Depth use and thermal environment of adult male Mediterranean loggerhead sea turtles | Discover Conservation
Lutcavage ME, Plotkin P, Witherington B, Lutz PL. Human impacts on sea turtle survival. In: Lutz PL, Musick JA, editors. The Biology of Sea Turtles. Boca Raton, FL: CRC; 1997. pp. 387–409.
FAO (Food and Agriculture Organization of the United Nations). Classification and Illustrated Definition of Fishing Gears, FAO Technical Paper No. 382. Rome: FAO; 2003.
Casale P. Sea turtle by-catch in the Mediterranean. Fish Fish. 2011;12:299–316. https://doi.org/10.1111/j.1467-2979.2010.00394.x.
Kooyman GL. Diverse divers: Physiology and Behaviour. Berlin: Springer; 1989. p. 200.
Hochscheid S, Bentivegna F, Speakman JR. The dual function of the lung in chelonian turtles: buoyancy control and oxygen store. J Exp Mar Biol Ecol. 2003;297:123–40. https://doi.org/10.1016/j.jembe.2003.07.004.
Minamikawa S, Naito Y, Sato K, Matsuzawa Y, Bando T, Sakamoto W. Maintenance of neutral buoyancy by depth selection in the loggerhead turtle Caretta caretta. J Exp Biol. 2000;203:2967–75. https://doi.org/10.1242/jeb.203.19.2967.
Hatase H, Omuta K, Tsukamoto K. Bottom or midwater: alternative foraging behaviours in adult female loggerhead sea turtles. J Zool. 2007;273:46–55. https://doi.org/10.1111/j.1469-7998.2007.00298.x.
DeRuiter SL, Larbi Doukara K. Loggerhead turtles dive in response to airgun sound exposure. Endang Species Res. 2012;16:55–63. https://doi.org/10.3354/esr00396.
Hounslow JL, Fossette S, Byrnes EE, Whiting SD, Lambourne RN, Armstrong NJ, Tucker AD, Richardson AR, Gleiss AC. Multivariate analysis of biologging data reveals the environmental determinants of diving behaviour in a marine reptile. R Soc Open Sci. 2022;9:211860. https://doi.org/10.1098/rsos.211860.
Stokes KL, Esteban N, Casale P, Chiardia A, Kaska Y, Kato A, Luschi P, Ropert-Coudert Y, Stokes HJ, Hays GC. Optimization of swim depth across diverse taxa during horizontal travel. Proc Nat Acad Sci. 2024;121:e2413768121. https://doi.org/10.1073/pnas.2413768121.
Wilson RP, Shepard ELC, Liebsch N. Prying into the intimate details of animal lives: use of a daily diary on animals. Endang Species Res. 2008;4:123–37. https://doi.org/10.3354/esr00064.
Hochscheid S, Maffucci F, Bentivegna F, Wilson RP. Gulps, wheezes and sniffs: how measurement of beak movement in sea turtles can elucidate their behaviour and ecology. J Exp Mar Biol Ecol. 2005;316:45–53. https://doi.org/10.1016/j.jembe.2004.10.004.
Eckert SA, Nellis DW, Eckert KL, Kooyman GL. Diving patterns of two leatherback sea turtles (Dermochelys coriacea) during internesting intervals at Sandy Point, St. Croix, U. S. Virgin Islands. Herpetologica. 1986;42(3):381–8.
Seminoff JA, Jones TT, Marshall GJ. Underwater behaviour of green turtles monitored with video-time-depth recorders: what’s missing from dive profiles? Mar Ecol Prog Ser. 2006;322:269–80. https://doi.org/10.3354/meps322269.
Hochscheid S. Why we mind sea turtles’ under water business: A review on the study of diving behavior. J Exp Mar Biol Ecol. 2014;450:118–36. https://doi.org/10.1016/j.jembe.2013.10.016.
Kishida N, Okuyama J, Arita M, Kume N, Fujita K, Nishizawa H, Shinsuke T, Mitsunaga Y. A validation of abstracted dive profiles relayed via the Argos satellite system: a case study of a loggerhead turtle. Anim Biotel. 2022;10:21. https://doi.org/10.1186/s40317-022-00292-0.
