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The heart development, form, and functional specializations of chelonians, squamates, crocodilians, and birds characterize how diverse structure and specializations arise from similar foundations. This review aims to summarize the morphologic diversity of sauropsid hearts and present it in an integrative functional and phylogenetic context. Besides the detailed morphologic descriptions, the integrative view of function, evolution, and development will aid understanding of the surprising diversity of sauropsid hearts. This integrated perspective is a foundation that strengthens appreciation that the sauropsid hearts are the outcome of biological evolution; disease often is linked to arising mismatch between adaptations and modern environments.

Variations in the number and arrangement of scutes often are used for species identification in hard-shelled sea turtles. Despite the conserved nature of scute arrangements, anomalous arrangements have been noted in the literature for over a century, with anomalies linked to sub-optimal environmental conditions in the nest during development. Long-held assumptions suggest that anomalous scute arrangements are indicative of underlying physiological or morphological anomalies, with presumed long-term survival costs to the individual. Here, we examined a 25-year photo database of two species of sea turtle (Caretta caretta and Chelonia mydas) captured incidentally and non-selectively on the eastern coast of Florida. Our results suggest that C. mydas is substantially more variable with respect to the arrangement of carapacial scutes, while C. caretta had a relatively higher proportion of individuals with anomalous plastron scute arrangements. We also show evidence that (a) the forms and patterns of anomalous scutes are stable throughout growth; (b) there is limited evidence for selection against non-modal arrangements in the size classes that were examined; and (c) that their frequency has remained stable in juvenile cohorts from 1994 until present. These findings indicate that there may not be a survival cost associated with anomalous scute arrangements once the turtles reach juvenile size classes, and that variation in scute arrangements within populations is relatively common.

Synopsis Here we describe examples of studies that have contributed both to a basic understanding of the biology of imperiled marine turtles, and to their management and conservation. Key elements include, first and foremost, correctly identifying species that differ strikingly in their morphology at different life stages because with growth, they change size by several orders of magnitude and have accompanying shape changes. We also review comprehensive field studies documenting the need for management actions to correct abnormal shifts in sex ratios caused by climate change. We highlight the need to describe those perturbations in terms that are clear to regulators and personnel responsible for management and conservation policies. Finally, we review several basic studies that enhance our understanding of how selection has shaped morphological, functional, and performance attributes, and describe how that knowledge can be applied to the tasks required for enhancing species recovery.

The goal of this study was to assess the consequences of single versus multiple paternity by identifying paternity of clutches per female to identify whether there were detectable costs or benefits. Multiple mating can occur when the benefits of mating outweigh the costs, but if costs and benefits are equal, no pattern is expected. Previous research on loggerhead sea turtle (Caretta caretta) populations found male‐biased breeding sex ratios and multiple mating by many females nesting in southwestern Florida. A sample of nesting loggerhead females who laid more than one nest over the course of the season and a subset of their hatchlings were examined from 36 clutches in 2016 on Sanibel Island, Florida. Males that fathered hatchlings in the first clutch sampled were identified in subsequent clutches. Interestingly, 75% of the females analyzed had mated singly. No male was represented in more than one female's clutches. The results suggest that females likely mate at the beginning of the season and use stored sperm for multiple clutches. Evidence for mating between laying events was limited. There was no consistent pattern across the subsequent multiple paternity clutches, suggesting benefits to loggerhead females likely equal their costs and subsequent mating is likely determined by female preference.

Loggerhead Caretta caretta and leatherback Dermochelys coriacea sea turtles are caught frequently as longline bycatch. Loggerhead turtles are often hooked in the mouth, while leatherbacks often are hooked in the shoulder or flippers. Comparisons of feeding behavior and accuracy in biting surrogate ‘prey’ targets, in the presence of waterborne food odors, identified species-specific differences that may predispose the turtles to be hooked differently. Additionally, the visual fields of post-hatchling loggerhead and leatherback sea turtles were measured and compared to determine if field of view could further contribute to feeding behavior differences and refining the explanation as to why the 2 species are hooked differently. We found that loggerhead turtles, which have a larger visual field and a small area of binocular vision, were more accurate when biting a target than leatherbacks. These traits may explain their greater incidence of mouth, throat, and stomach hookings. Leatherback turtles have a smaller field of view and no overlap of visual fields, and they were less accurate in apprehending targets. Leatherbacks lack binocular vision, which might explain why they are prone to entanglement or hooking external to the mouth more often than loggerheads. Together, visual field and biting accuracy may help explain why leatherbacks and loggerheads differ in how they interact with fishing lines and the locations of their hookings. Such understanding of the species’ behavior and limits of visual fields is essential for formulating further bycatch reduction approaches.

