Thalassas: An International Journal of Marine Sciences

https://doi.org/10.1007/s41208-019-00166-y

Seagrass and Fish in Los Petenes Biosphere Reserve, Campeche,

Mexico: Spatial and Temporal Biomass Patterns
A. M. Rosas-Valdez 1 & L. A. Ayala-Pérez 2 & M. G. Figueroa-Torres 2 & I. E. Roldán-Aragón 2

Received: 12 November 2018 / Revised: 30 June 2019 /

# Springer Nature Switzerland AG 2019

Abstract
Los Petenes Biosphere Reserve (LPBR), in Campeche, Mexico, is a coastal strip that integrates a land portion -where petenes are
distinguished as unique ecosystems-, and a marine portion that highlights the presence of mangroves and seagrasses as key
ecosystems. In LPBR, seagrasses have been identified as the largest and best conserved ecosystem in the Gulf of Mexico. There is
a diverse biotic community associated with seagrasses, where fish are the most abundant macrofauna. The objective of this
research is to describe the spatial and temporal behavior of the biomass of seagrasses and fish in the littoral portion of the reserve.
A database of temperature and precipitation from three meteorological stations was built to describe the regional climatic
behavior. Four sampling campaigns were conducted, visiting 12 previously located sites. At each site, environmental variables
were measured, and samples of seagrasses were gathered by free immersion; also, fish samples were taken with an experimental
trawl. The general weather in the reserve has two defined seasons (rainy and dry). This weather regulates the environmental
variables registered in the sampling sites, and it is possible to identify associations between the abundance of both grasses and
fish. Grass biomass reaches maximum values of 664.9 g/m2, and fish biomass 1.61 g/m2. We identified 12 dominant fish among
which Lagodon rhomboides and Haemulon plumierii stand out. The high productivity of seagrasses in LPBR is confirmed, in
relation with climate seasonality and the optimization of the environmental conditions of the area.

Keywords Fish . Seagrasses . Biomass . Gulf of Mexico . Spatial and temporal patterns

Introduction Seagrasses provide key ecological services: habitat for

Los Petenes Biosphere Reserve is located in the north coast of
Campeche, Mexico and is distinguished by the breadth of its
protected surface and because it connects a terrestrial coastal
region with a portion of sea. It also combines distinctive struc-
tural elements such as mangroves, petenes (Maya word that
means round or circular and refers to islands of arboreal veg-
etation around a freshwater spring) and the largest and best
conserved seagrass meadows in the Gulf of Mexico
(Villalobos-Zapata and Mendoza-Vega 2010).
* L. A. Ayala-Pérez
luayala13@icloud.com
1
Division of Biological Sciences and Health, Metropolitan
Autonomous University Xochimilco, Calz. del Hueso 1100 Col.
Villa Quietud, Coyoacán, 04920 Ciudad de México, México
2 Barbosa and Santamaría 2013).
Department The man and his environment, Division of Biological
Sciences and Health, Metropolitan Autonomous University
Xochimilco, Ciudad de México, México
multiple species and their capacity to capture carbon dioxide
and the production of biomass that sustains the chain of detri-
tus are the most important (Shing Yip et al. 2014). Also, they
contribute to the development of various nutrient cycles and
have a fundamental role in stabilizing sediments and increas-
ing water transparency (Ondiviela et al. 2014; Cervantes-
Maldonado and Quintero 2016). According to Lara-Lara
et al. (2008) six genera and nine species of these angiosperms
are recognized in Mexico: marine grasses (Zostera marina,
Phyllospadix scouleri, Phyllospadix torreyi, Halodule
wrightii, Halodule beaudettei, Halophila decipiens), turtle
grass (Thalassia testudinum), manatee grass (Syringodium
filiforme) and star grass (Halophila engelmanni).
For the Caribbean region, H. wrightii, S. filiforme,
T. testudinum, Halophila baillonis and H. decipiens have been
reported (Millán et al. 2016), on the other hand, Ruppia
marítima, is reported as exotic specie that lives in shallow,
low salinity waters in estuarine systems of Mexico (Bonilla-
Fauna associated with seagrasses are diverse and abundant.
However, some fish stand out because they develop all or part

