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Herpetology Notes, volume 10: 601-609 (2017) (published online on 26 October 2017)

Turtle survey of State Park reveals more than mermaids

Eric C. Munscher1,2,*, Andrew D. Walde1,3, Elizabeth M. Walton1,4, Nicole Salvatico1,5, Brian P. Butterfield 1,6, Wayne Osborne1,7, Collin McAvinchey1,8 and J. Brian Hauge1,9

Abstract. During a survey of in Weeki Wachee Springs State Park, , USA, in March 2015, 182 turtles representing ten were captured, marked, and released, including the Florida Softshell ( ferox), North American Snapping Turtle ( serpentina), Florida (Deirochelys reticularia chrysea), ( baurii), ( concinna suwanniensis), Florida Red-bellied Cooter (P. nelsoni), (P. peninsularis), ( minor minor), Common Musk Turtle (S. odoratus), and ( scripta). Two of T. scripta (Red-eared Slider, T. s. elegans, and Yellow-bellied Slider, T. s. scripta) were captured, and both are new county records. The most abundant species was S. odoratus with 101 individuals captured and a relative abundance of 0.555. Among the three Pseudemys species, the presence of P. c. suwanniensis is important because it is currently listed as a species of special concern in Florida. Weeki Wachee Springs had the highest species richness (ten species) and the highest overall diversity (H’ = 1.33) of turtles among Florida spring-fed lotic habitats that have been surveyed over the past thirty years.

Keywords: Florida, freshwater springs, mermaids, Pseudemys, Sternotherus, Trachemys

Introduction Hernando County, Florida just off of U.S. Highway 19, approximately 115 km north of Tampa Bay, Florida The state of Florida has over 600 freshwater springs, (Fig. 1). The name Weeki Wachee is derived from the the largest concentration of this particular habitat in the world (FitzPatrick, 2010; Florida Department of Environmental Protection [FDEP], 2014). These unique and beautiful habitats have been revered, worshiped, explored, and enjoyed by people visiting 1 Turtle Survival Alliance, North American Freshwater Turtle or inhabiting Florida for over 10,000 years (Scott et al., Research Group, 1989 Colonial Parkway, Fort Worth, Texas 2004; Stamm, 2008). In addition, Florida’s freshwater 76110, USA springs offer essential and, at times, critical habitat for 2 SWCA Environmental Consultants, 10245 West Little York, many freshwater organisms, such as rare Cave Crayfish Suite 600, Houston, Texas, USA 3 (Procambarus sp.) and the endangered West Indian Walde Research & Environmental Consulting, 8000 San Manatee (Trichechus manatus) (Florida Springs Task Gregorio Road, Atascadero, California 93422, USA 4 School of Geosciences, University of South Florida, 4202 East Force, 2000). Today, many of these sites are under state Fowler Avenue, Tampa, Florida 33620, USA protection as part of Florida’s extensive, nationally 5 Florida Department of Health in Seminole County, 400 W renowned, and award winning state park system. The Airport Boulevard, Sanford, Florida 32773, USA Florida State Park and Trail System is the first state park 6 Freed-Hardeman University, 158 East Main Street, Henderson, system in the to receive a Gold Medal 38340, USA 7 Award three times (FDEP, 2013). As part of the state Pine Ridge High School, 926 Howland Boulevard, Deltona, system, these parks contribute to Florida’s economy and Florida 32738, USA 8 , 600 East Park Avenue, Port Angeles, provide recreational opportunities, such as swimming, Washington 98362, USA canoeing, and wildlife watching for millions of people 9 Department of Biology, Peninsula College, Port Angeles, every year (Stamm, 2008). Washington 98362, USA Weeki Wachee Springs State Park is located in * Corresponding author e-mail: [email protected] 602 Eric C. Munscher et al.

