Artículo

 

Composition and spatio-temporal variation of the fish community in the Chacmochuch Lagoon system, Quintana Roo, Mexico

 

Composición y variación espacio-temporal de la comunidad de peces en el Sistema Lagunar Chacmochuch, Quintana Roo, México

 

J. Adán Caballero-Vázquez¹, Héctor C. Gamboa-Pérez², and Juan J. Schmitter-Soto¹

 

¹ El Colegio de la Frontera Sur. A.P. 424, 77000 Chetumal, Quintana Roo, México.

² Instituto de Impacto y Riesgo Ambiental de Quintana Roo. Calle 7 de Enero # 105, 77000 Chetumal, Quintana Roo, México. E-mail: adan7@mda.cinvestav.mx

 

Recibido: 27 de abril 2004.
Aceptado: 04 de octubre de 2005.

 

Abstract

The composition, distribution and abundance of fishes were determined in the Chacmochuch Lagoon System, a natural protected area located on the north coast of Quintana Roo. Monthly diurnal sampling was carried out at 10 sampling stations in March, June and September, 2002. During each field trip, bottom type, submerged and surrounding vegetation, salinity, temperature, dissolved oxygen and other variables were measured. Distribution and abundance of fishes were examined by means of a principal component analysis. Fifty species were recorded; Gerres cinereus, Floridichthys polyommus, Eugerres plumieri, Harengula jaguana, Sphyraena barracuda, and Archosargus rhomboidalis were dominant, and no significant difference in species composition was found between seasons. Apart from a single site where turbidity and nutrient concentration were particularly high, the other sites could be classified mainly according to salinity. Fish composition changed along this gradient.

Key words: Ichthyofauna, distribution, abundance, natural protected area, Caribbean.

 

Resumen

Se determinó la composición, distribución y abundancia de los peces en el Sistema Lagunar Chacmochuch, un área natural protegida en la costa norte de Quintana Roo. El trabajo se realizó durante un ciclo anual (marzo, junio y septiembre de 2002). Las 10 estaciones representaron los diversos hábitats del ecosistema. Los muestreos fueron diurnos. Se tomó nota del tipo de fondo, la vegetación circundante y sumergida, así como la salinidad, temperatura y oxígeno disuelto, entre otras variables ambientales. La distribución y abundancia de los peces se examinaron mediante un análisis de componentes principales. Se registraron 50 especies, siendo Gerres cinereus, Floridichthys polyommus, Eugerres plumieri, Harengula jaguana, Sphyraena barracuda y Archosargus rhomboidalis las dominantes. No hubo diferencia significativa en composición entre temporadas. Aparte de un sitio de turbidez y concentración de nutrientes particularmente alta, el resto de las estaciones de muestreo fue clasificable en función de la salinidad, principalmente. La composición íctica varió a lo largo de dicho gradiente.

Palabras clave: Ictiofauna, distribución, abundancia, área natural protegida, Caribe.

 

Introduction

The coastal zone is the most productive area of the oceans, both in terms of primary productivity and fisheries production (Holt, 1975). Coastal lagoons and estuarine environments are ecotones between the mainland and the sea, and are connected to the sea in a permanent or ephemeral manner. These ecosystems are shallow water bodies, semiclosed, with variable volumes of water, depending on the local climate and hydrology.

The life cycle of many fishes takes place partially or wholly within lagoons or estuaries (Castro-Aguirre et al., 1999). These sites are complex and variable biotopes, with an intricate web of trophic interactions and changing biogeochemical cycles (Toledo, 1991; Benedetti-Cecchi et al., 2001).

Fishes are the most diverse group of vertebrates, with more than 24,600 species known (Nelson, 1994). Mexico has about 2,800 described species, 14% of the World total (Lara-Domínguez et al., 1993). Many fishes use lagoon systems for feeding, breeding, or as shelter and recruiting areas.

The Chacmochuch lagoon system has many different habitats, where more than 194 species of Yucatan Peninsula flora and fauna can be found. There are subsistence fisheries, mainly mojarra, tarpon, snook, chihua, and barracuda. Local government and entrepreneurs plan to increase tourist activity in the system, which is a Natural Protected Area, established in 1999.

