Taxonomic Notes
The species has been recognized as distinct since its description and no subspecies have been described.
Justification
We assess Apalone ferox as Least Concern (LC) at the present time, as it was previously assessed on the Red List in 2011 (van Dijk 2011), although it may possibly be approaching Near Threatened (NT) status. However, in our opinion, A. ferox may instead meet the definition of Data Deficient (DD), as the available literature on its life history, particularly population demographics, is quite scarce. A solitary population demographic study is present in the literature, encompassing a population model along with biomass and density estimates, but it pertained only to a small population in a protected state park habitat. Wild populations of this species remain largely unstudied, with limited information available. A relatively recent past history of massive commercial harvests and current susceptibility to viral disease are documented, though the lasting and current impacts of these threats are unclear. Despite these uncertainties, the species is quite common in Florida and is often encountered crossing roads or seen in wetlands, and the population study noted above indicated high densities of animals in an optimal habitat. Further demographic field research, natural history and reproductive parameters, and population and trade monitoring of this species are all clearly needed.
Geographic Range Information
Apalone ferox occurs from southern South Carolina (vicinity of Charleston) through eastern and southern Georgia, southeastern Alabama and essentially all of Florida (Iverson 1992, Meylan and Moler 2006, TTWG 2021). In Florida, the species also occurs on Big Pine Key, but there is some debate over whether this population is native or introduced (Somma 2023). In Louisiana, there is an apparently established introduced population in Rockefeller Wildlife Refuge (Selman et al. 2016). In South Carolina, introduced populations appear to be established in the Edisto and Salkehatchie River drainages (Somma 2023). Other non-native records of this species (see iNaturalist, Somma 2023, VertNet) likely represent released individuals or captive escapees rather than established populations. The estimated historical indigenous range (area of occupancy, AOO) is 247,090 sq. km, and the estimated historical indigenous extent of occurrence (EOO) is 532,015 sq. km (TTWG in press).
Population Information
Apalone ferox is generally considered common to very common throughout most of Florida (Meylan and Moler 2006); however, demographic population structure is poorly understood even though this species was once the most important commercially harvested turtle in Florida (Meylan and Moler 2006). Before 2000, the species suffered unlimited harvest due to a lack of regulations. Resultant localized population declines have been attributed to this intensive collection, but only partial quantitative population estimates or trend data are available, and although preliminary indications were that a depleted population might be able to recover in about 20 years (Enge 1993, Meylan and Moler 2006), this rate of potential recovery is not supported by sufficient quantitative evidence (B. O'Hanlon pers. comm.). Only three studies have attempted to analyze and collect population data for this species (Bancroft et al. 1983, Aresco 2003, Munscher et al. 2015). Neither Bancroft et al. (1983) nor Aresco (2003) captured enough turtles to provide population demographics. Munscher et al. (2015) documented a male:female sex ratio of 1:2.6, showing a highly skewed female to male ratio and a population estimate of 92 adults and 49 juveniles with a total estimate of 141 turtles in the 2.67-ha study site.
The current population trend for the species would most likely be considered Stable. However, there is very little known regarding population demographics such as survivability over time, density, and biomass. The species is ranked 38th out of the 58 recognized species in the U.S. and Canada as having the fewest citations of published research articles (Lovich and Ennen 2013). The only study that presents these variables is Munscher et al. (2015) with a density for adults of 34.5/ha and juvenile density of 18.4/ha. The apparent survivorship in the studied population was high at 0.96 for both adult males and females, and slightly lower for juveniles at 0.92, however, this population is in a protected spring habitat, so these variables may be different in other more vulnerable habitat types and locations (Munscher et al. 2015).
Habitat and Ecology Information
Apalone ferox is a habitat generalist that has been documented occupying a variety of freshwater habitats, including shallow vegetated lake and wetland areas, freshwater streams and rivers, man-made canals, ponds, as well as freshwater spring ecosystems (Bancroft et al. 1983, Aresco 2003, Meylan and Moler 2006, Munscher et al. 2015). A population living in a spring-fed lagoon and river system characterized with thick layers of silt and detritus up to 1 m thick was described by Munscher et al. (2015).
The species is highly carnivorous and has been documented preying on various species of snails, insects, crayfish, fish, frogs, birds, and even small alligators (Meylan and Moler 2006, Flaherty and Friers 2013, Moler and Epstein 1994). Vegetation has also been found in the stomachs of A. ferox; however, this is believed to be due to incidental ingestion (Meylan and Moler 2006).