Figure 1. Weeki Wachee Springs State Park is located in Hernando County, Florida, just off of U.S. Highway 19, approximately 115 km north of Tampa Bay, Florida. The spring opens into a 0.5-ha lagoon that drains into a 19-km long spring run. Map created using Esri’s ArcGIS Version 10.3.1; data obtained from Esri Data Products, Southwest Florida Water Management District imagery (www.swfwmd.state.fl.us), U.S. Department of Commerce, U.S. Census Bureau, Geography Division, 2011 Florida Roads Census MAF / TIGER / Line data, and U.S. Geological Survey National Hydrography Dataset (www.datagateway.nrcs. usda.gov).

language of the Seminole Tribe and means “little spring” bank. The run eventually empties into the Gulf of or “winding river” (FitzPatrick, 2010). The spring is Mexico (Scott et al., 2004). located in a karst environment with an upper quartz Weeki Wachee Springs has a rich and unique history. In sand and residual clay layer formed during the recent 1946 the site was purchased by Newton Perry, a former Pliocene Epoch, and a lower layer primarily composed member of the U.S. Navy, who trained Navy Seals to of microfossiliferous carbonate and dolomitic-limestone swim underwater (FitzPatrick, 2010). Perry restored the interspersed with intergranular evaporates that was spring habitat and installed underwater breathing tubes formed during the Epoch (Hill et al., 2010). with free-flowing air. He then built the first 18-seat Weeki Wachee Springs is a first magnitude spring theatre at the spring boil that was situated nearly 2 m that expels more than 430 million litres of crystal clear below the water’s surface so the public could view the water per day at a constant temperature of 23°C. The natural beauty of the spring. Shortly thereafter, he began current in this spring run has been documented at over training young women to swim and perform aquatic 8 km/h (FitzPatrick, 2010). The basin of the spring boil ballets and other underwater exercises while using the is over 30 m wide, but the actual depth of the spring hidden breathing tubes. The first actual “mermaid” cavern leading to the Upper Floridan aquifer system show at Weeki Wachee Springs opened on 13 October is still unknown (Scott et al., 2004; FitzPatrick, 2010). 1947. The site and show became a huge hit and the The spring opens into a 0.5-ha lagoon that winds down spring was purchased by the American Broadcasting into a 19-km long spring run that, within the park, is Company (ABC) in 1959 (FitzPatrick, 2010). ABC built surrounded by low-lying densely forested swamp, and the current underwater amphitheatre that seats over 400 which outside of the park boundary becomes more people, and the company helped develop themes for urban with housing developments occurring on either underwater shows. An estimated one million people per Turtle survey of Weeki Wachee Springs State Park 603 year visited during the park’s heyday in the 1950s. In 1982, a water park called Buccaneer Bay, with a large sandy beach and swimming area, was added down from the springhead at the opening of the . In November 2008, Weeki Wachee Springs was purchased by the State of Florida and became one of the newest state parks (FitzPatrick, 2010). Although attendance is down considerably from the 1950s, the average number of visitors is still estimated to be over 390,000 people per year since becoming a state park (Park Manager Toby Brewer, pers. comm.). Anthropogenic inputs have caused an increase in nitrate concentrations by over 1000% during the last