In-water behaviour and long-term movements of oceanic-stage juvenile sea turtles are not well described or quantified. This is owing to technological or logistical limitations of tracking small, fast-growing animals across long distances and time periods within marine habitats. Here, we present, to our knowledge, the first long-term offshore tracks of oceanic green turtles ( Chelonia mydas ) in western North Atlantic waters. Using a tag attachment technique developed specifically for young (less than 1 year old) green turtles, we satellite-tracked 21 oceanic-stage green turtles (less than 19 cm straight carapace length) up to 152 days using small, solar-powered transmitters. We verify that oceanic-stage green turtles: (i) travel to and remain within oceanic waters; (ii) often depart the Gulf Stream and North Atlantic Subtropical Gyre currents, orienting towards waters associated with the Sargasso Sea; (iii) remain at the sea surface, using thermally beneficial habitats that promote growth and survival of young turtles; and (iv) green turtles orient differently compared to same stage loggerhead turtles ( Caretta caretta ). Combined with satellite tracks of oceanic-stage loggerhead turtles, our work identifies the Sargasso Sea as an important nursery habitat for North Atlantic sea turtles, supporting a growing body of research that suggests oceanic-stage sea turtles are behaviourally more complex than previously assumed.

Temperature-dependent sex determination, present in most turtle species, is a mechanism that uses temperature to direct the sex of the embryo. The rapid increase of global temperatures highlights the need for a clear assessment of how sex ratios of organisms with TSD are affected. In turtles with TSD, quantifying primary sex ratios is challenging because they lack external dimorphism and heteromorphic sex chromosomes. Here we describe a new technique used to identify sex in neonate turtles of two TSD species, a freshwater turtle (Trachemys scripta) and a marine turtle (Caretta caretta) via analysis of small blood samples. We used an immunoassay approach to test samples for the presence of several proteins known to play an important role in sex differentiation. Our results show that Anti-Mullerian Hormone (AMH) can be reliably detected in blood samples from neonate male turtles but not females and can be used as a sex-specific marker. Verification of sex via histology or laparoscopy revealed that this method was 100% reliable for identifying sex in both T. scripta and C. caretta 1–2 day-old hatchlings and 90% reliable for identifying sex in 83–177 day-old (120–160 g) loggerhead juveniles. The method described here is minimally invasive, and for the first time, greatly enhances our ability to measure neonate turtle sex ratios at population levels across nesting sites worldwide, a crucial step in assessing the impact of climate change on imperiled turtle species.

