
National Park), the southern Indian River drainage (St. Sebastian River
to the Loxahatchee River), and the Atlantic coastal drainages south of the Loxahatchee
River (now primarily canals). Historic species distributions were difficult
to assess because few collections were made prior to widespread canal construction
and drainage activities (Loftus and Kushlan 1987, McVoy et al. 2011). Modern
distributions and abundance patterns, as well as habitat preferences, have been
helpful in this regard. We used the following categories to assess species distributions:
primary or secondary freshwater species native to drainage (N-f), primary or
secondary freshwater species for which native status to drainage is uncertain (U-f),
euryhaline species collected in freshwater and native to drainage (N-e), euryhaline
species collected in freshwater for which native status to drainage is uncertain
(U-e), euryhaline species native to drainage, but not recorded from freshwater (e*),
and non-indigenous species (X). We categorized species as uncertain if their native
status within a drainage was questionable. We employed the uncertain designation
in several instances. For example, we assigned euryhaline species that occurred in
the Kissimmee River an uncertain native status because the drainage was not connected
to marine waters prior to canal construction. Additionally, we designated as
uncertain species that occurred primarily in canals because that habitat is a relatively
new construct on the landscape. The non-indigenous species included were
those we considered to be reproducing in southern Florida according to Shafland
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et al. (2008), Schfield and Loftus (2014) and W.F. Loftus (pers. observ.). We also
included in the non-indigenous group Ctenopharyngodon idella (Grass Carp) because
it is regularly stocked in many areas, although it is not known to reproduce
Figure 3. The major freshwater drainages of southern Florida.
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in southern Florida (Shafland et al. 2008). We performed cluster analysis on Bray-
Curtis similarities calculated from presence/absence of native species within the
different drainages.
Comparison of swamp ecosystems
Fish family-level comparisons were made between BCS and other large swampwetland
systems of the southeastern US including Great Dismal Swamp, VA and
NC (Jenkins and Burkhead 1994, Jenkins et al. 1975), Okefenokee Swamp, GA
(Freeman et al. 1984, Laerm et al. 1980), and the lower Mississippi floodplain
swamps (Baker et al. 1991, Bart et al. 1998, Guillory 1979). We separated the Mississippi
floodplain-swamp species list into permanent residents of floodplain ponds
and species that were present only seasonally during annual flooding events (Baker
et al. 1991). Non-native species were omitted from the swamp comparisons.
Results
We collected or observed 63 species of fishes representing 31 families including
9 non-indigenous species (Appendix 1). The most species-rich habitat type was
freshwater canal, from which we recorded 47 species. By contrast, the most depauperate
habitats were coastal marsh (20 species), and the short-hydroperiod cypress
savanna and freshwater wet prairie (21 and 22 species, respectively). We recorded
a total of 35 species from all freshwater wetlands combined.
We found 5 species—Gambusia holbrooki (Eastern Mosquitofish), Lucania
goodei (Bluefin Killifish), Lepomis marginatus (Dollar Sunfish), Fundulus chrysotus
(Golden Topminnow), and Poecilia latipinna (Sailfin Molly)—in at least 50%
of all sites from which frequency was calculated (Appendix 1). The only non-indigenous
species that occurred in >50% of sites within any freshwater wetland habitat
were Belonesox belizanus (Pike Livebearer), from both wet prairies and cypress
savannas, and Cichlasoma bimaculatum (Black Acara), from wet-season samples
in hardwood swamp and freshwater-marsh habitats. Two additional non-indigenous
species were present in >50% of samples within other habitat types—Cichlasoma
urophthalmus (Mayan Cichlid) in freshwater canals, coastal canals, and coastal
marshes, and Tilapia mariae (Spotted Tilapia) in both freshwater canals and the
Turner River.
The 2 most speciose families were Centrarchidae and Cichlidae, members of
which were most frequently collected in freshwater canals. Among centrarchids,
Enneacanthus gloriosus (Bluespotted Sunfish), Lepomis gulosus (Warmouth),
Dollar Sunfish, and L. punctatus (Spotted Sunfish) were common in all freshwater
wetland types. The cichlids Black Acara, Mayan Cichlid, and to a lesser degree,
Spotted Tilapia, were also frequent in wetland habitats.
Most native freshwater species had been documented during earlier studies;
however, our survey added 3 previously unrecorded species to the fauna: Esox
americanus (Grass Pickerel), Noturus gyrinus (Tadpole Madtom), and Pomoxis
nigromaculatus (Black Crappie). We collected 1 Grass Pickerel specimen during
this study; this species was only recently recorded from the southern Everglades
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(Chick et al. 2004, Kline et al. 2013), although it is well known from the northern
Everglades (Dineen 1984). We encountered Tadpole Madtom uncommonly, but
this fish was widespread within BCNP wetlands. We collected Black Crappie only
from the L-28 Interceptor canal near the northeastern border; this species probably
did not occur in BCNP prior to the construction of deep-water canals. We recorded
2 non-indigenous species from BCNP for the first time: Hoplosternum littorale
(Brown Hoplo) and Hemichromis letourneuxi (African Jewelfish).
We collected most euryhaline species from coastal marshes and coastal canals
below salinity barriers, although a few ranged more widely. Several ascended
Turner River into its upper reaches, and a few such as Anguilla rostrata (American
Eel), Centropomus undecimalis (Common Snook), Megalops atlanticus (Tarpon),
and Mugil cephalus (Black Mullet) moved up to 50 km inland via the canal system.
Notably, populations of Common Snook, Tarpon, and Black Mullet occurred in
Deep Lake, a 29-m-deep sinkhole (Hunt 1958) within a cypress swamp connected
to Barron River canal and located >14 km from the nearest brackish water; the presence
of Tarpon in Deep Lake was first reported by Kushlan and Lodge (1974). Of
the 3 euryhaline species we collected in freshwater wetlands, Marsh Killifish and
Sailfin Molly were both widespread and abundant, whereas Cyprinodon variegatus
(Sheepshead Minnow) was restricted to the southern portion of BCNP.
