COASTAL AND MARINE ECOLOGY

The thermal impacts of beach nourishment across a regionally

important loggerhead sea turtle (Caretta caretta) rookery
KAITLYNN M. SHAMBLOTT,1 JAYMIE L. RENEKER,2 AND STEPHANIE J. KAMEL1,
1
Department of Biology and Marine Biology, Center for Marine Science, University of North Carolina Wilmington, Wilmington, North
Carolina 28409 USA
2
Ecological Associates, Jensen Beach, Florida 34597 USA

Citation: Shamblott, K. M., J. L. Reneker, and S. J. Kamel. 2021. The thermal impacts of beach nourishment across a
regionally important loggerhead sea turtle (Caretta caretta) rookery. Ecosphere 12(3):e03396. 10.1002/ecs2.3396

Abstract. Beach nourishment is a common coastal management practice used to protect and maintain
infrastructure, tourist revenue, and sandy beach habitats. Coastal erosion and increased development
along the coast of North Carolina, USA, have resulted in increased use of beach nourishment as an envi-
ronmentally friendly alternative to hard structures, such as groins and jetties. Despite its advantages, beach
nourishment can alter the thermal properties of a beach, potentially impacting the incubation environment
of species that utilize this habitat during reproduction. Importantly, in organisms with temperature-depen-
dent sex determination, the incubation environment plays a key role in determining offspring sex ratios,
hatchling survival, and fitness. Here we investigate how beach nourishment influences thermal properties
and sand characteristics of eight beaches in the high-density loggerhead sea turtle (Caretta caretta) nesting
region of North Carolina. We find that, despite considerable spatial and temporal variation, nourishment is
a significant predictor of mean monthly sand temperatures in both univariate and multivariable predictive
models. Across a season, nourished beach sections are, on average, 0.4°C (range 0.3–0.8°C) warmer than
their unnourished counterparts. Nourishment is also a significant predictor of the mean and variance of
sand grain size. Furthermore, variation in mean grain size, the relative percent of small, medium and large
grain sizes and albedo are mainly responsible for differences in mean monthly sand temperatures. As such,
the coarser and darker sand often used in nourishment projects may exacerbate climate driven increases in
surface temperature.

Key words: beach erosion; climate change; nesting beach; sand grain characteristics; sand temperature; sea turtle.

Received 24 June 2020; revised 1 October 2020; accepted 21 October 2020; final version received 16 December 2020.
Corresponding Editor: Sean P. Powers.
Copyright: © 2021 The Authors. This is an open access article under the terms of the Creative Commons Attribution
License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
E-mail: kamels@uncw.edu

INTRODUCTION current and future projected concentrations of

atmospheric greenhouse gases and indicators
The impacts of climate change, primarily such as ocean acidity and ice mass coverage in
increased land and ocean surface temperatures the poles, the warming trend is expected to con-
(Hansen et al. 2010), sea level rise (Spencer et al. tinue throughout the next century (IPCC 2014).
2016, Schuerch et al. 2018) and increased severity With global mean sea level expected to rise by
and occurrence of precipitation events (Trenberth 0.3–0.8 m over this time (IPCC 2014), sandy, low-
2011, Emanuel 2013, Hanse et al. 2016) threaten lying coastal and island beaches are at special
coastal environments on a global scale. Given risk (Feagin et al. 2005, Baker et al. 2006,

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

FitzGerald et al. 2008, Hinkel et al. 2013), particu- All sea turtles exhibit temperature-dependent
larly when landward retreat is restricted by sex determination (TSD), where temperatures
human infrastructure and development (Gal- determine offspring sex during the thermosen-
braith et al. 2002, Fish et al. 2008, Noss 2011). sitive period in the middle third of incubation
Moreover, likely increases in the occurrence of (Yntema and Mrosovsky 1982, Mrosovsky et al.
major storm events as a result of increased ocean 1984). Female offspring are produced at higher
surface temperatures (Goldenberg et al. 2001, temperatures and males at lower temperatures
Trenberth 2011, IPCC 2014) further threaten within a thermal tolerance range of 25–35°C
these ecosystems as well as the species that rely (Ackerman 1997). The pivotal temperature,
on them (Reid and Trexler 1992, Sala et al. 2000, which results in a 1:1 ratio of male to female
Coombes et al. 2008, Van Houtan and Halley hatchlings, is 29°C for loggerheads (Caretta car-
2011, Schooler et al. 2019). etta) in the USA (Mrosovsky 1988) and is
Coastal management practices utilize a vari- bounded by a narrow 2–3°C window known
ety of techniques to protect infrastructure, tour- as the transitional range of temperatures in
ist dollars, and natural habitat from the which both sexes are produced (Pieau and
impacts of global climate change (Phillips and Mrosovsky 1991). This sensitivity leaves sea
Jones 2006, Fish et al. 2008). In the United turtles vulnerable to sub-lethal temperature
States, beach nourishment has emerged as a shifts in the incubation environment (Booth
potentially environmentally friendly solution to 2017). Specifically, increasing sand temperatures
combat coastal erosion and is currently the could produce female-biased primary sex
main approach used in such projects (Valverde ratios, contribute to decreased survivorship of
et al. 1999, Dean 2005, Speybroeck et al. 2006). clutches, alter nesting phenology and impact
Obvious advantages and benefits of beach hatchling characteristics, performance and fit-
nourishment include increasing or restoring ness (Poloczanska et al. 2009, Hamann et al.
habitat for endangered and threatened dune 2010, Hays et al. 2017).
plants, shorebirds and sea turtles (Greene 2002, North Carolina is the third most nourished
Dean 2005). However, there are concerns state in the United States and has completed over
regarding the quality of material used in nour- 300 nourishment projects along the coast since
ishment events and its effects on the wildlife 1939 (National Beach Nourishment Database
that rely on sandy beach ecosystems (Crain 2019). North Carolina also represents a region-
et al. 1995, Milton et al. 1997, Rumbold et al. ally important loggerhead rookery within the
2001, Peterson and Bishop 2005). For example, Northwest Atlantic Ocean Distinct Population
nourishment often creates steeper beach pro- Segment (DPS), which encompasses nesting bea-
files and scarps at the shoreline (Greene 2002). ches along the Atlantic coast of Florida though
Sand characteristics can also be altered with southern Virginia and is responsible for about
respect to color (Hawkes et al. 2005, Peterson 40% of all loggerhead nests laid annually
and Bishop 2005, Peterson et al. 2014), grain (68,000–90,000 nests; NOAA Fisheries). Here, we
shape, size, and porosity (Roman-Sierra et al. provide a comprehensive analysis of changes in
2014), as well as sediment minerology, density, sand temperature and grain size of eight beaches
and compaction (Speybroeck et al. 2006). Main- along the coast of North Carolina, USA, to inves-
taining proper sedimentary characteristics on tigate how beach nourishment influences ther-
beaches is especially important for species such mal properties and sand characteristics of
as sea turtles, since the nest incubation envi- loggerhead sea turtle nests. We use univariate
ronment directly impacts critical offspring char- and multivariable models to determine whether
acteristics (Packard and Packard 1988, Naro- nourishment is a significant predictor of sand
Maciel et al. 1999, Carthy et al. 2003) such as temperatures, in addition to other spatial and
offspring sex ratios (Yntema and Mrosovsky temporal variables. We also explore whether dif-
1982, Mrosovsky 1988), hatchling size (Atkin- ferences in sand grain size, composition, and
son 1994), and locomotor performance (Glen albedo directly influence sand temperature and
et al. 2003, Witt et al. 2010, reviewed in Booth whether these variables are significantly altered
2017). by nourishment events.

