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Home , Kinosternidae, Loggerhead musk turtle, Sternotherus, Sternotherus odoratus

Herpetology Notes, volume 14: 1043-1049 (2021) (published online on 03 August 2021)

Movements of minor (Testudines: ) in a Central Spring-run

Stephen J. Enders1,2, Andrew S. Weber1,2,3, Brian P. Butterfield4, Andrew Walde1, Ayla Ross5, Brian Hauge1,6, Tabitha Hootman1, Wayne Osborne1,7, and Eric C. Munscher1,8,*

Abstract. Movement patterns of can vary depending on natural and anthropogenic pressures and can correlate with sex, size, food availability, reproduction, territorial behaviour, seasonality, location, and habitat. We analysed the movements of Loggerhead Musk Turtles (Sternotherus minor) based on body size, sex, time between captures, and capture segments in Wekiwa Springs State Park, Florida, of America, over a period of six years using capture, mark, recapture sampling. To record movements, we divided the 1 km spring-run into ten 100 m long segments. Individuals with two or more data entries of segment locations were used for this study, with each paired capture/recapture constituting one movement event. A total of 133 movement events were recorded, and the majority, 60.9% (81), showed no inter-segment movement; 88.0% (117) of the movement events were one segment or less, a maximum distance of 200 m. Using canonical discriminant analysis we found no relationship between the movement of S. minor and carapace length, sex, time between captures, and capture segments. More research is needed into the movement ecology of this . Based on the observed paucity of movements, future work may focus around dividing the spring-run into shorter segments. The potential for territoriality and dominance hierarchies should also be examined. We strongly recommend the use of radio telemetry to learn more about the nuanced movements of this small turtle.

Keywords. , Sternotherus minor, movement, spring-run

Introduction environment. There are many natural reasons for movement of an , such as food and water Movement is a core behaviour necessary to the availability, seasonal shifts, temperature regulation, and survival of many animal species. For the purposes reproduction (Heape, 1931). Anthropogenic factors, of this research, we define movement as a change such as artificial light, noise, and development are also in position of the entire body of an organism in the known to influence animal movement (Thums et al., 2016; Cayuela et al., 2020). The same triggers, both natural and human-induced, have also been shown to 1 Turtle Survival Alliance, 1030 Jenkins Road, Suite D, motivate movement in turtles (Cagle, 1944). Studies Charleston, South Carolina 29407, USA. have shown a wide variation in magnitude and types 2 TheTurtleRoom, P.O. Box 521, Lititz, Pennsylvania 17543, of chelonian movements which include: sea turtles USA. that can travel thousands of kilometres annually (Hays 3 National Park Service, 274 River Road, Beach Lake, Pennsylvania 18405, USA. and Scott, 2013); Mojave Desert ( 4 Freed-Hardeman University, 158 East Main Street, Henderson, agassizii [Cooper, 1861]) with home ranges between 38340, USA. 5-236 ha (Harless et al., 2009, 2010); Box Turtles 5 Department of Biology, Indiana University of Pennsylvania, (Terrapene spp.) with a home range of ca. 4 ha (Habeck Indiana, Pennsylvania 15705, USA. et al., 2019); Smooth Softshell Turtles ( mutica 6 Department of Biology, Peninsula College, 1502 East [LeSueur, 1827]) with a linear home range of ca. 500 Lauridsen Boulevard, Port Angeles, Washington 98362, m (Plummer and Shirer, 1975); and Florida Red-bellied USA. 7 Pine Ridge High School, 926 Howland Boulevard, Deltona, Cooters ( nelsoni Carr, 1938) with a home Florida 32738, USA. range of only 120 m, despite the large size of the turtle 8 SWCA Environmental Consultants, 10245 West Little York (Kramer, 1995). These species-specific movements are Road, Suite 600, Houston, 77040, USA. important characteristics to take into account for further * Corresponding author. E-mail: [email protected] research activities or for effective, targeted conservation © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. measures. 1044 Stephen J. Enders et al.