Habitat differences
Coastal canals and coastal marshes had a high number of euryhaline species (Appendix
1) and formed a species assemblage distinct from those inhabiting waters that
remain fresh throughout the year (Fig. 4). There was also an important distinction
between species assemblages found in permanent freshwater (freshwater canal and
Turner River) versus impermanent freshwater habitats (hardwood swamp, cypress
swamp, cypress savanna, freshwater marsh, and wet prairie) (Fig. 4). The permanent
freshwater habitats had some euryhaline species and also species that were infrequent
or absent in impermanent freshwater habitats (Appendix 1). Overall, species
assemblages in impermanent freshwater habitats were quite similar to one another.
We identified 4 assemblages from the results of our analyses: dry-season hardwood
swamp, dry-season cypress savanna, a group comprised of wet-season
cypress swamp and hardwood swamp, and a group consisting of the remainder of the
habitats sampled (Fig. 4). The latter group was most interesting because it included
both short-hydroperiod wet-season samples (wet prairie and cypress savanna) and
long-hydroperiod dry-season samples (freshwater marsh and cypress swamp), suggesting
that the assemblage may shift from short-hydroperiod habitats in the wet
season to long-hydroperiod habitats in the dry season.
Comparison to regional drainages
A total of 155 fish species have been recorded from freshwater in southern Florida,
including 34 established non-indigenous species (Appendix 2). The greatest
species richness of native freshwater fishes occurred in the Kissimmee River Basin
with 39 known species, whereas the southeastern coast had the lowest richness
with only 22 species. The only other drainages that exceeded 30 native freshwater
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2015 Vol. 14, No. 3
species were Charlotte Harbor and Lake Okeechobee. When we included species
of uncertain native status (native to Florida, but perhaps not to the drainage), the
northern Everglades, southern Everglades and the southern Indian River drainages
had >30 species. In considerations of native species and those of uncertain status,
BCS was relatively low in species richness, with only 26 native freshwater species
and 3 species of uncertain native status. In contrast, the richness of non-indigenous
species was greatest in the southeastern coastal drainages, with 28 species currently
reproducing, and lowest in the Kissimmee River basin with 7 species. Cluster analysis
of native species revealed 2 major groupings among the drainages: an inland
cluster including Kissimmee River, Lake Okeechobee, and northern Everglades;
and a coastal cluster with the remaining drainages (Fig. 5). The main difference
between the 2 groups was the greater presence of euryhaline species in drainages
directly connected to marine waters.
Figure 4. The major fish-species assemblages within BCNP by habitat and season displayed
on a non-metric multi-dimensional scaling (NMDS) plot. Bray-Curtis similarities
were calculated from a matrix of frequency of occurrence for each species and analyzed
through cluster analysis. The frequency matrix was also used for NMDS to visually display
distances between assemblages. Cut-off values were chosen from the cluster analysis and
circled on the NMDS plot. Circles group habitat types that have Bray-Curtis similarities
on or above the level given in the figure legend. Habitat types are as follows: CM = coastal
marsh, CC = coastal canal, FC = freshwater canal, TR = Turner River, HD = hardwood
swamp dry season, HW = hardwood swamp wet season, CD = cypress forest dry season,
CW = cypress forest wet season, SD = cypress savanna dry season, SW = cypress savanna
wet season, PD = wet prairie dry season, PW = wet prairie wet season, MD = freshwater
marsh dry season, and MW = freshwater marsh wet season.
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There were several notable patterns in species distribution that may be zoogeographically
relevant. Three native species—Elassoma okefenokee (Okefenokee
Pygmy Sunfish), Lepomis auritus (Redbreaset Sunfish), and Percina nigrofasciata
(Blackbanded Darter)—reached the southern limit of their distribution within the
Kissimmee River drainage (Appendix 2), whereas several additional native species
extended from the Kissimmee River Basin to Lake Okeechobee and the Charlotte
Harbor drainages, including Lepisosteus osseus (Longnose Gar), Dorosoma
cepedianum (American Gizzard Shad), Notropis chalybaeus (Ironcolor Shiner),
Opsopoedus emiliae (Pugnose Minnow), Ameiurus catus (White Bullhead), Ictalurus
punctatus (Channel Catfish), Grass Pickerel, Chain Pickerel (not recorded
in Charlotte Harbor drainages), Aphredoderus sayanus (Pirate Perch), and Black
Crappie (Appendix 2). These species, with the exception of White Bullhead, Ironcolor
Shiner, and Pugnose Minnow, have been recorded as far south as the southern
Everglades (Kline et al. 2013, Loftus 2000); however, records are few, scattered,
and mainly from canals, leaving their native status in these areas in question.
Additionally, Fundulus lineolatus (Lined Topminnow), Fundulus rubrifrons (Redface
Topminnow), and Leptolucania ommata (Pygmy Killifish) had rather patchy
distributions, and were absent from large areas of southern Florida (Appendix 2).
A notable pattern for many native euryhaline species was their primarily Atlantic
coastal distribution in southern Florida; among those species rarely recorded or
absent from the southern Gulf coast were Microphis lineatus (Opossum Pipefish),
Centropomus ensiferus (Swordspine Snook), C. parallelus (Fat Snook), C. pectinatus
(Tarpon Snook), Pomadasys crocro (Burrow Grunt), Agonostomus monticola
(Mountain Mullet), Eleotis amblyopsis (Large-scaled Spinycheek Sleeper), Erotelus
smaragdus (Emerald Sleeper), Gobiomorus dormitor (Bigmouth Sleeper),
Awaous banana (River Goby), Ctenogobius pseudofasciatus (Slashcheek Goby),
and Gobioides broussonetii (Violet Goby) (Appendix 2).
Comparisons among swamp ecosystems
Of the 4 swamp ecosystems examined, Great Dismal Swamp had the lowest species
richness, whereas the transient community in Mississippi floodplain swamps
Figure 5. Dendrogram of Bray-Curtis similarities calculated from the presence/absence of
all native fish species among major drainage units of southern F lorida.
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2015 Vol. 14, No. 3
was by far the richest (Table 1). Several families from the Mississippi were absent
from other faunas including Polyodontidae, Hiodontidae, and Moronidae. Great
Dismal Swamp had 1 family that was absent from the rest, the Amblyopsidae,
and shared the Umbridae with the Okefenokee Swamp. Big Cypress Swamp had
representatives from several euryhaline families not shared with the other swamp
systems (Table 1). With the exception of the Mississippi floodplain-swamp transient
community, there was low species richness of Cyprinidae, Catostomidae, and
Percidae in all southeastern swamps.