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

METHODS 6.4
5.9 nest/yr; maximum Topsail Island,

86.4
47.1 nests/yr).
Study sites
We collected data on sand composition and Data collection
temperature from eight beaches spanning the Sand temperature.—We collected sand tempera-
North Carolina coast: Pea Island National Wild- ture data in two different years: Pea Island NWR,
life Reserve (NWR), Cape Hatteras National Sea- Emerald Isle Beach, Topsail Beach, Bald Head
shore (CHNS) located in the town of Buxton Island, and Ocean Isle Beach in 2015 and
(hereafter, Buxton), Emerald Isle Beach, Topsail Wrightsville Beach, Holden Beach, and Buxton in
Island Beach, Wrightsville Beach, Bald Head 2018. At each beach, we placed HOBO Pendant
Island, Holden Beach, and Ocean Isle Beach Temperature data loggers (Onset Computer Cor-
(Fig. 1). The northernmost beach was Pea Island poration, Bourne, Massachusetts, USA) along
National Wildlife Reserve (NWR), located four transects. These loggers have a guaranteed
approximately 400 kilometers from Ocean Isle accuracy of
0.2°C from 0° to 50°C and a resolu-
Beach, the southernmost beach. We selected bea- tion of 0.1°C at 25°C. Two transects were located
ches with a history of loggerhead sea turtle nest- in areas that had been nourished within the last
ing activity and recent nourishment (i.e., within two years, and the other two transects were
3 yr of data collection; Table 1). All beaches were placed in areas that had not received nourished
monitored for sea turtle nesting activity. Between sand within the last three years (hereafter, natu-
2014 and 2018, beaches had an average of ral). The distance between the two furthest tran-
39
33.8 loggerhead nests/yr laid collectively on sects varied across beaches, but the average
all 8 study beaches (minimum Wrightsville, midpoint distance between these transects was

Fig. 1. Sampling sites for loggerhead sea turtle (Caretta caretta) nesting beaches along the coast of North Caro-
lina, USA. Sites indicated by a square were sampled in 2015, and those indicated by a circle were sampled in
2018.

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Table 1. Description of key parameters of the most recent nourishment event and past nourishment events on all
eight sea turtle nesting beaches in North Carolina.

Most recent nourishment event All nourishment events (as of 2018)

Total Most
length of recent Volume Length Volume Length
beach event of sand of beach No. of sand of beach
Beach Lat. Long. (km) (yr) (m3) (km) Cost ($US) events (m3) (km) Total cost ($US)

PI NWR 35.723 −75.496 20.9 2013 444,149 1.77 7,222,368 19 7,383,922 27.8 56,154,185
BUX 35.267 −75.542 11.26 2018 1,987,842 4.67 22,000,000 1 1,987,842 4.67 22,000,000
EIB 34.666 −77.013 20.9 2013 38,227 0.61 623,660 19 3,992,098 49 44,500,219
TOP 34.370 −77.625 41.8 2015 638,497 0.73 10,347,748 14 2,856,826 23 41,069,545
WB 34.211 −77.798 6 2018 583,355 2.44 10,500,000 28 13,133,473 51.3 61,334,290
BHI 33.867 −78.006 7.72 2015 1,017,414 3.43 12,700,000 15 9,806,033 42.2 114,200,661
HB 33.915 −78.286 13 2017 1,001,566 6.46 15,000,000 29 3,646,063 32.35 32,433,725
OIB 33.887 −78.436 11.26 2014 586,728 2.44 8,569,514 20 3,927,633 19 28,024,183
Note: Abbreviations represent the following beaches: Pea Island National Wildlife Reserve (PINWR), Cape Hatteras
National Seashore in Buxton (BUX), Emerald Isle Beach (EIB), Topsail Beach (TOP), Wrightsville Beach (WB), Bald Head Island
(BHI), Holden Beach (HB), Ocean Isle Beach (OIB).

4 km. Each transect contained one temperature 500 μm, 250 μm, 125 μm, and 63 μm) on a Ro-Tap
logger placed at mean mid-nest depth (45 cm for sieve shaker (W.S. Tyler, Mentor, Ohio, USA) for
loggerheads) in the vegetated landward zone 10 min. All particles remaining in the bottom bin
and one logger placed in the open beach. We were categorized as <63 μm. We weighed the
deployed a total of 64 loggers (n = 8 per beach) contents of each weight class to the nearest one-
set to record hourly sand temperatures from hundredth gram on a portable electronic balance
May until October. The arrival of Hurricane Flor- (VWR International, Radnor, Pennsylvania,
ence on September 14, 2018 resulted in the 2018 USA). The percentage of each size class with
data loggers being collected early, on September respect to the total sample was then calculated.
9 and September 11. We used GPS coordinates Following Folk (1980), data were standardized to
obtained when loggers were deployed to aid in units of μm to calculate the arithmetic mean, mid-
recovery. We recovered a total of 46 data loggers point deviation, and standard deviation of the
across the two years. Comparisons were made mean grain size. We further categorized sand
from temperatures recorded between June and samples into the following size classes: extra
September, except for Buxton for which tempera- small <63–63 μm, small 125–250 μm, medium
ture data collection began in July. Mean seasonal 500–710 μm, large 1–2 mm, and extra-large
temperatures were calculated by averaging mean 4 mm. The percentage of each size class was cal-
daily temperatures throughout the months of culated relative to the rest of the sample.
July, August, and September. 2. Albedo.—We obtained albedo measurements
for each surface sand sample. Albedo is a unitless
Sand characteristics quantification of the amount of solar energy a
1. Sand grain size.—In June of the 2015 and 2018 surface reflects or an inverse indication of the
loggerhead nesting seasons, we collected two amount of solar energy a substance absorbs
sand samples within a 1 m radius of the tempera- (Hays et al. 2001). Lighter, more reflective sur-
ture logger sites: at a depth of 45 cm and from the faces will have high albedo whereas darker sur-
surface. Prior to analysis, 100 g of each sample faces will have low albedo. In 2015, we measured
was dried in a Shake n’ Bake Rocking Hybridiza- albedo in the field using a Litemaster Pro L-478D
tion Laboratory Oven (Boekel Scientific, Feaster- (Sekonic, Tokyo, Japan). We recorded the amount
ville, Pennsylvania, USA) at 60°C for 48 h or until of incoming solar radiation (RADINC) and the
the weight no longer changed. Dried samples amount of light reflecting off the sand
taken from 45 cm were poured through a set of 8 (RADSAND) in lux. We calculated albedo (p350–800)
mesh sieves (4 mm, 2 mm, 1 mm, 710 μm, as the fraction of solar radiation reflected relative