40 years. While NO3 concentrations have increased from 0.07 mg/l in 1974 to approximately 0.90 mg/l in 2013, there was no significant increase in either orthophosphates (inorganic phosphates typically used in 3- 3- 3- fertilisers, including PO4 , HPO4 , and H2PO4 ) or total phosphates (TP; orthophosphates; organic phosphates found in human/ faecal matter, as well as naturally decaying organic matter; and condensed 5- phosphates, such as water additives of P3O10 to prevent scaling and corrosion) (Scott et al., 2004; Johnson et al., 2007; FDEP, 2014). Additional changes in the spring system include a 75% reduction in native submerged aquatic vegetation throughout the Weeki Wachee River (Frazer et al., 2006). By 2008, algal mats of invasive Figure 2. (A) Volunteers from the TSA-NAFTRG sampling Lyngbya wollei, a filamentous cyanobacterium, had Weeki Wachee Springs Mermaid Lagoon and Buccaneer Bay caused a significant decrease in native submerged Water Park area. (B) A typical view of Weeki Wachee Springs aquatic vegetation, prompting the Southwest Florida run. Photos by Nicole Salvatico. Water Management District to begin a management plan that included the removal of algal mats and excess sediment with plans to replant native vegetation in the main spring area (FDEP, 2014). Little is known about the turtle populations inhabiting Turtles are among the most conspicuous and the Weeki Wachee River. Enge and Wood (2000) ecologically important components of spring ecosystems captured Florida Chicken Turtles (Deirochelys in Florida (Meylan et al., 1992; Lagueux et al., 1995; reticularia chrysea), Striped Mud Turtles (Kinosternon Jackson and Walker, 1997; Hrychyshyn, 2006; Chapin baurii), Florida Mud Turtles (K. steindachneri), and and Meylan, 2011; Munscher et al., 2015a, b; Johnston Eastern Musk Turtles () in drift et al., 2016). Ease of capture while snorkelling in clear fence arrays in wetlands approximately 4 km north of water coupled with the ease of long-term identification the Weeki Wachee River. Ballou et al. (2015, 2016) through a unique marking system on their shells makes conducted a basking turtle survey in the Weeki Wachee turtles ideal candidates for monitoring in this type of River and documented the occurrence of Suwannee ecosystem. Because of the diverse diets and habitat Cooters (Pseudemys concinna suwanniensis), Peninsula requirements of different species in turtle assemblages Cooters (P. peninsularis), and Florida Red-bellied (Meylan, 2006), turtle populations may serve as Cooters (P. nelsoni). In March 2015, the Turtle Survival indicators of ecosystem health and stability. Thus, Alliance North American Freshwater Turtle Research documenting their numbers may serve as a barometer Group (TSA-NAFTRG) added Weeki Wachee Springs not only for turtle species, but for a host of other species to its list of study sites. The TSA-NAFTRG has been existing within the ecosystem as well. conducting turtle population assessments for the past 604 Eric C. Munscher et al.

Figure 3. All turtles were captured by hand, placed in kayaks and canoes and brought to a central location in the spring run, where they were processed and released as close to their point of capture as possible. Shown here, a volunteer with a haul of turtles, including several species of Pseudemys and some Sternotherus odoratus captured in the Mermaid Lagoon. Photo by Jessy Wayles.

19 years in other Florida spring locations including Materials and Methods Wekiwa Springs State Park (Munscher et al., 2015a, b; We surveyed Weeki Wachee Springs from 25–27 Walde et al., 2016), Volusia Blue Spring (Riedle et al., March 2015. Twelve divers were in the spring from 2016; Walde et al., 2016), Peacock Springs (Walde et 0700–0900 h before the park opened to the public. The al., 2016), and Manatee Springs (Walde et al., 2016). To Mermaid Lagoon, Buccaneer Bay, and the Water Park our knowledge, no previous turtle population research Lagoon were surveyed first (Figs. 1, 2), followed by involving the capture and marking of freshwater turtles approximately 4.8 km of the spring run. Two capture has ever been conducted in the park, or the Weeki sessions, from 0800–1000 h and 1800–2000 h, were Wachee River. TSA-NAFTRG’s goal was to conduct an conducted each day. Researchers have time limitations initial survey of the Weeki Wachee Springs boil, lagoon, to be in the water due to daily mermaid shows and boat and as much of the spring run as possible, with a focus tours that begin at 1100 h and run on the hour, every on turtle species richness and relative abundance, and to hour. All turtles were captured by hand, placed in kayaks put this in context of other spring systems throughout or canoes, and brought to a central location in the spring the state. This survey is intended to provide a baseline run, where they were processed and released as close to which future surveys may be compared. to their point of capture as possible (Fig. 3). Trachemys scripta is not native to this region of Florida and the Turtle survey of Weeki Wachee Springs State Park 605

Table 1. ��������������������������������������������������������������������������������������������������������������������� of individuals (relative abundance). References for the different localities include a = ����������������������������������������� ChapinTable and 1. Meylan Comparison (2011), of turtled = Riedle species et composition al. (2016), andamong e = five Johnston spring-fed et al. lotic (2016). ecosystems All studies in Florida. used the Data method presented of hand-capture as number of whileindividuals snorkeling. (relative abundance). References for the different localities include a = this study, b = Meylan et al. (1992), c = Chapin and Meylan (2011), d = Riedle et al. (2016), and e = Johnston et al. (2016). All studies used the method of hand-capture while snorkeling.

