Natural Beach Vegetation Coverage and Type Influence the Nesting

Habitat of Hawksbill Turtles (Eretmochelys imbricata) in Campeche,

Mexico
Authors: Juan Alberto Hernández-Cortés, Enrique Núñez-Lara, Eduardo Cuevas, and
Vicente Guzmán-Hernández
Source: Chelonian Conservation and Biology, 17(1) : 94-103
Published By: Chelonian Research Foundation and Turtle Conservancy
URL: https://doi.org/10.2744/CCB-1280.1

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 Chelonian Conservation and Biology, 2018, 17(1): 94–103
doi:10.2744/CCB-1280.1
Ó 2018 Chelonian Research Foundation

Natural Beach Vegetation Coverage and Type Influence the Nesting Habitat of
Hawksbill Turtles (Eretmochelys imbricata) in Campeche, Mexico

JUAN ALBERTO HERNÁNDEZ-CORTÉS1, ENRIQUE NÚÑEZ-LARA2, EDUARDO CUEVAS3,*, AND

VICENTE GUZMÁN-HERNÁNDEZ4
1
Centro de Investigación de Ciencias Ambientales, Universidad Autónoma del Carmen, Calle 56 No. 4, Esq. Avenida Concordia, Col. Benito
Juárez, 24180 Ciudad del Carmen, Campeche, México [j.a.hernandez0877@gmail.com];
2
Universidad Autónoma del Carmen, Facultad de Ciencias Naturales y Exactas, Ciudad del Carmen, Campeche, México
[quique_nunezlara@hotmail.com];
3
Pronatura Penı́nsula de Yucatán, A.C., Calle 32 x 47 y 47A No. 269, Col. Pinzón II, 97205, Mérida, Yucatan, México;
4
Área de Protección de Flora y Fauna Laguna de Términos, Comisión Nacional de Áreas Naturales Protegidas, Av. Adolfo López Mateos,
Prolongación Playa Norte, C.P. 24140, Ciudad del Carmen, Campeche, México [vguzman@conanp.gob.mx]
*Present address of corresponding author: Centro de Investigación de Ciencias Ambientales, Universidad Autónoma del Carmen,
Calle 56 No. 4, Esq. Avenida Concordia, Col. Benito Juárez, 24180 Ciudad del Carmen, Campeche, México
[amir.cuevas@gmail.com]

ABSTRACT. – The hawksbill turtle (Eretmochelys imbricata) is a critically endangered species with a
global distribution and is broadly distributed along the Yucatan Peninsula in the Gulf of Mexico.
To complete its life cycle, this species uses sandy beaches with particular environmental conditions
that facilitate nesting and hatching. This study aimed to identify if beach physical characteristics
influence biological reproductive parameters (hatching and emergence success). Nesting activity
was monitored along 18 km of beach in the state of Campeche, Mexico, during 2014. Seventy-eight
nests were recorded, 12 physical and environmental variables were measured in the nests and
contiguous areas, and the hatching and emergence success of each nest was determined. Beach
slope and width had no significant relationship to nest site selection. Nest depth and nest distance
to highest tide and vegetation, including vegetation type, did influence hatching and emergence
success. Herbaceous and bushy plants were positively correlated with reproductive parameters,
highlighting the importance of preserving beach vegetation cover. This parameter was a central
structural component of hawksbill nesting habitat, possibly because it influenced nest shading,
preventing egg overheating and possible embryo death. Preservation and restoration of vegetation
structure on hawksbill nesting beaches is clearly vital to enhancing the reproductive success of this
species.
KEY WORDS. – biological reproductive parameters; emergence success; hatching success; marine
turtle; nesting process, site selection

