Ninja Turtles: high mobility and successful passage through common barriers to movement in a semi-terrestrial freshwater turtle

Ninja Turtles: high mobility and successful passage through common barriers to movement in a semi-terrestrial freshwater turtle | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Ninja Turtles: high mobility and successful passage through common barriers to movement in a semi-terrestrial freshwater turtle James M Dowling, Eric J Nordberg, Deborah S Bower This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3855993/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Anthropogenic landscape change due to urbanisation, agriculture and resource extraction inevitably results in linear barriers within the landscape. Artificial linear structures such as roads, fences, levees, and dams limit the movement of some species and further fragment residual habitat. In this study, we investigated the ability of Eastern long-necked turtles ( Chelodina longicollis ) to cross various terrestrial obstacles commonly encountered in their habitat. We tested two types of fences (chicken wire and exclusion fencing) commonly used in agricultural systems and three sizes of rocks (gravel, gabion, and large boulders) often used for road construction, erosion control, and waterway stabilisation. We examined the success rates of turtles in crossing obstacles, the effect of fatigue on crossing attempts, and the impact of individual boldness on movement behaviour. Turtles displayed high success rates in crossing gravel (85.4%), gabion (86%), boulders (73.3%) and hinged joint exclusion fencing (94.7%). Chicken wire style wire netting had no successful crossings (0%) despite 276 attempts. A significant fatigue effect was observed throughout the experiment, with turtles making an average of 3.94 (± 4.93 SE) fewer attempts at the end of the experiment (day eighteen) as opposed to day one. Bolder turtles were faster at crossing obstacles, however, boldness had no bearing on obstacle-crossing success. These results highlight the need for thoughtful selection of waterway and wetland infrastructure and the fatiguing impact of constant exposure to anthropogenic barriers for wildlife. Agriculture Chelodina longicollis Fence Livestock Riparian Roads Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Changing land use for agriculture, urbanisation and resource extraction threaten species by fragmenting suitable habitats (Aronson et al. 2014 ; Liu et al. 2019 ; McKinney 2002 ; Pattison et al. 2016 ). Drastic land conversion for cropping (Serneels and Lambin 2002 ), livestock grazing (Wegner and Merriam 1979 ), and open-cut mining (Cristescu et al. 2016 ; Young et al. 2018 ) create inhospitable habitats that many species cannot cross. In addition to habitat fragmentation, barriers in the form of roads (Forman 1998 ; Forman and Alexander 1998 ; Colino-Rabanal and Lizana 2012 ; Rytwinski and Fahrig 2015 ) and fences create obstacles for many terrestrial species. For species averse to urbanisation, the negative population-level impacts of roads and fences are more pronounced, often further endangering already threatened species (Taylor and Goldingay 2004 ; Seidler et al. 2015 ; Jakes et al. 2018 ). Species that disperse or migrate, such as semi-aquatic species using a complex of wetlands, are particularly sensitive to barriers (Roe et al. 2006 ; Glista et al. 2007 ; Balčiauskas et al. 2022 ). In particular, freshwater turtles often travel frequently between water bodies and are negatively impacted by these artificial barriers (Aresco 2005a ; Clinton Gooley 2010 ; Hamer et al. 2015 ; Proulx et al. 2014 ; Santori et al. 2018 ). Many freshwater turtles have large home ranges (Slavenko et al. 2016 ) and use water-adjacent land for nesting (Bodie 2001 ; Petrov et al. 2018 ; Van Dyke et al. 2018 ), thermoregulation (Chessman 2020 ), and to travel between water bodies (Graham et al. 1996 ; Serrano et al. 2020 ). This dependence on terrestrial habitat exposes these slow-moving and ungainly species to obstacles that may prevent movement or create ecological traps (Ferronato et al. 2014 ). Despite a preference for aquatic habitats, many turtle species climb terrestrial obstacles such as logs, concrete slabs (Scott et al. 2023 ), and other individuals (e.g., turtles and crocodiles; Lindeman 1999 ; McKnight et al. 2023 ). As such, the impact of fences on turtle populations is a nuanced issue, providing some positive outcomes for species while threatening others. Fences can benefit populations in urbanised areas by funnelling individuals towards safe road crossing structures (Aresco 2005b ) or protecting turtles or nesting beaches from introduced predators, like red foxes ( Vulpes vulpes ; Streeting et al. 2023 ). However, fences can also kill turtles due to entanglement and overheating (Ferronato et al. 2014 ) or indirectly through prey funnelling for predators (Little et al. 2002 ). Additionally, linear barriers dissuade turtles from moving between seasonal habitats and may alter movement routes and increase metabolic costs while reducing available resources (Paterson et al. 2019 ). Widespread agricultural and urban development in south-eastern Australia may create many obstacles for preferentially aquatic turtles, such as the Eastern long-necked turtles ( Chelodina longicollis ). These turtles travel over 3 km overland to exploit resource-rich ephemeral waterbodies away from main river channels (Roe and Georges 2008 ), where their risk of encountering anthropogenic barriers, such as roads, fences, or rocky erosion mitigation strategies is high. However, despite the widespread introduction of quarried rock used for erosion control, levees, and retaining walls along waterways along wetland habitats, the ability of Eastern long-necked turtles to navigate these obstacles is unknown. These structures effectively prevent erosion (Keller 2004 ) and provide additional habitat for invertebrate prey species (Shields 1991 ; Fischenich 2003 ), but may act as a barrier for terrestrially moving species. Here, we test the ability of C. longicollis to cross two common styles of livestock fencing and three sizes of rock, commonly used around waterways for road and levee construction and erosion mitigation. Specifically, we compared 1) the success rates of C. longicollis crossing fence and rock obstacles to an obstacle-free environment, and 2) tested the effect of fatigue and boldness on the willingness of turtles to approach obstacles. Methods Study area Our experiment occurred at the University of New England’s Newholme SMART Farm (-30.419, 151.637) in the Armidale region of NSW, Australia between 5–22 March 2023 (Fig. 1 ). Eighteen wild-caught adult Eastern long-neck turtles (Chelodina longicollis; straight carapace lengths > 150 mm) with no physical abnormalities (e.g., missing limbs) were used in the experiment. Morphometric data was collected on all individuals, including straight carapace length and weight. Experimental design The experimental mesocosms comprised of outdoor, fully shaded, enclosures covering an area of 180 m2 with eighteen 5 m x 2 m arenas, each equipped with a polypropylene pool (Foundation Products Clam Shell Sandpit (945 x 210 x 1100 mm) at the southern end (Fig. 2 ). The individual pens were constructed using 0.9 m wide silt fencing material for the walls, providing a wall height of 0.8 m once entrenched to prevent turtles from burrowing beneath the fence. Additional fencing material was placed above intersecting fences to prevent turtles from climbing the corners. Each arena was allocated as control or fitted with one of five obstacle treatments based on common features in the landscape (Fig. 3 ). Obstacles consisted of 1) aggregate gravel, a common road base in rural areas; 2) gabion, a medium (60–150 mm) crushed rock; 3) large boulders (> 300 mm), which are both used for erosion mitigation and riverbank stabilisation; 4) hinged joint exclusion fencing (15/150/15); or 5) “chicken wire” wire mesh netting due to their prevalence and popularity in agricultural system and ecosystem rehabilitation areas. Daily rainfall data was collected via a rain gauge attached to the enclosure and temperatures from a Bureau of Meteorology observation station 12.2 km southeast (Armidale Tree Group BOM observation station). At 09:00 h, turtles were randomly allocated to an initial arena without replacement where they were placed in a pond of fresh water (Fountain Products Clam Shell