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to the incoming radiation and multiplied by 100 Monthly and seasonal mean temperatures within
to convert to a percentage of solar radiation. and among beaches were calculated for natural
(p350–800 = (RADSAND/RADINC) × 100. In 2018, and nourished areas as well. We performed one-
surface sand samples were collected and brought and two-way ANOVA to determine whether sig-
back to UNCW’s Center for Marine Science for nificant spatial (beach) or temporal (month) varia-
analysis to minimize variation in the amount of tion existed among beaches and areas. Finally, for
incoming solar radiation from daily and monthly each beach, we tabulated the number of days
weather patterns. Referencing the validation in where the daily average temperatures reached or
methodology for collecting albedo in field and exceeded the pivotal temperature for loggerhead
laboratory environments published by Hays sea turtles regionally (29°C) and compared values
et al. (2001), our approach combined aspects of among natural and nourished areas within bea-
both field and laboratory protocols to produce ches.
data that could be compared with 2015 measure- We calculated the mean difference between
ments. We took all albedo measurements on the mean daily temperatures at nourished and natu-
same day under natural light conditions with the ral sites for each zone within each beach. This was
Litemaster Pro L-478D. We calculated albedo done monthly and seasonally for each zone sepa-
using following the equation provided by Hays rately and for the overall beach (except for
et al. (2001) to calculate albedo as. Wrightsville and Topsail for which we did not
have open beach zone temperatures). We per-
p350800 ¼ ðLS =LG Þ  18%
formed a one-tailed t-test between mean daily
where LS the amount of light (lux) reflecting off temperatures on natural and nourished areas and
the surface sand sample, and (LG) was the a regression analysis to evaluate the effect of lati-
amount of reflected light off of a photographic tude on temperatures. We ran univariate fixed-ef-
gray card with a known albedo of 18%. We fects models with mean and CV of monthly
repeated measurements of reflectance from the temperature as well as mean and CV of grain size
sand and gray card three times; albedo values as the response variables and BEACH, MONTH,
we report represent the mean of the three ZONE (open vs. vegetated), and AREA (natural
repeated measurements. Before each set of reflec- vs. nourished) as the predictor variables. Before
tance values were recorded, the amount of performing univariate fixed-effects analyses, we
incoming solar radiation was observed to ensure logit transformed the relative percent of each size
consistency of measurements across beaches. class as well as albedo measurements. We ana-
Across all beaches, the average incoming solar lyzed models with mean and CV of monthly tem-
radiation value observed throughout the 2018 perature as the response variables and mean
albedo data collection was 75,120
3,650 (range grain size (MGS), CV of grain size (CVGS), indi-
70,000–80,000 lux). vidual grain size classes (%XL, %L, %M, %S, %
XS), and albedo as the predictor variables. Finally,
Data analysis we built fixed-effect multivariable models from
We focused our analyses on mean daily and the predictors used in our univariate models. We
monthly sand temperatures as current literature performed all analyses using JMP Pro, Version 13
on reproductive success in sea turtles most often (SAS Institute, Cary, North Carolina, USA).
uses mean incubation temperature as a predictor
of incubation duration, and therefore offspring RESULTS
sex ratio (Mrosovsky et al. 1999). Using mean
daily temperatures from each data logger, we cal- Sand temperature
culated the mean, standard deviation, and coeffi- We found significant spatial variation in mean
cient of variation (CV) of monthly sand seasonal temperatures across all eight study sites
temperatures. We also calculated mean seasonal (ANOVA F7,3,123 = 59.26, P < 0.0001). Bald Head
temperatures for each beach using mean daily Island was the warmest beach (30.1°
1.2°C)
temperatures from July through September, as which was 1.4°C warmer than the coolest (Topsail
June temperatures in Buxton were not recorded. Beach 28.7°
1.2°C). There was also significant

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

spatial variation in seasonal temperatures among significantly influenced both seasonal and
both natural (ANOVA F7,1,398 = 51.39, P < 0.0001) monthly mean temperatures. Overall, Emerald
and nourished (ANOVA F7,1,717 = 21.84, P < 0.0001) Isle Beach had the largest difference
areas. In the univariate analysis, BEACH was a signif- (1.6°
0.4°C) in mean daily sand temperature
icant predictor of the mean and CV of monthly tem- between natural and nourished areas. We
peratures (P < 0.0001 and P = 0.001, respectively). observed the smallest difference between areas in
There was no correlation between mean or CV of Buxton (0.2°
0.3°C). Nourished areas were con-
monthly temperature and latitude (r2 = 0.12, P = 0.4; sistently warmer on all beaches except for Holden
r2 = 0.02, P = 0.7, respectively). Beach where the seasonal mean difference
When beaches were combined, we found signifi- was −0.5°C. This pattern was mirrored in the dif-
cant temporal variation in mean monthly tempera- ferences between mean monthly temperatures
tures (ANOVA F2,3,128 = 18.21, P < 0.0001). July (Fig. 2). In the univariate analysis, AREA was also
was the warmest month (29.6°
1.4°C) while a significant predictor of the mean but not the CV
September was the coolest (29.2°
1.5°C). There of monthly temperatures (P = 0.03 and P = 0.79,
was also significant temporal variation in tempera- respectively). Overall, we found a significant dif-
ture among both natural (ANOVA F7,53 = 5.13, ference between the mean daily temperatures of
P = 0.0002) and nourished (ANOVA F7,65 = 2.91, natural and nourished areas (29.3°
1.4°C and
P = 0.01) areas. In the univariate analysis, 29.7°
1.2°C, respectively; t-test t1,2800 = 9.01,
MONTH was also a significant predictor of mean P < 0.0001). Nourished areas were significantly
and CV of monthly temperatures (P < 0.0001 and warmer than natural on six out of the eight bea-
P < 0.0001, respectively). ches (Fig. 3a).
Despite significant spatial and temporal varia- The number of days above the pivotal temper-
tion among beaches, beach nourishment ature varied within and among beaches (Fig. 4).