For thousands of years, marine turtles have exhibited
high plasticity in adapting to environmental variations,
making them globally resilient species (Fuentes et al.
2013). However, the International Union for Conservation
of Nature (IUCN; http://www.iucnredlist.org/) classifies
the hawksbill turtle (Eretmochelys imbricata) as Critically
Endangered (Mortimer and Donnelly 2008). The hawksbill
is also considered as an indicator species because of its
sensitivity and vulnerability to changes in the habitats it
occupies. Hawksbill populations’ adaptations to past and
present environmental changes make this species an
interesting case study for ecological and habitat restoration
assessments. The hawksbill is also a Priority Species for
Conservation in Mexico, making it the focus of many
conservation efforts (Comisión Nacional de Áreas Natu-
rales Protegidas 2009).
Nest selection on nesting beaches is among the most
important aspects of hawksbill reproductive biology (Witt
et al. 2010; Huerta-Rodrı́guez et al. 2013; Kelly et al.
2017). Temperature is one of several environmental
parameters reported as determining reproductive success
in this species; it affects embryo survival and regulates the
relative number of male and female offspring produced
(Kamel and Mrosovsky 2006a).
Other nesting beach physical parameters that influ-
ence sea turtle reproductive success include vegetation
community type and structure on the beach and in adjacent
ecosystems, sand organic content, and sand sedimento-
logical characteristics (Moreno-Casasola and Travieso-
Bello 2006; Bolongaro-Crevenna-Recaséns et al. 2010;
Ditmer and Stapleton 2012). Vegetation is particularly
important because it shades nests, consequently affecting
nest temperature (Mrosovsky et al. 1992; Kolbe and
Jansen 2002; Kamel 2013). Beach extent and slope can
also impact reproductive success because both parameters
are associated with the capacity of female turtles to locate