Sandpit) at the southern end of the arena for a one-hour acclimatisation period and provided with food (commercial turtle food; Fish fuel Co. “Turtle Food” 3 g frozen blocks). At 10:00 h, each turtle was moved and released on the northern side of the obstacle and given 23 hours to cross the obstacle and return to the pond containing the food and water. Turtles were exposed to each testing arena only once, but they encountered each obstacle (in random order) three times throughout the whole trial (18 testing areas, with 3 replicates each of 6 different treatments). Assessing movement, boldness, and fatigue Testing arenas were monitored via time-lapse wildlife cameras (Campark T85; 1-min photo interval) to record the movement of turtles throughout the experiment. The time stamp on the photos allowed for me to calculate the time and the duration of each crossing attempt. We considered a turtle to have made an "attempt" at crossing when it moved to within a body length of the obstacle. Attempts were considered successful if the individual crossed and exited the opposite side of the obstacle. Unsuccessful attempts were recorded if an individual engaged with the obstacle but failed to reach the other side. The outcome of each trial was determined as either successful or unsuccessful, dependent on whether the turtle crossed over the obstacle to the other side. If no attempt was made, the trial was classified as "no attempt". Boldness was measured as the time taken, in minutes, for turtles to make their first movement in each trial. The number of attempts made per trial was used to examine the effect of fatigue on turtles willingness to attempt obstacles. Statistical analysis We used generalised linear mixed effect models to analyse relationships between various response variables and the fixed effects: obstacle treatment (grass, gravel, gabion, rock, boulder, exclusion fencing, chicken wire), trial number (1–18), and straight carapace length (SCL). Individual turtle ID was included as a random effect to account for variation among individuals. To analyse obstacle crossing success we used a binary response variable (success = 1, failure = 0) with the logit link function. To evaluate the impact of fatigue on the turtles' performance we used the number of attempts per trial and the logit link function. To explore the influence of individual boldness on the turtles' movement patterns, we used the time (latency) to the first movement as an indicator of boldness (Kabelik et al. 2021 ; Talavera et al. 2021 ) and the logit link function. The significance of variation in individual boldness was assessed using the ANOVA function in the car package (v3.0.12) with a type II sum of squares. We used estimated marginal means for post hoc testing for all GLMM and GLMER. All statistical analyses were conducted using R statistical software (version 2023.03.0 + 386). The specific packages used for analysis included "stats", “emmeans”, and "lme4" for the GLMM, GLMER and post hoc testing. All results are presented as mean ± SE unless otherwise stated. Results Obstacle crossing Collectively, individual turtles attempted to cross the obstacles 1,545 times throughout the 18-day study. Non-attempts occurred in seven instances from 6 of the 18 individuals. Turtles successfully crossed a range of obstacles. Gravel, gabion, boulders, and exclusion fencing were all crossed by all turtles at least once, with varying degrees of success. Attempts to cross the control arenas were successful in 100% of attempts. In contrast, chicken wire was attempted 298 times with a 0% success rate. Exclusion fencing had the highest success rate among the obstacles with 94.7% crossings from 228 attempts. While crossed by all turtles at least once, boulders exhibited the second lowest rate of success per attempt with 73.3% crossings from 236 attempts. Gravel and gabion had notably high success rates, with an 85.4% success rate from 308 gravel attempts and 86% of the 215 gabion attempts resulting in success. Turtle body size (SCL) did not predict obstacle-crossing success (R2 = 0.002, F 1,82 = 0.721, p = 0.230). Obstacles that were crossed more often, were also crossed more quickly. The time taken to cross obstacles was lowest for exclusion fencing, taking an average of 1.56 ± 2.21 min (mean ± SE) followed by gravel (1.91 ± 2.09 min), gabion (2.84 ± 4.39 min), and boulders (6.85 ± 13.40 min). Despite having no success, turtles attempted to cross chicken wire for 35.64 ± 47.91 min per attempt before resting, with the longest continuous attempt lasting 595 min. Recognition of mesh fences as an obstacle remains somewhat uncertain for C. longicollis. Turtles appeared to routinely “follow their head” through obstacles, resulting in unnecessary climbing through the second row of exclusion fencing despite the turtle’s body size permitting easy movement through fence gaps at ground level. For example, C. longicollis typically walk with their head elevated above the ground so, when they encounter a fence, if the head enters the mesh gap on the second row, the turtle will climb through that opening rather than ducking its head to easily walk through the lowest mesh gap (Fig. 4 ). This tendency became more pronounced for chicken wire obstacles, where turtles would place their head and neck through the chicken wire and attempt to push their body through the mesh (which was far too small for their whole body), often for several hours or several attempts. Turtles were also observed using their heads and necks to routinely climb boulder obstacles and cross large gaps. Turtles extended their necks before using their heads as a point of contact, allowing for the bridging of obstacles. In addition to crossing obstacles, turtles climbed the walls of their enclosure. Two individuals scaled the 800 mm high woven polyester silt fencing and entered the pools of neighbouring arenas during a pilot study. Turtles wedged themselves in the vertex of the corners of the silt fencing and climbed walls using their claws to penetrate the weave of the silt fencing fabric (Fig. 5 ). The fence was modified with an additional overhang to prevent this (not pictured). Fatigue effect There was a negative linear relationship between number of attempts per trial and trial number. Turtles made 0.218 fewer attempts per trial or approximately one less attempt every five days (t = -4.121, p < 0.001). Turtles attempted obstacles an average of 3.94 ± 4.93 times on day eighteen as opposed to 8.39 ± 5.12 on day one. Boldness Individual boldness varied significantly between turtles (X2 = 431.96, p < 0.001). Boldness was negatively correlated with time taken per attempt (β = -0.14547, p = 0.009), with bolder turtles dwelling on or near obstacles for longer. However, boldness did not significantly affect the success rate (β = 0.007, p = 0.11). Obstacle type, trial number and arena number did not affect the time to first move. Discussion Our study demonstrates that while Eastern long-necked have a remarkable ability to navigate many structures, barriers exist from common landscape features (Coffin 2007 ; Forman & Alexander 1998 ; Jakes et al. 2018 ; Jones et al. 2022 ; McInturff et al. 2020 ; Smith et al. 2020 ; Trombulak & Frissell 2000 ). The obstacle materials we tested here are commonly used in road, stream, and wetland restoration or common fence types used by commercial and private landholders and therefore are likely to impact Eastern-long necked turtles throughout much of their range. Chicken wire or similar small-diameter mesh fences presented unpassable obstacles for the sampled eastern long-necked turtles, and this reflects the barrier created for many other wildlife groups as well (Baines & Andrew 2003 ; Bock & Bock 1994 ; Moseby & Read 2006 ; Smith et al. 2020 ). Although other obstacle types were successfully crossed by turtles, the time required to successfully navigate through obstacles was often substantial and this likely affects an energy cost. Crossing success Rocks Turtles crossed rock obstacles with seemingly little trouble, regardless of the size of rock encountered. We expected the sharp, irregular angularity of the quarried boulders to pose a challenge to turtles, particularly smaller individuals with carapace lengths below 160 mm (33.3% of turtles in our study) as boulders were generally taller than this. Yet, 73% of boulder crossing attempts were successful, and multiple crossings of the obstacle in each trial were common. Small turtles in the