Sea turtles are internationally threatened and as a result are protected by regional, federal, and international legislations (National Marine Fisheries Service 2008; Casale and Tucker 2015). This status reflects drastic population declines caused by threats such as fisheries bycatch, poaching, and habitat loss (Koch et al. 2006; Ryder et al. 2006; Fish et al. 2008; National Marine Fisheries Service 2008). Several management strategies are used to minimize these threats at both their terrestrial (e.g., nest relocation, caging against predation) and oceanic life-stages (e.g., use of turtle excluder devices, development of marine protected areas; Turtle Expert Working Group 1998; Hart et al. 2013; Revuelta et al. 2015). Nest relocation is one of the most widely employed management strategies used to mitigate threats during sea turtles' terrestrial stage, and historically has been used to minimize impacts to the reproductive output of turtles caused by inundation (Whitmore and Dutton 1985; McGehee 1990; Foley et al. 2006; Tuttle and Rostal 2010), erosion (Boulon 1999; Dellert et al. 2014; Ahles and Milton 2016), vehicular traffic (National Marine Fisheries Service 2011), artificial lighting (Trindell et al. 2008; Wilson 2009), and poaching (Mortimer 1999; Garcia et al. 2003; Liles et al. 2015a). If conducted properly, nest relocation can have a minimal impact on embryonic development and hatching success (Boulon 1999; Abella et al. 2007; McElroy et al. 2015; Ahles and Milton 2016). In some cases, relocated nests have demonstrated greater hatching success than have their in situ counterparts (Wyneken et al. 1988; Hoekert et al. 1998; Tuttle and Rostal 2010). Indeed, the long-term use of nest relocation to save otherwise “doomed” eggs is a potential component in the recovery of nesting populations in the United States (Hopkins-Murphy and Seithel 2005; National Marine Fisheries Service 2015), St. Croix (Eckert and Eckert 1990; Dutton et al. 2005), French Guiana, Suriname, and Gabon (Fossette et al. 2008), and other locations globally.

Despite the fact that nests may be “saved,” there are several concerns associated with nest relocation. When eggs are relocated after the start of embryonic attachment, movement of eggs can kill the embryo and reduce hatching success (Limpus et al. 1979; Parmenter 1980; Grand and Beissinger 1997; but see Abella et al. 2007 and Candan 2018). Further, incubating environment influences egg development; therefore, concern exists over potential alterations to this environment. Environmental conditions influence embryonic growth rate (Mrosovsky 1980; Reid et al. 2009), morphological development (Whitmore and Dutton 1985; Glen et al. 2003; Türkozan and Yilmaz 2007), sexual development (Mrosovsky and Yntema 1980; Mortimer 1999; Sari and Kaska 2017), and hatching and emergence success (Eckert and Eckert 1990; Wood and Bjorndal 2000; Glen et al. 2005; Maulany et al. 2012). If nest site selection in the adults is genetically driven, nest relocation may be relieving selective pressures that would otherwise reduce maladaptive traits (i.e., nesting in areas of frequent inundation) and could reduce future population viability (Kamel and Mrosovsky 2005; Mrosovsky 2006, 2008; Pike 2008).

Given potential issues inherent in relocating nests, several studies have explored the effects of relocation on embryonic development, sex ratios, and hatching and emergence success across sea turtle species and nesting locations (Eckert and Eckert 1990; Pintus et al. 2009; Tuttle and Rostal 2010; Ahles and Milton 2016). While these studies often compare reproductive output and environmental conditions from relocated and in situ nests, a key component to consider is information pertaining to the original location of the nest (i.e., thermal profile, sand characteristics, exposure to threats). This comparison of environmental conditions between the original and relocated location is needed to assess whether relocations are altering sex ratios, nest productivity, and mitigating threats (e.g., inundation). To address this, we explored 1) how sand temperature, inundation exposure, and sand grain size and moisture content change between original and relocated location pairs; and 2) how relocation affects nest productivity.