Table 1. Number of species by family for 4 major swamp systems, which include Big Cypress Swamp
(BC), Okefenokee Swamp (Okef), Great Dismal Swamp (Dism), and the lower Mississippi floodplain
swamps. The lower Mississippi floodplain swamps are separated into 2 categories representing permanent
residents of the floodplain (Miss-P) and transient species (Miss-T) following Baker et al. (1991).
Families are ordered taxonomically following Nelson et al. (2004).
BC Okef Dism Miss-P Miss-T
Polyodontidae - - - - 1
Lepisosteidae 2 1 1 4 4
Amiidae 1 1 1 1 1
Hiodontidae - - - - 2
Elopidae 1 - - - -
Megalopidae 1 - - - -
Anguillidae 1 1 1 - 1
Clupeidae 1 - - - 3
Cyprinidae 3 - 1 5 15
Catostomidae 1 2 2 1 8
Ariidae 1 - - - -
Ictaluridae 3 5 4 2 7
Esocidae 1 2 2 2 2
Umbridae - 1 1 - -
Aphredoderidae - 1 1 1 1
Amblyopsidae - - 1 - -
Mugilidae 1 - - - -
Atherinopsidae 2 1 - 1 2
Belonidae 3 - - - -
Fundulidae 8 4 1 4 4
Cyprinodontidae 3 - - - -
Poeciliidae 3 2 1 1 1
Centropomidae 1 - - - -
Moronidae - - - - 3
Centrarchidae 8 11 8 9 13
Percidae 1 2 4 4 6
Carangidae 1 - - - -
Lutjanidae 1 - - - -
Gerreidae 2 - - - -
Sparidae 2 - - - -
Sciaenidae 1 - - - 1
Elassomatidae 1 2 - 1 1
Eleotridae 1 - - - -
Gobiidae 3 - - - -
Achiridae 1 - - - -
Total Richness: 59 36 29 36 76
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Discussion
During the current study, we recorded 63 fish species (9 of these non-indigenous)
from fresh water within BCNP, substantially more than the 32 species recorded by
Loftus and Kushlan (1987). The primary reasons for the larger number we detected
were our study’s increased spatial extent and sampling effort, and the recent invasions
by several non-indigenous fishes into BCNP. Distinct species assemblages
occurred in seasonally brackish coastal habitats, permanent freshwater habitats,
and impermanent freshwater wetlands. The ichthyofaunal assemblage of BCS was
similar to other coastal drainages in southern Florida, particularly the southern Everglades
(Appendix 2; Rehage and Loftus 2007), but it included several euryhaline
species and thereby differed from inland drainages and other major swamp systems
of the southeastern US.
Based on our results and surveys of others, a total of 70 species (12 non-indigenous)
have been recorded from fresh water in BCS, with 67 of these (10 nonindigenous)
collected in BCNP. Species that we did not collect in BCNP but were
recorded there by others include Adinia xenica (Diamond Killifish) from coastal
ditches (Fowler 1945) and Turner River (Loftus and Kushlan 1987), Dormitator
maculatus (Fat Sleeper) off Loop Road (W.F. Loftus, unpubl. data), and Lepisosteus
osseus (Longnose Gar) in the L-28 canal (Kline et al. 2013). A non-indigenous species,
Parachromis managuense (Jaguar Cichlid), has also been reported since the
completion of our survey (W.F. Loftus, unpubl. data; Shafland et al. 2008). Species
not recorded in BCNP but present in other parts of BCS include Redface Topminnow
collected near Immokalee (Gilbert et al. 1992) and the non-indigenous species
Pterygoplichthys disjunctivus (Vermiculated Sailfin Catfish) and P. multiradiatus
(Many-rayed Pleco) collected in Corkscrew Swamp (S.E. Clem, unpubl. data). Several
species collected from the adjacent Everglades, including Dorosoma petenense
(Threadfin Shad), Channel Catfish, Chain Pickerel, and Pirate Perch, appear to have
spread from the north via the canal system (Gandy et al. 2012, Kline et al. 2013,
Loftus and Kushlan 1987) and may also be present in BCNP in low numbers.
The longest-established non-indigenous species in BCNP, Black Acara and
Clarias batrachus (Walking Catfish), have been present since the 1970s (Kushlan
1972, Loftus and Kushlan 1987), whereas Astronotus ocellatus (Oscar) and Spotted
Tilapia became established in the early 1980s (Loftus and Kushlan 1987). More
recent additions were Pike Livebearer, Mayan Cichlid, and Oreochromis aureus
(Blue Tilapia) in the 1990s (Fuller et al. 1999). We first collected Brown Hoplo in
2002, and its overall distribution in Florida indicates that it probably spread into
BCNP from the north (Nico 2005, Nico et al. 1996). African Jewelfish was first
collected in 2003 and because most specimens were collected along the eastern portions
of Tamiami Trail and Loop Road likely spread westward into BCNP via the
Tamiami canal. Both Brown Hoplo and African Jewelfish continue to expand their
ranges in southern Florida (Idelberger et al. 2011, Kline et al. 2013)
Most non-indigenous species were more frequent in permanent habitats such as
canals and the Turner River, but we captured all species in wetland habitats. A few
species were common in wetlands. Pike Livebearer was notable because of its high
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2015 Vol. 14, No. 3
frequency of occurrence in shallow, short-hydroperiod wetlands; this species is an
efficient predator on small-bodied fishes (Belshe 1961, Loftus and Kushlan 1987) and
seems to be exploiting habitats in which native piscivorous fishes are scarce (Trexler
et al. 2000). Other non-indigenous species that occurred frequently in freshwater
wetlands included Black Acara, Mayan Cichlid, and Spotted Tilapia. Both Walking
Catfish and Brown Hoplo may have been more abundant in wetlands than our data indicate
because our sampling methods did not always collect them effectively.