Fig. 2 . Mean
SD of the difference in mean daily sand temperatures (°C) between natural and nourished
beach areas between June and September. Positive values indicate that nourished areas are warmer; negative val-
ues indicate that natural areas are warmer.

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Fig. 3. Comparisons between natural and nourished beach areas in (a) mean seasonal sand temperatures (°C),
(b) mean grain size (µm), and (c) albedo. * indicate significant differences between values.

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Fig. 4. Number of days observed throughout the season where daily sand temperatures were above the piv-
otal temperature (29°C) of loggerhead turtles for natural and nourished areas on all eight beaches.

Among beaches, natural areas exceeded 29°C for univariate analyses, AREA as well as BEACH and
an average of 33 d. Among nourished areas, ZONE were significant predictors of mean grain
sand temperatures exceeded the pivotal tempera- size (Table 3). AREA was the second-best predictor,
ture for an average of 50 d. The largest difference explaining 13% of the variation in mean grain size.
in the number of days above the pivotal was The distribution of grain size classes largely
observed on Wrightsville Beach. In the natural mirrored the patterns of mean grain size, with
area, we documented only six days above 29°C. significant differences between natural and nour-
However, within nourished areas we recorded ished areas. Overall, nourished areas had a
50 d above the pivotal temperature. We observed greater proportion of extra-large and large size
the smallest difference between areas in the num- classes, and natural areas had a greater propor-
ber of days above the pivotal in Buxton, where tion of small and extra small size classes (see
we observed 48 and 51 d, respectively. Appendix S1: Fig. S1 for overall grain size class
distribution and grain size classes broken down
Sand characteristics for each beach). Indeed, when classes were con-
Mean grain sizes were not significantly different sidered separately, AREA was a significant pre-
among beaches (ANOVA F7,51 = 1.92, P = 0.08). dictor of grain size in all univariate analyses
Among natural areas, there were no significant dif- (data not shown). With the exception of the
ferences in mean grain size (ANOVA F7,21 = 2.38, extra-large grain size class, in which every beach
P = 0.06). However, among nourished areas, we we sampled had more %XL grain sizes on nour-
observed significant differences in mean grain size ished areas than natural, we consistently found
(F7,22 = 10.14, P < 0.0001), with the largest and the opposite patterns in grain size distribution
smallest mean grain sizes on Wrightsville Beach (less of the larger/medium and more of the smal-
(1519.6
103.2 μm) and Topsail Beach (519.3 ler grain sizes) in nourished areas than natural

287.2 μm), respectively (Table 2). Mean grain on Holden and Ocean Isle beaches (Table 2).
size was larger in nourished areas on six out of In the univariate analyses, BEACH and AREA
eight beaches; however, only two of these compar- were significant predictors for CV of grain size
isons were significant (Fig. 3b). Holden Beach and (P < 0.0001, P = 0.01, respectively; Table 3).
Pea Island NWR both had larger mean grain sizes Nourished areas also had lower albedo than nat-
on natural areas as opposed to nourished, though ural areas, though only three of the eight com-
the differences were not significant. In the parisons were significant (Fig. 3c).

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Table 2. Descriptive statistics of temperature and sand variables across all eight sea turtle nesting beaches in
North Carolina.
Pea Island Buxton Emerald Topsail Wrightsville Bald Head Holden Ocean
Variable Beach type NWR (CHNS) Isle Beach Beach Beach Island Beach Isle Beach

Mean Natural 29.0
1.1 29.7
1.7 28.6
1.4 28.3
1.3 28.5
1.1 29.8
1.1 30.2
1.2 29.4
1.2
seasonal Nourished 29.3
0.8 29.9
1.5 29.5
1.5 29.1
1.0 30.0
1.3 30.2
1.3 29.3
1.2 29.7
1.0
temp (°C)
CV seasonal Natural 0.04 0.06 0.05 0.05 0.04 0.04 0.04 0.04
temp Nourished 0.03 0.05 0.04 0.04 0.04 0.04 0.03 0.03
Mean grain Natural 574.9
238.7 446.8
79.2 403.8
129.9 381.6
18.1 347.0
125.7 436.0
99.1 837.8
342.0 660.7
307.2
size (µm) Nourished 520.9
43.9 552.5
38.5 536.7
285.5 519.3
287.1 1519.6
103.2 819.2
119.9 678.1
73.3 689.2
399.2
CV grain Natural 0.42 0.16 0.32 0.05 0.36 0.23 0.41 0.46
size Nourished 0.08 0.07 0.53 0.55 0.07 0.15 0.11 0.58
%XS grain Natural 0.004
0.004 0.002
0.003 0.004
0.002 0.007
0.006 0.014
0.006 0.004
0.001 0.005
0.007 0.008
0.004
size (µm) Nourished 0.001
0.00 0.002
0.001 0.006
0.003 0.015
0.001 0.004
0.003 0.004
0.005 0.013
0.007 0.004
0.002
%S grain Natural 0.57
0.31 0.61
0.30 0.88
0.16 0.92
0.02 0.94
0.05 0.80
0.2 0.45
0.15 0.63
0.16
size (µm) Nourished 0.49
0.06 0.44
0.19 0.80
0.15 0.83
0.18 0.28
0.14 0.32
0.20 0.59
0.06 0.67
0.18
%M grain Natural 0.32
0.28 0.35
0.26 0.10
0.12 0.05
0.01 0.01
0.01 0.17
0.17 0.31
0.20 0.27
0.08
size (µm) Nourished 0.49
0.05 0.49
0.16 0.14
0.08 0.07
0.07 0.21
0.08 0.50
0.17 0.27
0.05 0.23
0.09
%L grain Natural 0.10
0.14 0.04
0.04 0.02
0.04 0.01
0.00 0.02
0.03 0.02
0.02 0.22
0.20 0.07
0.06
size (µm) Nourished 0.02
0.01 0.07
0.05 0.04
0.05 0.07
0.08 0.39
0.11 0.16
0.06 0.11
0.02 0.06
0.05
%XL grain Natural 0.001
0.001 0.002
0.002 0.003
0.006 0.004
0.004 0.015
0.024 0.004
0.004 0.012
0.010 0.031
0.046
size (µm) Nourished 0.002
0.002 0 0.019
0.038 0.017
0.022 0.115
0.046 0.018
0.006 0.021
0.009 0.046
0.074
Albedo Natural 29.5
1.0 33.6
2.4 33.0
1.0 33.5
0.7 45.6
1.5 29.0
1.2 28.7
1.5 28.3
1.5
Nourished 29.0
1.2 29.8
2.4 31.5
1.7 29.0
1.4 39.7
5.7 27.5
0.6 27.1
0.5 28.3
1.3