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 HERNÁNDEZ-CORTÉS ET AL. — Nesting Habitat of Hawksbill Turtles in Campeche, Mexico
Figure 1. Hawksbill turtle (Eretmochelys imbricata) nesting beach at Chenkan, Campeche, Mexico.
adequate nesting sites (Fish et al. 2005; Cuevas et al.
2010). These nesting habitat physical and environmental
characteristics influence sea turtle reproductive success
and therefore the degree to which populations are
sustained (Kamel and Mrosovsky 2006a; Pfaller et al.
2008).
The western coast of the Yucatan Peninsula, Mexico,
has extensive sandy beaches and is a critical hawksbill
turtle nesting and hatching habitat. However, it is expected
to experience strong impacts from the sea level rise
predicted for coming decades (Grupo Intergubernamental
de Expertos sobre el Cambio Climático [IPCC] 2013) and
from increasing exposure to pressure from human
activities (Ortı́z-Pérez and Méndez-Linares 1999). As the
main nesting area for hawksbill turtles in the Western
Atlantic (Mortimer and Donnelly 2008), this region is
considered strategic for marine turtle conservation by
national governmental and international agencies (Nation-
al Fish and Wildlife Foundation 2010; Campbell 2014;
Guzmán and Garcı́a 2015). Mexico has signed the Inter-
American Convention for the Protection and Conservation
of Sea Turtles (IAC 1996) and the Convention on
International Trade in Endangered Species of Wild Fauna
and Flora (CITES 1973). Both conventions constitute
commitments to contribute to strengthening conservation
and protection actions throughout the country.
Chenkan beach in the state of Campeche is a Ramsar
site (Convention on Wetlands [Ramsar Convention] 1971)
which supports the third-largest hawksbill nesting popu-
lation in the state (. 350 hawksbill nests per yr) (Huerta-
Rodrı́guez and Muñoz 2009; Guzmán 2017). Despite the
efforts of numerous nongovernmental organizations and
government authorities to protect marine turtles, the
habitats hawksbills use, such as Chenkan beach, are being
modified by several anthropogenic threats including
fishing and urban infrastructure development. This leads
to habitat fragmentation and loss of vegetation cover on
the sandy dunes backing nesting beaches (Comisión
Nacional de Áreas Naturales Protegidas 2009). Dune
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95
vegetation cover is a critical component of beach
ecosystems which contributes to stabilizing the sediment
sea turtles use to build their nests and to regulating a
nesting area’s thermal regime (Nordstrom et al. 2000;
Martı́nez et al. 2004; Hannan et al. 2007).
For several decades, most of the conservation efforts
aimed at protecting marine turtles and their habitat have
focused on the sandy beaches where they nest. The goal is
to prevent population decline by guaranteeing offspring
viability through in situ and ex situ beach management
strategies and the release of emerged hatchlings to the sea
(Comisión Nacional de Áreas Naturales Protegidas 2009).
However, the aforementioned ongoing and predicted
threats to nesting beaches along the Yucatan Peninsula
threaten the long-term viability of the local hawksbill turtle
nesting population.
The present study’s objective was to assess the
influence of nesting beach physical and environmental
factors on reproductive parameters including nesting
success, hatching success, and hatchling emergence
success. Because preservation or restoration of natural
habitat conditions on nesting beaches is expected to
positively influence marine turtle reproductive success, the
importance of protecting the natural beach habitat
(encompassing the beach, dunes, and associated vegeta-
tion) is discussed in terms of how it may help decision-
makers and authorities to design optimum conservation
and restoration programs for this endangered species.
METHODS
Study Area. — This study was conducted on the
nesting beach of Chenkan, located on the western coast of
the Yucatan Peninsula in Campeche state in the south-
eastern Gulf of Mexico. This beach is an important marine
turtle nesting ground and has been a Ramsar site since
2003 (Huerta-Rodrı́guez and Munoz 2009). The beach is
also adjacent the Laguna de Términos Flora and Fauna
Protection Area (Fig. 1). Chenkan Beach is a highly
dynamic area subject to beach erosion. The dominant
 96 CHELONIAN CONSERVATION
beach profile consists of a pronounced slope and reduced
beach width, with sand grain coarseness ranging from
thick (2–5 mm) to intermediate (0.08–1.99 mm) (Bolon-
garo-Crevenna-Recaséns et al. 2010). Three substratum
layers have been defined on the beach: a superficial layer
(1–30-cm depth) of light, low-humidity sandy grains; an
intermediate layer (30–180-cm depth) of high-humidity
grains; and a bottom layer (. 2-m depth) molded by
coarse biogenic conch remains (Bolongaro-Crevenna-
Recaséns et al. 2010).
During an annual cycle, the region experiences 3
different climatic seasons: dry (March–June), rainy (June–
October), and norte (sequential storms out of the north;
November–February) (Álvarez and Barrios 1999). Dune
vegetation on Chenkan Beach forms a fringe of varying
width along the beach’s inland side. It consists mainly of