of their life cycles, and also play a fundamental role for the
services they provide to the ecosystem. They are also valued
for the economic benefits of fisheries (Allen Peña et al. 2004;
Ordóñez-López and García-Hernández 2005; Morales-López
et al. 2007).
The use of seagrasses as breeding, rearing and feeding
areas or, where appropriate, for the development of the entire
life cycle of many fish species, depends on the habitat’s envi-
ronmental stability, the possibilities of protection against pred-
ators and the availability of food (Gillanders 2007).
Regardless of its evident ecological importance, published
information on the composition, diversity and abundance of
the fish community in the marine portion of Los Petenes
Biosphere Reserve (LPBR) is scarce (Ayala-Pérez et al.
2012; Muñoz-Rojas et al. 2013). For this reason, the objective
of this research is to describe the spatial and temporal variabil-
ity of biomass (g/m2) of seagrasses and the composition and
biomass behavior of the fish community within a framework
of climatic variability in the coastal marine portion of LPBR,
Campeche, Mexico.
Methods
Los Petenes Biosphere Reserve is located on the northeast
coast of the state of Campeche. It has an area of
282,857 ha; more than half corresponds to the marine
zone up to 12 nautical miles from land. It extends through
the municipalities of Calkiní, Hecelchakán, Tenabo and
Campeche and is located in 15Q between 740,610–
779,347 and 2,308,198–2,193,398 UTM. The predomi-
nant climate in the center-south zone is AW (warm sub-
humid with summer rains); in the northern area, it is
BS’h’w (semi-dry and hot dry) (CONANP 2006).
Temperature and precipitation data of three weather sta-
tions: 1) Campeche (code 004038) location −90,544 W,
19,838 N, altitude 5 m, period 1982–2013; 2) Calkini (code
004002) location-90,054 W, 20,360 N, altitude 50 m, period
1985–2013; and 3) Poc Boc (code 004087) location
−90,835 W, 20,300 N, altitude 10 m, period 2002–2009, were
obtained from National Meteorological Service.
Four sampling campaigns were conducted during
September 2015, February, May and October 2016, visiting
12 sites distributed in the coastal marine strip of the LPBR, the
location of each site was established considering the geomor-
phological conditions, the connections with the continental
portion through estuaries and channels, the depth gradient
and the potential presence of seagrasses. Figure 1 shows the
location of the reserve, as well as the geospatial position of the
sampling sites.
At each site the environmental variables of salinity, tem-
perature, dissolved oxygen and pH were measured at two
depth levels (surface and bottom), using a YSI multiparameter
Thalassas
probe (556 MPS). Depth and transparency values were also
recorded with a Secchi disk.
The percentage of seagrass cover was calculated with the
help of a 1 × 1 m quadrant randomly positioned in each sam-
pling site. The quadrant was subdivided into four spaces of
0.5 m per side, in such a way that four replicas were taken at
each location. Depending on the density of seagrasses, the
total of coverage of only one sub quadrant was taken with
the help of a shovel, extracting the entire plants and storing
it in a plastic bag. The total biomass of quadrant was calculat-
ed considering the coverage percentage and the average
weight.
The fish samples were obtained with an experimental
trawl net of 5 m long, with a working opening of 2.5 m
and a mesh of 19 mm, operated for 12 min from a boat
with an outboard motor at a speed of 2.5 knots, covering
an approximate area of 1800 m2. This capture technique
was chosen considering previous work in the area, as well
as the order of magnitude of the fish biomass it provides.
The trawls were made in a circular way around the sam-
pling site. The organisms obtained were placed in plastic
bags, fixing them with 10% formaldehyde solution for
their preservation. Grasses and fish samples were
transported to the laboratory for processing.
Seagrasses were washed to remove adhering organisms
and their wet weight was recorded; afterwards, they were