sample group also used their head and necks as a point of contact to both climb and cross the large boulder obstacles. We expected the smallest obstacles such as gravel and gabion to be crossed far more often, however, gabion had only 12 more crossings (185 of 215 attempts) than boulders (173 of 236 attempts), a more difficult obstacle. Multiple attempts were unsuccessful for both gravel and gabion, where turtles either approached the obstacle and chose not to cross or aborted crossing part-way through. This may be attributed to several factors that were not measured in our experiment. Warm surface temperatures of rocks may explain aborted crossings, as burns and eventual death from hot rocks have been recorded in turtles attempting to gabion railway track beds (Abinesh et al. 2022 ; Kornilev et al. 2006 ). However, the effect of dangerously high temperatures is unlikely to have had a major impact on turtles in our experiment due to the use of shade cloth which shaded the entire mesocosm. Fences Turtle's success at crossing fences was highly dependent on the fence design. The chicken wire style mesh, which was completely impassable for all turtles, is prolific in agricultural regions with sheep. The wire mesh usually extends to ground level to prevent the separation of lambs from ewes or enforce weaning (Galeana et al. 2007 ; McGregor 1990 ) and exclude predators (Kondinin Group 2016 ; Long & Robley 2004 ). Despite the complete failure to cross, chicken wire had the third highest number of overall attempts and the longest amount of time attempting to cross. The average time per crossing attempt was far higher than other obstacles at 35.6 minutes per attempt, likely imparting a high energy cost. Increased energy expenditure may place turtles at elevated risk of mortality due to exhaustion, desiccation and overheating (Ferronato et al. 2014 ). While turtles with roads in their home range have only small increases in energy expenditure (Patterson et al. 2019), the presence of chicken wire-type fences without suitable crossing structures would likely result in turtle exhaustion and death, as has been witnessed in other ’turtle-proof fences’ (Ferronato et al. 2014 ). Exclusion fencing (prefabricated hinge joint) may still pose a risk to C. longicollis which was not identified by our study. Chelodina longicollis of the New England region, particularly Armidale NSW, exhibit irregularly small adult body sizes, (average body sizes (SCL) of females = 239 mm; male = 199 mm; Parmenter 1976 ), which is far smaller than in other regions where SCL may exceed 260 mm (Chessman 1984 ). As such, turtles used in our experiment passed through the ~ 150 mm wide spaces of the exclusion fencing with relative ease. In other regions, the carapace width of C. longicollis may exceed this width, making passage through unmodified exclusion fencing difficult or impossible (Kennett et al. 2009 ). However, simple cost-effective solutions, such as turtle gates (Waltham et al. 2022 ), that permit turtle movement while also maintaining fence integrity have been developed and trialled for other, larger-bodied, freshwater turtles. Although fences may prevent the movement of non-target species, fences are rarely completely intact, with breaks in fences acting as potential crossing points (Jones et al. 2018 ). Gates often act as weak points in fences, with substantial gaps beneath gates existing either through the gate design or wheel ruts from repeated vehicle traffic (Long & Robley 2004 ). Even in high-cost reinforced fences, holes may be created by burrowing species such as rabbits and foxes, compromising fence integrity (Moseby & Read 2006 ). While intermittent gaps may be beneficial for more motile species, freshwater turtles show a significant reduction in their ability to locate gaps within the space of 3m (Waltham et al. 2022 ), making the likelihood of encountering gaps in turtle proof fences unlikely in all but the most dilapidated of cases. The ability of C. longicollis to climb the silt fencing used to divide the arenas demonstrates a remarkable capacity to tolerate certain linear barriers. The loose weave of the silt fencing allowed the turtles to use their claws to climb in the vertex where the edges met at right angles. Turtles were not seen climbing parallel edges, likely due to their sprawling posture and limited mobility from hips sitting inside their shells (Vitt & Caldwell 2013 ). Australian turtles are not typically known for their ability to scale vertical surfaces; however, a considerable ability to climb fences (Aresco 2005a ; Griffin 2005 ), logs (Lindeman 1999 ), and rocks (Davenport et al. 1984 ) have been recorded in several North American turtle species. Fatigue The number of attempts per trial decreased consistently throughout the experiment, resulting in turtles making just over half (46%) of the attempts on the final day as compared to day one. Mean daily movement lengths for C. longicollis are generally below 10 m-1 day − 1 (Roe & Georges 2008 ), yet our subjects exceeded this distance in 60% of trials, likely expending significant energy that led to reduced attempts as trials progressed. Additionally, the tendency for turtles to climb their enclosures reduced with the number of treatments, with no climbing of the mesocosm walls observed after trial five. Therefore, we believe that fatigue rather than habituation towards food supplied in pools was the primary cause of reduced crossing attempts as trials progressed. Boldness As turtles retract into their shells when exposed to potentially threatening stimuli (Greene 1988 ), higher degrees of boldness may reduce the risk of mortality to turtles attempting to cross busy roads because bold individuals cross more quickly. Bold turtles crossed obstacles faster than turtles with lower boldness scores, which suggests that less bold individuals may be more vulnerable to threats such as vehicle strikes or predation as they linger exposed on obstacles. Implications for management Our study highlights the need for selecting wildlife-friendly fence designs when planning and installing fences around wetlands and waterways. Exclusion fencing is manufactured in a range of sizes and designs and populations with larger-bodied turtles may not be as successful at crossing exclusion fencing as in our study. However, simple, cost-effective mitigation methods exist for reducing the threat posed to turtles by exclusion fencing in these scenarios. For example, clipping small sections of vertical exclusion fence wires at regular intervals to permit the movement of larger turtles through exclusion fences incurs minimal cost to landholders while maintaining the functionality of the exclusion fence (Waltham et al. 2022 ). The agricultural sector, particularly the sheep industry, encourages the use of wire mesh fencing, similar to that tested here, for the exclusion of canines and foxes to limit livestock depredation (Kondinin Group 2016 ). Many of these designs encourage wire mesh to be anchored at ground level or installed below the surface to prevent canids from digging under (Green & Gipson 1994 ; Kondinin Group 2016 ). Alternatively, the implementation of species-specific one-way gates, such as those currently used for small mammals to transit through conservation areas (Coates 2013 ), may warrant further investigation for the possibility of turtle-specific modification. Conclusion Our study underscores the significant impact of artificial barriers on freshwater turtle terrestrial spatial use. While turtles can traverse rock barriers with relative ease, depending on rock size, the need to continuously cross rock barriers lead to fatigue. Additionally, rocks larger than those tested here may be impassable completely. Similarly, the impact of fences is dependent on design, with small gauge wire mesh fences preventing the movement of turtles completely. While these results focus on turtles, these trends likely impact numerous small and medium vertebrates across freshwater systems. Wire mesh fences are often used to protect vulnerable species and fragile ecological areas, yet disruptions to movements and mortality due to entanglement or collision are common across birds, reptiles, and mammals. As such, design-specific considerations are crucial for effective wetland and waterway management, necessitating collaboration for wildlife-friendly adaptations to infrastructure in riparian zones to limit unintended negative ecological impacts. In light of this growing body of evidence, it is imperative that wildlife-friendly fencing solutions gain broader acceptance among both land managers and industry stakeholders. Declarations Author Contribution All authors project design and planning.J.D. and E.N. data collection.J.D. and E.N. data analysis.J.D. initial manuscript drafting.All authors editing. Acknowledgements We acknowledge the Traditional Custodians of the land where this research was conducted, the Anaiwan People. We pay our respects to their cultures and Elders, past, present and emerging. Funding for this project was supported through the University of New England. All experiments were conducted under the Animal Ethics permit ARA23-010 through the University of New England and the New South Wales Department of Planning and Environment Scientific License SL102308. Open access publishing facilitated by University of New England, as part of the Wiley - University of New England agreement via the Council of Australian University Librarians. We thank Dr Zenon Czenze for helpful insights and troubleshooting during data analysis and Remo Boscarino-Gaetano for assistance in the construction of the mesocosm. 