Swimmer Y, Arauz R, McCracken M, McNaughton L, Ballestero J, Musyl M, Bigelow K, Brill R. Diving behavior and delayed mortality of olive ridley sea turtles Lepidochelys olivacea after their release from longline fishing gear. Mar Ecol Prog Ser. 2006;323:253–61. https://doi.org/10.3354/meps323253.
Southwood AL, Darveau CA, Jones DR. Metabolic and cardiovascular adjustments of juvenile green turtles to seasonal changes in temperature and photoperiod. J Exp Biol. 2003;206:4521–31. https://doi.org/10.1242/jeb.00689.
Hochscheid S, Bentivegna F, Speakman JR. Long-term cold acclimation leads to high Q10 effects on oxygen consumption of loggerhead sea turtles, Caretta caretta. Physiol Biochem Zool. 2004;77(2):209–22. https://doi.org/10.1086/381472.
Stokes KL, Esteban N, Stokes HJ, Hays GC. High dive efficiency in shallow water. Mar Biol. 2023;170:45. https://doi.org/10.1007/s00227-023-04179-3.
Wallace BP, Posnik ZA, Hurley BJ, DiMatteo AD, Bandimere A, Rodriguez I, Maxwell SM, Meyer L, Brenner H, Jensen MP, LaCasella E, Shamblin BM, Abreu-Grobois FA, Stewart KR, Dutton PH, Barrios-Garrido H, Dalleau M, Dell’Amico F, Eckert KL, FitzSimmons NN, Garcia-Cruz M, Hays GC, Kelez S, Lagueux CJ, Madden Hof CA, Marco A, Martins SLT, Mobaraki A, Mortimer JA, Nel R, Phillott AD, Pilcher NJ, Putman NF, Rees AF, Rguez-Baron JM, Seminoff JA, Swaminathan A, Türkozan O, Vargas SM, Vernet PD, Vilaça S, Whiting SD, Hutchinson BJ, Casale P, Mast RB. Marine turtle regional management units 2.0: an updated framework for conservation and research of wide-ranging megafauna species. End Species Res. 2023;52:209–23. https://doi.org/10.3354/esr01243.
Margaritoulis D, Argano R, Baran I, Bentivegna F, Bradai MN, Caminas JA, Casale P, De Metrio G, Demetropoulos A, Gerosa G, Godley BJ, Haddoud DA, Houghton J, Laurent L, Lazar B. Loggerhead turtles in the Mediterranean Sea: present knowledge and conservation perspectives. In: Bolten AB, Witherington B, editors. Biology and conservation of loggerhead sea turtles. Washington, DC: Smithsonian Institution; 2003. pp. 175–98.
Hochscheid S, Bentivegna F, Hays GC. First records of dive durations for a hibernating sea turtle. Biol Lett. 2005;1(1):82–6. https://doi.org/10.1098/rsbl.2004.0250.
Hochscheid S, Bentivegna F, Hamza A, Hays GC. When surfacers do not dive: multiple significance of extended surface times in marine turtles. J Exp Biol. 2010;213:1328–37. https://doi.org/10.1242/jeb.037184.
Chimienti M, Blasi MF, Hochscheid S. Movement patterns of large juvenile loggerhead turtles in the Mediterranean Sea: Ontogenetic space use in a small ocean basin. Ecol Evol. 2020;10:6978–92. https://doi.org/10.1002/ece3.6370.
Harvey-Carroll J, Menéndez-Blázquez J, Crespo-Picaz JL, Sagarminaga R, March D. Unlocking sea turtle diving behaviour from low-temporal resolution time-depth recorders. Sci Rep. 2025;15:19934. https://doi.org/10.1038/s41598-025-05336-y.
Margaritoulis D, Rees AF, Riggall TE. Exponential increase in a loggerhead sea turtle nesting population: Investigating the role of multi-decadal nest protection in Kyparissia Bay, Greece. Zool Stud. 2025;64:29. https://doi.org/10.6620/ZS.2025.64-29.