Table 3. Univariate analysis of the relationship between temperature and sand characteristics and various spatial
and temporal variables.

Mean monthly CV mean monthly Mean grain size

temperature (°C) temperature (μm) CV mean grain size Albedo
2 2 2 2
Variable P r P r P r P r P r2

Month <0.0001 0.14 <0.0001 0.27 0.73 0.01 0.35 0.03 <0.0001 0.36
Beach <0.0001 0.25 0.001 0.13 0.01 0.14 <0.0001 0.31 <0.0001 0.90
Area 0.03 0.03 0.99 3.42E−07 <0.0001 0.13 0.01 0.05 0.2 0.01
Zone 0.53 0.002 0.97 9.62E−06 0.02 0.04 0.89 2.00E−04 0.26 0.01
Note: Significant values are bolded, and the predictor variable with the highest r2 value is italicized.

Mixed-effect models model that explained the most variation in mean

We ran a mixed-effect model controlling for monthly temperature included mean grain size,
the natural spatial and temporal variation by CV of mean grain size, and the three size classes
including BEACH and MONTH as random identified as significant predictors in univariate
effects. AREA was a significant predictor of mean analyses (r2 = 0.21, P = 0.004; Appendix S1:
monthly sand temperatures (r2 = 0.4, P = 0.02), Table S2).
but not of CV. Moreover, the best multivariable
model for mean monthly temperature included DISCUSSION
AREA (Appendix S1: Table S1). AREA was also
included as a term in the best multivariable Coastal ecosystems are estimated to contribute
model explaining 30% and 36% of variation in approximately 25% ($28 trillion/yr) to the total
the mean and CV of grain size, respectively $125 trillion/yr provided by global ecosystem
(Appendix S1: Table S1). In the univariate analy- services (Costanza et al. 2014). In the United
ses, albedo, the relative percentage of small, States alone, beaches generate approximately
medium, and large grain size classes and mean $225 billion/yr for the national economy (Hous-
grain size were also significant predictors of mean ton 2013). Since the 1920s there have been
monthly temperature (Table 4). The multivariable approximately 3400 nourishment events that

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Table 4. Univariate analysis of the relationship nourishment projects and the sand that naturally
between temperature and sand characteristics. accumulates on the beach. On most beaches,
nourished sand was characterized by consider-
Mean monthly CV mean monthly
temperature temperature able amounts of extra-large and large grain sizes
Variable P r2 P r2
and a distinct lack of small grain sizes. Sedimen-
tary features such as grain size, distribution, and
Mean grain size 0.04 0.06 0.85 5.00E−04 albedo can exert influence over thermal conduc-
CV grain size 0.84 6.00E−04 0.89 2.00E−04
tivity as well as water and gas diffusion pro-
%XL 0.49 0.01 0.45 0.01
%L 0.02 0.07 0.99 8.00E−07 cesses within the incubation environment of sea
%M 0.0004 0.15 0.84 6.00E−04 turtles (Ackerman 1997), indirectly influencing
%S 0.0004 0.15 0.67 2.00E−03 incubation duration and hatchling success
%XS 0.15 0.03 0.88 3.00E−04 (Fadini et al. 2011, Ditmer and Stapleton 2012).
Albedo 0.001 0.14 <0.0001 0.30 Previous work has reported significant effects
Note: Significant values are bolded, and the predictor vari- of nourishment on the thermal properties of nest-
able with the highest r2 value is italicized. ing beaches. In the Delaware Bay, a combination
of darker, finer, borrowed sand increased sand
have placed nearly one trillion cubic yards of temperatures, and oxygen limitation, leading to
sand over 1500 km of coastline, costing over $7 a lower habitat suitability index with respect to
trillion (National Beach Nourishment Database horseshoe crab nesting areas (Avissar 2006).
2019) to protect these economically and ecologi- These results highlighted the need to match the
cally valuable habitats. However, even under the fill sediment to the natural grain size and color of
most stringent scenario limiting greenhouse gas the original sand, in order to minimize adverse
emissions, the global mean sea level is expected impacts on horseshoe crab nests. Similarly, Mil-
to rise for centuries to come (IPCC 2014). By the ton et al. (1997) found that the aragonite sand
end of the 21st century, 70% of global coastlines imported from the Bahamas Banks (Sealy 1994)
will experience a change in mean sea level within and deposited on Fisher Island, Florida was sig-
20% of the global mean (IPCC 2014). Climate nificantly cooler than the native silicate sand. In
warming is thus one of the most significant haz- fact, nests laid in aragonite sand incubated at
ards to endangered sea turtles (Rees et al. 2016, temperatures as much as 3°C cooler than nests
Nalovic et al. 2019), which critically rely on the laid in silica sand (Milton et al. 1997). While tem-
availability of nesting habitat. Currently, beach peratures increased in both sand types as the sea-
nourishment is among the most common coastal son progressed, the slope of the increase in
management tools used to protect these habitats aragonite was 66% that of the silicate sand, indi-
(Greene 2002, Peterson and Bishop 2005, Spey- cating a lower thermal capacity for the Bahamian
broeck et al. 2006). It is a practice that inherently sand. As such, changes in the thermal regime are
requires continued maintenance and replenish- clearly dependent on the characteristics of
ment events and therefore is likely to persist, if dredged material used and the natural morpho-
not increase, on a global scale (Dean 2003). dynamic and sand grain properties found on
Though effective, beach nourishment potentially each nourished beach, emphasizing the ecologi-
operates at a cost to the reproductive output of cally and biologically relevant impacts of beach
nesting females. nourishment (Peterson and Bishop 2005). Fur-
We found a significant effect of beach nourish- ther, monitoring the morphological changes of
ment on sand temperature in at least one month nesting habitat is a key factor in assessing the
throughout the season of every beach we stud- impact of climate change on sea turtles, as dis-
ied, with differences most frequently observed in cussed in several reviews highlighting manage-
July and August. We observed significantly war- ment and conservation priorities (Hamann et al.
mer mean seasonal sand temperatures in nour- 2010, Rees et al. 2016).
ished areas on six of the eight beaches we Differences in sand temperature on nesting
studied. We also found that temperature differ- beaches raises concerns about the future produc-
ences resulted from differences in sand character- tion of male hatchlings from regionally and glob-
istics between the borrowed sand used in ally important rookeries (Hays et al. 2014). The