Species Weeki Wachee Springs a Rainbow Runb Ichetucknee Riverc Volusia Blue Springd Santa Fe Rivere Apalone ferox 4 (0.022) 1 (0.002) 1 (0.003) 3 (0.006) 3 (0.003) Chelydra serpentina 2 (0.011) 0 (0.0) 4 (0.01) 6 (0.011) 30 (0.027) Chrysemys picta1 0 (0.0) 0 (0.0) 0 (0.0) 1 (0.002) 0 (0.0) Deirochelys reticularia 1 (0.006) 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) pseudogeographica1 0 (0.0) 0 (0.0) 0 (0.0) 1 (0.002) 0 (0.0) Kinosternon baurii 1 (0.006) 0 (0.0) 0 (0.0) 1 (0.002) 4 (0.004) suwanniensis 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 9 (0.008) Pseudemys nelsoni 20 (0.11) 1 (0.002) 3 (0.008) 63 (0.116) 17 (0.016) Pseudemys peninsularis 41 (0.225) 15 (0.034) 0 (0.0) 214 (0.395) 18 (0.016) Pseudemys concinna suwanniensis 8 (0.044) 22 (0.05) 79 (0.2) 0 (0.0) 625 (0.570) Sternotherus minor 2 (0.011) 294 (0.662) 222 (0.566) 230 (0.424) 228 (0.208) Sternotherus odoratus 101 (0.555) 111 (0.25) 19 (0.048) 5 (0.009) 2 (0.002) Trachemys scripta 2 (0.011)2 0 (0.0) 60 (0.153) 18 (0.033)2 160 (0.146) Total 182 444 388 542 1096

1 Species not native to Florida 2 Individuals collected outside of their native ranges

Florida Fish and Wildlife Conservation Commission Munscher et al. (2015a). Capture and handling protocols requires us to remove invasive Red-eared Sliders (T. were approved by FDEP (District III, Orlando, Florida) s. elegans), but release Yellow-bellied Sliders (T. s. and conform to those promulgated by the American scripta). Society of Ichthyologists and Herpetologists (ASIH), the Turtles were measured with tree callipers to obtain Herpetologists League, and the Society for the Study of 1 straight-line measurements of maximum carapace Amphibians and animal use guidelines (ASIH, length (CL), maximum plastron length (PL), carapace 2004). width (CW), and shell height (SH), recorded to the We counted the number of species captured to nearest mm. Turtles were then sexed based on secondary determine species richness and calculated relative sexual characteristics, such as tail and claw length abundance (RA) of each species by determining the (Ernst and Lovich, 2009). Unique identifying features, proportion of each species relative to the total number such as damage, scars, or colouration, were noted and of individual turtles captured. We calculated a natural photographed for each turtle and will aid in confirming log Shannon Index (H’) to describe diversity and identity during future surveys. All turtles were weighed Equitability (J’) to estimate heterogeneity. to the nearest g using Ohaus top-loading scales (Ohaus, follows the Turtle Taxonomy Working Group (2017). Parsippany, New Jersey, USA) or to the nearest 50 g for turtles over 5 kg using Pesola spring scales (Pesola Results AG, Schindellegi, Switzerland). The carapace of each turtle was marked by filing a unique notching pattern, a We captured and processed a total of 182 turtles variation of the technique described by Cagle (1939). representing ten species and 11 taxa at Weeki Wachee All captured turtles with CL > 70 mm were injected Springs State Park (Table 1). Two taxa constitute new with a passive integrated transponder (PIT) tag records for Hernando County according to records of (Biomark, Boise, Idaho, USA). PIT tags were inserted the Florida Museum of Natural History (FLMNH). under the right bridge of the shell, anterior to the right Photographic vouchers were submitted to the FLMNH rear leg. This area is considered an acceptable site for to be catalogued. New records include one T. s. scripta PIT tag retention (Runyan and Meylan, 2005). PIT tag (FLMNH 175364) and one T. s. elegans (FLMNH insertion methods were described in greater detail by 175366). The most frequently captured species was 606 Eric C. Munscher et al.