Because of these development plans, we felt it was important to carry out ichthyological studies, which can help set a baseline to monitor man-made changes in Chacmochuch. It is known, for example, that selective fishing for large piscivores such as snappers, with high demand in tourism-oriented restaurants, may bias the trophic structure of the fish community (cf. Loreto et al., 2003). Several attributes of fish communities have been proven as useful indicators of biotic integrity in other aquatic systems in Mexico (e.g. Lyons et al., 1995).

Doing this type of research is especially important in tropical regions, where understanding about the ecology and management of the natural resources may be scant, and development decisions may lack an adequate scientific background.

The objectives of the present study were to describe the composition, distribution, and abundance of fishes in the Chacmochuch lagoon system in different seasons, and to ascertain the influence of physicochemical variables.

 

Materials and methods

The Chacmochuch lagoon system is located northwest of the city of Cancún (21^o10'53"N, 86^o48'45"W; 21^o15'14"N, 86^o51'29"W), with an area of 11,527 ha. It is comprised of Chacmochuch Bay, the Ría or channel Nagigo, and nine other main water bodies (Figure 1). The climate is subhumid and warm, with a mean temperature of 26^oC, and a mean annual rainfall of 1300 mm (Comisión Nacional del Agua (CNA), unpubl. data). Tides have little influence.

Field work was carried out in March 10-14, June 10-15, and September 10-15, 2002, attempting to represent climatic variability: March has been in average the driest month (32.2 mm) during the last 30 years; June has been the warmest month (27.1°C, daily mean), and September-October, the rainiest (240.2-242.5 mm) (CNA, unpubl. data). Ten stations were established in representative areas of the system, based on bottom type, depth, and distance from the Caribbean Sea at the mouth of Chacmochuch Bay (Figure 1). Quantitative sampling was done with a 2.5-m high, 70-m long, 1.5-cm mesh seine net, a sampling gear that proved to be useful at more sites than other devices (but which, however, has a bias towards small fishes). To obtain the greatest possible diversity of organisms in each site, samples were taken at different hours, and other fishing gear was also used. At the Ría Nagigo, Larga and Manatí lagoon stations, the seine net did not work properly, so those data were not analyzed quantitatively.

Physicochemical conditions were determined in situ with a multiparametric sonde YSI 6600, measuring temperature, salinity, dissolved oxygen, pH, turbidity, depth, chlorophyll, TDS, ammonium, nitrates, and transparency. Organic matter was measured by the Walkley-Black method (Jackson, 1976). Submerged vegetation was measured by a quadrant method (percent cover in a square meter), and granulometric analysis was performed with sieves (12, 14, 16, 18, 20, 22, 45, 60, 80, 100, 140 and 200 mm) (Folk, 1969).

All fishes collected were fixed in 10% formaldehyde and preserved in 70% ethanol (Cailliet et al., 1986) for later identification in the laboratory, using methods described by Guitart (1977, 1978), Randall (1983), Humann (1994), Schmitter-Soto (1998), and Castro-Aguirre et al. (1999).

Each specimen was measured (standard length, SL) to the nearest mm, using a digital vernier caliper, and weighed to the nearest tenth of gram, with an electronic balance. All specimens were deposited as vouchers in the fish collection at ECOSUR-Chetumal (acronym ECOCH).

To check the reliability of our richness estimate, an accumulation curve of fishes to predict the total number of species in an area was adjusted, following the model S(t)= a/b (1 - exp(-bt)), where S(t) is the expected richness, t= is the effort (man-hours), and a and b are parameters. This model is especially adequate when one is sampling a relatively small area, a taxonomically well-known group, or both (Soberón & Llorente, 1993).

Community attributes measured were diversity by abundance (H'n) and biomass (H'w) (Shannon-Wiener index, bits·individual⁻¹ or bits·g⁻¹) (Ludwig & Reynolds, 1988), richness (Margalef, 1968), and evenness (Pielou, 1975). Species dominance was measured with the Sanders biological value index (BVIS) (Subrahmanyam & Drake, 1975). All indices were calculated with the ANACOM software (de la Cruz, 1994). Two-way ANOVA tests (Zar 1984) were used to test differences for each variable between the sample sites and sampling periods; data were log-transformed for this analysis. For all tests, P > 0.05 was used.