Habitat differences
Our analyses indicated differences in species assemblages among habitat types
in BCNP. The largest distinction was between the euryhaline-dominated communities
in seasonally fresh-water habitats in the coastal region and those further
inland in permanently fresh waters. The second-largest distinction was between
the temporally permanent waters of freshwater canals and Turner River and the
temporally impermanent waters of wetland habitats. The third distinction was
between wetland habitats. The most notable wetland species assemblage spanned
both short-hydroperiod wet-season and long-hydroperiod dry-season samples, suggestive
of the movement of fishes from wet prairies and savanna in the wet season,
to freshwater marsh and cypress swamp in the dry season. Numerous studies within
southern Florida wetlands have noted the movement of fishes to long-hydroperiod
wetlands as shorter-hydroperiod wetlands dried (Carlson and Duever 1977, Kobza
et al. 2004, Parkos et al. 2011). Our study was not designed to examine differences
between seasons; thus, our results could be an artifact of sample-size issues; this is
an important caveat because species richness among the habitat types and seasonal
categories generally increased with increased number of samples, so our differences
between wet season and dry season could merely reflect differences in sampling
effort. As such, our conclusions in this regard must be considered preliminary.
Comparison to regional drainages
The fish fauna of BCS was highly similar to the southern Everglades, indicating
that the habitat differences between them did not affect fish distributions. Although
BCS, the southern Everglades, and the northern Everglades border each other and
share direct water connections, the fish community of the northern Everglades was
distinctly different from the other 2 in the cluster analysis. The distinction between
these drainages reflects the limited invasion of euryhaline species into freshwater
in southern Florida. Numerous euryhaline species occurred in drainages that abut
estuarine habitats, but we found relatively few of them far inland; this was the
primary reason why all drainages with direct access to estuarine waters clustered
together in our analysis separately from the 3 inland drainages (Fig. 3). When we
compared the number and identity of native freshwater fishes between BCS, the
southern Everglades, and the northern Everglades, they were nearly identical, as
found in Loftus and Kushlan (1987).
The lower numbers of native freshwater fishes in BCS compared to the Kissimmee
River, Lake Okeechobee, and Charlotte Harbor drainages are most likely
attributable to the seasonally harsh abiotic conditions and the paucity of both fluvial
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habitats and permanent open-water habitats in BCS (Kushlan and Lodge 1974,
Loftus 2000, Loftus and Kushlan 1987).
Comparisons among swamp ecosystems
The species richness of the BCS was among the highest of the swamps in
southeastern US with which we compared it and was second only to the transient
community of the Mississippi River. When we excluded canals and seasonally
brackish habitats, BCS had 31 species, exceeding only the Great Dismal Swamp.
Big Cypress Swamp and Great Dismal Swamp are similar in that they are parts
of small coastal drainages, whereas the other 2 occur within substantial river systems
with larger potential species pools. Notably, despite being part of the vast
Mississippi drainage with its large and diverse fish community, the permanent
floodplain assemblage of the lower Mississippi had no more species than the Okefenokee
Swamp, and only a few more than the other 2 swamps in the comparison.
The disparity in species richness between transient and permanent Mississippi
drainage-floodplain assemblages indicates that a rather restricted subset of the fish
community is able to maintain populations within lentic swamp ecosystems. Fish
species present in swamp habitats are not restricted to lentic swamps, and most are
widespread in both lentic and lotic habitats throughout the southeastern coastal
plain (Boschung and Mayden 2004, Marcy et al. 2005). The most notable difference
among the fish community of BCS and the other swamp systems examined was the
prevalence of the marine-derived families Fundulidae, Cyprinodontidae, and Poeciliidae
and the presence of other euryhaline species. In both respects, BCS bears
some resemblance to the karstic wetlands of Quintana Roo, Mexico (Zambrano et
al. 2006). Quintana Roo wetlands display many physical similarities to wetlands in
the greater Everglades ecosystem, but the Characidae and Cichlidae are dominant,
in addition to the Cyprinodontidae and Poeciliidae. Both Pike Livebearer and Mayan
Cichlid are common in the Quintana Roo wetlands (Zambrano et al. 2006) and
well-established in BCS and the Everglades (Kline et al. 2013).
Summary
During the current study, we collected 63 fish species in fresh water in BCNP,
9 of which were non-indigenous. We collected 35 fish species from freshwater
wetlands. Species assemblages were most dissimilar between coastal habitats,
permanent freshwater habitats, and impermanent freshwater habitats; species composition
varied little among freshwater-wetland habitat types. The fish fauna of
the BCS is very similar to the southern Everglades and consists largely of species
widespread throughout Florida and the southeastern Coastal Plain. The main factor
that distinguishes the ichthyofauna of BCS from other swamp systems, as well as
drainages outside of the Florida peninsula, is the importance of euryhaline members
of the Poeciliidae, Fundulidae, and Cyprinodontidae.
Acknowledgments
We wish to thank Ron and Christine Clarke, Robert Sobzack, Deborah Jansen, Peg
Kohl, and Bill Evans of BCNP for their assistance and advice during this project. We would
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2015 Vol. 14, No. 3
also like to thank James Snyder, Brian Jeffery, and J. Hardin Waddle of the US Geological
Survey in BCNP for transportation to remote sites and for logistical help. We gratefully
acknowledge the support of G. Ronnie Best, coordinator of the Greater Everglades Priority
Ecosystems Science Program, and Matt Patterson, Coordinator of the NPS Southeast Inventory
and Management Program, for funding this project. We thank Nancy Russell at
Everglades National Park for assistance with documentation of voucher specimens.
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Appendix 1. List of species collected in BCNP from October 2002 to May 2004 by habitat type (CM = coastal marsh, CC = coastal canal, FC = freshwater
canal, TR = Turner River, H = mixed hardwood swamp, C = cypress forest, S = cypress savanna, P = wet prairie, M = freshwater marsh), and season (-D
= dry season, -W = wet season). Frequency is given for each species by habitat and in total. + = species collected or observed from samples for which
frequency was not calculated. Species status is as follows: N-f = primary or secondary freshwater species native to BCNP, U-f = primary or secondary
freshwater species for which native status is uncertain, N-e = native euryhaline species collected in freshwater, U-e = euryhaline species collected in
freshwater for which native status is uncertain, and X = non-indigenous species. Species richness by habitat type and number of samples by gear type are
given at the bottom of the table. Families are ordered taxonomically following Nelson et al. (2004).