bushy and creeping species such as seagrape (Coccoloba
uvifera), siricote (Cordia sebestana), bayhops (Ipomea
pes-caprae), jacquinia (Jacquinia flammea), and coconut
(Coccus nucifera) (Bravo 1985; Reyes-Gómez and
Vásquez-Lule 2009). Fauna on the beach and in dune
zones include ghost crabs (Ocypode sp.), raccoons
(Procyon lotor), white-nosed coati (Nasua narica), gray
fox (Urocyon cinereoargenteus), green iguana (Iguana
iguana), black iguana (Ctenosaura similis), anteater
(Myrmecophaga sp.), opossum (Didelphis marsupialis),
common long-nosed armadillo (Dasypus novemcinctus),
pelican (Pelecanus occidentalis), common frigate (Fregata
magnificens), gulls (Laurus spp.), and boa (Boa constric-
tor) (Escanero-Figueroa et al. 1990). The marine habitats
neighboring the beach are shallow (, 5 m) and covered
mainly by seagrass and algae beds that support a variety of
sponges, echinoderms, crustaceans, and mollusks.
Nesting Activity. — Biological monitoring of turtle
nesting activity on this beach is done by personnel of the
Laguna de Términos Flora and Fauna Protection Area of
the National Commission of Natural Protected Areas
(Consejo Nacional de Areas Naturales Protegidos [CON-
ANP]). Night patrols (from about 2100 to 0600 hrs the
next day) are done throughout the turtle nesting season
(May–October) during which personnel search for nests
and nesting females. Confirmed nests are recorded every
night, and a set of biological and physical data are
collected including the geographic coordinates of each
nest. After the incubation period, 2 biological parameters
associated with incubation success were calculated: 1)
hatching success, expressed as the percentage of hatchling
turtles that broke through the eggshell; and 2) emergence
success, which is the percentage of hatchlings that left the
nest unaided (this is estimated by subtracting the number
of live and dead hatchlings remaining in the nest from the
total number of recorded eggs).
Environmental Variables. — Beach slope and width
were measured at 15 sites along the nesting beach with
approximately 700–2000 m between each measured nest.
Beach profile was characterized for 3 zones: intertidal,
berm, and dune (Guzmán and Garcı́a 2010). Profile data
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AND BIOLOGY, Volume 17, Number 1 – 2018
were recorded with a hand level (CST model 17621) for
slope and a conventional plastic tape measure for width
(50 m in length). The beach profile data were used to
calculate slope in degrees using the following equation
(Márquez-Garcı́a 2002):
DV DV
tg ¼ ’ ¼ arctg
DH DH
where DV is vertical distance and DH is horizontal
distance.
For each nest, measurements were taken of its depth,
its distance to the highest tidemark (based on the highest
location of algae washed up on the beach), and the shortest
distance to vegetation. Vegetation community data were
collected by photographing vegetation cover in a 1.5-m2
area surrounding the nest, focusing the camera from the
top of the nest at a distance of 1.8 m. Each photo was
analyzed using the Coral Point Count CPCe V3.5 tool to
calculate vegetation cover per square meter (Kohler and
Gill 2006). Counts were done per nest, and percentages
calculated for each vegetation type. Recorded vegetation
was identified to the species level using illustrated guides
for Yucatan Peninsula Flora (Guadarrama et al. 2014).
This approach only evaluates the effect of the vegetation
cover in the area immediately around an incubating nest;
the temporal dynamics of vegetation cover exceed the
present study scope, although its potential effect warrants
further research.
Temperature data loggers (UA-001-08 Hobo pendant,
Onset Computer Corporation) were used to record nest
internal temperature. Sensors were placed in the central
portion of the nests during egg laying. The loggers were
programmed to take temperature readings every hour
during the entire incubation period (approximately 55 d).
Complete temperature measurements were obtained from
17 nests, which corresponded to the number of available
loggers. Nests that predated logger placement were
excluded from the analysis.
Statistical Analyses. — Linear regression analyses
were run to assess the relationship between beach
morphology (width and slope) and the number of
hawksbill nests (reproductive parameters). Data for the
entire nesting beach was used; each analyzed site produced
a description of spatial variation in physical characteristics.
Partial regressions were also done for 3 separate months
(May, July, and September), considered representative of
the 3 regional climatic seasons, to evaluate whether beach
width and slope influenced the number of nests.
A redundancy analysis (RDA) was used to analyze
how hatching success (HATsuc) and emergence success
(EMEsuc) correlated to the environmental variables
measured on all recorded nests. Graphic ordination plots
were produced using 2 different scaling methods: scaling 1
to enhance the relationships between biological parameter
response variables, and scaling 2 to enhance the
correlations between the independent environmental
variables. These diagrams helped to identify the most-
 HERNÁNDEZ-CORTÉS ET AL. — Nesting Habitat of Hawksbill Turtles in Campeche, Mexico 97