dried in an ARSA oven for 72 h at a temperature of 74 °C.
The fish samples were washed and rehydrated for 24 h. They
were classified with the help of specialized literature (Fischer
1978; Castro-Aguirre 1999) and were measured individually
in standard length (cm) and total length (cm). To determine
biomass of grasses (dry weight by square meter) and fish
(weight by square meter), a digital scale of 2160 g with a
precision of 0.1 g was used.
The temporal and spatial environmental behavior, as
well as the biomass results of both grasses and fish are
described by box and whisker plots with the help of the
Systat program (Systat Software Inc. 2009). In both ana-
lyzes the obtained graphs express values of the median,
first and third quartile, minimum and maximum values in
addition to the outliers. The statistical criteria to define
the outliers were the following: Outliers mild: <Q1–1.5
* IQR, >Q3 + 1.5 * IRQ; and Outliers extreme: <Q1–3 *
IQR, >Q3 + 3 * IQR.
The dominant fish species were identified according to the
Relative Importance Index (RII) modified by Koranteng
(2001), which is defined as: RII = (% N) * (% W) * (% F).
Where: % N = Percentage of the contribution in number of
individuals of species i to the total catch; % W = Percentage
of contribution by weight of species i to the total catch; % F =
Percentage frequency of appearance of species i in relation to
the total number of sampling sites. Species with RII values
≥20 were considered dominant species.
Thalassas
Fig. 1 Location of sampling sites
in the marine littoral portion of
Los Petenes. Google earth. Image
Lansat/Copernicus. US Dept of
State Geographer. INEGI 2018
Results
The general behavior of the climate obtained from the daily
records of three meteorological stations of the National
Meteorological Service distributed along LPBR is presented
in Fig. 2, which shows the data grouped by month and the
trends of the average values of temperature and precipitation.
For the PocBoc weather station, there were 2795 records of
temperature with an interval of 12.5 °C to 34.5 °C. The central
tendency values were: average: 27.9; median: 28.5 and mode
29. The minimum average value was observed in January and
the maximum in May. With the same number of precipitation
records, the range of variation was from 0 to 210 mm with an
average of 4.26, and median and mode of zero. The rainiest
months were August, September and October.
In the case of the Calkini weather station, 10,382 tempera-
ture records were analyzed. The range of variation was from 8
to 35 °C; average was 24.2, median 24.5 and mode 28. The
lowest average temperature was observed in December and
the highest in May. Regarding precipitation, there were 11,288
records with a range of 0 to 200 mm, an average of 3.3, and a
median and mode of zero. The rainiest months were August
and September.
Finally, from the Campeche weather station, 11,566 tem-
perature records were used, which showed an average of
27.3 °C with an interval of 10.1 to 40 °C. The median was
27.7 and the mode was 28.5. The coldest month was January
and the warmest May. For precipitation, there were 11,863
records with an average of 2.9 mm and a range of 0 to
254.3; the rainiest months were September and October.
In general terms, there is a tendency for an increase in
temperature from January to May–June and a decreasing trend
from July to December. The coldest months are December and
January and the warmest May and June. Regarding precipita-
tion, there was substantial variability throughout the year,
however, between November and May, the values are minimal
and from June to October, the highest values. According to
this behavior, two climatic seasons are identified: rainy (June
to October) and dry (November to May). The difference in
temperature and precipitation between the two climatic sea-
sons determines the degree of salinity in the coastal-littoral
portion of LPBR.
Regarding on-site records, the temperature ranged from
19.7 °C (February, site 4) to 32.9 °C (September, site 8);
Salinity had a variation interval of 26.9 (September, site 8)
to 48.6 (May, site 3); dissolved oxygen had values between
 Thalassas