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COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

observed differences between the number of Reneker and Kamel 2016b). However, we have
days that exceeded the pivotal temperature shown that beach nourishment can cause signifi-
among areas within beaches strongly indicate cant changes to the thermal profile of loggerhead
that beach nourishment can directly influence nesting beaches and we support a directed effort
the incubation environment and therefore critical to incorporate these impacts into species assess-
fitness characteristics as well as primary sex ments. Moreover, it is critical to take a regional
ratios of loggerhead sea turtle hatchlings. Nour- approach to not only assess temperature at all
ished areas within beaches had a greater number rookeries and identify male-producing beaches
of days above the pivotal temperature on each for protection, but also to identify beaches where
beach we observed except for Holden Beach nourishment projects may be exacerbating sex
(Fig. 4). These data suggest that nests were more ratio distortions and thus require additional
likely to produce female-biased primary sex intervention (Klein et al. 2017, Esteban et al.
ratios on nourished compared to natural areas. 2018).
The incubation duration of loggerhead nests has
already decreased an average of seven days over
ACKNOWLEDGMENTS
the past 25 yr at an important North Carolina Thank you to the following people for allowing
loggerhead rookery, which has led to a concomi- access and sampling on the beaches: Bald Head (Cal-
tant increase in the estimated percentage of vin Peck and the Bald Head Island Conservancy),
female hatchlings produced, from a mean of 55% Wrightsville Beach (Katie Ryan), Holden Beach
in 1991 to a mean of 88% in 2015 (Reneker and (Christy Ferguson and David Hewett), Buxton (Will
Kamel 2016a). Thompson, Sabrina Henry and the National Parks Ser-
This pattern of extreme female bias in the pri- vice), Topsail (Michael Moore and Karen Beasley Res-
mary sex ratios of loggerheads is extensive (see cue and Rehabilitation Center), Pea Island National
Fig. 3 in Hays et al. 2014). Potential mitigation Wildlife Reserve (National Parks Service), Ocean Isle
techniques such as the relocation, shading, and (Daisy Ivey and the Ocean Isle Beach Sea Turtle Protec-
tion Organization), and Emerald Isle (Frank A. Rush
watering of nests have been investigated in sev-
Jr. and the Emerald Isle Sea Turtle Program). We also
eral sea turtle populations. For example, shading thank Dr. Matthew Godfrey and the North Carolina
sand with fence mesh in Playa Grande, Costa Wildlife Resources Commission for their support of
Rica, resulted in about a 2°C decrease in temper- the sea turtle monitoring programs. JLR was funded
ature and was more effective than watering (Hill by a Judith C. Bryan, Holden Beach Turtle Watch Fel-
et al. 2015). Decreases of similar magnitude were lowship and KMS was funded by an NSF grant (BIO-
also reported when sand was sprinkled with OCE 1459384) to SJK.
water or shaded in an experimental setting in
Australia (Jourdan and Fuentes 2015). More
LITERATURE CITED
recently, a combination of relocation and shading Ackerman, R. A. 1997. The nest environment and the
with palm trees was predicted to reduce primary embryonic development of sea turtles. In P. L. Lutz
sex ratios from a current range of 97–100% and J. A. Musick, editors. The biology of sea turtles.
female to 60–90% female (Esteban et al. 2018). CRC Press, Boca Raton, Florida, USA.
This low-cost, low technology method might Atkinson, D. 1994. Temperature and organism size - a
therefore be useful to mitigate the immediate biological law for ectotherms. Advances in Ecologi-
effects of warming due to beach nourishment. A cal Research 25:1–58.
decision science approach would be a useful tool Avissar, N. G. 2006. Modeling potential impacts of
to identify the most cost-effective, feasible and beach replenishment on horseshoe crab nesting
habitat suitability. Coastal Management 34:427–441.
beneficial management interventions relevant to
Baker, J. D., C. L. Littnan, and D. W. Johnston. 2006.
North Carolina’s sea turtle populations (Klein
Potential effects of sea level rise on the terrestrial
et al. 2017). habitats of endangered and endemic megafauna in
Whether these long-lived species have the the Northwestern Hawaiian Islands. Endangered
potential to adapt to global climate change via Species Research 4:1–10.
shifts in nesting phenology or nest-site selection Booth, D. T. 2017. Influence of incubation temperature
behaviors is still unknown (Hawkes et al. 2007, on sea turtle hatchling quality. Integrative Zoology
Schwanz and Janzen 2008, Refsnider et al. 2013, 12:352–360.