Figure 2. Number of hawksbill turtle (Eretmochelys imbricata) nests during the 2014 season in addition to beach width and slope
measurements along the 18-km study area (error bars indicate standard errors).

important environmental variables affecting turtle repro- points established at onset of monitoring and used
ductive parameters. throughout sampling period); 9) creeping vegetation cover;
Twelve variables were considered in the redundancy 10) herbaceous vegetation cover; 11) bushy vegetation
analyses: 1) slope of the intertidal beach sector (slope A); cover; and 12) arboreal vegetation cover. All analyses
2) slope of the beach berm (slope B); 2) slope of the beach were done using the statistics package Vegan (Oksanen et
segment between the berm and vegetation line (slope C); al. 2013) for the R numerical environment (R Develop-
4) total slope (average of the three independent slope ment Core Team 2013).
measurements); 5) distance from nest to the high-tide
mark; 6) distance from nest to nearest vegetation; 7) nest RESULTS
depth; 8) beach width (measured from high-tide mark to Nesting Activity. — A total of 78 hawksbill turtle
dune reference points, i.e., wooden stakes or permanent nests were recorded along the 18 km of Chenkan Beach
from April to October 2014. Mean hatching success was
66% (SD = 25.4) whereas emergence success was 60%
Table 1. Results of the regression analysis between number of
clutches and beach physical environment variables. r2 = coeffi- (SD = 26.8).
cient of determination, m = slope, and p = statistical probability Beach Physical Parameters and Nest Distribution. —
value. Beach slope ranged from 48 to 128 (x̄ = 6.99, SD = 2.04)
Parameter r2 m p in the 15 studied sections. Beach width ranged from 12 to
28 m (x̄ = 21.62, SD = 6.92). The steepest slope and
May
Beach width 0.0190 0.265 0.621 shortest width values occurred in the beach’s middle
Total slope 0.0090 0.453 0.741 section (section 7) while the gentlest slope and largest
Slope A 0.0810 0.734 0.303 width were in the eastern extreme of the study area
Slope B 0.0960 1.126 0.26
Slope C 0.0620 0.714 0.372 (section 15) (Fig. 2). Nest spatial distribution did not
Jul significantly correlate to 2 physical parameters (Table 1).
Beach width 0.0010 0.168 0.902 The regression analysis results showed no significant
Total slope 0.0040 0.722 0.829
Slope A 0.0490 2.361 0.427 relationship between beach slope or width and the number
Slope B 0.0220 1.088 0.597 of nests during the 2014 nesting season for the overall
Slope C 0.0090 0.518 0.74 length of beach (Table 1; Fig. 3).
Sep
Beach width 0.0290 0.133 0.544 Relationship Between Habitat Variables and Turtle
Total slope 0.0510 0.489 0.418 Reproductive Parameters. — The plant community
Slope A 0.0520 0.404 0.412
Slope B 0.0060 0.399 0.791 associated with the hawksbill nests included 11 species,
Slope C 0.0630 0.227 0.368 all typical of coastal dunes on the Yucatan Peninsula. Total
Total mean surface area data for the nests showed 50% to have no
Width/beach 0.0020 0.336 0.877
Total slope 0.0020 0.863 0.876 vegetation and the rest to be covered by creeping,
Slope A 0.0003 0.294 0.952 herbaceous, bushy, and arboreal vegetation (or some
Slope B 0.0030 0.564 0.842 combination) (Table 2). The distance from the nests to the
Slope C 0.0430 2.142 0.459
nearest vegetation ranged from 0.1 to 2.5 m (x̄ = 0.46 m,

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 98 CHELONIAN CONSERVATION AND BIOLOGY, Volume 17, Number 1 – 2018

Figure 3. Topographic profiles of each of the 15 studied sections of Chenkan Beach during the 3-mo hawksbill turtle nesting season.