Fig. 2 Yearly trend of

temperature and precipitation
from three meteorological stations
of the State of Campeche (data
from 1983 to 2013). Outliers mild
are represented by circles and
Outliers extreme are represented
by asterisks

1.1 (October, site 6) and 7.3 mg/L (February, site 3), and pH
varied between 7.6 (October, site 8) and 9.9 (September, site
2). According to the Kolmogorov-Smirnov/Lilliefor test, there
Fig. 3 Spatial and temporal behavior of environmental variables. Boxes„
is sufficient evidence against normality for all registered pa- in red are surface records and the blue boxes correspond to those from the
rameters (p = 0.004–0.019) and, the nonparametric Kruskal- bottom. Outliers mild are represented by circles and Outliers extreme are
Wallis test for comparison of two independent samples represented by asterisks
Thalassas

indicates that there are no significant differences between var-
iables at the surface and the bottom (p > 0.766) (Fig. 3).
The range of depth variation at the sampling sites was from
0.7 to 4.1 m. The sites closest to the coast (1, 4, 7 and 10) had
an average depth of 1.67 m, the intermediate ones (2, 5, 8 and
11) an average of 2.12 m and the furthest (3, 6, 9 and 12) an
average depth of 2.34 m. the deepest site was 6. In all cases the
water transparency was 100%.
Thalassia testudinum is the dominant marine flowering
plant in LPBR. Its spatial and temporal biomass shows the
highest values in September and minimum in May. On a spa-
tial scale, biomass in several sites is minimal and sites 3, 5 and
7 are distinguished by having the highest average values of the
study area (630.5, 664.9 and 612.3 g/m2 respectively) (Fig. 4).
A total of 48 trawls were made gathering 3466 fish with a
total weight of 63.6 Kg. They were classified into 38 species
and 19 families (Table 1).
In October, biomass is highest (1.036 g/m 2 ) and in
September minimum (0.353 g/m2). Spatially, sites 1, 4 and 7
had the highest biomass values (1.61, 1.07 and 0.98 g / m2
respectively), and the minimum values were documented at
sites 11 and 12 (0.29 g / m2). It is necessary to highlight the
outliers in February, given the biomass at site 7 (Fig. 4).
Fig. 4 Temporal and spatial
behavior of the biomass of fish
and seagrasses (g/m2) in LPBR.
Outliers mild are represented by
circles and Outliers extreme are
represented by asterisks
Thalassas
From the RII, 12 dominant species were identified,
representing 92.07% of the total catch in number of individ-
uals and 88.58% in weight. Lagodon rhomboids and
Haemulon plumierii were the species with the highest RII
value (Table 2).
The spatial and temporal behavior of the biomass of
L. rhomboides and H. plumierii are presented in Fig. 5.
Discussion
In a context of tropical environmental variability, Los Petenes
Biosphere Reserve, together with Ría Celestún Biosphere
Reserve and El Palmar State Reserve, form an ecoregion in
Mexico’s Yucatan peninsula, with high diversity of flora and
fauna and unique critical habitats such as petenes, mangroves
and communities of seagrass (Espinosa-Carreón et al. 2001).
In accordance with a report by CONANP (2006), weather
in the region is defined by two periods: dry (November to
April) and rainy (May to October). The average values of
water temperature are the same as those of the report: 24 °C
in winter and 28 °C in summer, lowest temperatures are re-
corded in February (19.7–22.9 °C). Ayala-Pérez et al. (2003)
Thalassas
Table 1 Species list of fish collected in the Los Petenes Biosphere
Reserve, according to Nelson et al. (2016)
Family Species
Clupeidae Harengula jaguana Poey, 1865
Synodontidae Synodus foetens (Linnaeus, 1766)
Batrachoididae Opsanus beta (Goode & Bean, 1880)
Fundulidae Lucania parva (Baird & Girard, 1855)
Syngnathidae Syngnathus scovelli (Evermann & Kendall, 1896)
Scorpaenidae Scorpaena plumieri Bloch, 1789
Serranidae Diplectrum formosum (Linnaeus, 1766)
Lutjanidae Lutjanus analis (Cuvier, 1828)
Lutjanus apodus (Walbaum, 1792)
Lujtanus griseus (Linnaeus, 1758)
Lutjanus synagris (Linnaeus, 1758)
Ocyurus chrysurus (Bloch, 1791)
Gerridae Eucinostomus argenteus Baird & Girard, 1855
Eucinostomus gula (Quoy & Gaimard, 1824)
Gerres cinereus (Walbaum, 1792)
Haemulidae Haemulon aurolineatum Cuvier, 1830
Haemulon bonariense Cuvier, 1830
Haemulon plumierii (Lecepède, 1801)
Orthopristis chrysoptera (Linnaeus, 1766)
Sparidae Archosargus probatocephaus (Walbaum, 1792)