v www.esajournals.org 11 March 2021 v Volume 12(3) v Article e03396

COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Carthy, R. R., A. M. Foley, and Y. Matsuzawa. 2003. sea turtle nesting beach loss. Ocean & Coastal
Incubation environment of loggerhead turtle nests: Management 51:330–341.
effects on hatching success and hatchling character- FitzGerald, D. M., M. S. Fenster, B. A. Argow, and I. V.
istics. Pages 144–153 in A. B. Bolten and B. E. Buynevich. 2008. Coastal impacts due to sea-level
Witherington, editors. Loggerhead sea turtles. rise. Annual Review of Earth and Planetary
Smithsonian Press, Washington, DC, USA. Sciences 36:601–647.
Coombes, E. G., A. P. Jones, and W. J. Sutherland. Folk, R. L. 1980. Petrology of sedimentary rocks. Hem-
2008. The biodiversity implications of changes in phill Publishing, Austin, TX, USA.
coastal tourism due to climate change. Environ- Galbraith, H., R. Jones, R. Park, J. Clough, S. Herrod-
mental Conservation 35:319–330. Julius, B. Harrington, and G. Page. 2002. Global cli-
Costanza, R., R. de Groot, P. Sutton, S. van der Ploeg, mate change and sea level rise: potential losses of
S. J. Anderson, I. Kubiszewski, S. Farber, and R. K. intertidal habitat for shorebirds. Waterbirds
Turner. 2014. Changes in the global value of 25:173–183.
ecosystem services. Global Environmental Change- Glen, F., A. C. Broderick, B. J. Godley, and G. C. Hays.
Human and Policy Dimensions 26:152–158. 2003. Incubation environment affects phenotype of
Crain, D. A., A. B. Bolten, and K. A. Bjorndal. 1995. naturally incubated green turtle hatchlings. Journal
Effects of beach nourishment on sea turtles - of the Marine Biological Association of the United
review and research initiatives. Restoration Ecol- Kingdom 83:1183–1186.
ogy 3:95–104. Goldenberg, S. B., C. W. Landsea, A. M. Mestas-
National Beach Nourishment Database. 2019. Ameri- Nunez, and W. M. Gray. 2001. The recent increase
can Shore and Beach Preservation Association. in Atlantic hurricane activity: causes and implica-
National Beach Nourishment Database, Washing- tions. Science 293:474–479.
ton, D.C., USA. Greene, K. 2002. Beach nourishment: a review of
Dean, R. G. 2003. Beach Nourishment: theory and biological and physical impacts. Atlantic State Marine
Practice. World Scientific Publishing Co., Pte. Ltd, Fisheries Commission, Washington, D.C., USA.
Toh Tuk Link, Singapore. Hamann, M., et al. 2010. Global research priorities for
Dean, R. G. 2005. Beach nourishment: benefits, theory sea turtles: informing management and conserva-
and case examples. Environmentally Friendly tion in the 21st century. Endangered Species
Coastal Protection 53:25–40. Research 11:245–269.
Ditmer, M. A., and S. P. Stapleton. 2012. Factors Affect- Hansen, J., R. Ruedy, M. Sato, and K. Lo. 2010. Global
ing Hatch Success of Hawksbill Sea Turtles on surface temperature change. Reviews of Geo-
Long Island, Antigua, West Indies. PLOS ONE physics 48:29.
7:12. Hansen, J., et al. 2016. Ice melt, sea level rise and
Emanuel, K. A. 2013. Downscaling CMIP5 climate superstorms: evidence from paleoclimate data, cli-
models shows increased tropical cyclone activity mate modeling, and modern observations that 2°C
over the 21st century. Proceedings of the National global warming could be dangerous. Atmospheric
Academy of Sciences of the United States of Amer- Chemistry and Physics 16:3761–3812.
ica 110:12219–12224. Hawkes, L. A., A. C. Broderick, M. H. Godfrey, and B.
Esteban, N., J. O. Laloe, F. Kiggen, S. M. Ubels, L. E. J. Godley. 2005. Status of nesting loggerhead turtles
Becking, E. H. Meesters, J. Berkel, G. C. Hays, and Caretta caretta at Bald Head Island (North Carolina,
M. J. A. Christianen. 2018. Optimism for mitigation USA) after 24 years of intensive monitoring and
of climate warming impacts for sea turtles through conservation. Oryx 39:65–72.
nest shading and relocation. Scientific Reports 8:8. Hawkes, L. A., A. C. Broderick, M. H. Godfrey, and B.
Fadini, L. S., A. G. Silva, and P. D. Ferreira. 2011. Sedi- J. Godley. 2007. Investigating the potential impacts
mentary characteristics and their effects on hatch- of climate change on a marine turtle population.
ing success and incubation duration of Caretta Global Change Biology 13:923–932.
caretta (Testudines: Cheloniidae) in Espirito Santo, Hays, G. C., J. S. Ashworth, M. J. Barnsley, A. C. Brod-
Brazil. Zoologia 28:312–320. erick, D. R. Emery, B. J. Godley, A. Henwood, and
Feagin, R. A., D. J. Sherman, and W. E. Grant. 2005. E. L. Jones. 2001. The importance of sand albedo
Coastal erosion, global sea-level rise, and the loss for the thermal conditions on sea turtle nesting
of sand dune plant habitats. Frontiers in Ecology beaches. Oikos 93:87–94.
and the Environment 3:359–364. Hays, G., A. Mazaris, and G. Schofield. 2014. Different
Fish, M. R., I. M. Cote, J. A. Horrocks, B. Mulligan, A. male vs. female breeding periodicity helps mitigate
R. Watkinson, and A. P. Jones. 2008. Construction offspring sex ratio skews in sea turtles. Frontiers in
setback regulations and sea-level rise: mitigating Marine Science 1:43.