Despite the importance of the study of movement S. minor movements in a spring-run system, and lead patterns for chelonian conservation and survival, to a better understanding of the requirements of the many species are still relatively understudied (Lovich species. and Ennen, 2013; Munscher et al., 2020), including those within the Sternotherus (musk turtles), Materials and Methods with the possible exception of the Common Musk Field Site. This study took place in Wekiwa Springs Turtle ( [Latreille in Sonnini Run (28.714°N, 81.453°W; WGS 84) within Wekiwa and Latreille, 1801]). Most movement and home range Springs State Park in Orange and Seminole Counties, research on Sternotherus has focused on the S. odoratus in Florida, as part of a larger monitoring effort (see with a few studies on the Razor-backed Musk Turtle (S. Munscher et al., 2013, 2015a, b, 2020; Walde et al., carinatus [Gray, 1856]) and the Stripe-necked Musk 2016). Wekiwa Springs Run is a 1 km section of spring- Turtle (S. peltifer Smith and Glass, 1947), the latter was run habitat connecting the main lagoon and Rock Springs formerly considered a of the Loggerhead Run to the Wekiva River. We divided the spring-run into Musk Turtle (S. minor [Agassiz, 1857]). Examination ten 100 m long segments (Fig. 1). of the movement and home range of S. odoratus has Data Collection. We sampled from August 2010 to shown it to typically exhibit limited movement in lentic March 2016, twice per year (spring and summer) for (Andres and Chambers, 2006; Holinka et al., 2003; a time period of 3–5 days. Loggerhead Musk Turtles Smar and Chambers, 2005) as well as lotic habitats (Fig. 2) were captured via hand while snorkelling and (Mahmoud, 1969; Belleau, 2008; Wilhelm and Plummer, placed in containers marked with the number of the 2012). Sternotherus carinatus also demonstrates corresponding segment of capture. Snorkelling occurred limited movements, with a preference for lotic habitats intermittently during daylight hours. Captured turtles (Mahmoud, 1969; Kavanagh and Kwiatkowski, 2016). were marked using a variation of the Cagle (1939) While home range of the S. peltifer showed that most method. Turtles with straight-line maximum carapace individuals used multiple habitats within their home length (CLmax) ≥ 70 mm were implanted with 8.5 range (Ennen and Scott, 2013) and displayed site mm passive integrated transponders (BIOMARK, fidelity within daily movement patterns (Ennen and Boise, Idaho, USA) containing unique identification Scott, 2008). In the case of S. minor, to the best of our numbers. Turtles were sexed using secondary sexual knowledge, this is the first study describing movement characteristics of size and cloacal vent location. patterns or home ranges of S. minor. Only turtles less than 50 mm CLmax were coded as Sternotherus minor is a small musk turtle, with juvenile if sex could not be otherwise determined. If we similarly sized sexes, that ranges throughout most of coded an individual as juvenile upon initial capture, but south of the Appalachian Mountains, across sex could be determined at recapture, the sex variable much of the northern part of Florida, and far southeastern was recoded with the recapture sex. (Ernst and Lovich, 2009). In Florida, S. minor Analysis. A “movement event” was defined as the occurs in a variety of habitats including freshwater distance moved, in segments (used as a unit), between springs, spring-runs, lagoons, lakes, , and river two capture events of an individual. If the variable days systems (Zappalorti and Iverson, 2006; Reidle et al., between captures was less than five, those events were 2016; Munscher et al., 2020) showing preference for removed. This left us with 133 movement events with lotic habitats (Tinkle, 1958; Zappalorti and Iverson, the shortest time between capture at 225 days and the 2006). longest at 715 days. Florida is high in turtle diversity, second to only Canonical discriminant analysis was chosen because Alabama worldwide, and spring-run systems are known we desired a categorical response (did the turtle move to provide habitat for a variety of species, making them segments or not?) with both categorical and continuous well-suited for sustained research efforts (Meylan et al., input variables. Variables examined included: days 1992; Mittermeier et al., 2015; Munscher et al., 2020). between captures (days), sex, carapace length at initial Populations of Sternotherus have been sampled as part capture (CL1), carapace length at recapture (CL2), of a long-term, multispecies monitoring effort over the change in carapace length (CL change), segment of last 20 years by the Turtle Survival Alliance–North capture (CS1–9), and segment of recapture (RCS1– American Freshwater Turtle Research Group (Munscher 9). Statistical tests were performed using Statistical et al., 2020). The purpose of this study is to characterise Analysis Systems software (SAS Institute, 2018). Movements of Sternotherus minor in a Central Florida Spring-run 1045