SD = 0.35). Nest distance to the highest tide mark ranged containing fewer nests (Fig. 4). A high proportion of the
from 4.3 to 48.8 m (x̄ = 15.6 m, SD = 6.98). Nest depth total variation in the response variables (HATsuc and
ranged from 0.41 to 0.67 m (x̄ = 0.51 m, SD = 0.06). EMEsuc) was clearly associated with the group containing
The ordination analysis showed that the main the most nests (left side). All the analyzed environmental
ordination axis runs approximately parallel to the first variables explained 49% of total variation in the measured
axis (RDA1), distinguishing 2 groups: one on the left side reproductive parameters; an analysis of variance (AN-
containing most of the nests, and one on the right side OVA) permutation test showed this to be significant
(p , 0.05). The habitat variables that contributed to
explaining high variation in hatching and emergence
Table 2. Plant species, type of structural vegetation, and mean success included nest depth (NestDep), distance to
percentage associated with turtle nests within a 1.5-m2 area
surrounding each nest. NV = no vegetation. vegetation (DisVeg), and coverage of beach substratum
by different vegetation types, e.g., bushy (BushVeg),
Species Structure % arborous (ArboVeg), and herbaceous (HerbVeg) plants but
Coccoloba uvifera Shrub 7.4 not creeping plants (CreeVeg). Creeping plants apparently
Scaevola plumierii Shrub compete with other vegetation types and were inversely
Suriana marı́tima Shrub related to turtle reproductive success and other vegetation
Canavalia rosea Herbaceous 24.6
Croton punctatus Herbaceous types (Fig. 4).
Passiflora foetida Herbaceous Nest Thermal Regime. — Average temperature in the
Panicum maximum Herbaceous 17-nest sample was 31.788C (SD = 0.74) and average
Cocos nucifera Tree 0.6
Ipomoea alba Creeping 16.7 vegetation cover area surrounding the nest was 61%. The
Ipomoea pes-caprae Creeping presence of vegetation around the nest influenced nest
Sesuvium portulacastrum Creeping temperature; for instance, Nest 7 had no surrounding
NV NV 50.7
vegetation and an average temperature of 31.978C,

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 HERNÁNDEZ-CORTÉS ET AL. — Nesting Habitat of Hawksbill Turtles in Campeche, Mexico 99

Figure 4. Ordination diagram produced by redundancy analysis displaying the relationships between the hawksbill turtle (Eretmochelys
imbricata) reproductive parameters of hatching success (HATsuc) and emergence success (EMEsuc) and the 12 habitat variables
measured in the turtle nesting area (arrows). Habitat variables: no vegetation (WOVeg), creeping vegetation (CreeVeg), average slope
on the first beach sector (SloAave), herbaceous vegetation (HerbVeg), nest depth (NestDep), distance from nest to vegetation (DisVeg),
bushy vegetation (BushVeg), arboreal vegetation (ArboVeg), distance to tide (DisTide), average slope on the second beach sector
(SloBave), average slope on the third beach sector (SloCave), and mean average beach slope (SloTave).

whereas Nest 6 had 100% vegetation cover and a
temperature of 29.478C, the lowest temperature of all
evaluated nests (Table 3).
The highest temperature was recorded in Nest 11
(33.058C), which had 29% vegetation cover dominated by
creeping vegetation. The same was the case for Nests 8, 9,
12, and 16, where temperature was higher than the mean
temperature for all nests (Table 3). In contrast, nests covered
by herbaceous and shrub type vegetation had a lower
temperature than the mean average (e.g., Nests 1, 2, 5, 6).
Only Nest 4 was adjacent to trees and its temperature was
30.468C, below the mean temperature. Hatching and
emergence success was highest in Nest 4 and lowest in
Nests 11 and 12, both covered by creeping vegetation. Of
note is that Nest 10 was a relocated nest. The female had
made her nest in the intertidal zone, leaving her eggs at risk
of flooding. We relocated the nest to a dune area dominated
by creeping and herbaceous vegetation, where its average
temperature was 32.078C, with a 95.8% hatching success
and a 56.9% emergence success (Table 3).
DISCUSSION
Hawksbill turtle females showed no nesting prefer-
ences associated with beach slope and width. Previous
studies have demonstrated that narrow beaches are
vulnerable to sea level rise, forcing turtles to modify
nesting site selection due to limited suitable habitat (Fish et
al. 2005; Bolongaro-Crevenna-Recaséns et al. 2010).
Turtles may be able to adapt to small-scale changes in
beach characteristics over short time periods (Hamann et
al. 2007; Hawkes et al. 2009), so it is important to
accommodate conservation planning for hawksbill turtle
nesting to local nesting beach characteristics (Liles et al.
2015).
The lack of a relationship between absolute beach
slope and width and the number of nests suggests that
turtles may be responding to changes in local beach
morphology dynamics (Fuentes et al. 2013). Some studies
have shown that hawksbills preferentially nest on steep
beaches in the Caribbean Sea (Fish et al. 2005) while
others suggest they nest at a certain elevation above sea
level (Horrocks and Scott 1991). The present findings
coincide with Cuevas et al. (2010), who suggested that
hawksbills exhibit plasticity in nesting site preference in
response to frequent changes in local beach morphology,
as observed at Chenkan Beach.
Turtle fidelity to particular microhabitats changes in
response to patterns in biological processes, and the main
structural components of such microhabitats include