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Tail autotomy is associated with boldness in male but not female water anoles. Behavioral Ecology and Sociobiology , 75 (2), 44. https://doi.org/10.1007/s00265-021-02982-w Taylor, B. D., & Goldingay, R. L. (2004). Wildlife road-kills on three major roads in north-eastern New South Wales. Wildlife Research , 31 (1), 83–91. https://doi.org/10.1071/WR01110 Trombulak, S. C., & Frissell, C. A. (2000). Review of Ecological Effects of Roads on Terrestrial and Aquatic Communities. Conservation Biology , 14 (1), 18–30. https://doi.org/10.1046/j.1523-1739.2000.99084.x Van Dyke, J. U., Ferronato, B. de O., & Spencer, R.-J. (2018). Current conservation status of Australian freshwater turtles. Australian Journal of Zoology , 66 (1), 1. https://doi.org/10.1071/zov66n1_in Vitt, L. J., & Caldwell, J. P. (2013). Turtles. In K. Gomez & P. Gonzalez (Eds.), Herpetology (Fourth, pp. 523–543). Elsevier. https://doi.org/10.1016/B978-0-12-386919-7.00018-6 Waltham, N. J., Schaffer, J., Walker, S., Perry, J., & Nordberg, E. (2022). Simple fence modification increases land movement prospects for freshwater turtles on floodplains. Wildlife Biology , e01012. https://doi.org/10.1002/wlb3.01012 Wegner, J. F., & Merriam, G. (1979). Movements by Birds and Small Mammals Between a Wood and Adjoining Farmland Habitats. The Journal of Applied Ecology , 16 (2), 349. https://doi.org/10.2307/2402513 Young, M. E., Ryberg, W. A., Fitzgerald, L. A., & Hibbitts, T. J. (2018). Fragmentation alters home range and movements of the Dunes Sagebrush Lizard (Sceloporus arenicolus ). Canadian Journal of Zoology , 96 (8), 905–912. https://doi.org/10.1139/cjz-2017-0048 Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3855993","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":266802250,"identity":"8db9f65d-0492-4ea6-8142-a8fe11247981","order_by":0,"name":"James M Dowling","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6ElEQVRIiWNgGAWjYBACxoYEBoYHBgxy/HChA8RoSTBgMJZsIFYLUD0YJRrAVRLSwtyefEwioeBOgvG1M2bSBRUMcnw3Ehg/8+BzWM+zNIkEg2d5ZrdzzKRnnAG68EYCszReLTNyzG4kGBwuBmvhbWNI3HAjgYEoLYmbZ4O0/GOoB2ph/k2Ulg3SIC0NwNC7kcCG35aeZ+k/gFqMJW6nFVvzHJMwnHnmYZvlHDxaDNuTDxt8+HNYjn928sbbPDU28nzHkw/feINPSwOcyWEAJCRANjfgUAwB8ggm+wO8KkfBKBgFo2DkAgAZIU8tYPbENAAAAABJRU5ErkJggg==","orcid":"","institution":"University of New England","correspondingAuthor":true,"prefix":"","firstName":"James","middleName":"M","lastName":"Dowling","suffix":""},{"id":266802251,"identity":"486a04b4-7bfe-4497-a244-87f903df0a81","order_by":1,"name":"Eric J Nordberg","email":"","orcid":"","institution":"University of New England","correspondingAuthor":false,"prefix":"","firstName":"Eric","middleName":"J","lastName":"Nordberg","suffix":""},{"id":266802252,"identity":"6a4dec3c-6049-4988-a170-f24e5b1736d2","order_by":2,"name":"Deborah S Bower","email":"","orcid":"","institution":"University of New England","correspondingAuthor":false,"prefix":"","firstName":"Deborah","middleName":"S","lastName":"Bower","suffix":""}],"badges":[],"createdAt":"2024-01-12 06:59:11","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3855993/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3855993/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":49712986,"identity":"d02fd8f4-3a81-4bf6-91e8-091911ae473c","added_by":"auto","created_at":"2024-01-16 20:35:57","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1835482,"visible":true,"origin":"","legend":"\u003cp\u003eLocation of trapping sites and mesocosm where this experiment was conducted. Turtles were trapped in Sandy Creek (western red star) and Duval Creek (eastern red star). Mesocosm was located at the University of New England’s Newholme Field Laboratory (purple asterisks).\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-3855993/v1/f0d5f8d56d2a9a453ee2f697.png"},{"id":49712985,"identity":"36b9da2a-0ce7-4a11-bebe-28fa7c94c810","added_by":"auto","created_at":"2024-01-16 20:35:57","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":727201,"visible":true,"origin":"","legend":"\u003cp\u003eThe shaded mesocosm that was constructed for the purpose of this study. Trail cameras were angled to capture the entirety of each arena. Additional material was added to corners to prevent turtles from escaping due to their tendency to climb arena walls.\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-3855993/v1/3eb4cb8efa744f708ed56ba6.jpeg"},{"id":49712988,"identity":"66201e24-3ea4-4bc6-b83f-f06dd1a3cb9d","added_by":"auto","created_at":"2024-01-16 20:35:58","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":837014,"visible":true,"origin":"","legend":"\u003cp\u003eObstacle materials were used to test the ability of Chelodina longicollis to cross several common obstacles encountered throughout their distribution. Three replicate arenas were constructed for each treatment type, being, clockwise from top left: (a) 20 mm aggregate gravel, (b) exclusion fence, (c) chicken wire, (d) boulders and (e) gabion rock.\u003c/p\u003e","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-3855993/v1/f42c653868c0335f266d433f.jpeg"},{"id":49712989,"identity":"e6d02b7a-0e3a-4055-b5eb-19efd695ec8a","added_by":"auto","created_at":"2024-01-16 20:35:58","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":442194,"visible":true,"origin":"","legend":"\u003cp\u003eAn Eastern long-necked turtle (Chelodina longicollis) moving through the second row of exclusion fencing gaps. Despite being small enough to easily move through fence gaps at ground level, turtles moved with their heads elevated and routinely climbed up and over the second row of the fence if their head passed through the gaps first.\u003c/p\u003e","description":"","filename":"floatimage4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-3855993/v1/0b00d80a19ea4e3e2d71a8ad.jpeg"},{"id":49712987,"identity":"542b7520-5f59-4424-8f54-90f228a50516","added_by":"auto","created_at":"2024-01-16 20:35:58","extension":"jpeg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":540634,"visible":true,"origin":"","legend":"\u003cp\u003eAn Eastern long-necked turtle (Chelodina longicollis) was observed climbing the walls of its enclosure, which were constructed of polyester silt fencing. Turtles were able to climb the silt fence by sprawling between perpendicular walls. To mitigate against turtles potentially escaping, I added material across the corners to block turtles from escaping (not pictured).\u003c/p\u003e","description":"","filename":"floatimage5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-3855993/v1/d00753fbc0e53556a87e0dd9.jpeg"},{"id":49936819,"identity":"807415d8-c23a-4fc9-976f-09a89a1490d0","added_by":"auto","created_at":"2024-01-21 20:37:24","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2666669,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3855993/v1/7926fa9c-be8d-49e1-b128-64d5d4ffca18.