METHODS

Study Area

This study was conducted at Fort Morgan, Alabama, including the Bon Secour National Wildlife Refuge (BSNWR; Fig. 1, Global Positioning System [GPS]: 30.2262°N, 88.0279°W–30.2307°N, 87.8009°W). The refuge is divided into 2 Gulf of Mexico–facing units separated by residential Fort Morgan. Fort Morgan has a 22.2-km-long (total), south-facing microtidal dissipative-to-intermediate beach where loggerhead (Caretta caretta) sea turtles nest as part of the Northern Gulf of Mexico Recovery Unit of the Northwest Atlantic Distinct Population Segment (National Marine Fisheries Service 2008). This Recovery Unit is among the smallest in the United States, with 323–634 individuals estimated in the population (Richards et al. 2011). A representative beach profile is provided in Fig. 2 based on 10 light detection and ranging (LiDAR) surveys conducted from 1998 to 2016, which were retrieved from the National Oceanic and Atmospheric Administration Data Access Viewer (https://coast.noaa.gov/dataviewer/, accessed 20 September 2017).

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Monitoring

From 1 May through 31 August 2016, morning nesting surveys were conducted in collaboration with Share the Beach volunteers and US Fish and Wildlife Service personnel. The deposition of new nests occurs during this interval, with hatchling emergence occurring as late as October. Surveys consisted of driving utility vehicles along the beach looking for sea turtle activity (e.g., crawls, thrown sand, body pitting). Suspected nests were confirmed by excavating to the top of the egg chamber. For each nest encountered, data regarding the nest's deposition date, GPS location, distance to the high tide line (HTL), and nearest dune or obstruction were recorded following the procedures of the Alabama Sea Turtle Conservation Manual (ASTCM; US Fish and Wildlife Service 2008). Wire screens were placed over all nests to reduce depredation. In an effort to increase nest productivity by reducing inundation exposure, nests were relocated to higher elevations near the dune toe if they satisfied the requirements under the ASTCM (e.g., nest laid near or below the mean HTL). All relocations were performed within 12 hrs of nest deposition. Nest location was recorded for the new location of relocated nests, along with distance moved, number of eggs, and the reason for relocation (see Fig. 3 for a visual depiction of original–relocated–in situ terminology).

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Each nest was monitored for up to 75 d for signs of disturbance (e.g., depredation, inundation, erosion) or hatching per the ASTCM. In the event a disturbance was identified during a survey, the date, type of disturbance, causal agent, fate of the nest, and any remedial action taken was recorded. Nest inventories were conducted 3 d after hatchling emergence or at 75 d postoviposition, whichever came first, to determine nest productivity. Hatching success was defined as the number of open eggs divided by the total number of eggs. Emergence success was calculated as the number of hatchlings successfully leaving the nest divided by the number of hatched (i.e., empty) shells.

Thermal Profile

Information on the sand thermal profile of nest locations (in situ, relocated, and original) were collected hourly with DS1922L-F5# iButton temperature loggers for the duration of the nest's incubation. Mean sand temperature and mean daily amplitude (maximum daily temperature minus minimum daily temperature) during the middle third of incubation were determined. For nests left in situ, one iButton was placed at chamber depth 0.5 m adjacent to the egg chamber, as per Tuttle and Rostal (2010). For relocated nests, temperature loggers were deployed at nest chamber depth 0.5 m adjacent to both the original and relocated locations. Ibuttons were retrieved during the nest excavation. The iButtons were placed in the sand for location comparisons, as opposed to the nest chamber, in order to remove the effect of metabolic heating, which would not be present at the original nest location.

Exposure to Inundation

Inundation was monitored using an inundation measuring device, which consisted of a series of polyvinyl chloride (PVC) tubes deployed 0.5 m from the egg chamber parallel to the HTL following the methodologies of Foley et al. (2006) and Ware and Fuentes (2018). Three 1.27-cm-diameter PVC tubes, each with a hole drilled either 30, 40, or 50 cm from the sand surface, were stored inside a larger 5.08-cm-diameter PVC pipe. The distances correspond to the average depth to the top, middle, and bottom of the egg chamber for loggerhead nests at Fort Morgan (D. Ingram, US Fish & Wildlife Service, pers. comm., January 2016). For nests left in situ, one inundation device was deployed 0.5 m adjacent to the egg chamber. For relocated nests, devices were deployed 0.5 m adjacent to the original and relocated egg chambers.

Four metrics were used to assess inundation stress: 1) frequency, which corresponds to number of days a site recorded inundation throughout the site's incubation period; 2) severity, the percentage of the nest exposed to groundwater during an inundation event; and 3) duration, the number of consecutive days of groundwater inundation observed. The frequency, severity, and duration were then combined into a fourth metric—a cumulative inundation stress index (CISI). The CISI score was calculated by multiplying the length of inundation (in days) at a particular depth by the proportion of eggs exposed to inundation at that depth, summed across all inundation events during incubation (see Supplemental Material for an example calculation; all supplemental material is available online at http://doi.org/10.2744/CCB-1306.1.s1).