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 100 CHELONIAN CONSERVATION AND BIOLOGY, Volume 17, Number 1 – 2018

Table 3. Mean temperature and percentage of vegetation and reproductive parameters measured for 17 turtle nests on Chenkan Beach,
Campeche, Mexico.

Temperature (8C) %
Nest
Nest Mean SD Creeping Herbaceous Shrub Tree No vegetation Hatching success Emergence success depth (cm)
1 31.24 4.39 30 55 0 0 15 83.5 77.7 0.53
2 31.30 1.93 10 65 0 0 25 94.8 91.3 0.49
3 31.53 1.65 33 15 0 0 52 92.5 77.4 0.56
4 30.46 2.32 0 0 0 29 71 97.1 96.0 0.56
5 31.62 2.25 10 25 53 0 12 72.2 63.2 0.51
6 29.47 1.62 5 5 90 0 0 94.9 94.9 0.46
7 31.97 2.11 0 0 0 0 100 77.4 70.8 0.55
8 32.04 2.60 35 15 20 0 30 74.3 46.5 0.61
9 32.04 1.26 40 10 0 0 50 87.8 85.7 0.53
10 32.07 1.93 30 35 0 0 35 95.8 56.9 0.57
11 33.05 1.70 27 2 0 0 71 63.8 58.0 0.49
12 32.72 1.42 45 20 0 0 35 58.6 58.6 0.56
13 31.89 1.21 12 0 68 0 20 96.2 96.2 0.49
14 32.31 1.04 16 17 32 0 35 93.0 90.1 0.43
15 32.28 1.50 12 8 50 0 30 85.4 84.8 0.55
16 32.40 1.34 35 30 0 0 35 90.7 89.8 0.51
17 32.25 1.58 5 5 50 0 40 69.2 68.6 0.58
Mean 31.80 0.78 20 18 21 29 39 84.0 76.9 0.53