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Ninja Turtles: high mobility and successful passage through common barriers to movement in a semi-terrestrial freshwater turtle","fulltext":[{"header":"Introduction","content":"\u003cp\u003eChanging land use for agriculture, urbanisation and resource extraction threaten species by fragmenting suitable habitats (Aronson et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; McKinney \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Pattison et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Drastic land conversion for cropping (Serneels and Lambin \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), livestock grazing (Wegner and Merriam \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1979\u003c/span\u003e), and open-cut mining (Cristescu et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Young et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) create inhospitable habitats that many species cannot cross. In addition to habitat fragmentation, barriers in the form of roads (Forman \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Forman and Alexander \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Colino-Rabanal and Lizana \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Rytwinski and Fahrig \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and fences create obstacles for many terrestrial species. For species averse to urbanisation, the negative population-level impacts of roads and fences are more pronounced, often further endangering already threatened species (Taylor and Goldingay \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Seidler et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Jakes et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Species that disperse or migrate, such as semi-aquatic species using a complex of wetlands, are particularly sensitive to barriers (Roe et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Glista et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Balčiauskas et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In particular, freshwater turtles often travel frequently between water bodies and are negatively impacted by these artificial barriers (Aresco \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2005a\u003c/span\u003e; Clinton Gooley \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Hamer et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Proulx et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Santori et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMany freshwater turtles have large home ranges (Slavenko et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and use water-adjacent land for nesting (Bodie \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Petrov et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Van Dyke et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), thermoregulation (Chessman \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and to travel between water bodies (Graham et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Serrano et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This dependence on terrestrial habitat exposes these slow-moving and ungainly species to obstacles that may prevent movement or create ecological traps (Ferronato et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Despite a preference for aquatic habitats, many turtle species climb terrestrial obstacles such as logs, concrete slabs (Scott et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and other individuals (e.g., turtles and crocodiles; Lindeman \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; McKnight et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). As such, the impact of fences on turtle populations is a nuanced issue, providing some positive outcomes for species while threatening others. Fences can benefit populations in urbanised areas by funnelling individuals towards safe road crossing structures (Aresco \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2005b\u003c/span\u003e) or protecting turtles or nesting beaches from introduced predators, like red foxes (\u003cem\u003eVulpes vulpes\u003c/em\u003e; Streeting et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, fences can also kill turtles due to entanglement and overheating (Ferronato et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) or indirectly through prey funnelling for predators (Little et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Additionally, linear barriers dissuade turtles from moving between seasonal habitats and may alter movement routes and increase metabolic costs while reducing available resources (Paterson et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWidespread agricultural and urban development in south-eastern Australia may create many obstacles for preferentially aquatic turtles, such as the Eastern long-necked turtles (\u003cem\u003eChelodina longicollis\u003c/em\u003e). These turtles travel over 3 km overland to exploit resource-rich ephemeral waterbodies away from main river channels (Roe and Georges \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), where their risk of encountering anthropogenic barriers, such as roads, fences, or rocky erosion mitigation strategies is high. However, despite the widespread introduction of quarried rock used for erosion control, levees, and retaining walls along waterways along wetland habitats, the ability of Eastern long-necked turtles to navigate these obstacles is unknown. These structures effectively prevent erosion (Keller \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and provide additional habitat for invertebrate prey species (Shields \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Fischenich \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), but may act as a barrier for terrestrially moving species. Here, we test the ability of \u003cem\u003eC. longicollis\u003c/em\u003e to cross two common styles of livestock fencing and three sizes of rock, commonly used around waterways for road and levee construction and erosion mitigation. Specifically, we compared 1) the success rates of \u003cem\u003eC. longicollis\u003c/em\u003e crossing fence and rock obstacles to an obstacle-free environment, and 2) tested the effect of fatigue and boldness on the willingness of turtles to approach obstacles.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eOur experiment occurred at the University of New England\u0026rsquo;s Newholme SMART Farm (-30.419, 151.637) in the Armidale region of NSW, Australia between 5\u0026ndash;22 March 2023 (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Eighteen wild-caught adult Eastern long-neck turtles (Chelodina longicollis; straight carapace lengths\u0026thinsp;\u0026gt;\u0026thinsp;150 mm) with no physical abnormalities (e.g., missing limbs) were used in the experiment. Morphometric data was collected on all individuals, including straight carapace length and weight.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eExperimental design\u003c/h2\u003e \u003cp\u003eThe experimental mesocosms comprised of outdoor, fully shaded, enclosures covering an area of 180 m2 with eighteen 5 m x 2 m arenas, each equipped with a polypropylene pool (Foundation Products Clam Shell Sandpit (945 x 210 x 1100 mm) at the southern end (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The individual pens were constructed using 0.9 m wide silt fencing material for the walls, providing a wall height of 0.8 m once entrenched to prevent turtles from burrowing beneath the fence. Additional fencing material was placed above intersecting fences to prevent turtles from climbing the corners. Each arena was allocated as control or fitted with one of five obstacle treatments based on common features in the landscape (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Obstacles consisted of 1) aggregate gravel, a common road base in rural areas; 2) gabion, a medium (60\u0026ndash;150 mm) crushed rock; 3) large boulders (\u0026gt;\u0026thinsp;300 mm), which are both used for erosion mitigation and riverbank stabilisation; 4) hinged joint exclusion fencing (15/150/15); or 5) \u0026ldquo;chicken wire\u0026rdquo; wire mesh netting due to their prevalence and popularity in agricultural system and ecosystem rehabilitation areas. Daily rainfall data was collected via a rain gauge attached to the enclosure and temperatures from a Bureau of Meteorology observation station 12.2 km southeast (Armidale Tree Group BOM observation station).