Although sea turtles have received substantial focus worldwide, research on the immature life stages is still relatively limited. The latter is of particular importance, given that a large proportion of sea turtle populations comprises immature individuals. We set out to identify knowledge gaps and identify the main barriers hindering research in this field. We analyzed the perceptions of sea turtle experts through an online survey which gathered their opinions on the current state of affairs on immature sea turtle research, including species and regions in need of further study, priority research questions, and barriers that have interfered with the advancement of research. Our gap analysis indicates that studies on immature leatherback Dermochelys coriacea and hawksbill Eretmochelys imbricata turtles are lacking, as are studies on all species based in the Indian, South Pacific, and South Atlantic Oceans. Experts also perceived that studies in population ecology, namely on survivorship and demography, and habitat use/behavior, are needed to advance the state of knowledge on immature sea turtles. Our survey findings indicate the need for more interdisciplinary research, collaborative efforts (e.g. data-sharing, joint field activities), and improved communication among researchers, funding bodies, stakeholders, and decision-makers. Fil: Wildermann, Natalie E.. Florida State University; Estados Unidos Fil: Gredzens, Christian. Florida State University; Estados Unidos Fil: Avens, Larisa. National Ocean And Atmospheric Administration; Estados Unidos Fil: Barrios Garrido, Héctor A.. Universidad del Zulia; Venezuela. Florida State University; Estados Unidos Fil: Bell, Ian. Aquatic Species Program. Conservation And Biodiversity Operations Branch; Australia Fil: Blumenthal, Janice. Cayman Islands Department Of Environment; Islas Caimán Fil: Bolten, Alan B.. University of Florida; Estados Unidos Fil: McNeill, Joanne Braun. National Ocean And Atmospheric Administration; Estados Unidos Fil: Casale, Paolo. Università degli Studi di Pisa; Italia Fil: Di Domenico, Maikon. Universidade Federal do Paraná; Brasil Fil: Domit, Camila. Universidade Federal do Paraná; Brasil Fil: Epperly, Sheryan P.. National Ocean And Atmospheric Administration; Estados Unidos Fil: Godfrey, Matthew H.. North Carolina Wildlife Resources Commission; Estados Unidos. University of North Carolina; Estados Unidos. University of Duke; Estados Unidos Fil: Godley, Brendan J.. University of Exeter; Reino Unido Fil: González Carman, Victoria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Subsede Instituto Nacional de Investigación y Desarrollo Pesquero; Argentina Fil: Hamann, Mark. James Cook University; Australia Fil: Hart, Kristen M.. United States Geological Survey; Estados Unidos Fil: Ishihara, Takashi. Sea Turtle Association of Japan; Japón. Suma Aqualife Park; Japón Fil: Mansfield, Kate L.. University Of Central Florida; Estados Unidos Fil: Metz, Tasha L.. No especifica; Fil: Miller, Jeffrey D.. No especifica; Fil: Pilcher, Nicolas J.. Marine Research Foundation; Malasia Fil: Read, Mark A.. Great Barrier Reef Marine Park Authority; Australia Fil: Sasso, Christopher. National Ocean And Atmospheric Administration; Estados Unidos Fil: Seminoff, Jeffrey A.. National Ocean And Atmospheric Administration; Estados Unidos Fil: Seney, Erin E.. University Of Central Florida; Estados Unidos Fil: Williard, Amanda Southwood. University of North Carolina; Estados Unidos Fil: Tomás, Jesús. Universidad de Valencia; España Fil: Vélez Rubio, Gabriela M.. Universidad de la República; Uruguay. Karumbé Ngo; Uruguay Fil: Ware, Matthew. Florida State University; Estados Unidos Fil: Jessica L. Williams. Tartarugas para o Amanhã; Mozambique Fil: Wyneken, Jeanette. Florida Atlantic University; Estados Unidos Fil: Fuentes, Mariana M. P. B. Florida State University; Estados Unidos

Marine turtles exhibit temperature-dependent sex determination (TSD). During critical periods of embryonic development, the nest's thermal environment directs whether an embryo will develop as a male or female. At warmer sand temperatures, nests tend to produce female-biased sex ratios. The rapid increase of global temperature highlights the need for a clear assessment of its effects on sea turtle sex ratios. However, estimating hatchling sex ratios at rookeries remains imprecise due to the lack of sexual dimorphism in young marine turtles. We rely mainly upon laparoscopic procedures to verify hatchling sex; however, in some species, morphological sex can be ambiguous even at the histological level. Recent studies using immunohistochemical (IHC) techniques identified that embryonic snapping turtle (Chelydra serpentina) ovaries overexpressed a particular cold-induced RNA-binding protein in comparison to testes. This feature allows the identification of females vs. males. We modified this technique to successfully identify the sexes of loggerhead sea turtle (Caretta caretta) hatchlings, and independently confirmed the results by standard histological and laparoscopic methods that reliably identify sex in this species. We next tested the CIRBP IHC method on gonad samples from leatherback turtles (Dermochelys coriacea). Leatherbacks display delayed gonad differentiation, when compared to other sea turtles, making hatchling gonads difficult to sex using standard H&E stain histology. The IHC approach was successful in both C. caretta and D. coriacea samples, offering a much-needed tool to establish baseline hatchling sex ratios, particularly for assessing impacts of climate change effects on leatherback turtle hatchlings and sea turtle demographics. Anat Rec, 300:1512-1518, 2017. © 2017 Wiley Periodicals, Inc.