Archosargus rhomboidalis (Linnaeus, 1758)
Calamus penna (Valenciennes, 1830)
Lagodon rhomboides (Linnaeus, 1766)
Sciaenidae Bairdiella chrysoura (Lecepède, 1802)
Bairdiella ronchus (Cuvier, 1830)
Corvula batabana (Poey, 1860)
Cichlidae Cichlasoma urophthalmus (Günther, 1862)
Scaridae Nicholsina usta usta (Valenciennes, 1840)
Achiridae Achirus lineatus (Linnaeus, 1758)
Monacanthidae Aluterus shoepfii (Walbaum, 1792)
Monocanthus ciliatus (Mitchill, 1818)
Stephanolepis hispidus (Linnaeus, 1766)
Ostraciidae Acanthostracion quadricornis (Linnaeus, 1758)
Tetraodontidae Sphoeroides nephelus (Goode & Bean, 1882)
Sphoeroides spengleri (Bloch, 1785)
Sphoeroides testudineus (Linnaeus, 1758)
Diodontidae Chilomycterus schoepfii (Walbaum, 1792)
Diodon hystrix Linnaeus, 1758
report cold fronts during the dry season with winds and rain
causing an overall decrease in the temperature; however, the
effect is of short duration and no special behavior is identified
in climatic records.
Authors such as Sosa-López et al. (2009), Gutiérrez-Ayala
et al. (2012), Muñoz-Rojas et al. (2013) and Toro-Ramírez
et al. (2017) report values of some environmental variables
for LPBR that are useful to compare with the results in this
research. In general, the intervals of variation are similar; how-
ever, a difference is observed in February when there was a
stationary cold front in the Yucatan Peninsula that caused a
decrease in temperature, as well as an increase in the concen-
tration of dissolved oxygen. On the other hand, it is important
to note the negative trend of the pH values, which would
suggest the need for continuous monitoring to explain the
biogeochemical processes that take place under the different
environmental and climatic conditions of the area.
The spatial and temporal distribution of seagrasses depends
on variations in local environmental conditions, where salini-
ty, transparency, dissolved oxygen, as well as depth and the
harmonic movement of the tides are fundamental (Nielsen-
Muñoz and Cortés 2008). According to Gómez-López et al.
(2003), temperature and light penetration contribute to the
dispersal of T. testudinum seagrass meadows. In the LPBR
the continental shelf is very extensive with a very low slope
where the tidal regime is diurnal with an amplitude of 0.3 m
between low tide and high tide. These conditions allow
seagrasses to be established and developed properly, particu-
larly in shallower areas where water transparency is total.
Another correlation that becomes evident in these results is
between grasses biomass and salinity. Considering what was
reported by Hill et al. (2014), seagrasses have a tolerance interval
between 24 and 35 salinity practical units; in LPBR the highest
biomass values were documented in September when salinity
ranged from 27.42 to 30.7 SPU and the lowest biomass was
registered in May when salinity varied from 47.9 to 48.58 SPU.
Seagrass ecosystems are productive because of the amount
of organic matter they contribute in the form of detritus
(Millán et al. 2016). Foliar biomass (dry matter) of
T. testudinum in seagrass beds of the Colombian Caribbean
reaches average values of 150 g/m2 (Barrios et al. 2003); in
comparison, the results of this work report an average biomass
of 346.57 g/m2 of dry matter.
Regarding the fish community, there are some previous
studies that allow us to contrast our results. Firstly, the re-
search of Villalobos-Zapata (2004) that reports 68 fish species
for the LPBR; from those, 38 were registered in the present
study. In the work of Muñoz et al. (2013) 46 species were
classified, from which 32 matches; Ayala-Pérez et al. (2014)
reported 44 species from which 30 coincide and, finally, the
research by Toro-Ramírez et al. (2017) indicates 31 species of
which 22 are reported in this investigation. With this set of
observations, species richness of the fish community is evi-
dent and, furthermore, the close connection of fish with
seagrass meadows is emphasized because all the reports cited
were developed in the coastal zone of LPBR.
The total biomass of fish ranged between 0.01 and 2.84 g/
m2, the highest abundance was found in the sites closest to the
coast (1, 4, 7 and 10) with an average of 1.06 g/m2 followed
by the intermediate sites (2, 5, 8 and 11) with an average of
0.6 g/m2, and finally the sites furthest from the coast (3, 6, 9
 Thalassas