Coastal Deepwater Freshwater wetlands Total
Status CM CC FC TR H-D H-W C-D C-W S-D S-W P-D P-W M-D M-W freq.
Lepisosteidae
Lepisosteus platyrhincus DeKay N-f - 0.13 0.59 0.36 0.33 0.07 0.04 - - 0.08 - 0.04 0.05 0.10 0.201
Amiidae
Amia calva L. N-f - - 0.17 0.09 - - 0.03 - - - - - - - 0.048
Elopidae
Elops saurus L. N-e - + + - - - - - - - - - - - -
Megalopidae
Megalops atlanticus Valenciennes N-e - 0.06 + - - - - - - - - - - - 0.003
Anguillidae
Anguilla rostrata (Lesueur) N-e - - 0.02 0.09 - - - - - - - - - - 0.008
Clupeidae
Dorosoma cepedianum (Lesueur) U-e - - 0.04 - - - - - - - - - - - 0.008
Cyprinidae
Notemigonus crysoleucas (Mitchill) N-f - - 0.20 0.36 0.11 - 0.03 - - - - - 0.08 - 0.076
Notropis maculatus (Hay) N-f - - 0.01 - - - - - - - - - - - 0.003
Notropis petersoni Fowler N-f - - 0.02 - - - - - - - - - - - 0.006
Catostomidae
Erimyzon sucetta (Lacepède) N-f - - 0.09 - - - - + - - - - - - 0.025
Callichthyidae
Hoplosternum littorale (Hancock) X - - 0.01 - - - + - - - - - - - 0.003
Clariidae
Clarias batrachus (L.) X 0.10 - 0.17 0.09 0.06 - 0.01 - - - - - - - 0.051
Ariidae
Ariopsis felis (L.) N-e - 0.06 - - - - - - - - - - - - 0.003
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Coastal Deepwater Freshwater wetlands Total
Status CM CC FC TR H-D H-W C-D C-W S-D S-W P-D P-W M-D M-W freq.
Ictaluridae
Ameiurus natalis (Lesueur) N-f - - 0.20 + 0.06 - 0.16 0.06 - 0.08 - 0.15 0.16 0.05 0.116
Ameiurus nebulosus (Lesueur) N-f - - 0.01 - - - - - - - - - - - 0.003
Noturus gyrinus (Mitchill) N-f - - - - 0.17 0.07 0.04 - - - - - 0.03 - 0.025
Esocidae
Esox americanus Gmelin U-f - - - - - - 0.01 - - - - - - - 0.003
Mugilidae
Mugil cephalus L. N-e - 0.19 - 0.18 - - - - - - - - - - 0.014
Belonidae
Strongylura marina (Walbaum) N-e - + - - - - - - - - - - - - -
Strongylura notata (Poey) N-e - 0.06 - - - - - - - - - - - - 0.003
Strongylura timucu (Walbaum) N-e - + - - - - - - - - - - - - -
Atherinopsidae
Labidesthes sicculus (Cope) N-f - - 0.23 0.18 0.39 - 0.14 - 0.11 0.08 - - - 0.05 0.116
Menidia beryllina (Cope) N-e 0.50 0.50 0.02 - - - - - - - - - - - 0.042
Fundulidae
Fundulus chrysotus (Günther) N-f 0.20 0.13 0.68 0.27 0.50 0.20 0.55 0.24 0.67 0.77 0.38 0.56 0.53 0.67 0.520
Fundulus confluentus Goode & Bean N-e 0.60 0.13 0.23 - 0.22 0.13 0.45 0.18 0.67 0.31 0.38 0.41 0.47 0.24 0.322
Fundulus grandis Baird & Girard N-e 0.30 + - - - - - - - - - - - - 0.008
Fundulus seminolis Girard N-f - + 0.01 0.18 - - - - - - - - 0.03 - 0.011
Lucania goodei Jordan N-f 0.20 0.06 0.67 0.55 0.83 0.80 0.49 0.53 0.56 0.31 0.13 0.52 0.61 0.86 0.559
Lucania parva (Baird & Girard) N-e 0.50 0.19 - 0.09 - - - - - - - - - - 0.025
Cyprinodontidae
Cyprinodon variegatus Lacepède N-e 0.80 0.25 + - - - - - - - - 0.07 0.05 - 0.045
Jordanella floridae Goode & Bean N-f 0.30 0.06 0.51 0.18 0.39 0.20 0.55 0.24 0.67 0.62 0.50 0.70 0.58 0.67 0.486
Poeciliidae
Belonesox belizanus Kner X 0.20 0.19 0.26 0.09 0.33 0.07 0.42 0.35 0.44 0.62 0.75 0.67 0.37 0.29 0.353
Gambusia holbrooki Girard N-f 0.80 0.44 0.86 0.36 1.00 0.73 0.91 0.53 0.78 0.77 0.75 0.96 0.92 0.81 0.825
Heterandria formosa Girard N-f - 0.06 0.53 - 0.39 0.27 0.38 0.12 0.33 0.08 - 0.15 0.37 0.62 0.333
Poecilia latipinna (Lesueur) N-e 0.90 0.56 0.60 0.36 0.50 0.40 0.39 0.24 0.78 0.46 0.50 0.44 0.50 0.52 0.497
Centropomidae
Centropomus undecimalis (Bloch) N-e - 0.06 0.01 0.09 - - - - - - - - - - 0.008
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Coastal Deepwater Freshwater wetlands Total
Status CM CC FC TR H-D H-W C-D C-W S-D S-W P-D P-W M-D M-W freq.