The goal of this study was to assess the consequences of single versus multiple paternity by identifying paternity of clutches per female to identify whether there were detectable costs or benefits. Multiple mating can occur when the benefits of mating outweigh the costs, but if costs and benefits are equal, no pattern is expected. Previous research on loggerhead sea turtle (Caretta caretta) populations found male‐biased breeding sex ratios and multiple mating by many females nesting in southwestern Florida. A sample of nesting loggerhead females who laid more than one nest over the course of the season and a subset of their hatchlings were examined from 36 clutches in 2016 on Sanibel Island, Florida. Males that fathered hatchlings in the first clutch sampled were identified in subsequent clutches. Interestingly, 75% of the females analyzed had mated singly. No male was represented in more than one female's clutches. The results suggest that females likely mate at the beginning of the season and use stored sperm for multiple clutches. Evidence for mating between laying events was limited. There was no consistent pattern across the subsequent multiple paternity clutches, suggesting benefits to loggerhead females likely equal their costs and subsequent mating is likely determined by female preference., The focus of this paper is to examine and describe the costs and benefits of multiple mating across taxa and then focus on a case study by examining repeating nesting of loggerhead sea turtles (Caretta caretta) nesting in the Gulf of Mexico. We identified in all nests, males that were present in the first nest were also present in subsequent nests (confirming sperm storage). We also found instances of possible sperm depletion and weak evidence for remating events, but no evidence of male genotypes in multiple female's nests (polygyny).

All species of sea turtles exhibit a cooler male/warmer female temperature-sex ratio response. Field and experimental studies on loggerhead sea turtle sex ratios suggest that increased sand moisture impacts sea turtle sex ratios with, and perhaps beyond, a cooling effect. This study examines how varying sand moisture impacts the embryo’s response to temperature. Across three years, loggerhead (Caretta caretta) sea turtle eggs were incubated at temperatures ranging from 28.0 °C to 33.0 °C. Groups of eggs were incubated in one of three volumetric moisture contents: low, medium, or high. Temperatures inside the group of eggs were recorded throughout incubation. Hatchlings were raised for 2–3 months and sex was identified laparoscopically. We calculated temperature response curves for groups of eggs incubated at each moisture level. Pivotal temperatures did not different among eggs incubated in different sand moistures. The transitional ranges of temperatures (TRT) for eggs incubated in high moisture and low moisture were narrower than the TRT for eggs incubated in medium moisture. The results of this study are crucial for understanding how sea turtle embryos respond to temperature directly or indirectly under different moisture conditions. Current sex ratio predictions rely on the embryos response to temperature only and may inaccurately estimate sex ratios especially during periods of heavy rainfall or drought.

International audience; Phenological shifts, by initiating reproductive events earlier, in response to advanced seasonal warming is one of the most striking effects currently observed in wild populations. For sea turtles, phenological adjustment to warming conditions could be the most effective short-term adaptation option against climate change. We calculated future phenological changes required in seven important loggerhead (Caretta caretta) nesting populations to continue achieving a high hatching success and a sex ratio that lies within current ranges. Considering temperature-mediated phenological changes, we found that most populations (six out of seven) will not be able to keep pace with a warming climate. Under an optimistic climate warming scenario (RCP4.5), these populations will face a climatic debt, that is, a difference between required and expected phenological changes, and warming will substantially reduce hatching success and induce a feminization of hatchlings, which may jeopardize their reproductive sustainability. Our approach offers the possibility to quantify the efficiency of phenological shifts in oviparous reptiles by considering physiological, developmental and phenological processes.