Inundation devices were deployed for the duration of the nest's incubation and were monitored daily as part of morning surveys. The date and cause of wash-over events of any monitoring site were also recorded. Wave-related wash-over events differ from inundation events in that wash-over events are concentrated on the nest surface. These events are responsible for the erosion or accretion of wet sand and do not trap water within the inundation devices, except under extreme circumstances (e.g., a hurricane), owing to the design of the device. Wash-overs were identified by alterations to the nest's surface (i.e., deposition of wrack material, deposition or removal of wet sand). Inundation events submerge eggs from below the surface due to increases in the height of the water table (e.g., high tide, storm surge) and may leave no trace of their occurrence on the surface of the nest.

Sand Characteristics

Sand samples (∼ 500 cm³) were collected from the surface and at nest chamber depth 0.5 m adjacent to the nest chamber for original, relocated, and in situ nest locations. Grain size was determined by sequential dry-sieving through 1-mm-, 500-μm-, 250-μm-, 180-μm-, 150-μm-, and 125-μm-diameter screens, with the relative contribution given by dry mass of the given size fraction divided by the total dry mass (Foley et al. 2006). Mean particle size, grain size distribution, and moisture content were calculated following Foley et al. (2006) in GRADISTAT (GRADISTAT version 8.0, http://www.kpal.co.uk/gradistat.html, accessed 12 September 2016).

Statistical Analyses

Thermal profiles, inundation history, and sand characteristics were compared between original and relocated locations within relocated nests, as well as between in situ and relocated nests, to determine any differences in the incubation environment. Hatching and emergence success were evaluated between in situ and relocated nests. Data were assessed for normality by Shapiro-Wilks Tests. Student's t-tests were used for normally distributed data, while Wilcoxon tests were used for nonnormal, continuous data, and Poisson and chi-square tests for count data.

Differences in environmental parameters and nest productivity between the BSNWR units were assessed before forming a pooled “Refuge” data set, as were “Refuge” and “Residential” data before forming a combined-location data set. The combined-location data were used unless otherwise noted. All statistical analyses were performed in Program R (R Version 3.4.3).

RESULTS

In 2016, 94 nests were observed, of which 20 were relocated because of their proximity to the HTL (Supplemental Table S1). The mean distance to the daily HTL for the original location of a relocated nest was 10.2 m (± 1.2 m SE) with a minimum of 0 m and a maximum of 22.4 m. Relocated nest locations ranged from 19.0 to 66.0 m with an average of 33.3 m (± 7.5 m SE). In situ nests averaged 19.9 m (± 1.1 m SE; range, 3.0–50.0 m) from the daily HTL. There were significantly more relocations off-refuge than on-refuge (n = 15 and 5, respectively, χ² p = 0.005) despite no difference in the mean distance to the HTL for on- vs. off-refuge nests (Refuge: 17.2 m ± 1.4 m SE, Residential: 18.7 m ± 1.4 m SE, Wilcoxon test p = 0.44). Monitoring sites (inundation device, temperature sensor, or both) were established covering 70 different nests, including all 20 original–relocated pairs. Not all the nests laid could be included in the study because of limitations in available equipment.

Thermal Profile

Across the full incubation duration, original locations were 0.2°C warmer than relocated locations, and in situ nests were 0.4°C warmer than relocated nests, on average, when measured 0.5 m adjacent to the egg chamber (Table 1). However, these differences were not significant (original–relocated paired t-test p = 0.169, in situ–relocated Wilcoxon test p = 0.077). During the middle third of incubation, original and relocated locations had identical mean temperatures and daily amplitudes (Table 1 and Supplemental Fig. S1; mean temperature: 29.6°C, paired t-test p = 0.864; daily amplitude: 1.6°C, paired t-test p = 0.387). In situ nests had slightly warmer mean temperatures and larger daily amplitudes than relocated nests during the middle third of incubation (Table 1), although these differences were not statistically significant (t-test p = 0.313 and 0.386, respectively). Incubation duration, however, was significantly different—in situ nests averaged 2.7 d longer than relocated nests (Wilcoxon test p = 0.009).