A cluster analysis (flexible sorting, Euclidean distance) determined sites associations, based on abundance of fishes. A principal components analysis (PCA) (Pla, 1986) was performed, separately for each season and also for the annual cycle, to explore the relationship between the presence and abundance of dominant species and environmental variables. These analyses were performed with MVSP (Kovach, 1995), and Statgraphics 7.0 software (Statistical Graphics Corporation, 1993).

 

Results

A total of 3124 specimens, 50 species, 39 genera, and 31 families were obtained; 14 species represented 95% of the total abundance. Total weight was 36.9 kg; 20 species represented 99% of the total biomass (Table 1). Most species are peripheral: 26 are euryhaline marine and 18 stenohaline marine (according to Castro-Aguirre et al., 1999). Six were freshwater secondary, and none were primary (sensu Myers, 1938). Two species, Apogon aurolineatus and Chaetodon capistratus, are new records for epicontinental waters of Quintana Roo.

The species-effort curve showed that the observed number of species was close to the estimated true richness (Figure 2).

Gerres cinereus was the dominant species throughout the year (Table 2). Juvenile fishes (considered as such based on lengths at first maturity from Claro, 1994) were collected in high proportion in all samples (Table 1).

The most widespread species were G. cinereus, Floridichthys polyommus, and Eugerres plumieri, which are bottom-feeders. Also common were Harengula jaguana, Archosargus rhomboidalis, Sphyraena barracuda, and Strongylura notata, among others. This includes carnivores as well as filter feeders (Claro, 1994).

In September, 1527 specimens in 36 species were obtained. G. cinereus, F. polyommus, E. plumieri, H. jaguana, A. rhomboidalis, and S. barracuda were the dominant species, representing 88% of the total abundance in this season. In April, 997 specimens in 21 species were recorded; total weight was 10.1 kg, with the same dominant species, plus Anchoa parva. In June, 600 individuals and 21 species were collected, with a total weight of 17.2 kg and the same dominant species, plus Lactophrys quadricornis and S. notata (Table 3).

There was no significant difference among seasons and sites in abundance, diversity, biomass, richness or evenness, perhaps because of the continuous dominance of the above-mentioned species. On the other hand, salinity, pH, TDS, dissolved oxygen, nitrate, and turbidity all had significant differences between seasons and sites. Temperature, ammonium, chlorophyll a, and depth showed significant differences among sampling sites, but not between seasons. Abiotic data are shown in Table 4; ANOVA results for these variables, in Table 5.

Cluster analyses of sites by species in every month presented nearly pectinate patterns, with no clear-cut groupings. On the other hand, a cluster analysis of the data pooled for the whole year produced three rather distinct groups (Figures 3 and 4):

Group I, characterized by stenohaline and marine species, captured mainly in sites 1, 3, 4 and 7 of Chacmochuch Bay. These fishes are temporary and occasional visitors: Hyporhamphus unifasciatus, Hippocampus erectus, and Caranx latus. They grouped together with L. quadricornis, Mugil curema, and Chilomycterus schoepfi, which are euryhaline species. Others, as A. aurolineatus, Haemulon aurolineatum and H. plumieri, are stenohaline and sporadic.

Group II, estuarine and euryhaline fishes captured mainly in sites 2 and 5 of Chacmochuch Bay. These permanent residents or common visitors include Atherinomorus stipes, H. jaguana, S. notata, A. rhomboidalis, E. plumieri, G. cinereus, A. parva, Centropomus undecimalis, Sphoeroides testudineus, S. barracuda, and F. polyommus. Together with Opsanus beta and Megalops atlanticus, these are peripheral species.

Group III, freshwater fishes, in the smaller water bodies Esperanza, Manatí, Zapatito, Larga and Ría Nagigo. These fishes are resident species, found all year long in fresh to oligohaline waters: "Cichlasoma" urophthalmus, Gambusia yucatana, G. sexradiata, Poecilia mexicana, P. velifera, and P. orri. The association of sampling sites, the salinity gradient and turbidity values are present in Chacmochuch lagoon system throughout the year (Figure 4).

Group IV, consisting only of site 6 in Chacmochuch Bay, separates clearly both in the species-based dendrogram and the environment-based PCA (Figures 3 and 5, see below). All the most abundant and frequent species in the system are included here, a fact attributable to the high productivity of that site, as shown by the values of nitrate concentration and turbidity.