The influence of environmental conditions during sea turtle egg development makes the ongoing threat of climate change particularly concerning. The nest environment greatly influences several aspects of sea turtle hatchling biology, including developmental rate and sexual differentiation. Various studies examine the effectiveness of watering nests as a mitigation strategy for high temperatures and dry conditions but few have rigorously identified the impacts across multiple nesting seasons and weather conditions. This study experimentally examined the impact of increased moisture on loggerhead turtle (Caretta caretta) hatchling sex ratios and hatching success across three nesting seasons. Supplemental water was added to experimental treatments while control treatments received only ambient water as rainfall. Sand moisture and nest temperature were measured throughout incubation. Overall, watering nests lowered nest temperatures and increased nest sand moisture. Watered nests had significantly higher hatching success; however, nests that received the highest treatment amounts had similar hatching success as controls. After hatchlings emerged, nests were inventoried and a sample of hatchlings from each nest was collected and their sex was verified. Only nests watered with the highest water volume treatment produced significantly more males. Watering nests may mediate the effects of high temperatures and desiccation associated with climate change. However, as temperatures continue to increase, watering nests may not prove effective in mitigating for entirely female-producing beaches.

Marine turtles produce hundreds of precocial offspring (“hatchlings”) that are virtually defenseless. Many are consumed by predators. Hatchlings improve their survival prospects by migrating to offshore “nursery” areas with lower predator densities and, as they grow, by developing morphological defenses. The flatback turtle (Natator depressus), however, remains in the predator-rich coastal waters of Australia. To gain insights into how they survive there, we compared patterns of early growth and morphological development in flatbacks to their closest relative, the green turtle (Chelonia mydas), which migrates offshore. We found that morphological structures likely to be used in defense are better developed in juvenile flatbacks than in juvenile green turtles. Those structures probably represent one of a suite of characters that enable young flatbacks to survive in coastal habitats where interactions with predators are likely to be more frequent.

For turtles, the thermal environment experienced during development plays critical roles in many biological processes. While the temperature inside an egg is assumed to match the substrate temperature, many factors such as evaporative cooling, metabolic heating and the insulating properties of extra-embryonic components can lead to thermal differences. However, no method developed to date has allowed for measurement of the embryonic temperature in live chelonian eggs. We designed a thermocouple-based technique to measure embryonic temperature, achieving 94% survival in Trachemys scripta. This methodology may be applicable to other reptile species. We found that, while the temperature in the substrate adjacent to the eggshell accurately reflects the internal egg temperature, it differs from air temperature (∼2°C) in a moisture-dependent manner. Our results demonstrate that external egg temperature, but not air temperature, is suitable for assessing the effects of temperature on biological processes, which could be critical when considering that processes such as temperature-dependent sex determination in turtles occurs within a 4°C window.

Species that display temperature-dependent sex determination are at risk as a result of increasing global temperatures. For marine turtles, high incubation temperatures can skew sex ratios towards females. There are concerns that temperature increases may result in highly female-biased offspring sex ratios, which would drive a future sex ratio skew. Studying the sex ratios of adults in the ocean is logistically very difficult because individuals are widely distributed and males are inaccessible because they remain in the ocean. Breeding sex ratios (BSR) are sought as a functional alternative to study adult sex ratios. One way to examine BSR is to determine the number of males that contribute to nests. Our goal was to evaluate the BSR for loggerhead turtles (Caretta caretta) nesting along the eastern Gulf of Mexico in Florida, from 2013-2015, encompassing three nesting seasons. We genotyped 64 nesting females (approximately 28% of all turtles nesting at that time) and up to 20 hatchlings from their nests (n = 989) using 7 polymorphic microsatellite markers. We identified multiple paternal contributions in 70% of the nests analyzed and 126 individual males. The breeding sex ratio was approximately 1 female for every 2.5 males. We did not find repeat males in any of our nests. The sex ratio and lack of repeating males was surprising because of female-biased primary sex ratios. We hypothesize that females mate offshore of their nesting beaches as well as en route. We recommend further comparisons of subsequent nesting events and of other beaches as it is imperative to establish baseline breeding sex ratios to understand how growing populations behave before extreme environmental effects are evident.

