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The Ichthyofauna of Big Cypress National Preserve, Florida
Marcus Zokan, Greg Ellis, Shawn E. Clem, Jerome Lorenz, and William F. Loftus

Southeastern Naturalist, Volume 14, Issue 3 (2015): 517–550

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Southeastern Naturalist 517 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 518 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). Southeastern Naturalist 519 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 520 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 Southeastern Naturalist 521 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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. Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 522 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 Southeastern Naturalist 523 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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. Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 524 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 Southeastern Naturalist 525 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 526 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. Southeastern Naturalist 527 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 528 (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 Southeastern Naturalist 529 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 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. Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 530 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. Southeastern Naturalist 531 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 532 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 Southeastern Naturalist 533 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 534 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 Southeastern Naturalist 535 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 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. 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Pp. 523–556, In J.N. Rinne, R.M. Hughes, and B. Calamusso (Eds.). Historical Changes in Large River Fish Assemblages of the Americas. American Fisheries Society Symposium 45, Bethesda, MD. 612 pp. Nico, L.G., S.J. Walsh, and R.H. Robins. 1996. An introduced population of the South American callichthyid catfish Hoplosternum littorale in the Indian River Lagoon system, Florida. Florida Scientist 59:189–200. Obaza, A., D.L. DeAngelis, and J.C. Trexler. 2011. Using data from an encounter sampler to model fish dispersal. Journal of Fish Biology 78:495–513. O’Donnell, P. 2000. Fish species of the upper Ten Thousand Islands and Rookery Bay. Available online at http://www.evergladesplan.org/pm/projects/project_docs/pdp_30_ sgge/pir_final/sgge_pir_sd_fish_species.pdf. Accessed 30 December 2012 Paperno, R., and R.B. Brodie. 2004. Effects of environmental variables upon the spatial and temporal structure of a fish community in a small, freshwater tributary of the Indian River Lagoon, Florida. Estuarine, Coastal and Shelf Science 61:229–241. Parkos III, J.J., C.R. Ruetz III, and J.C. Trexler. 2011. Disturbance regime and limits on benefits of refuge use for fishes in a fluctuating hydroscape. Oik os 120:1519–1530. Pearlstine, E., W.M. Bear, F.J. Mazzotti, and K.G. Rice. 2007. Checklist of fish in Rice and Sugarcane fields of the Everglades Agricultural Area. Florida Scientist 70:113–119. Poulakis, G.R., R.E. Matheson Jr., M.E. Mitchell, D.A. Blewett, and C.F. Idelberger. 2004. Fishes of the Charlotte Harbor estuarine system, Florida. Gulf of Mexico Science 22:117–150. Rehage, J.S., and W.F. Loftus. 2007. Seasonal fish-community variation in the upper stretches of mangrove creeks in the southwestern Everglades: The role of creeks as drydown refuges. Bulletin of Marine Science 80:625–645. Rivas, R. 1986. Systematic review of the perciform fishes of the genus Centropomus. Copeia 1986:579–611. Southeastern Naturalist 539 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 Schofield, P.J., and W.F. Loftus. 2014. Non-native fishes in Florida freshwaters: A literature review and synthesis. Reviews in Fish Biology and Fisheries 25:117–145. Shafland, P.L., K.B. Gestring, and M.S. Stanford. 2008. Florida’s exotic freshwater fishes: 2007. Florida Scientist 71:220–245. Smith, H.M. 1895. Notes on Biscayne Bay, Florida with reference to its adaptability as the site of a marine hatching and experiment station. Reports of the US Fishery Commission 21:169–181. Trexler, J.C. 1995. Restoration of the Kissimmee River: A conceptual model of past and present fish communities and its consequences for evaluating restoration success. Restoration Ecology 3:195–210 Trexler, J.C., W.F. Loftus, F. Jordan, J.J. Lorenz, J.H. Chick, and R.M. Kobza. 2000. Empirical assessment of fish introductions in southern Florida: An evaluation of contrasting views. Biological Invasions 2:265–277. Trexler, J.C., W.F. Loftus, C.F. Jordan, J. Chick, K.L. Kandl, T.C. McElroy, and O.L. Bass. 2002. Ecological scale and its implications for freshwater fishes in the Florida Everglades. Pp. 153–181, In J.W. Porter and K.G. Porter (Eds.). The Everglades, Florida Bay, and Coral Reefs of the Florida Keys: An Ecosystem Sourcebook. CRC, Boca Raton, FL. 1024 pp. Woolman. A.J. 1892. A report upon the rivers of central Florida tributary to the Gulf of Mexico, with lists of fishes inhabiting them. Bulletin of the United States Fish Commission for 1890 10:293–302. Zambrano, L., E. Vazquez-Dominguez, D. Garcia-Bedoya, W.F. Loftus, and J.C. Trexler. 2006. Fish-community structure in freshwater karstic water bodies of the Sian Ka’an Reserve in the Yucatan peninsula, Mexico. Ichthyological Exploration of Freshwaters 17:193–206. Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 540 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 Southeastern Naturalist 541 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 542 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 Southeastern Naturalist 543 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 544 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 Southeastern Naturalist 545 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 546 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 Southeastern Naturalist 547 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 KR LO NE SE BC CH CR IR SC 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 548 KR LO NE SE BC CH CR IR SC 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 Southeastern Naturalist 549 M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 KR LO NE SE BC CH CR IR SC 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 Southeastern Naturalist M. Zokan, G. Ellis, S.E. Clem, J. Lorenz, and W.F. Loftus 2015 Vol. 14, No. 3 550 KR LO NE SE BC CH CR IR SC 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.