Rees AF, Dimalexis T, Mikoniatis G, Vavassis Y. Further evidence supporting internesting habitat boundaries for loggerhead turtles nesting in Kyparissia Bay, Greece. MedTurtle Bull. 2025;7:29–33.
Rees AF, Dimalexis T, Mikoniatis G, Vavassis Y. Dispersal and foraging strategies of adult loggerhead sea turtles (Caretta caretta) breeding in Kyparissia Bay, Greece: Implications for conservation. Zool Stud. 2025;64:61. https://doi.org/10.6620/ZS.2025.64-61.
CLS (Collecte Localisation Satellites). 2016. Argos User’s Manual. https://www.argos-system.org/support/. Accessed 17 May 2025.
Ehrhart LM, Ogren LH. Studies in foraging habitats: capturing and handling turtles. In: Eckert KL, Bjorndal KA, Abreu-Grobois FA, Donnelly M, editors. Research and management techniques for the conservation of sea turtles. IUCN/SSC Marine Turtle Specialist Group Publication No; 1999. pp. 61–4. 4.
Rees AF, Papathanasopoulou N, Papoulias P, Samlidou G, Theodorou P, Margaritoulis D. Internesting habitat preference and updated clutch frequency values for loggerhead turtles (Caretta caretta) nesting in Kyparissia Bay, Greece. Herp Cons Biol. 2023;18(3):551–9.
R Core Team. (2025). _R: A Language and Environment for Statistical Computing_. R Foundation for Statistical Computing, Vienna, Austria. <https://www.R-project.org/
Jonsen ID, Grecian WJ, Phillips L, Carroll G, McMahon C, Harcourt RG, Hindell MA, Patterson TA. aniMotum, an R package for animal movement data: Rapid quality control, behavioural estimation and simulation. Methods Ecol Evol. 2023;14:806–16. https://doi.org/10.1111/2041-210X.14060.
Felger RS, Cliffton K, Regal PJ. Winter dormancy in sea turtles: independent discovery and exploitation in the Gulf of California by two local cultures. Science. 1976;191(4224):283–5. https://doi.org/10.1126/science.191.4224.283.
Hochscheid S, Bentivegna F, Bradai MN, Hays GC. Overwintering behaviour in sea turtles: dormancy is optional. Mar Ecol Prog Ser. 2007;340:287–98. https://doi.org/10.3354/meps340287.
Reynolds RW, Zhang HM, Smith TM, Gentemann CL, Wentz F. Impacts of in situ and additional satellite data on the accuracy of a sea-surface temperature analysis for climate. Int J Climatol. 2005;25(7):857–64. https://doi.org/10.1002/joc.1168.
Dalleau M, Benhamou S, Sudre J, Ciccione S, Bourjea J. The spatial ecology of juvenile loggerhead turtles (Caretta caretta) in the Indian Ocean sheds light on the ‘lost years’ mystery. Mar Biol. 2014;161:1835–49. https://doi.org/10.1007/s00227-014-2465-z.
Iverson AR, Fujisaki I, Lamont MM, Hart KM. Loggerhead sea turtle (Caretta caretta) diving changes with productivity, behavioral mode, and sea surface temperature. PLoS ONE. 2019;14(8):e0220372. https://doi.org/10.1371/journal.pone.0220372.
Minamikawa S, Naito Y, Uchida I. Buoyancy control in diving behavior of the loggerhead Turtle. Caretta caretta. J Ethol. 1997;15:109–18. https://doi.org/10.1007/BF02769396.
Hochscheid S, Godley BJ, Broderick AC, Wilson RP. Reptilian diving: highly variable dive patterns in the green turtle Chelonia mydas. Mar Ecol Prog Ser. 1999;185:101–12. https://doi.org/10.3354/meps185101.
Fedak MA, Lovell P, Grant SM. Mar Mamm Sci. 2001;17(1):94–110. https://doi.org/10.1111/j.1748-7692.2001.tb00982.x. Two approaches to compressing and interpreting time-depth information as collected by time-depth recorders and satellite-linked data recorders.