This species has marked sexual dimorphism with females growing to be substantially larger than males (Meylan and Moler 2006, Munscher et al. 2015). Females can be differentiated from males based on tail structure. Males have large thick tails with vents that can stretch past the carapace rim, while the tails of females are very short and small in comparison (Moler and Berish 1995, Ernst and Lovich 2009, Munscher et al. 2015). Average sizes reported for each sex by Munscher et al. (2015) were 44.9 cm carapace length and 9.43 kg for females and 31.3 cm and 3.29 kg for males. The maximum straightline carapace length record for males is 32.4 cm and for females 67.3 cm, with the largest known individual being a female with a curved carapace length of 73.8 cm and mass of 41.4 kg (Pritchard 2001, Ernst and Lovich 2009, TTWG 2021).
Meylan and Moler (2006) reported that females can reach maturity between 25 and 40 cm carapace length while males reach sexual maturity between 15 and 21 cm carapace length, but the ages at which they reach maturity is unknown, and generation time remains unknown.
Nesting season occurs from late March to early August (Iverson and Moler 1997). While females are believed to produce up to six clutches per year, Iverson and Moler (1997) found evidence that not all females produce eggs each year. Typically, the clutch size for this species ranges between nine and 38 eggs, with 20.6 being the average (Iverson and Moler 1997, Iverson 1985) also reported a maximum egg diameter range between 25.9 and 32.0 mm. The incubation period is inversely correlated to temperature and lasts between 64 and 79 days (Iverson 1985). Assuming five clutches are laid per year, Iverson (1985) estimated an average annual reproductive output of approximately 78 eggs per female. When comparing the relative wet clutch mass as a percentage of female body mass of 1.70%, as calculated by Goff and Goff (1935), it appears that this species produces some of the lowest values for reproductive effort per clutch among turtles (Iverson 1985). Average hatchling size is 41 mm (range, 36–44 mm) with an average mass of 9.7 g (range, 8–11 g) (Meylan and Moler 2006).
This species has been documented utilizing excavated material from other species as nest sites. Heinrich and Richardson (1993) reported finding a nest within the abandoned burrow apron of a Gopher Tortoise (Gopherus polyphemus), while multiple sources have noted the use of nests of American Alligators (Alligator mississippiensis) by this species as well (Deitz and Jackson 1979, Enge et al. 2000).
Nest predation by Fish Crows (Corvus ossifragus), foxes, raccoons, and skunks has been documented (Meylan and Moler 2006). Additional natural predators include birds of prey (Beissinger 1990, Meylan and Moler 2006, Walde and Saumure 2006), River Otters (Stacy et al. 2014), and occasionally American Alligators (Meylan and Moler 2006).
Threats Information
Before the 2009 Florida freshwater commercial turtle harvest ban, Apalone ferox was intensively harvested, with thousands of kilograms of turtles being taken out of freshwater systems (Enge 1993, Meylan and Moler 2006); however, it is unknown how current non-commercial capture rates are impacting the species.
A significant threat currently faced by this species is collision with motor vehicles (van Dijk 2011). This primarily occurs when females are moving overland during the nesting season; however, fluctuations in weather conditions, such as drought, can also cause individuals to move across roadways or other areas where contact with vehicles is more likely (Aresco 2005, van Dijk 2011). Additionally, many roadways and other developments break up habitats and disrupt normal movement patterns for the species (Aresco 2005). While lack of data makes it difficult to quantify the role development and motor vehicles have played in the population metrics over time, it is reasonable to assume based on information gathered from other species that it is a substantial threat to A. ferox (review by Aresco 2005).
The explosion of mesopredator populations, such as raccoons, opossums, skunks, foxes, and armadillos, have resulted in dramatic increases in nest and hatchling predation for most turtle species (Munscher et al. 2012). There currently are no data available on depredation rates for A. ferox populations, as the species is lacking data in many aspects of its natural history. Other freshwater turtle species have suffered depredation rates of over 90% in nesting areas across the country, including ecosystems within the range of A. ferox (Munscher et al. 2012, Riley and Litzgus 2014).