Centrarchidae
Enneacanthus gloriosus (Holbrook) N-f 0.10 - 0.12 0.27 0.33 0.47 0.20 0.18 - 0.08 0.25 0.04 0.13 0.33 0.169
Lepomis gulosus (Cuvier) N-f 0.10 - 0.77 0.91 0.50 0.53 0.45 0.29 - 0.38 0.38 0.37 0.34 0.43 0.469
Lepomis macrochirus Rafinesque N-f - 0.19 0.57 0.09 0.06 - 0.06 - - - - - 0.08 0.10 0.172
Lepomis marginatus (Holbrook) N-f 0.20 0.06 0.65 0.27 0.56 0.60 0.68 0.53 0.78 0.85 0.50 0.37 0.50 0.62 0.556
Lepomis microlophus (Günther) N-f - 0.13 0.28 0.27 0.06 - 0.06 - - - - - 0.11 - 0.107
Lepomis punctatus (Valenciennes) N-f - 0.13 0.68 0.64 0.39 0.20 0.54 0.24 - 0.31 0.25 0.15 0.45 0.38 0.421
Micropterus salmoides (Lacepède) N-f - 0.31 0.65 0.18 0.22 - 0.10 - - 0.15 - 0.04 0.13 0.05 0.226
Pomoxis nigromaculatus (Lesueur) U-f - - + - - - - - - - - - - - -
Percidae
Etheostoma fusiforme (Girard) N-f - - 0.02 - - - + - - - - - - - 0.006
Carangidae
Caranx hippos (L.) N-e - + - - - - - - - - - - - - -
Lutjanidae
Lutjanus griseus (L.) N-e - 0.06 - - - - - - - - - - - - 0.003
Gerreidae
Eucinostomus harengulus Goode & Bean N-e - 0.06 + - - - - - - - - - - - 0.003
Eugerres plumieri (Cuvier) N-e - 0.13 0.02 0.09 - - - - - - - - - - 0.014
Sparidae
Archosargus probatocephalus (Walbaum) N-e - 0.06 - - - - - - - - - - - - 0.003
Lagodon rhomboides (L.) N-e - 0.06 - - - - - - - - - - - - 0.003
Sciaenidae
Sciaenops ocellatus (L.) N-e - + - - - - - - - - - - - - -
Elassomatidae
Elassoma evergladei Jordan N-f - - 0.16 0.09 0.78 0.20 0.32 0.18 0.22 - - 0.07 0.18 0.24 0.203
Cichlidae
Astronotus ocellatus (Agassiz) X - 0.06 0.07 0.09 - - 0.01 - - - - 0.04 - - 0.028
Cichlasoma bimaculatum (L.) X 0.30 0.13 0.23 0.09 0.22 0.67 0.26 0.29 0.22 0.54 0.25 0.30 0.34 0.62 0.302
Cichlasoma urophthalmus (Günther) X 0.70 0.69 0.74 0.45 0.22 0.13 0.42 0.18 0.22 0.38 0.38 0.37 0.37 0.38 0.458
Hemichromis letourneuxi Sauvage X - - 0.02 - - - 0.06 - 0.11 - 0.13 0.04 0.08 - 0.034
Oreochromis aureus (Steindachner) X 0.30 - 0.20 0.18 - - - 0.06 - - - - 0.03 - 0.065
Tilapia mariae Boulenger X 0.10 0.38 0.51 0.64 0.06 0.27 0.13 0.06 - 0.15 0.13 0.22 0.16 0.33 0.260
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Coastal Deepwater Freshwater wetlands Total
Status CM CC FC TR H-D H-W C-D C-W S-D S-W P-D P-W M-D M-W freq.
Gobiidae
Bathygobius soporator (Valenciennes) N-e - + - - - - - - - - - - - - -
Lophogobius cyprinoides (Pallas) N-e - 0.44 0.04 - - - - - - - - - - - 0.028
Microgobius gulosus (Girard) N-e 0.10 0.06 - - - - - - - - - - - - 0.006
Achiridae
Trinectes maculatus (Bloch and Schneider) N-e - 0.38 0.01 0.18 - - - - - - - - - - 0.025
Species richness by habitat 20 42 47 32 25 18 30 18 14 19 15 22 27 21 63
Number of sample sites 10 16 81 11 18 15 69 17 9 13 7 28 38 21 353
Number of sample sites by primary gear type
Trap sites 10 4 42 6 5 15 50 15 9 12 7 27 28 19 249
Electrofishing sites 0 8 28 4 0 0 3 0 0 0 0 0 2 0 45
Dip-net sites 0 4 11 1 13 0 16 2 0 1 0 1 8 2 59
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Appendix 2. List of fish species recorded from major drainage areas of south Florida, abbreviated (citationsA in parentheses—see footnote at end of appendix)
as follows: KR (1, 2) = Kissimmee River; LO (2, 3, 4, 5, 6, 7, 8, 9, 10) = Lake Okeechobee; NE (11, 12, 13, 14, 15) = northern Everglades; SE
(12, 14, 15, 16, 17, 18, 44, 45 = southern Everglades; BC (7, 14, 16, 19, 20, 21, 22, 23, 24, 25, 44, 45) = Big Cypress Swamp; CH (7, 19, 26, 27, 28, 29,
30, 31) = Charlotte Harbor; CR (4, 7, 19, 28, 29, 31, 32) = Caloosahatchee River; IR (14, 33, 34, 35, 36, 37, 38, 39, 40, 45) = Indian River drainages; and
SC (14, 18, 41, 42, 43, 45) = southeast coast. Species categories are as follows: N-f = native primary or secondary freshwater species, U-f = primary or
secondary freshwater species for which native status to drainage is uncertain, N-e = native euryhaline species collected in freshwater, U-e = euryhaline
species collected in freshwater for which native status to drainage is uncertain), *e = native euryhaline species not yet collected from freshwater, and X =
non-native species. Families are ordered taxonomically following Nelson et al. (2004). Authorities provided for species not listed in Appendix 1.