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eAt 09:00 h, turtles were randomly allocated to an initial arena without replacement where they were placed in a pond of fresh water (Fountain Products Clam Shell Sandpit) at the southern end of the arena for a one-hour acclimatisation period and provided with food (commercial turtle food; Fish fuel Co. \u0026ldquo;Turtle Food\u0026rdquo; 3 g frozen blocks). At 10:00 h, each turtle was moved and released on the northern side of the obstacle and given 23 hours to cross the obstacle and return to the pond containing the food and water. Turtles were exposed to each testing arena only once, but they encountered each obstacle (in random order) three times throughout the whole trial (18 testing areas, with 3 replicates each of 6 different treatments).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eAssessing movement, boldness, and fatigue\u003c/h2\u003e \u003cp\u003eTesting arenas were monitored via time-lapse wildlife cameras (Campark T85; 1-min photo interval) to record the movement of turtles throughout the experiment. The time stamp on the photos allowed for me to calculate the time and the duration of each crossing attempt. We considered a turtle to have made an \"attempt\" at crossing when it moved to within a body length of the obstacle. Attempts were considered successful if the individual crossed and exited the opposite side of the obstacle. Unsuccessful attempts were recorded if an individual engaged with the obstacle but failed to reach the other side. The outcome of each trial was determined as either successful or unsuccessful, dependent on whether the turtle crossed over the obstacle to the other side. If no attempt was made, the trial was classified as \"no attempt\". Boldness was measured as the time taken, in minutes, for turtles to make their first movement in each trial. The number of attempts made per trial was used to examine the effect of fatigue on turtles willingness to attempt obstacles.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eWe used generalised linear mixed effect models to analyse relationships between various response variables and the fixed effects: obstacle treatment (grass, gravel, gabion, rock, boulder, exclusion fencing, chicken wire), trial number (1\u0026ndash;18), and straight carapace length (SCL). Individual turtle ID was included as a random effect to account for variation among individuals.\u003c/p\u003e \u003cp\u003eTo analyse obstacle crossing success we used a binary response variable (success\u0026thinsp;=\u0026thinsp;1, failure\u0026thinsp;=\u0026thinsp;0) with the logit link function. To evaluate the impact of fatigue on the turtles' performance we used the number of attempts per trial and the logit link function. To explore the influence of individual boldness on the turtles' movement patterns, we used the time (latency) to the first movement as an indicator of boldness (Kabelik et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Talavera et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and the logit link function. The significance of variation in individual boldness was assessed using the ANOVA function in the car package (v3.0.12) with a type II sum of squares. We used estimated marginal means for post hoc testing for all GLMM and GLMER. All statistical analyses were conducted using R statistical software (version 2023.03.0\u0026thinsp;+\u0026thinsp;386). The specific packages used for analysis included \"stats\", \u0026ldquo;emmeans\u0026rdquo;, and \"lme4\" for the GLMM, GLMER and post hoc testing. All results are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE unless otherwise stated.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\n\u003ch2\u003eObstacle crossing\u003c/h2\u003e\n\u003cp\u003eCollectively, individual turtles attempted to cross the obstacles 1,545 times throughout the 18-day study. Non-attempts occurred in seven instances from 6 of the 18 individuals. Turtles successfully crossed a range of obstacles. Gravel, gabion, boulders, and exclusion fencing were all crossed by all turtles at least once, with varying degrees of success. Attempts to cross the control arenas were successful in 100% of attempts. In contrast, chicken wire was attempted 298 times with a 0% success rate. Exclusion fencing had the highest success rate among the obstacles with 94.7% crossings from 228 attempts. While crossed by all turtles at least once, boulders exhibited the second lowest rate of success per attempt with 73.3% crossings from 236 attempts. Gravel and gabion had notably high success rates, with an 85.4% success rate from 308 gravel attempts and 86% of the 215 gabion attempts resulting in success. Turtle body size (SCL) did not predict obstacle-crossing success (R2\u0026thinsp;=\u0026thinsp;0.002, F 1,82\u0026thinsp;=\u0026thinsp;0.721, p\u0026thinsp;=\u0026thinsp;0.230).\u003c/p\u003e\n\u003cp\u003eObstacles that were crossed more often, were also crossed more quickly. The time taken to cross obstacles was lowest for exclusion fencing, taking an average of 1.56\u0026thinsp;\u0026plusmn;\u0026thinsp;2.21 min (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE) followed by gravel (1.91\u0026thinsp;\u0026plusmn;\u0026thinsp;2.09 min), gabion (2.84\u0026thinsp;\u0026plusmn;\u0026thinsp;4.39 min), and boulders (6.85\u0026thinsp;\u0026plusmn;\u0026thinsp;13.40 min). Despite having no success, turtles attempted to cross chicken wire for 35.64\u0026thinsp;\u0026plusmn;\u0026thinsp;47.91 min per attempt before resting, with the longest continuous attempt lasting 595 min.\u003c/p\u003e\n\u003cp\u003eRecognition of mesh fences as an obstacle remains somewhat uncertain for C. longicollis. Turtles appeared to routinely \u0026ldquo;follow their head\u0026rdquo; through obstacles, resulting in unnecessary climbing through the second row of exclusion fencing despite the turtle\u0026rsquo;s body size permitting easy movement through fence gaps at ground level. For example, C. longicollis typically walk with their head elevated above the ground so, when they encounter a fence, if the head enters the mesh gap on the second row, the turtle will climb through that opening rather than ducking its head to easily walk through the lowest mesh gap (Fig.\u0026nbsp;\u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003e). This tendency became more pronounced for chicken wire obstacles, where turtles would place their head and neck through the chicken wire and attempt to push their body through the mesh (which was far too small for their whole body), often for several hours or several attempts. Turtles were also observed using their heads and necks to routinely climb boulder obstacles and cross large gaps. Turtles extended their necks before using their heads as a point of contact, allowing for the bridging of obstacles.\u003c/p\u003e\n\u003cp\u003eIn addition to crossing obstacles, turtles climbed the walls of their enclosure. Two individuals scaled the 800 mm high woven polyester silt fencing and entered the pools of neighbouring arenas during a pilot study. Turtles wedged themselves in the vertex of the corners of the silt fencing and climbed walls using their claws to penetrate the weave of the silt fencing fabric (Fig.\u0026nbsp;\u003cspan class=\"InternalRef\"\u003e5\u003c/span\u003e). The fence was modified with an additional overhang to prevent this (not pictured).\u003c/p\u003e\n\u003c/div\u003e\n\u003ch2\u003eFatigue effect\u003c/h2\u003e\n\u003cp\u003eThere was a negative linear relationship between number of attempts per trial and trial number. Turtles made 0.218 fewer attempts per trial or approximately one less attempt every five days (t = -4.121, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Turtles attempted obstacles an average of 3.94\u0026thinsp;\u0026plusmn;\u0026thinsp;4.93 times on day eighteen as opposed to 8.39\u0026thinsp;\u0026plusmn;\u0026thinsp;5.12 on day one.\u003c/p\u003e\n\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\n\u003ch2\u003eBoldness\u003c/h2\u003e\n\u003cp\u003eIndividual boldness varied significantly between turtles (X2\u0026thinsp;=\u0026thinsp;431.96, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Boldness was negatively correlated with time taken per attempt (\u0026beta; = -0.14547, p\u0026thinsp;=\u0026thinsp;0.009), with bolder turtles dwelling on or near obstacles for longer. However, boldness did not significantly affect the success rate (\u0026beta;\u0026thinsp;=\u0026thinsp;0.007, p\u0026thinsp;=\u0026thinsp;0.11). Obstacle type, trial number and arena number did not affect the time to first move.