Physiology as Integrated Systems Amanda Southwood Williard Vision Kerstin A. Fritsches and Eric J. Warrant Natal Homing and Imprinting in Sea Turtles Kenneth J. Lohmann, Catherine M.F. Lohmann, J. Roger Brothers, and Nathan F. Putman The Skeleton: An In Vivo View of Structure Jeanette Wyneken Age and Age Estimation in Sea Turtles Larisa Avens and Melissa L. Snover Molecular Genetics of Sea Turtles Michael P. Jensen, Nancy N. FitzSimmons, and Peter H. Dutton Oceanic Habits and Habitats: Dermochelys coriacea Vincent S. Saba Oceanic Habits and Habitats: Caretta caretta Katherine L. Mansfield and Nathan F. Putman Feeding Biology: Advances from Field-Based Observations, Physiological Studies, and Molecular Techniques T. Todd Jones and Jeffrey A. Seminoff Predators, Prey, and the Ecological Roles of Sea Turtles Michael R. Heithaus Exposure to and Effects of Persistent Organic Pollutants Jennifer M. Keller Fisheries Bycatch of Marine Turtles: Lessons Learned from Decades of Research and Conservation Rebecca Lewison, Bryan Wallace, Joana Alfaro-Shigueto, Jeff Mangel, Sara Maxwell, and Elliott Hazen Climate Change and Marine Turtles Mark Hamann, Mariana M.P.B. Fuentes, Natalie C. Ban, and Veronique J.L. Mocellin Free-Ranging Sea Turtle Health Mark Flint Sea Turtle Epibiosis Michael G. Frick and Joseph B. Pfaller Parasites of Marine Turtles Ellis C. Greiner Index

Marine turtles deposit their eggs in underground nests where they develop unattended and without parental care. Incubation temperature varies with environmental conditions, including rainfall, sun/shade and sand type, and affects developmental rates, hatch and emergence success, and embryonic sex. We documented (1) rainfall and sand temperature relationships and (2) rainfall, nest temperatures and hatchling sex ratios at a loggerhead turtle (Caretta caretta) nesting beach in Boca Raton, Florida, USA, across the 2010 to 2013 nesting seasons. Rainfall data collected concur- rently with sand temperatures at different depths showed that light rainfall affected surface sand; effects of the heaviest rainfall events tended to lower sand temperatures but the temperature fluctu- ations were small once upper nest depths were reached. This is important in understanding the po- tential impacts of rainfall as a modifier of nest temperatures, as such changes can be quite small. Nest temperature profiles were synchronized with rainfall data from weather services to identify re- lationships with hatchling sex ratios. The sex of each turtle was verified laparoscopically to provide empirical measures of sex ratio for the nest and nesting beach. The majority of hatchlings in the samples were female, suggesting that across the 4 seasons most nest temperatures were not suffi- ciently cool to produce males. However, in the early portion of the nesting season and in wet years, nest temperatures were cooler, and significantly more males hatched.

Marine turtles are large reptiles that compensate for high juvenile mortality by producing hundreds of hatchlings during a long reproductive lifespan. Most hatchlings are taken by predators during their migration to, and while resident in, the open ocean. Their survival depends upon crypticity, minimizing movement to avoid detection, and foraging efficiently to grow to a size too difficult for predators to either handle or swallow. While these behavioral antipredator tactics are known, changes in morphology accompanying growth may also improve survival prospects. These have been only superficially described in the literature. Here, we compare the similarities and differences in presumed morphological defenses of growing loggerhead (Caretta caretta) and Kemp's ridley (Lepidochelys kempii) posthatchlings, related species that differ in growth rate, timing of habitat shift (the return from oceanic to neritic locations), and size at maturity. In both species, vertebral spination and carapace widening increase disproportionally as small turtles grow, but later in ontogeny, the spines regress, sooner in ridley than in loggerhead turtles. Carapace widening occurs in both species but loggerheads are always longer than they are wide whereas in Kemp's ridley turtles, the carapace becomes as wide as long. Our analysis indicates that these changes are unrelated to when each species shifts habitat but are related to turtle size. We hypothesize that the spines function in small turtles as an early defense against gape-limited predators, but changes in body shape function throughout ontogeny—initially to make small turtles too wide to swallow and later by presenting an almost flat and hardened surface that large predators (such as a sharks) are unable to grasp. The extremely wide carapace of the Kemp's ridley may compensate for its smaller adult size (and presumed greater vulnerability) than the loggerhead. J. Morphol. 276:929–940, 2015. © 2015 Wiley Periodicals, Inc.