KR LO NE SE BC CH CR IR SC
Carcharhinidae
Carcharhinus leucas (Müller and Henle) N-e *e *e *e N-e *e
Dasyatidae
Dasyatis sabina (Lesueur) N-e *e *e *e N-e *e
Acipenseridae
Acipenser oxyrinchus Mitchell *e
Lepisosteidae
Lepisosteus osseus (L.) N-f N-f U-f U-f U-f N-f U-f U-f
Lepisosteus platyrhincus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Amiidae
Amia calva N-f N-f N-f N-f N-f N-f N-f N-f N-f
Notopteridae
Chitala ornata (Gray) X
Elopidae
Elops saurus U-e N-e N-e *e *e N-e
Megalopidae
Megalops atlanticus U-e U-e N-e N-e *e *e N-e N-e
Anguillidae
Anguilla rostrata U-e U-e U-e N-e N-e N-e *e N-e N-e
Clupeidae
Brevoortia patronus Goode *e *e *e
Brevoortia smithi Hildebrand *e *e *e *e N-e
Brevoortia tyrannus (Latrobe) N-e
Dorosoma cepedianum N-e N-e U-e U-e N-e *e U-e U-e
Dorosoma petenense (Günther) U-e U-e U-e U-e U-e U-e U-e U-e
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KR LO NE SE BC CH CR IR SC
Engraulidae
Anchoa mitchilli (Valenciennes) N-e *e *e *e N-e *e
Cyprinidae
Ctenopharyngodon idella (Valenciennes) X X X
Cyprinus carpio L. X X
Notemigonus crysoleucas (Mitchell) N-f N-f N-f N-f N-f N-f N-f N-f N-f
Notropis chalybaeus (Cope) N-f N-f N-f
Notropis maculatus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Notropis petersoni N-f N-f N-f N-f N-f N-f N-f N-f
Opsopoeodus emiliae Hay N-f N-f N-f
Pteronotropis metallicus (Jordan & Meek) N-f
Catostomidae
Erimyzon sucetta N-f N-f N-f N-f N-f N-f N-f N-f N-f
Characidae
Metynnis sp. X
Callichthyidae
Hoplosternum littorale (Hancock) X X X X X X X X X
Loricariidae
Ancistrus sp. X
Pterygoplichthys disjunctivus (Weber) X X X X X
Pterygoplichthys multiradiatus (Hancock) X X X X X X
Hypostomus sp. X
Clariidae
Clarias batrachus X X X X X X X X X
Ariidae
Ariopsis felis N-e N-e *e *e N-e *e
Bagre marinus (Mitchell) N-e *e *e *e N-e *e
Ictaluridae
Ameiurus catus (L.) N-f N-f U-f N-f N-f U-f
Ameiurus natalis N-f N-f N-f N-f N-f N-f N-f N-f N-f
Ameiurus nebulosus N-f N-f N-f N-f N-f N-f N-f N-f
Ictalurus punctatus Rafinesque N-f N-f U-f U-f N-f N-f U-f
Noturus gyrinus N-f N-f N-f N-f N-f N-f N-f N-f
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KR LO NE SE BC CH CR IR SC
Esocidae
Esox americanus Gmelin N-f N-f U-f U-f U-f N-f U-f
Esox niger Lesueur N-f N-f U-f U-f
Aphredoderidae
Aphredoderus sayanus (Gilliams) N-f N-f U-f U-f N-f U-f
Mugilidae
Agonostomus monticola (Bancroft) N-e N-e
Mugil cephalus U-e U-e N-e N-e N-e *e N-e N-e
Mugil curema Valenciennes *e *e *e *e N-e N-e
Mugil gyrans (Jordan & Gilbert) N-e *e *e *e *e *e
Atherinopsidae
Labidesthes sicculus (Cope) N-e N-e N-e N-e N-e N-e N-e N-e N-e
Menidia beryllina (Cope) U-e U-e N-e N-e N-e N-e N-e N-e
Belonidae
Strongylura marina U-e U-e U-e N-e N-e *e *e N-e N-e
Strongylura notata N-e N-e *e *e N-e *e
Strongylura timucu N-e N-e *e *e *e N-e
Rivulidae
Kryptolebias marmoratus (Poey) N-e *e *e *e *e
Fundulidae
Adinia xenica (Jordan & Gilbert N-e N-e *e *e
Fundulus chrysotus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Fundulus confluentus N-e N-e N-e N-e N-e N-e N-e N-e
Fundulus grandis N-e N-e *e *e N-e N-e
Fundulus lineolatus (Agassiz) N-f N-f N-f N-f
Fundulus rubrifrons (Jordan) N-f N-f N-f N-f N-f N-f
Fundulus seminolis N-f N-f N-f N-f N-f N-f N-f N-f
Fundulus similis (Baird & Girard) N-e *e *e *e N-e N-e
Leptolucania ommata (Jordan) N-f N-f
Lucania goodei N-f N-f N-f N-f N-f N-f N-f N-f N-f
Lucania parva N-f N-f N-f *e N-f N-f
Cyprinodontidae
Cyprinodon variegatus U-e U-e N-e N-e N-e *e N-e N-e N-e
Floridichthys carpio (Günther) N-e *e *e *e *e N-e
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Jordanella floridae N-f N-f N-f N-f N-f N-f N-f N-f N-f
Poeciliidae
Belonesox belizanus Kner X X X X
Gambusia holbrooki Girard N-f N-f N-f N-f N-f N-f N-f N-f N-f
Gambusia rhizophorae Rivas N-e
Heterandria formosa Girard N-f N-f N-f N-f N-f N-f N-f N-f N-f
Poecilia latipinna (Lesueur) N-e N-e N-e N-e N-e N-e N-e N-e N-e
Syngnathidae
Microphis brachyurus (Bleeker) U-e N-e N-e
Syngnathus louisianae Günther *e *e *e *e N-e *e
Syngnathus scovelli (Evermann and Kendall) *e *e *e *e N-e *e
Synbranchidae
Monopterus albus (Zuiew) X X
Mastacembelidae
Macrognathus siamensis (Günther) X X
Centropomidae
Centropomus ensiferus Poey N-e N-e
Centropomus parallelus Poey U-e *e *e N-e N-e
Centropomus pectinatus Poey N-e *e *e N-e N-e
Centropomus undecimalis U-e U-e N-e N-e N-e *e N-e N-e
Serranidae
Epinephelus itajara N-e *e *e *e *e *e
Centrarchidae
Enneacanthus gloriosus (Holbrook) N-f N-f N-f N-f N-f N-f N-f N-f N-f
Lepomis auritus (L.) N-f