vegetation type and beach width (Kamel and Mrosovsky
2006b). Of the habitat variables measured in the present
study, nest depth and distance to adjacent vegetation
(particularly shrubby, herbaceous, and arboreal plants)
positively affected hatching and emergence success.
Unlike other marine turtle species, the hawksbill tends to
select nesting sites with particular vegetation characteris-
tics (Kamel 2013). Kolbe and Janzen (2002) and Kamel
and Mrosovsky (2005) showed that hawksbill turtles
primarily nest in heterogeneous habitats composed of a
patchwork arrangement of areas with and without
vegetation. Other studies indicate that pioneer herbaceous
vegetation species have a consistent effect between nesting
seasons on hawksbill turtle nest site choice along with
other variables such as the vegetation line, distance of nest
from current waterline, and the highest spring tide (Santos
et al. 2016).
The hawksbill population nesting on Chenkan Beach
primarily nested at sites containing vegetation, supporting
a previous study on other beaches in the state of Campeche
which reported that more that 60% of nests were shaded by
vegetation (Guzmán et al. 1995). The present results
suggest that vegetation, mainly trees and shrubs, regulates
the temperature around nests, maintaining it at levels that
facilitate hatching and emergence. It is thus clearly
important to maintain natural vegetation structure, type,
and coverage patterns to optimize hawksbill turtle nesting
habitat.
The inverse relationship between creeping vegetation
and the other vegetation types shown in the ordination
diagram is explained by the effect plants had on variation
in reproductive variables, i.e., creeping vegetation pro-
duced a negative effect on variation in hatching and
emergence success compared with the positive effect of
the other vegetation types. This agrees with previous
studies indicating the presence of creeping plants to be
negatively correlated to nest depth (e.g., Kamel 2013),
perhaps due to the presence of roots, which prevent
females from forming adequate nests and thus slows egg
development. This highlights the importance of carefully
selecting plant species when restoring dune habitat on
nesting beaches. Native species are generally preferred
when restoring dune habitat (Moreno-Casasola et al.
2008), but any plant species that potentially facilitate nest
temperature regulation should be considered. Future
studies should determine if the presence of creeping plants
promotes the expansion of other vegetation types through
succession which could, in turn, increase the number of
turtle nests and improve reproductive output.
The incubation success variables of hatching and
emergence success were higher in dune zones with
vegetation in which nests had lower mean temperatures.
A negative relationship was observed between the
presence of creeping vegetation and hatching and
emergence success. This supports previous reports that
creeping plant species such as Ipomoea pes-caprae reduce
hatching success because roots enter the nest, often
harming the eggs (Conrad et al. 2011), trapping emerging
hatchlings, or both.
The present results coincide with the widely accepted
observation that vegetation is an important beach structural
component influencing the hawksbill turtle nesting
process. Hatching success was higher in nests surrounded
by vegetation, which supports the findings of Kamel
(2013) but contrasts with those of Ditmer and Stapleton
(2012). In addition to the spatial pattern of nest
arrangement on beaches, the percentage of plant coverage
and vegetation type are both important habitat factors that
contribute to successful hawksbill turtle reproductive
output. Novel visual techniques might reveal new insights
into how the spatial pattern of vegetation growing on

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 HERNÁNDEZ-CORTÉS ET AL. — Nesting Habitat of Hawksbill Turtles in Campeche, Mexico
beaches influences the nesting process and reproductive
success of marine turtles.
Overall, hawksbill turtles did not show preferences for
particular nesting beach slopes and widths, suggesting they
may be flexible users of different sites if severe changes in
beach morphology occur. For instance, turtles were
observed to avoid a beach sector where breakwater
structures had been installed, meaning turtles that had
consistently nested in this location were forced to move to
alternative sites. Breakwater structures are particularly
controversial because they reduce beach erosion in the
immediate vicinity but may represent an obstacle for
transit of female turtles to the beach.
Vegetation was a central structural component of
hawksbill nesting habitat, far more so than beach physical
characteristics. Vegetation limits beach erosion and
degradation as well as preventing temperature increases
in nests. Hatching and emergence success were generally
higher in sectors where the dune zone had high vegetation
coverage. The present findings indicate that vegetation
restoration programs on turtle nesting beaches should
consider vegetation spatial distribution patterns and the
vegetation type present to enhance hatching production on
beaches where high temperatures are an issue; these can
compromise embryo survival and lead to highly female-
biased sex ratios. This issue could become more salient in
the coming decades as climate change raises overall global
temperatures.
ACKNOWLEDGMENTS
The authors thank the Comisión Nacional de Áreas
Naturales Protegidas (CONANP) for providing informa-
tion and data, as well as staff, to carry out this project. We
also thank the Consejo Nacional de Ciencia y Tecnologı́a
(CONACYT) and the Universidad Autónoma del Carmen
for providing the support necessary to complete the study
as part of a research thesis for the Ecological Restoration
Master Science Program (J.A.H.-C.). The applicable
monitoring protocol for this project was the Official
Mexican Norm (NOM-162-SEMARNAT-2012) for col-
lection of standardized marine turtle nesting data (Diario
Oficial de la Federación [DOF] 2013). Finally, we thank
the editor and reviewers for their contributions to
improving manuscript quality.
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Received: 6 September 2017
Revised and Accepted: 6 March 2018
Published Online: 31 May 2018
Handling Editor: Jeffrey A. Seminoff

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