As with other turtle species, diseases are of increasing concern for A. ferox. For example, recent research by Waltzek et al. (2022) outlined a novel group of negative sense RNA viruses that were found to infect numerous freshwater turtle species in Florida, including A. ferox. The proposed name for the bunyavirus detailed in their article is Turtle Fraservirus (TFV1); however, earlier reports of the same infection refer to the infectious agent as Turtle Bunyavirus (TBV) (Jones 2022, Waltzek et al. 2022). According to a 2022 report by the Florida Fish and Wildlife Conservation Commission, infection by this virus since 2018 has been linked to finds of moribund and dead turtles across the state of Florida in the following counties: Putnam, Lake, Seminole, Orange, Polk, Osceola, Brevard, Indian River, and Collier (Jones 2022). Symptoms of infection include lethargy, irregular swimming patterns, reddened or splotchy skin on the plastron or appendages, as well as irregular appearance of the eyes (Jones 2022, Waltzek et al. 2022).
Additionally, climate change has the potential to negatively impact this species through higher temperatures during the nesting season. Specifically, there is concern that elevated temperatures will create unsuitable environments for eggs to incubate within the species’ current range (Sullivan 2020, Sullivan et al. 2022). While scientists have noted A. ferox sometimes utilizing cooler nesting sites, and thereby probably maintaining better nest incubation temperatures, it is still unknown to what extent this species will be able to withstand the effects of warming temperatures on incubation (Sullivan 2020). Turtle species of the genus Apalone do not have temperature-dependent sex determination, but A. spinifera has sex chromosomes that are thermosensitive that could affect hatchling development at higher nesting temperatures (Bista et al. 2021), and this may affect A. ferox as well.
Use and Trade Information
Apalone ferox was previously the most important commercially harvested turtle in Florida (Meylan and Moler 2006). Before 2000, Florida softshell turtles suffered unlimited harvest due to a lack of regulations. Meylan and Moler (2006) described details of the fishery, amounting to a minimum of 7,500 adults annually (van Dijk 2011).
Ranching of the species, specifically egg collection from confined wild-caught animals, has produced substantial quantities of hatchlings for the international pet trade and for rearing facilities in China. Declared exports of A. ferox are recorded in the USFWS Law Enforcement Management Information System (LEMIS) database, but available numbers do not reliably allow separating the data by area of origin or whether collected from the wild or produced in captive conditions. Numbers of live specimens exported from the USA were under 100,000 per year during 1999–2003, but then rose rapidly and significantly to a peak of nearly 470,000 in 2012, after which trade volumes declined to 237,000 in 2013 and 213,000 in 2014, and then declined further, back to under 100,000 annually for the years 2015–2019. In total, nearly 3.12 million live specimens of A. ferox were declared to have been exported during 2000–2019 (see Figure S1 in Supplementary Information for annual declared numbers of A. ferox, A. spinifera, and A. mutica), in addition to some quantities of meat and eggs that were also recorded as exported.
Conservation Actions Information
In 2000, the Florida Fish and Wildlife Conservation Commission (FWC) enacted a harvest closure on all softshell turtles (Apalone spp.) and their eggs during the prime breeding and nesting season (1 May until 31 July: Interim FWC Rule 68A-25.002) (Munscher et al. 2015). In 2008, the FWC enacted an interim rule limiting the daily possession of softshell turtles to 5 turtles per person per day. During this interim period, licensed commercial fishermen could still remove up to 20 softshell turtles per day (Interim FWC Rule 68A- 25.002). In 2009, commercial harvest was ended and at that time the legal limit of take became one turtle per person per day, with some exceptions. Furthermore, no softshell turtles may be taken from 1 May to 31 July, which reflects protection during the nesting season (Munscher et al. 2015). These particular take permits have not been renewed (Interim FWC Rule 68A-25.002); thus, all commercial harvest of Florida softshell turtles has effectively ceased (Munscher et al. 2015). In 2021, the FWC enacted a new Executive Order 21-19 (EO 21-19 (myfwc.com)) that prohibits the taking, handling, and transporting of the species due to the recent issues with the turtle Fraservirus (B. O’Hanlon, Reptile and Amphibian Conservation Coordinator, FWC, pers. comm.).
Other states in the range of A. ferox, notably Georgia and South Carolina, have less stringent laws but still offer some protection. South Carolina regulations prohibit possession of more than five A. ferox or more than 10 native turtles in aggregate and possession of native species cannot be transferred (buy, sell, trade, etc) without a permit (A. Grosse, SC State Herpetologist, pers. comm.). In Georgia, no more than 10 freshwater turtles (any combination of species) may be possessed without a commercial turtle permit; with permitted turtle harvest still being allowed in the public waters of Georgia anyone with a permit can take up to 100 A. ferox on an annual basis. In Alabama, state law prohibits the sale of freshwater turtles, but allows for the taking of two turtles per day by means of hand capture, dip net, or hook and line for personal use of turtles not protected by the state.