Table 2. Comparison of Species Richness (S), Shannon Diversity (H’), and Equitability (J’) of turtles among five spring-fed lotic ecosystemsTable 2.in ComparisonFlorida. ���������������������������������������������������������������������������������������������������� of Species Richness (S), Shannon Diversity (H’), and Equitability (J’) of turtles among five spring-fed lotic and Meylanecosystems (2011), d in= Florida.Riedle et References al. (2016), for and the e =different Johnston localities et al. (2016). include All a =studies this study, used bthe = Meylanmethod etof al.hand-capture (1992), c = whileChapin and Meylan snorkeling.(2011), d = Riedle et al. (2016), and e = Johnston et al. (2016). All studies used the method of hand-capture while snorkeling.

Number of Individuals Location (Dates) S H’ J’ Captured (n) aWeeki Wachee Springs (2015) 182 10 1.33 0.58 bRainbow Run (1990) 444 6 0.91 0.51 cIchetucknee River (2007) 388 7 1.18 0.61 dVolusia Blue Spring (2007–2014) 542 10 1.25 0.54 eSanta Fe River (2006–2012) 1096 10 0.921 0.40

1 H’ = mean H’ of 10 sampling sessions as reported by Johnston et al. (2016)

S. odoratus (n = 101, RA = 0.555) followed by P. abundance for this species at Weeki Wachee Springs peninsularis (n = 41, RA = 0.225) and P. nelsoni (n = was 0.555. In comparison, relative abundance for S. 20, RA = 0.11) (Table 1). Species richness was 10, the odoratus in other habitats ranges from 0.002 in Santa Shannon Index (H’) was 1.33, and Equitability (J’) was Fe River to 0.25 in Rainbow Run (Table 2) (Meylan 0.5786 (Table 2). et al. 1992; Johnston et al. 2016). The highest reported density of S. odoratus in a Florida spring run is 106 Discussion turtles/ha in Rainbow Run (Meylan et al., 1992), but this species is consistently less abundant than the congeneric At Weeki Wachee Springs State Park, the most Loggerhead Musk Turtle (S. minor minor) at study sites abundant large river turtle was P. peninsularis, next where the two species co-occur (Carr, 1940; Meylan was P. nelsoni, and third was P. c. suwanniensis. Both P. peninsularis and P. nelsoni are considered common et al., 1992; Chapin and Meylan, 2011; Munscher et within their ranges (Jackson, 2006; Thomas and Jansen, al., 2015c; Johnston et al., 2016). Relative abundance for S. m. minor at Weeki Wachee Springs was 0.011, 2006), whereas P. c. suwanniensis has been a species 2 of special concern in Florida for many years and has while at our comparison sites relative abundance ranged suffered population declines (Carr, 1940; Heinrich et al., from 0.280–0.662 (Table 1). Munscher et al. (2015c) 2010, 2015). Interestingly, Carr (1940) observed that P. suggested that S. odoratus was unable to compete nelsoni “are rarely found in numbers where floridana with the larger, more aggressive S. m. minor, and as (= P. peninsularis and P. c. suwanniensis) is abundant,” such is typically found in reduced numbers or pushed which is contrary to our findings in this diverse spring to marginal habitat. Weeki Wachee Springs is near the habitat. All three Pseudemys species (P. c. suwanniensis, southern boundary of the known range of S. m. minor. P. nelsoni, P. peninsularis) occur together in the Santa We captured only two S. m. minor during this initial Fe River (north of Gainesville), sampling session, suggesting that the habitat here is not (northwest of Gainesville), and Rainbow Run (~120 km favourable for this species. north of Tampa Bay) (Meylan et al., 1992; Chapin and Weeki Wachee Springs has a rich freshwater turtle Meylan, 2011; Johnston et al., 2016). Published data on fauna. We were able to document the presence of nine of relative abundances of these species in these locations the ten freshwater turtle species whose native geographic differ from those observed at Weeki Wachee. Of note ranges include Weeki Wachee Springs (Krysko et al., is that each spring system seems to have a dominant 2011). We failed to find the (K. Pseudemys and Sternotherus species. steindachneri) during this initial survey, but this was One interesting result of this survey was that the most expected. Springs are not considered a preferred habitat frequently captured turtle was S. odoratus, a species that type for this turtle (Carr, 1952; Ashton and Ashton, 1985; is often captured in low numbers at other freshwater Meshaka and Gibbons, 2006), and we have never found spring sites in Florida (Marchand, 1942; Chapin and K. steindachneri in any Florida spring system during our Meylan, 2011; Munscher et al., 2015a, c). The relative 17 years of surveys. In addition, T. s. scripta is native to Turtle survey of Weeki Wachee Springs State Park 607