Lepomis gulosus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Lepomis macrochirus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Lepomis marginatus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Lepomis microlophus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Lepomis punctatus N-f N-f N-f N-f N-f N-f N-f N-f N-f
Micropterus salmoides N-f N-f N-f N-f N-f N-f N-f N-f N-f
Pomoxis nigromaculatus N-f N-f U-f U-f U-f N-f U-f U-f
Percidae
Etheostoma fusiforme N-f N-f N-f N-f N-f N-f N-f N-f
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Percina nigrofasciata (Agassiz) N-f
Carangidae
Caranx hippos N-e N-e *e *e N-e N-e
Caranx latus Agassiz *e *e *e N-e
Oligoplites saurus (Bloch and Schneider) N-e *e *e *e N-e *e
Lutjanidae
Lutjanus griseus N-e N-e *e *e N-e N-e
Gerreidae
Diapterus auratus Ranzani *e N-e *e N-e
Eugerres plumieri (Cuvier) N-e N-e *e *e N-e N-e
Eucinostomus argenteus Baird & Girard N-e *e *e *e N-e *e
Eucinostomus gula (Quoy & Gaimard) N-e *e *e *e N-e *e
Eucinostomus harengulus *e N-e *e *e N-e N-e
Haemulidae
Pomadasys crocro (Cuvier) N-e
Sparidae
Archosargus probatocephalus N-e N-e *e *e N-e *e
Lagodon rhomboides *e N-e *e *e N-e N-e
Sciaenidae
Bairdiella chrysoura (Lacepède) N-e *e *e *e N-e
Cynoscion arenarius Ginsburg *e *e *e *e
Cynoscion regalis (Bloch and Schneider) N-e
Leiostomus xanthurus Lacepède *e *e *e *e N-e *e
Micropogonias undulatus (L.) *e *e *e *e N-e *e
Pogonias cromis (L.) *e *e *e *e N-e *e
Sciaenops ocellatus N-e N-e *e *e N-e *e
Elassomatidae
Elassoma evergladei N-f N-f N-f N-f N-f N-f N-f N-f
Elassoma okefenokee Böhlke N-f
Cichlidae
Amphilophus citrinellus (Günther) X
Astatotilapia calliptera (Günther) X
Astronotus ocellatus X X X X X X
Cichla ocellaris Bloch and Schneider X X
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Cichlasoma bimaculatum X X X X X X
Cichlasoma urophthalmus X X X X X X X X
Geophagus sp. X
Hemichromis letourneuxi X X X X X X
Heros severus Heckel X X
Oreochromis aureus X X X X X X X X X
Oreochromis mossambicus (Peters) X X X
Oreochromis niloticus (L.) X X X
Parachromis managuensis (Günther) X X X
Parachromis salvini (Günther) X X
Rocio octofasciata (Regan) X
Sarotherodon melanotheron Rüppell X
Therops melanurus x T. zonatus X
Tilapia mariae X X X X X X X X
Tilapia buttikoferi (Hubrecht) X
Eleotridae
Dormitator maculatus (Bloch) N-e N-e *e *e N-e N-e
Eleotris amblyopsis (Cope) N-e N-e
Erotelis smaragdus (Valenciennes) N-e
Gobiomorus dormitor Lacepède U-e N-e N-e
Gobiidae
Awaous banana (Valenciennes) N-e
Bathygobius curacao (Metzelaar) N-e
Bathygobius soporator *e N-e *e *e *e
Ctenogobius boleosoma (Jordan & Gilbert) *e *e *e N-e
Ctenogobius pseudofasciatus (Gilbert & Randall) N-e
Ctenogobius shufeldti (Jordan & Eigenmann) *e N-e
Ctenogobius smaragdus (Valenciennes) N-e *e *e *e *e
Evorthodus lyricus (Girard) N-e
Gobioides broussonnetii Lacepède N-e
Gobionellus oceanicus *e *e *e N-e
Gobiosoma bosc (Lacepède) U-e N-e *e *e *e N-e *e
Lophogobius cyprinoides N-e N-e *e *e N-e N-e
Microgobius gulosus U-e N-e N-e *e *e N-e *e
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Sphyraenidae
Sphyraena barracuda (Edwards) N-e *e *e *e *e N-e
Osphronemidae
Trichopsis vittata (Cuvier) X
Channidae
Channa marulius (Hamilton) X
Paralichthyidae
Citharichthys spilopterus Günther *e *e *e *e N-e
Achiridae
Achirus lineatus (L.) N-e *e *e *e N-e *e
Trinectes maculatus U-e N-e N-e N-e N-e N-e
Number of species by category:
N-f 39 35 24 25 26 35 25 28 22
U-f 0 0 7 6 3 0 0 6 2
N-e 3 4 4 43 28 10 7 65 30
U-e 6 15 6 1 1 1 1 2 2
*e 0 0 0 14 32 51 57 8 21
X 7 10 10 17 12 10 8 13 28
A1 = Trexler (1995), 2 = Nico (2005), 3 = Ager (1971), 4 = Lee et al. (1980), 5 = Gilmore and Hastings (1983), 6 = Rivas (1986), 7 = Gilbert et al. (1992),
8 = Chick and McIvor (1994), 9 = Havens et al. (1996), 10 = James and Zhang (2011), 11 = Dineen (1984), 12 = Chick et al. (2004), 13 = Pearlstine et al.
(2007), 14 = Shafland et al. (2008), 15 = Parkos et al. (2011), 16 = Loftus and Kushlan (1987), 17 = Loftus (2000), 18 = Gandy et al. (2012), 19 = Fowler
(1945), 20 = Carter et al. (1973), 21 = Kushlan and Lodge (1974), 22 = Dunker (2003), 23 = Ceilly (2008), 24 = O’Donnell (2000), 25 = present study, 26
= Woolman (1892), 27 = Champeau (1990), 28 = Poulakis et al (2004), 29 = Fraser (2007), 30 = Champeau et al. (2009), 31 = Idelberger et al. (2011), 32
= Gunter and Hall (1965), 33 = Gunter and Hall (1963), 34 = Christensen (1965), 35 = Gilmore (1977), 36 = Gilmore and Hastings (1983), 37 = Gilmore
et al. (1983), 38 = Gilmore (1995), 39 = Paperno and Brodie (2004), 40 = Main et al. (2007), 41 = Smith (1895), 42 = Evermann and Kendall (1899), 43
= Ellis et al. (2003), 44 = Kline et al. (2013), 45 = W.F. Loftus, pers. observ.