Table 1. Summary of environmental variables and nest productivity during assessment of differences in sand temperature, inundation exposure, grain size, and moisture content between the original and final locations of relocated loggerhead turtle nests. Mean ± SE sand temperature metrics are relative to the middle third of incubation. CISI = cumulative inundation stress index. “Lost to inundation” refers to a location that was inundated and produced zero hatchlings, or was eroded away.

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Nest Inundation

Fifty-five percent of nests recorded some level of groundwater inundation. No difference was observed between in situ and relocated nests with respect to the number of inundated sites (χ² p = 1, Table 1). Overall, original locations were not more likely to be inundated with groundwater (χ² p = 0.112), nor washed over (χ² p = 0.201) than their relocated counterparts. However, of the original–relocated pairs that were inundated, original locations recorded a greater mean frequency of inundation (6.9 ± 1.5 inundations) than relocated locations (2.1 ± 0.6 inundations, Poisson p ≪ 0.001), along with a greater CISI (original: 3.9 ± 0.7, relocated: 1.6 ± 0.5, paired Wilcoxon p = 0.006). Of the pairs that were washed over, original locations recorded a greater mean frequency of wash-over (2.1 ± 0.4 wash-overs) than relocated locations (0.9 ± 0.3 wash-overs, Poisson p = 0.014). Fifty-four percent of nests were washed over at least once during their incubation, with no difference observed between in situ and relocated nests (χ² p = 0.632).

Sand Characteristics

Mean grain size at chamber depth at the original nest locations was 24.7 μm (± 7.9 μm SE; Table 2) larger than the relocated locations, a significant difference (paired Wilcoxon test p = 0.009). Significant differences between original and relocated locations were also found for median grain size (paired Wilcoxon test p = 0.012), sorting (paired Wilcoxon test p = 0.017), skewness (paired Wilcoxon test p = 0.043), and kurtosis (paired Wilcoxon test p = 0.001). No significant differences were observed between in situ and relocated nests in any sand grain comparison (mean grain size: Wilcoxon test p = 0.318; median grain size: Wilcoxon test p = 0.501; sorting: Wilcoxon test p = 0.302; skewness: Wilcoxon test p = 0.852; kurtosis: Wilcoxon test p = 0.347; Table 2). Mean initial moisture content for in situ nests at chamber depth was 2.66% (± 0.24% SE), whereas it was 2.90% (± 0.32% SE) for relocated nests. This difference was not significant (Wilcox test p = 0.703), nor was the 0.26% difference between original–relocated pairs at chamber depth (paired Wilcoxon test p = 0.266; Table 2). Mean initial moisture content in relocated location surface samples were approximately 54% that of original or in situ locations, although this difference was not significant (in situ–relocated Wilcoxon test p = 0.895, original–relocated paired Wilcoxon test p = 0.065).

Table 2. Sand grain distribution and moisture content reported as mean ± SE, during assessment of differences between the original and final locations of relocated loggerhead turtle nests.

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Nest Productivity

In situ nests averaged greater hatching success (+ 0.5%) and emergence success (+ 8.3%) than relocated nests (Table 1). However, only emergence success was significantly different (Wilcoxon p = 0.714 and 0.030, respectively). Fourteen nests (14.9%) were eroded away or did not hatch following inundation; only 3 of the 20 original locations washed away.

DISCUSSION

No differences were found between the original and final locations for relocated loggerhead nests with respect to sand temperature, likelihood of inundation, or sand moisture. Relocations did, however, reduce incubation duration and emergence success. In theory, the reduced incubation duration in relocated nests should indicate warmer temperatures (Marcovaldi et al. 1997; Mrosovsky et al. 2009; DeGregorio and Williard 2011). However, there were no significant differences in mean sand temperature or daily temperature amplitude between in situ and relocated nests. The observed difference in incubation duration may be related to sand compaction or other environmental variables associated with hatchling emergence not measured during this study (Crain et al. 1995; Miller et al. 2003). For example, reduced sand compaction above relocated nests would result in greater air-filled pore space, increasing gas exchange and thereby reducing incubation duration by reducing hypoxic stress late in embryonic development (Suss 2013).