The academic world is full of people with rich personalities, brilliant minds, and eclectic interests. Thomas (T. H.) Frazzetta certainly was a special one of these. He passed away on New Year's Eve 2015, following several months of ill health. Ironically, New Year's Eve was one of his favorite holidays, only exceeded by Halloween. Sadly, he was unable to celebrate either in his final year.

Experimental and field studies of different turtle species suggest that moisture influences embryonic development and sex ratios, wetter substrates tend to produce more males, and drier substrates produce more females. In this study, we used Trachemys scripta elegans to test the effect of moisture on embryonic development and sex ratios. T. s. elegans eggs were incubated under different temperature and moisture regimes. We monitored embryonic development until stage 22 (after sex determination) and, for the first time, we estimated sex ratios using a male-specific transcriptional molecular marker, Sox9. Among treatments, we found differences in developmental rates, egg mass, and sex ratio. Embryos developed slowly in cooler and wetter sand substrate while water uptake by the eggs was significantly greater on wetter substrates. Developmental differences were due to moisture interacting with temperature where increased water content of the sand resulted in temperatures that were 2-3°C lower than air temperatures. The coolest and the wettest substrates produced 100% males compared to 42% males from the warmest and driest treatment. Further, we found that embryonic growth appears to be more sensitive to temperature at earlier stages of development and to moisture at later stages. This study shows how moisture may change the incubation conditions inside nests by changing the temperature experienced by eggs, which affects development, growth and sex ratios. The results of this study highlight the importance of including moisture conditions when predicting embryo growth and sex ratios and in developing proxies of embryonic development.

Many reptiles have temperature-dependent sex determination (TSD). Sex determination in marine turtles is described by a cool-male, warm-female pattern. Nest sand temperature strongly influences sea turtle embryo development and sex differentiation. Yet, variation in hatchling sex ratios is explained only partially by nest temperature and can be predicted only at the warmest and coolest temperatures. Hence, other factors during development influence sex determination. Rainfall is a common environmental variable that may impact development and sex determination. We experimentally evaluated bias in sex ratio production associated with nest moisture. Conditions tested in surrogate nests were sand moisture in combination with (i) very restricted evaporation, (ii) moderate evaporation (allowing evaporative cooling), and (iii) evaporative cooling plus cooling from rain-temperature water. We collected eggs from 32 unique loggerhead (Caretta caretta L.) turtle clutches, distributed them among the three different conditions, and incubated the eggs until they hatched. All hatchlings were raised for several months and sex was verified laparoscopically to establish sex ratios for each treatment. The nests were expected to produce 50:50 sex ratios or a moderate female bias (∼70%) based on incubation temperatures. All experimental treatments produced high male bias (87-96%). These results support the hypothesis that moisture impacts sex ratios through evaporation and rainfall-based cooling. High male bias was observed in nests with and without restricted evaporative cooling and no direct cooling due to watering as well as those nests hydrated via cool (rainwater temperature) water. High moisture conditions may produce males through thermal or other mechanisms, highlighting the importance of examining other nest environmental factors on sex determination.

Leatherback sea turtles are globally distributed and endangered throughout their range. There are limited data available on disease in this species. Initial observations of solitary large intestinal diverticulitis in multiple leatherbacks led to a multi-institutional review of cases. Of 31 subadult and adult turtles for which complete records were available, all had a single exudate-filled diverticulum, as large as 9.0 cm in diameter, arising from the large intestine immediately distal to the ileocecal junction. All lesions were chronic and characterized by ongoing inflammation, numerous intralesional bacteria, marked attenuation of the muscularis, ulceration, and secondary mucosal changes. In three cases, Morganella morganii was isolated from lesions. Diverticulitis was unrelated to the cause of death in all cases, although risk of perforation and other complications are possible.