\u003c/p\u003e\n\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur study demonstrates that while Eastern long-necked have a remarkable ability to navigate many structures, barriers exist from common landscape features (Coffin \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Forman \u0026amp; Alexander \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Jakes et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Jones et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; McInturff et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Smith et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Trombulak \u0026amp; Frissell \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe obstacle materials we tested here are commonly used in road, stream, and wetland restoration or common fence types used by commercial and private landholders and therefore are likely to impact Eastern-long necked turtles throughout much of their range. Chicken wire or similar small-diameter mesh fences presented unpassable obstacles for the sampled eastern long-necked turtles, and this reflects the barrier created for many other wildlife groups as well (Baines \u0026amp; Andrew \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Bock \u0026amp; Bock \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Moseby \u0026amp; Read \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Smith et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Although other obstacle types were successfully crossed by turtles, the time required to successfully navigate through obstacles was often substantial and this likely affects an energy cost.\u003c/p\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eCrossing success\u003c/h2\u003e \u003cdiv id=\"Sec13\" class=\"Section3\"\u003e \u003ch2\u003eRocks\u003c/h2\u003e \u003cp\u003eTurtles crossed rock obstacles with seemingly little trouble, regardless of the size of rock encountered. We expected the sharp, irregular angularity of the quarried boulders to pose a challenge to turtles, particularly smaller individuals with carapace lengths below 160 mm (33.3% of turtles in our study) as boulders were generally taller than this. Yet, 73% of boulder crossing attempts were successful, and multiple crossings of the obstacle in each trial were common. Small turtles in the sample group also used their head and necks as a point of contact to both climb and cross the large boulder obstacles.\u003c/p\u003e \u003cp\u003eWe expected the smallest obstacles such as gravel and gabion to be crossed far more often, however, gabion had only 12 more crossings (185 of 215 attempts) than boulders (173 of 236 attempts), a more difficult obstacle. Multiple attempts were unsuccessful for both gravel and gabion, where turtles either approached the obstacle and chose not to cross or aborted crossing part-way through. This may be attributed to several factors that were not measured in our experiment. Warm surface temperatures of rocks may explain aborted crossings, as burns and eventual death from hot rocks have been recorded in turtles attempting to gabion railway track beds (Abinesh et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Kornilev et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). However, the effect of dangerously high temperatures is unlikely to have had a major impact on turtles in our experiment due to the use of shade cloth which shaded the entire mesocosm.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eFences\u003c/h2\u003e \u003cp\u003eTurtle's success at crossing fences was highly dependent on the fence design. The chicken wire style mesh, which was completely impassable for all turtles, is prolific in agricultural regions with sheep. The wire mesh usually extends to ground level to prevent the separation of lambs from ewes or enforce weaning (Galeana et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; McGregor \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1990\u003c/span\u003e) and exclude predators (Kondinin Group \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Long \u0026amp; Robley \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Despite the complete failure to cross, chicken wire had the third highest number of overall attempts and the longest amount of time attempting to cross. The average time per crossing attempt was far higher than other obstacles at 35.6 minutes per attempt, likely imparting a high energy cost. Increased energy expenditure may place turtles at elevated risk of mortality due to exhaustion, desiccation and overheating (Ferronato et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). While turtles with roads in their home range have only small increases in energy expenditure (Patterson et al. 2019), the presence of chicken wire-type fences without suitable crossing structures would likely result in turtle exhaustion and death, as has been witnessed in other \u0026rsquo;turtle-proof fences\u0026rsquo; (Ferronato et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eExclusion fencing (prefabricated hinge joint) may still pose a risk to C. longicollis which was not identified by our study. Chelodina longicollis of the New England region, particularly Armidale NSW, exhibit irregularly small adult body sizes, (average body sizes (SCL) of females\u0026thinsp;=\u0026thinsp;239 mm; male\u0026thinsp;=\u0026thinsp;199 mm; Parmenter \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1976\u003c/span\u003e), which is far smaller than in other regions where SCL may exceed 260 mm (Chessman \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). As such, turtles used in our experiment passed through the ~\u0026thinsp;150 mm wide spaces of the exclusion fencing with relative ease. In other regions, the carapace width of C. longicollis may exceed this width, making passage through unmodified exclusion fencing difficult or impossible (Kennett et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). However, simple cost-effective solutions, such as turtle gates (Waltham et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), that permit turtle movement while also maintaining fence integrity have been developed and trialled for other, larger-bodied, freshwater turtles.\u003c/p\u003e \u003cp\u003eAlthough fences may prevent the movement of non-target species, fences are rarely completely intact, with breaks in fences acting as potential crossing points (Jones et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Gates often act as weak points in fences, with substantial gaps beneath gates existing either through the gate design or wheel ruts from repeated vehicle traffic (Long \u0026amp; Robley \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Even in high-cost reinforced fences, holes may be created by burrowing species such as rabbits and foxes, compromising fence integrity (Moseby \u0026amp; Read \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). While intermittent gaps may be beneficial for more motile species, freshwater turtles show a significant reduction in their ability to locate gaps within the space of 3m (Waltham et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), making the likelihood of encountering gaps in turtle proof fences unlikely in all but the most dilapidated of cases.\u003c/p\u003e \u003cp\u003eThe ability of C. longicollis to climb the silt fencing used to divide the arenas demonstrates a remarkable capacity to tolerate certain linear barriers. The loose weave of the silt fencing allowed the turtles to use their claws to climb in the vertex where the edges met at right angles. Turtles were not seen climbing parallel edges, likely due to their sprawling posture and limited mobility from hips sitting inside their shells (Vitt \u0026amp; Caldwell \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Australian turtles are not typically known for their ability to scale vertical surfaces; however, a considerable ability to climb fences (Aresco \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2005a\u003c/span\u003e; Griffin \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), logs (Lindeman \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), and rocks (Davenport et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1984\u003c/span\u003e) have been recorded in several North American turtle species.