Figure 1. A map of a section of Wekiwa Springs State Park, showing the main lagoon and Wekiwa Springs Run divided into ten segments, each representing a scale of 100 m long (LABINS, 2019)

Results Of the total movement events sampled (n = 133), 60.9% (n = 81) involved a distance of zero segments (Table 1) and 88.0% (n = 117) involved a distance of zero or one segments, meaning that these movements were less than 200 meters in stream length (Table 1). The longest male movement was seven segments, and the longest female movement was five segments (Table 2). The F-value of the resulting discriminant function was 1.31 (df: 22, 106) with a p-value of 0.1848. Therefore, no single variable nor function involving any combination of the examined variables explained the observed movement patterns.

Discussion

Figure 2. A Loggerhead Musk Turtle (Sternotherus minor) In this study, the movements of individual S. minor basking on a branch in Wekiwa Springs State Park, Florida, were typically contained within one to two segments USA. Photograph by Joanne Boleman. of spring-run habitat. We did have one turtle with an 1046 Stephen J. Enders et al.

Table 1. Number of segments that individual Loggerhead survival, or reproduction may exist for unintentionally Musk Turtles (Sternotherus minor) moved between captures in translocated S. minor that are not released where they Wekiwa Springs Run in Wekiwa Springs State Park, Florida. were captured. Most (79%) S. odoratus, have been found to be recaptured in the same location as original

Number of segments capture (Holinka et al., 2003). They have also been found to move more often once displaced; however, 0 1 2 3 4 5 6 7 8 9 Total 29% were still recaptured at the site of release (Andres Male 30 12 3 1 0 1 0 1 0 0 48 and Chambers, 2006). Although Smar and Chambers Female 48 22 4 4 1 1 0 0 0 0 80 (2005) found that the majority (64%) of displaced Juvenile 3 2 0 0 0 0 0 0 0 0 5 individuals were recaptured at the site of most recent release, they found homing behaviour to be male-biased. Total 81 36 7 5 1 2 0 1 0 0 133 Similar results of increased movements for translocated

Table 2. and male-biased homing have been observed in other chelonians (Hinderle et al., 2015). We did not extreme movementLongest movement of seven Longest to eight time segmentsbetween Longestand two CL Shortest CL CL Mean; SD (segments) captures (days) (mm) observe(mm) differences(mm) in movement between the sexes in with movements between five and six segments, but our study. The larger movement patterns of male Sliders Male 7 2053 115 64 85.7; 14.7 clearly these are rare movement events for S. minor. We ( scripta [Thunberg in Schoepff, 1792]) are observedFemale no significant5 differences1577 between the sexes.129 thought59 to increase90.0; 13.1 opportunities for mating (Morreale InJuvenile both males and1 females, 88.0%1439 of events included62 et al.,48 1984; Parker,55; 5.3 1984; Tuberville et al., 1996) and movements of zero or one segments. decrease competition (Smar and Chambers, 2005). Movement activity within other members of the genus Males have also been found to be more likely to travel Sternotherus is relatively limited (Table 3), and our to new areas (Tuberville et al., 1996). study suggests similar results for S. minor. Exceptions The biggest limitation of this study is the coarse are the relatively large home ranges reported by Belleau resolution of the survey segments. Smaller survey (2008) and Bennett et al. (2015), near the northern range segments may have provided greater insight into limit for S. odoratus. Although S. minor movement in which variables led to movement or lack of movement. our study was limited, we did observe some individuals Considering the study methodology, it is possible that make movements of up to seven to eight segments. a movement of one segment was just several meters in These larger movements could have been made by length if a turtle lived near the segment border. It should dispersing individuals; however, none of the variables also be noted that specimens were observed to move we examined (as discussed in the methods section) into and out of the survey area and lagoon and could accounted for these movements. have done the same at the downstream end of the study Our study reveals the relatively sedentary nature of S. area; however, since the lagoon is relatively large, and minor and highlights the need for captured animals to be we only have that relatively coarse area as a capture released at the point of capture. A reduction in apparent location, we cannot make any furthers statements about survival has been observed for translocated Gopher these movements, other than to say they happened. A Tortoises (G. polyphemus [Daudin,Number 1801]; of segments Tuberville variable we did not use, and did not have data for, but et al., 2008), and disruption,0 1 2 or 3 exclusion4 5 6 from7 8 might9 Totalbe interesting to examine in future work would reproduction has been noted in another study (Mulder be population density of both the initial capture segment Male 30 12 3 1 0 1 0 1 0 0 48 et al., 2017). It is unknown if similar reductions in and recapture segment. It is possible that the observed Female 48 22 4 4 1 1 0 0 0 0 80 1 Juvenile 3 2 0 0 0 0 0 0 0 0 5