Many environmental variables that affect incubating turtle eggs in the nest may also affect hatchling development, following hatchling emergence. However, these effects may be subtle and are largely unexamined. In this study, we analyzed the effect of sand moisture content during incubation on the postemergence growth rates of loggerhead sea turtles (Caretta caretta) in southeastern Florida. We divided 10 clutches in halves, reburied them, and exposed them to 1 of 2 treatments. At emergence, 7 clutches met minimum criteria for inclusion in the study. One halfclutch received only ambient rainfall (“dry” treatment) while the other half-clutch received ambient rainfall plus daily watering (“wet” treatment). Data loggers were used to record incubation temperatures in both groups. Hatchlings were captured at emergence and laboratoryreared over a period of ∼ 3 mo. Mass, straight carapace length (SCL), and straight carapace width (SCW) were measured weekly to track growth. Initial measurements were larger for turtles from the wet nests in all metrics. Turtles from wet nests grew more in SCW than turtles from dry nests. Turtle growth from the 2 treatments did not differ in SCL or mass measurements. Larger initial sizes and faster SCW growth may enable the turtles to more quickly achieve a refuge size from their gape-limited predators. Moisture availability during nesting season is projected to decrease based on climate change models. If that change materializes, it could negatively affect hatchling sizes and neonate growth rates, survival, and hence the recovery of this imperiled species.

Significant gaps exist in our understanding of early sea turtle life stages. Dispersal and habitat use of young oceanic sea turtles are largely inferred. Historically, available tracking technology and tag attachment methods were limited by small body sizes and rapid growth of neonate sea turtles. We tested methods in the laboratory for attaching small solar-powered satel- lite tags to neonate loggerhead sea turtles Caretta caretta, including harnesses, hard epoxy and neoprene-silicone mounts. Non-harness attachments were tested on turtles with clean carapaces and carapaces treated with an acrylic base-coat. Turtle growth and condition were measured among treatment and control groups. We tested surrogate solar cells, coated with clear silicone antifoulant for biofouling and performance, and field tested the performance of 7 solar-powered satellite tags on neonate loggerheads released off southeastern Florida (USA). Attachments with acrylic base-coats remained affixed 4- to 8-fold longer than on untreated carapaces. Harness attachments resulted in long-term tag retention (>60 d). However, harness and hard epoxy attach- ments did not adjust for turtle growth; we do not recommend these methods for rapidly growing neonate turtles. The method with longest retention was a neoprene-silicone attachment on an acrylic base-coat. Growth and condition were comparable among treated and control turtles using this flexible neoprene-silicone-acrylic attachment. Field-tested tags transmitted for 38 to 172 d. There were significant differences in charge rates of tags treated with an antifoulant and un - treated tags; however, all tags charged optimally and transmitted high-quality locations. Our data suggest that small solar-powered tags are viable tools for monitoring the in-water behavior of oceanic sea turtles.

In numerous studies involving hatchling sea turtles, researchers have collected small numbers of hatchlings from nests a few hours before the turtles would otherwise have emerged naturally. This procedure makes it possible to do experiments in which the behavioral or physio- logical responses of numerous hatchlings must be tested in a limited period of time, and also allows hatchlings to be released back into the sea in time to migrate offshore before dawn. In prin- ciple, however, the procedure might inadvertently reduce nest productivity (the number of hatch- lings that successfully leave the nest), if digging into a nest prior to emergence somehow reduces the ability of the remaining turtles to emerge. We compared nest productivity in 67 experimental loggerhead nests, from which we removed 10 hatchlings before a natural emergence, to 95 control nests left undisturbed before a natural emergence. The 2 groups showed no statistical differences in productivity. We conclude that taking a few hatchlings from a loggerhead nest shortly before a natural emergence has no negative impact on hatchling production if sampling is done with care at locations where there are few nest predators, and at sites where an emergence can be predicted because nest deposition dates are known.