Florida, but not native as far south as Hernando County area offered to our group were amazing. Thanks to Curt Reilly (Krysko et al., 2011). It should therefore be considered at Paddling Adventures for the invaluable loan of canoes, which introduced at Weeki Wachee Springs. However, given made sampling of the Weeki Wachee River possible. We also thank the FDEP for permitting us to sample in the park (permit that this taxon is native to Florida, we did not remove # 06201310-C). it from the spring system. All non-native species, including the new county record for T. s. elegans, were removed from the site in accordance with Rule 68-5.001 References (Florida Administrative Code). Ashton, R.E., Jr., Ashton, P.S. (1985): Handbook of Reptiles and The species composition and relative abundances Amphibians of Florida. Part II, Lizards, Turtles and Crocodilians. (RA) of each species (Table 1) as well as the diversity Miami, Florida, USA, Windward Publishing. 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Tallahassee, Florida, USA, Florida Fish and Wildlife Conservation Commission. and J’ were similar among sites (Table 2). Weeki Wachee Ballou, A.R., Deyle, A.C., Jackson, D.R. (2016): Pseudemys Springs and Volusia Blue Spring had the highest species concinna suwanniensis. Geographic Distribution. Herpetological richness; however, species richness for both of these Review 47: 253–254. systems was inflated by the presence of one non-native Cagle, F.R. (1939): A system for marking turtles for future species at Weeki Wachee Springs and three non-native identification. Copeia 1939: 170–173. species at Volusia Blue Spring. Weeki Wachee had the Carr, A. (1940): A contribution to the herpetology of Florida. University of Florida Publications, Biological Science Series 3: highest diversity of native species (n = 9) (Table 2). 1–118. Clearly, more than mermaids inhabit this iconic Carr, A. (1952): Handbook of Turtles. Ithaca, New York, USA, habitat of Florida. 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Baltimore, Maryland, USA, Johns whether continued urbanisation, eutrophication, or Hopkins University Press. water loss have long-term effects on this diverse spring ESRI (2011): ArcGIS, version 10.3.1. Redlands, California, USA, Environmental Systems Research Institute. system. FitzPatrick, K. (2010): Weeki Wachee Springs: Analysis and Restoration Proposal. Available at http://floridaspringsinstitute. Acknowledgements. The Turtle Survival Alliance-North org/Resources/Documents/Weeki_FitzPatrick.pdf. Accessed on American Freshwater Turtle Research Group (TSA-NAFTRG), 11 December 2015. would like to express its gratitude to all of the volunteers who Florida Department of Environmental Protection (2013): Florida joined us for this survey, including Ben Williams, Wayne State Parks to Celebrate Third Gold Medal Award. Available Osborne, Jessy Wales, Ande Williams, Irene Gaz, Megan at https://www.content.govdelivery.com/accounts/FLDEP/ Burger, Derek Richards, Garrett Mueller, Jennifer Weir, Jennifer bulletins/8ee52a. Accessed on 11 December 2015. Beckett, and Nason Beckett. Funding was provided by Peninsula Florida Department of Environmental Protection (2014): Final College, the Peninsula College Foundation, Huxley College TMDL Report: Nutrient TMDLs for Weeki Wachee Spring and of the Environment, and Western Washington University. We Weeki Wachee River (WBIDs 1382B and 1382F). Available would like to specifically thank George L. Heinrich for his kind at http://www.dep.state.fl.us/water/tmdl/docs/tmdls/final/gp5/ and helpful review. We thank the employees of Weeki Wachee WeekiWachee-nutr-TMDL.pdf. Accessed on 11 December Springs State Park for their hospitality – the house and camping 2015. 608 Eric C. Munscher et al.

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Accepted by Hinrich Kaiser