Plot V, de Thoisy B, Georges J-Y. Dispersal and dive patterns during the post-nesting migration of olive ridley turtles from French Guiana. Endang Species Res. 2015;26:221–34. https://doi.org/10.3354/esr00625.
Casale P, Broderick AC, Camiñas JA, Cardona L, Carreras C, Demetropoulos A, Fuller WJ, Godley BJ, Hochscheid S, Kaska Y, Lazar B, Margaritoulis D, Panagopoulou A, Rees AF, Tomás J, Türkozan O. Mediterranean sea turtles: current knowledge and priorities for conservation and research. Endang Species Res. 2018;36:229–67. https://doi.org/10.3354/esr00901.
DiMatteo A, Cañadas A, Roberts J, Sparks L, Panigada S, Boisseau O, Moscrop A, Fortuna CM, Lauriano G, Holcer D, Peltier H, Ridoux V, Raga JA, Tomás J, Broderick AC, Godley BJ, Haywood J, March D, Snape R, Sagarminaga R, Hochscheid S. Basin-wide estimates of loggerhead turtle abundance in the Mediterranean Sea derived from line transect surveys. Front Mar Sci. 2022;9:930412. https://doi.org/10.3389/fmars.2022.930412.
Fuentes MMPB, Bell I, Hagihara R, Hamann M, Hazel J, Huth A, Seminoff JA, Fuentes MMPB, McMichael E, Kot CY, Silver-Gorges I, Wallace BP, Godley BJ, Brooks AML, Ceriani SA, Cortés -Gómez AA, Dawson TM, Dodge KL, Flint M, Jensen MP, Komoroske LM, Kophamel S, Lettrich MD, Long CA, Nelms SE, Patrício AR, Robinson NJ, Seminoff JA, Ware M, Whitman ER, Chevallier D, Clyde-Brockway CE, Korgaonkar SA, Mancini A, Mello-Fonseca J, Monsinjon JR, Neves-Ferreira I, Ortega AA, Patel SH, Pfaller JB, Ramirez MD, Raposo C, Smith CE, Abreu-Grobois FA, Hays GC. Key issues in assessing threats to sea turtles: knowledge gaps and future directions. Endang Species Res. 2023;52:303–41. https://doi.org/10.3354/esr01278.
Polovina J, Howell E, Parker DM, Balazs GH. Dive-depth distribution of loggerhead (Carretta carretta [sic]) and olive ridley (Lepidochelys olivacea) sea turtles in the central North Pacific: Might deep longline sets catch fewer turtles? Fish Bull. 2003;101(1):189–93.
Howell EA, Dutton PH, Polovina JJ, Bailey H, Parker DM, Balazs GH. Oceanographic influences on the dive behavior of juvenile loggerhead turtles (Caretta caretta) in the North Pacific Ocean. Mar Biol. 2010;157:1011–26. https://doi.org/10.1007/s00227-009-1381-0.
Patel SH, Morreale SJ, Panagopoulou A, Bailey H, Robinson NJ, Paladino FV, Margaritoulis D, Spotila JR. Changepoint analysis: a new approach for revealing animal movements and behaviors from satellite telemetry data. Ecosphere. 2015;6(12):291. https://doi.org/10.1890/ES15-00358.1.
Fuentes MMPB, Bell I, Hagihara R, Hamann M, Hazel J, Huth A, Seminoff JA, Sobtzick S, Marsh H. Improving in-water estimates of marine turtle abundance by adjusting aerial survey counts for perception and availability biases. J Exp Mar Biol Ecol. 2015;471:77–83. https://doi.org/10.1016/j.jembe.2015.05.003.
Patel SH, Barco SG, Crowe LM, Manning JP, Matazen E, Smolowitz RJ, Haas HL. Loggerhead turtles are good ocean-observers in stratified mid-latitude regions. Est Coast Shelf Sci. 2018;213:128–36. https://doi.org/10.1016/j.ecss.2018.08.019.