Irrespective of potential temperature changes, one of the most common reasons for nest relocation is to reduce losses due to inundation. The nature of the study site was microtidal, so significant wave and elevated tide exposure was only observed during cyclonic storm events. During these storms, the majority of the cross-shore beach area was affected, with waves inundating and/or washing over both the original and relocated locations simultaneously. Though relocation may not have reduced the likelihood of inundation in Fort Morgan, it did reduce the severity of the inundation event when it occurred. Inundation poses a serious threat because sea turtle embryonic tolerance to inundation is poorly understood (Foley et al. 2006; Caut et al. 2010; Pike et al. 2015; Ware and Fuentes 2018). Inundation is not inherently lethal. Observations made by Foley et al. (2006) in the Ten Thousand Islands, Florida, demonstrated hatching success ≥ 70% in several nests despite ≥ 3 inundations during incubation, though the nest responses were highly variable. Two days of inundation only reduced hatching success by 27% in Keewaydin Island, Florida (Shaw 2013). Significant inundation can restrict gas exchange across the shell membrane, but the variability in survival is likely a function of the frequency, severity, and timing of groundwater inundation. Any sublethal effects of inundation (e.g., neurological or muscular impairment, altered spatial orientation) also represent a significant knowledge gap (McGehee 1990; Pike et al. 2015). Relocation practices and decisions could be better informed by further knowledge of the tolerance of embryos to inundation and improved identification of areas within nesting beaches that are most prone to inundation (Ware and Fuentes 2018).

In addition to inundation, alterations to the physical sand characteristics of the incubation site due to relocation may affect embryonic development and hatchling emergence. Despite the short-distance relocations within the same beach, all metrics of grain size distribution between original and relocated nest locations were significantly different. Grain size influences moisture retention and gas exchange within the sediment (Mortimer 1990; Rumbold et al. 2001; Mota 2009). However, initial moisture content was not statistically different between original and relocated nest locations. This is contrary to Sönmez and Özdilek (2013), who noted significant declines in sand moisture associated with nest relocation in Turkey. Reduced sand moisture can increase the diffusion of oxygen (Mota 2009), but increases the risk of cave-ins as well (Mortimer 1990; Hewavisenthi and Parmenter 2002).

At the other extreme, excessive sand moisture has been correlated with reduced hatching success across species (McGehee 1990; Patino-Martinez et al. 2014; Wyneken and Lolavar 2015). If sand moisture is elevated, but at nonlethal levels, it may promote increased male hatchling production—a key consideration given climate predictions (Fuentes et al. 2012; Jourdan and Fuentes 2015; Lolavar and Wyneken 2015; Wyneken and Lolavar 2015). Wyneken and Lolavar (2015) and Lolavar and Wyneken (2015) documented increased male production in both natural nests incubated during years with above average rainfall and laboratory-incubated nests under typically female-producing temperatures but greater sand moisture. They hypothesized that either the pivotal temperature of South Florida loggerheads is higher than expected, increased moisture modifies the transitional range of temperatures that determines loggerhead hatchling sexual differentiation, or the mechanisms of sexual differentiation covary with moisture or other variables beyond temperature.

With respect to nest productivity, relocated nests had 8.3% lower emergence success relative to their in situ counterparts, with insignificant changes in other metrics of nest productivity. This change is consistent with several other studies, which documented reduced hatching and/or emergence success associated with nest relocation in Turkey, the US Virgin Islands, Ascension Island, and the United States (Eckert and Eckert 1990; Pintus et al. 2009; McElroy et al. 2015; Candan 2018). If relocated nests had 1) reduced sand compaction, and/or 2) reduced moisture in surface sands relative to in situ nests, these conditions would promote cave-ins as the hatchlings attempted to crawl to the surface (Mortimer 1990; Hewavisenthi and Parmenter 2002). Though the difference was not statistically significant, the initial surface-sand moisture content of relocated nests was roughly 54% that of in situ nests. Assuming these drier conditions persisted throughout the nesting and hatching seasons, this suggests that cave-ins may have played a role in emergence success in Fort Morgan. Sand compaction was not measured as part of this study, so reduced compaction at relocated nests remains speculation.

Given that relocation did not improve nest productivity nor reduce the likelihood of inundation, this practice conferred minimal net benefit to sea turtle nests in Fort Morgan, Alabama, in its current form. The only benefit occurred during cyclonic storms when the increased elevation and distance from the daily high tide line of the relocated nest location decreased the overall severity of the inundation. To improve the assessment of inundation stress experienced by the nests in our study, a unitless cumulative inundation stress index was developed that accounted for frequency, severity, and duration of groundwater inundation. Such an approach is useful because it creates a single unit for comparison of inundation stress between nests throughout their incubation. This is the first time such a stress index has been applied; therefore, additional development is required to improve the input parameters (e.g., higher resolution recording of inundation duration and severity, tolerance of embryos to inundation, effects of developmental stage on inundation tolerance, sublethal effects) and to calibrate the index relative to nest productivity before the index can be fully operationalized.