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eFatigue\u003c/h2\u003e \u003cp\u003eThe number of attempts per trial decreased consistently throughout the experiment, resulting in turtles making just over half (46%) of the attempts on the final day as compared to day one. Mean daily movement lengths for C. longicollis are generally below 10 m-1 day \u0026minus;\u0026thinsp;1 (Roe \u0026amp; Georges \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), yet our subjects exceeded this distance in 60% of trials, likely expending significant energy that led to reduced attempts as trials progressed. Additionally, the tendency for turtles to climb their enclosures reduced with the number of treatments, with no climbing of the mesocosm walls observed after trial five. Therefore, we believe that fatigue rather than habituation towards food supplied in pools was the primary cause of reduced crossing attempts as trials progressed.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eBoldness\u003c/h2\u003e \u003cp\u003eAs turtles retract into their shells when exposed to potentially threatening stimuli (Greene \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1988\u003c/span\u003e), higher degrees of boldness may reduce the risk of mortality to turtles attempting to cross busy roads because bold individuals cross more quickly. Bold turtles crossed obstacles faster than turtles with lower boldness scores, which suggests that less bold individuals may be more vulnerable to threats such as vehicle strikes or predation as they linger exposed on obstacles.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eImplications for management\u003c/h2\u003e \u003cp\u003eOur study highlights the need for selecting wildlife-friendly fence designs when planning and installing fences around wetlands and waterways. Exclusion fencing is manufactured in a range of sizes and designs and populations with larger-bodied turtles may not be as successful at crossing exclusion fencing as in our study. However, simple, cost-effective mitigation methods exist for reducing the threat posed to turtles by exclusion fencing in these scenarios. For example, clipping small sections of vertical exclusion fence wires at regular intervals to permit the movement of larger turtles through exclusion fences incurs minimal cost to landholders while maintaining the functionality of the exclusion fence (Waltham et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The agricultural sector, particularly the sheep industry, encourages the use of wire mesh fencing, similar to that tested here, for the exclusion of canines and foxes to limit livestock depredation (Kondinin Group \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Many of these designs encourage wire mesh to be anchored at ground level or installed below the surface to prevent canids from digging under (Green \u0026amp; Gipson \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Kondinin Group \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Alternatively, the implementation of species-specific one-way gates, such as those currently used for small mammals to transit through conservation areas (Coates \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), may warrant further investigation for the possibility of turtle-specific modification.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOur study underscores the significant impact of artificial barriers on freshwater turtle terrestrial spatial use. While turtles can traverse rock barriers with relative ease, depending on rock size, the need to continuously cross rock barriers lead to fatigue. Additionally, rocks larger than those tested here may be impassable completely. Similarly, the impact of fences is dependent on design, with small gauge wire mesh fences preventing the movement of turtles completely. While these results focus on turtles, these trends likely impact numerous small and medium vertebrates across freshwater systems. Wire mesh fences are often used to protect vulnerable species and fragile ecological areas, yet disruptions to movements and mortality due to entanglement or collision are common across birds, reptiles, and mammals. As such, design-specific considerations are crucial for effective wetland and waterway management, necessitating collaboration for wildlife-friendly adaptations to infrastructure in riparian zones to limit unintended negative ecological impacts. In light of this growing body of evidence, it is imperative that wildlife-friendly fencing solutions gain broader acceptance among both land managers and industry stakeholders.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll authors project design and planning.J.D. and E.N. data collection.J.D. and E.N. data analysis.J.D. initial manuscript drafting.All authors editing.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eWe acknowledge the Traditional Custodians of the land where this research was conducted, the Anaiwan People. We pay our respects to their cultures and Elders, past, present and emerging. Funding for this project was supported through the University of New England. All experiments were conducted under the Animal Ethics permit ARA23-010 through the University of New England and the New South Wales Department of Planning and Environment Scientific License SL102308. Open access publishing facilitated by University of New England, as part of the Wiley - University of New England agreement via the Council of Australian University Librarians. We thank Dr Zenon Czenze for helpful insights and troubleshooting during data analysis and Remo Boscarino-Gaetano for assistance in the construction of the mesocosm.\u003c/p\u003e\n \u003ch2\u003eDeclaration of interests\u003c/h2\u003e \u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAbinesh, A., Maharaja, R., \u0026amp; Vishnu, C. S. (2022). 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Fragmentation alters home range and movements of the Dunes Sagebrush Lizard \u003cem\u003e(Sceloporus arenicolus\u003c/em\u003e). \u003cem\u003eCanadian Journal of Zoology\u003c/em\u003e, \u003cem\u003e96\u003c/em\u003e(8), 905\u0026ndash;912. https://doi.org/10.1139/cjz-2017-0048\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Agriculture, Chelodina longicollis, Fence, Livestock, Riparian, Roads","lastPublishedDoi":"10.21203/rs.3.rs-3855993/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3855993/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAnthropogenic landscape change due to urbanisation, agriculture and resource extraction inevitably results in linear barriers within the landscape. Artificial linear structures such as roads, fences, levees, and dams limit the movement of some species and further fragment residual habitat. In this study, we investigated the ability of Eastern long-necked turtles (\u003cem\u003eChelodina longicollis\u003c/em\u003e) to cross various terrestrial obstacles commonly encountered in their habitat. We tested two types of fences (chicken wire and exclusion fencing) commonly used in agricultural systems and three sizes of rocks (gravel, gabion, and large boulders) often used for road construction, erosion control, and waterway stabilisation. We examined the success rates of turtles in crossing obstacles, the effect of fatigue on crossing attempts, and the impact of individual boldness on movement behaviour. Turtles displayed high success rates in crossing gravel (85.4%), gabion (86%), boulders (73.3%) and hinged joint exclusion fencing (94.7%). Chicken wire style wire netting had no successful crossings (0%) despite 276 attempts. A significant fatigue effect was observed throughout the experiment, with turtles making an average of 3.94 (\u0026plusmn;\u0026thinsp;4.93 SE) fewer attempts at the end of the experiment (day eighteen) as opposed to day one. Bolder turtles were faster at crossing obstacles, however, boldness had no bearing on obstacle-crossing success. These results highlight the need for thoughtful selection of waterway and wetland infrastructure and the fatiguing impact of constant exposure to anthropogenic barriers for wildlife.\u003c/p\u003e","manuscriptTitle":"Ninja Turtles: high mobility and successful passage through common barriers to movement in a semi-terrestrial freshwater turtle","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-01-16 20:35:53","doi":"10.21203/rs.3.rs-3855993/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"41f5c1b1-5686-48ee-9837-395992a856d3","owner":[],"postedDate":"January 16th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-01-21T20:29:17+00:00","versionOfRecord":[],"versionCreatedAt":"2024-01-16 20:35:53","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3855993","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3855993","identity":"rs-3855993","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}