Table 2 Dominant species

according to the Relative Species Ind. No. Weight (g) %N %W %F IIR
Importance Index (RII)
Lagodon rhomboides 954 15,662.1 27.5 25.1 54.2 37,397
Haemulon plumierii 794 15,459.9 22.9 24.8 54.2 30,723
Eucinostomus gula 503 3421.8 14.5 5.5 50.0 3976
Archosargus rhomboidalis 342 6214.9 9.8 9.9 29.2 2864
Orthopristis chrysoptera 228 4186.3 6.6 6.7 33.3 1470
Lutjanus synagris 129 2215.1 3.7 3.5 43.7 577
Nicholsina usta usta 96 2607.3 2.8 4.2 37.5 433
Acanthostracion quadricornis 38 2342.3 1.1 3.7 25.0 102
Opsanus beta 37 1226.7 1.1 1.9 25.0 52
Calamus penna 42 7 18.2 1.2 1.1 29.2 40
Stephanolepis hispidus 39 743.3 1.1 1.2 25.0 33
Synodus foetens 22 1566.4 0.6 2.5 14.6 23
Total 3147 56,364.3
Capture total 3466 63,627.3

and 12) with an average of 0.58 g/m2, which suggests that the and it also shows that the values are higher at the end of the
fish also they have a close relationship with the mangrove dry season (May). It is necessary to mention the atypical abun-
ecosystem. Muñoz et al. (2013) reports biomass values of dance value registered in February, which corresponded to an
the fish community for the LPBR from 0.2 to 1.072 g/m2; exceptional catch of a school of L. rhomboides and it is

Fig. 5 Biomass of dominant fish

species in the LPBR throughout
the sites (space) and the year
(time). Outliers mild are repre-
sented by circles and Outliers ex-
treme are represented by asterisks
Thalassas
assumed that it was due to the effect of a cold front in this
particular sampling campaign.
Twelve dominant species were identified, among which
L. rhomboides and H. plumierii stood out. Sosa-López et al.
(2009), Muñoz et al. (2013), Ayala-Pérez et al. (2014) and
Toro-Ramírez et al. (2017) mention that these species are rep-
resentatives of biological success in the region, reaching alto-
gether 25% of the abundance of coastal fish in this region.
L. rhomboides commonly called Xlavita is common in
bays, estuaries and coastal lagoons, especially in meadows
of T. testudinum and in mangrove roots; they are usually
grouped in schools, characterized as an omnivorous species
and has local fishing importance (Darcy 1985; Canto-Maza
and Vega-Cendejas 2008; Lamle 2015). The relationship of
this species with seagrasses has been discussed by Spitzer
et al. (2000) after evaluating the effect of the density of
T. testudinum on the growth rates of juvenile individuals; their
results suggest that the capture of their preys is energetically
more expensive in the treatments of greater density because
the prey were harder to capture.
On the other hand, H. plumierii, also known as Chac-Chí,
is common in prairie areas of T. testudinum. In the juvenile
stage, it feeds mainly on benthic crustaceans, worms and mol-
lusks, while adults feed on shrimps, crabs and mollusks, and it
has local and national commercial value (Mexicano-Cíntora
et al. 2007). Ordóñez-López and García-Hernández (2005)
point out relationships of H. plumierii with the structural char-
acteristics of T. testudinum and suggest that grasses are a vital
habitat for their survival since they use it for protection and
feeding purposes.
Conclusions
The climate behavior of LPBR, analyzed by the temperature
and precipitation records of three meteorological stations dis-
tributed throughout the extensive protected area, allows us to
understand the environmental framework of a coastal tropical
zone with two clearly defined climatic seasons.
The seagrass meadows in the LPBR are a fundamental
structural component in ecosystem dynamics since it is the
habitat of multiple species for the development of all or parts
of their life cycles, and it is also a source of food for the chain
of detritus. The seagrasses maintain a wide distribution in
large patches that are associated with the environmental be-
havior of the area, such as the amplitude and slope of the
continental shelf, the temperature of a tropical zone, the salin-
ity pattern conditioned by climatic seasons, the regime of tides
and a low anthropogenic impact.
Fish are also a structural component of LPBR and are inti-
mately linked with seagrasses either as permanent residents,
cyclical visitors or occasional visitors. They stand out for their
abundance and diversity and for the niches they develop.
Particularly, 12 dominant species are identified, among which
L. rhomboides and H. plumierii stand out for their numerical
abundance and frequency of appearance. Both species repre-
sent 50% of the total capture.
Undoubtedly, it is important to highlight the need to con-
tinue with monitoring studies in this protected area that allows
us to investigate the ecological interactions among the differ-
ent components of the ecosystem.
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