v www.esajournals.org 12 March 2021 v Volume 12(3) v Article e03396

COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Hays, G. C., A. D. Mazaris, G. Schofield, and J. O. Naro-Maciel, E., N. Mrosovsky, and M. A. Marcovaldi.
Laloe. 2017. Population viability at extreme sex- 1999. Thermal profiles of sea turtle hatcheries and
ratio skews produced by temperature-dependent nesting areas at Praia do Forte, Brazil. Chelonian
sex determination. Proceedings of the Royal Soci- Conservation and Biology 3:407–413.
ety B: Biological Sciences 284:20162576 Noss, R. F. 2011. Between the devil and the deep blue
Hill, J. E., F. V. Paladino, J. R. Spotila, and P. S. Tomillo. sea: Florida’s unenviable position with respect to
2015. Shading and watering as a tool to mitigate sea level rise. Climatic Change 107:1–16.
the impacts of climate change in sea turtle nests. Packard, G., and M. Packard. 1988. The physiological
PLOS ONE 10:e0129528. ecology of reptilian eggs and embryos.
Hinkel, J., R. J. Nicholls, R. S. J. Tol, Z. B. Wang, J. M. Pages 523–605 in A. R. B. Huey, editor. Biology of
Hamilton, G. Boot, A. T. Vafeidis, L. McFadden, A. the Reptilia. Alan R. Liss, New York, New York,
Ganopolski, and R. J. T. Klein. 2013. A global anal- USA.
ysis of erosion of sandy beaches and sea-level rise: Peterson, C. H., and M. J. Bishop. 2005. Assessing the
an application of DIVA. Global and Planetary environmental impacts of beach nourishment.
Change 111:150–158. BioScience 55:887–896.
Houston, J. R. 2013. The economic value of beaches: a Peterson, C. H., M. J. Bishop, L. M. D’Anna, and G. A.
2013 update. American Shore and Beach Preserva- Johnson. 2014. Multi-year persistence of beach
tion Association, Washington, D.C., USA. habitat degradation from nourishment using
IPCC. 2014. Climate Change 2014: Synthesis Report. coarse shelly sediments. Science of the Total Envi-
Contribution of Working Groups I, II and III to the ronment 487:481–492.
Fifth Assessment Report of the Intergovernmental Phillips, M. R., and A. L. Jones. 2006. Erosion and tour-
Panel on Climate Change. Page 151 in Core Writing ism infrastructure in the coastal zone: problems,
Team, R. K. Pachauri, and L. A. Meyer, editors. consequences and management. Tourism Manage-
IPCC, Geneva, Switzerland. ment 27:517–524.
Jourdan, J., and M. M. P. B. Fuentes. 2015. Effectiveness Pieau, C., and N. Mrosovsky. 1991. Transitional range
of strategies at reducing sand temperature to miti- of temperature, pivotal temperatures and ther-
gate potential impacts from changes in environ- mosensitive stages for sex determination in rep-
mental temperature on sea turtle reproductive tiles. Amphibia-Reptilia 12:169–179.
output. Mitigation and Adaptation Strategies for Poloczanska, E. S., C. J. Limpus, and G. C. Hays. 2009.
Global Change 20:121–133. Vulnerability of marine turtles to climate change.
Klein, C. J., J. Beher, M. Chaloupka, M. Hamann, C. Pages 151–211in D. W. Sims, editor. Advances in
Limpus, and H. P. Possingham. 2017. Prioritization Marine Biology. Elsevier Academic Press, San
of marine turtle management projects: a protocol Diego, California, USA.
that accounts for threats to different life history Rees, A., et al. 2016. Are we working towards global
stages. Conservation Letters 10:547–554. research priorities for management and conserva-
Milton, S. L., A. A. Schulman, and P. L. Lutz. 1997. The tion of sea turtles? Endangered Species Research
effect of beach nourishment with aragonite versus 31:337–382.
silicate sand on beach temperature and loggerhead Refsnider, J. M., B. L. Bodensteiner, J. L. Reneker, and
sea turtle nesting success. Journal of Coastal F. J. Janzen. 2013. Nest depth may not compensate
Research 13:904–915. for sex ratio skews caused by climate change in tur-
Mrosovsky, N. 1988. Pivotal temperatures for logger- tles. Animal Conservation 16:481–490.
head turtles (Caretta caretta) from northern and Reid, W. V., and M. C. Trexler. 1992. Responding to
southern nesting beaches. Canadian Journal of potential impacts of climate change on U.S. Coastal
Zoology 66:661–669. Biodiversity. Coastal Management 20:117–142.
Mrosovsky, N., C. Baptistotte, and M. H. Godfrey. Reneker, J. L., and S. J. Kamel. 2016a. The maternal
1999. Validation of incubation duration as an index legacy: female identity predicts offspring sex ratio
of the sex ratio of hatchling sea turtles. Canadian in the loggerhead sea turtle. Scientific Reports
Journal of Zoology 77:831–835. 6:29237.
Mrosovsky, N., P. H. Dutton, and C. P. Whitmore. Reneker, J. L., and S. J. Kamel. 2016b. Climate change
1984. Sex ratios of 2 species of sea turtle nesting increases the production of female hatchlings at a
in Surinam. Canadian Journal of Zoology northern sea turtle rookery. Ecology 97:3257–3264.
62:2227–2239. Roman-Sierra, J., J. J. Munoz-Perez, and M. Navarro-
Nalovic, M., E. Cuevas, and M. E. Godfrey. 2019. Pons. 2014. Beach nourishment effects on sand
North Atlantic and Wider Caribbean. MTSG Regio- porosity variability. Coastal Engineering 83:
nal Report, Ross, California, USA. 221–232.

v www.esajournals.org 13 March 2021 v Volume 12(3) v Article e03396

COASTAL AND MARINE ECOLOGY SHAMBLOTT ET AL.

Rumbold, D. G., P. W. Davis, and C. Perretta. 2001. sea-level rise and related stresses: the DIVA Wet-
Estimating the effect of beach nourishment on Car- land Change Model. Global and Planetary Change
etta caretta (loggerhead sea turtle) nesting. Restora- 139:15–30.
tion Ecology 9:304–310. Speybroeck, J., et al. 2006. Beach nourishment: an eco-
Sala, O. E., et al. 2000. Biodiversity - Global biodiver- logically sound coastal defence alternative? A
sity scenarios for the year 2100. Science 287: review. Aquatic Conservation: Marine and Fresh-
1770–1774. water Ecosystems 16:419–435.
Schooler, N. K., J. E. Dugan, and D. M. Hubbard. 2019. Trenberth, K. E. 2011. Changes in precipitation with
No lines in the sand: Impacts of intense mecha- climate change. Climate Research 47:123–138.
nized maintenance regimes on sandy beach ecosys- Valverde, H. R., A. C. Trembanis, and O. H. Pilkey.
tems span the intertidal zone on urban coasts. 1999. Summary of beach nourishment episodes on
Ecological Indicators 106:15. the U.S. East Coast Barrier Islands. Journal of
Schuerch, M., et al. 2018. Future response of global Coastal Research 15:1100–1118.
coastal wetlands to sea-level rise. Nature 561:231. Van Houtan, K. S., and J. M. Halley. 2011. Long-term
Schwanz, L. E., and F. J. Janzen. 2008. Climate Change climate forcing in loggerhead sea turtle nesting.
and temperature-dependent sex determination: can PLOS ONE 6:7.
individual plasticity in nesting phenology prevent Witt, M. J., L. A. Hawkes, M. H. Godfrey, B. J. Godley,
extreme sex ratios? Physiological and Biochemical and A. C. Broderick. 2010. Predicting the impacts
Zoology 81:826–834. of climate change on a globally distributed species:
Sealey, N. E. 1994. Bahamian landscapes: introduction the case of the loggerhead turtle. Journal of Experi-
to the Physical Geography of the Bahamas. Media mental Biology 213:901–911.
Publishing, Nassau, Bahamas. Yntema, C. L., and N. Mrosovsky. 1982. Critical peri-
Spencer, T., M. Schuerch, R. J. Nicholls, J. Hinkel, D. ods and pivotal temperatures for sexual differenti-
Lincke, A. T. Vafeidis, R. Reef, L. McFadden, and S. ation in loggerhead sea turtles. Canadian Journal
Brown. 2016. Global coastal wetland change under of Zoology 60:1012–1016.

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