In addition to the need to improve our understanding of inundation tolerance and thus better inform relocation decisions, relocation criteria and guidelines must also be clarified. For example, the Alabama Sea Turtle Conservation Manual states that nest relocation is authorized when the nest is laid “near or below the mean high tide line” (US Fish and Wildlife Service 2008, p. 25). Ambiguity in “near” and variability in high tide line position can result in different operational definitions under this authorization, as demonstrated by the difference in the number of relocations within the BSNWR and residential Fort Morgan despite comparable nesting locations. Changes in beach slope and elevation should be taken more explicitly into account in relocation decisions (Shaw 2013). Clear guidance is also required when deciding on a final location for a relocated nest. Close proximity to the dune may encourage mammalian predation (Fowler 1979; O'Connor et al. 2017) and root incursion from nearby vegetation into the nest (pers. obs.). It is important to note that relocation did not influence whether or not a nest was predated in Fort Morgan, although the dominant predator at our study site was the ghost crab (Ocypode quadrata) as opposed to a mammalian predator. When investigating ghost crab (Ocypode cursor) predation in Boa Vista, Marco et al. (2015) also noted no significant difference in predation between relocated and in situ nests. When choosing a relocation site, other variables of concern include sand moisture content, sand density, total organic content, and total calcium carbonate, each of which may affect gas transfer, embryonic development, and hatchling emergence (Milton et al. 1997; Mota and Peterson 2002; Mota et al. 2005; Mota 2009).

There is an active debate on the potential selective pressures (e.g., alterations to hatchling sexual and morphological development, fitness, nest site selection) and benefits (e.g., environmental outreach) of nest relocation (Kamel and Mrosovsky 2005; Mrosovsky 2006, 2008; Pike 2008). If nest site selection is heritable and individuals exhibit significant repeatability in site-selection preferences, nest relocation may preserve “poor location” nesters through artificial selection and be significantly detrimental to population longevity (Kamel and Mrosovsky 2005; Mrosovsky 2006, 2008). However, the argument has been made that the public conservation education opportunities offered by nest relocation in the short term may outweigh genetic costs of artificial selection (Tisdell and Wilson 2005; Pike 2008).

This study observed a significant difference in hatchling emergence associated with relocation. The literature does not agree on relocation-related mortality across nesting beaches (Limpus et al. 1979; Wyneken et al. 1988; Eckert and Eckert 1990; Grand and Beissinger 1997; Pintus et al. 2009; Tuttle and Rostal 2010; McElroy et al. 2015). Assumptions regarding nest-site selection preferences (Kamel and Mrosovsky 2004, 2005; Pfaller et al. 2009; Liles et al. 2015b), susceptibility to inundation and erosion (Whitmore and Dutton 1985; Foley et al. 2006; McElroy et al. 2015), and geophysical controls of incubation temperature (Mortimer 1990; Hays et al. 2001; Booth and Freeman 2006) may differ between nesting sites and species. These across- and within-site variations require the use of local data to minimize management and genetic costs and maximize ecological benefits with respect to nest relocation when applying population- or species-level management actions.

Conclusions

Inundation poses a serious risk to sea turtle nests, which is commonly addressed by nest relocation. There are concerns that use of this strategy may alter the incubating environment of the developing embryos, and thus proper hatchling development and fitness. Our study alleviated several of these concerns because the incubating environment between original and relocated nest locations were comparable. However, nest relocation in the dissipative beach conditions of Fort Morgan, Alabama, offered a minimal net benefit because it decreased emergence success and did not reduce the likelihood of inundation. Additional information is required in the relocation decision process to improve the selection of nests truly at risk of inundation, thus improving the efficiency and efficacy of this technique. This may not be the case in other locations where tidal ranges are much greater, relocations occur over greater distances, or where nests are placed into hatcheries (Morreale et al. 1982; Mortimer 1999; Kornaraki et al. 2006; Liles et al. 2015b). Local assessments of original and relocated incubation conditions, a better understanding of the tolerance of sea turtle embryos to inundation, and clarity regarding relocation criteria and guidelines are required to improve nest relocation as a conservation strategy.