Table 2. Assorted Totalminimum 81and 36maximum 7 5 data 1 of 2 Loggerhead 0 1 0 Musk 0 Turtles 133 (Sternotherus minor) in Wekiwa Springs Run in Wekiwa Springs State Park, Florida. CL (carapace length). Table 2.

Longest movement Longest time between Longest CL Shortest CL CL Mean; SD (segments) captures (days) (mm) (mm) (mm) Male 7 2053 115 64 85.7; 14.7

Female 5 1577 129 59 90.0; 13.1

Juvenile 1 1439 62 48 55; 5.3

1

Movements of Sternotherus minor in a Central Florida Spring-run 1047

Table 3. A selected review of movement variables for the genus Sternotherus, by species, sex, and habitat.

Species Sex Habitat Variable of Movement Citation Average distance Mean aquatic Mean home Mean maximum between captures linear home range range (ha) distance moved (m) (m) (m) Sternotherus odoratus female mixed -- -- 20.9 -- Belleau, 2008 male mixed -- -- 25.4 -- female lotic -- -- 25.5 -- Bennett et. al, 2015 male lotic -- -- 105.85 -- female mixed 89.5 -- 0.94 -- Ernst, 1986 male mixed 117.3 -- 1.75 -- female lotic 44.5 -- 0.05 -- Mahmoud, 1969 male lotic 67.6 -- 0.02 -- both lotic -- 176 0.15 -- Wilhelm and Plummer, 2012 Sternotherus carinatus female lotic1 -- 334.00 -- 234.23 Kavanagh and Kwiatkowski, 2016 male lotic1 -- 109.67 -- 82.96 female lotic2 -- 369.12 -- 307.24 male lotic2 -- 333.00 -- 287.95 female lotic 17.4 ------Mahmoud, 1969 male lotic 38.6 ------Sternotherus peltifer female lotic -- 346 0.52 -- Ennen and Scott, 2013 male lotic -- 335.2 0.52 -- 1 1 and 2 denote different sites reported on by Kavanagh and Kwiatkowski, 2016.

movements had the purpose to leave more densely Walton, and the many other students and biologists who make up populated areas. the Turtle Survival Alliance-North American Freshwater Turtle This study demonstrates that S. minor movements are Research Group. A special thank you to Deborah Shelly, Oros, and Barbara Howell from the FDEP Aquatics Preserve typically small and suggests the needs of an individual 1 for their years of constant support in and out of the field. Their – food sources, water sources, seasonality, temperature, contributions were truly impactful to this long-term study. We and breeding – are met within a limited home range. This thank James Fenwick from Millersville University for talking could mean that the habitat is particularly good, and thus through the options for statistical analysis. We thank Jeff Stein large movements are not necessary (Harless et al., 2010). from SWCA Environmental Consultants for his help in preparing Alternatively, well defined home range areas are often Figure 1. We thank the Friends of the Wekiva River Foundation, associated with territories and defined social hierarchies Wekiva Wild and Scenic Committee, Guy Marwick, The Felburn Foundation, and Keep Seminole Beautiful for providing much- but may not always be the primary determinants of space needed grant money over the many years of this study. Additional use (Harless et al., 2009). However, other studies have thanks go to SWCA Environmental Consultants for their shown the attributes of a turtle species that contribute constant support and to Alice Bard of the Florida Department of to home range size have been difficult to determine, Environmental Protection for issuing research permits for the past although energy loss associated with movement through 20 years. The study was conducted under permit #06240913 from different habitats likely plays a major role (Slavenko et the Florida Department of Environmental Protection and Permit al., 2016). For future research, we strongly recommend #LSSC-09-0411 from the Florida and Wildlife Conservation Commission. the use of radio transmitters to learn more about the nuanced movements of this small turtle. References

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Accepted by Jindřich Brejcha