As for environmental variables, a PCA for the pooled yearly data showed three groups of sites. One consists of sites 1-5 and 7 of Chacmochuch Bay, where high salinity, TDS, ammonia, and pH are determinant. A second group consisted of lagunas Zapatito, Esperanza and Garzas, with a high level of chlorophyll a, temperature homogeneity and low salinity. The last group consisted only of site 6 of Chacmochuch Bay, and it is set apart by its high turbidity, nitrate and chlorophyll a (Figure 5).

 

Discussion

Chacmochuch has 50 recorded fish species in an area of 11,527 ha; nearby lagoons Nichupté and Bojórquez have 37 species in 4452 ha (Reséndez-Medina, 1979). On the other hand, the richness of these systems in the northern coast of Quintana Roo is lower than that of Río Huach lagoon (southern coast), with 65 species in only 145 ha (Avilés-Torres, 2002). Even the predicted real richness of Chacmochuch, 59 species, fails to correct this large divergence. We do not see obvious differences in habitat complexity between the sites. As for the effect of regional faunas (cf. Caley and Schluter, 1997), we expected Chacmochuch to be more diverse because of the possible influence of fauna from the Gulf of Mexico. We speculate that the higher diversity of Río Huach might be explained because of its closer proximity to a well-developed coral reef (Avilés-Torres et al., 2001). Also, the extension of the wetlands bordering Río Huach is larger, a fact that can increase the productivity and as a consequence the diversity (Boero, 1994).

Numerical processes (i.e. abundance and ultimately diversity) and energetic processes (closely approximated by biomass) are two sides of the same phenomenon. For example, individual growth and condition, which are energetic processes, influence mortality, a numerical process. Habitat selection and recruitment are a compromise of both kinds of processes (Jones & McCormick, 2002; Gillanders et al., 2003).

The utilization of lagoon environments by marine and freshwater organisms is not random (Day & Yáñez-Arancibia, 1985); many species have evolved adaptations that optimize the use of estuarine and lagoon system habitat. One example are juvenile and preadult fishes that live in lagoons and migrate to the sea when they become adults, exporting energy and importing it again as larvae and juveniles (Deegan, 1993 — again, a linkage of numerical and energetic processes). Lagoons have a fundamental role in the storage and flow of energy to the sea; this fact explains the oligotrophy of reef systems. In Chacmochuch, water circulation patterns are generally complex and are strongly affected by wind, which, together with the tidal oscillation and geomorphology, helps keep larvae within the system (Stoner & Lewis, 1985; Joyeux, 1998; Jager & Mulder, 1999; Barleta-Bergan et al., 2002; Chittaro et al., 2005).

The influence of environmental parameters on estuarine fish communities has been verified many times. For example, Pauly (1975) found that biotic and abiotic variables controlled the composition of the community in a coastal semiclosed lagoon in West and South Africa. Apart from a single site separated because of its high turbidity and nutrient concentration, we found that our sampling sites were grouped by the PCA based mainly on salinity and temperature. This is also what Castro-Aguirre (1982) found in a coastal lagoon in the Mexican state of Oaxaca. In Southeastern Asia and Australia a gradient of several variables helps young fishes find breed ing areas (Blaber, 1985; Griffiths, 2001). Avilés-Torres (2002) found that salinity, bottom type and other abiotic variables controlled the composition of the community in the small coastal lagoon of Río Huach. On the other hand, in Chesapeake Bay, vegetation is more important than variables such as temperature (Orth & Heck, 1980). Similar conditions were found at two lakes in southeastern Brazil (Vono & Barbosa, 2001).

On the other hand, throughout the year there are species widely distributed in Chacmochuch, in spite of habitat heterogeneity. A possible explanation for this pattern is the tolerance of dominant species to environmental conditions; most of the fishes in Chacmochuch are either marine euryhaline or freshwater euryhaline (that is, secondary sensu Myers, 1938). On the other hand, the groupings of fish (marine stenohalyne, marine euryhalyne, and freshwater) maintain their spatial distribution in the lagoon throughout the year, in spite of the changes in the environmental variables, possibly because the gradient of the salinity and other parameters remain in spite of those variations.

Commercial species fished in the adjacent marine area (mojarras, jacks, snappers, snook, barracuda, among others) show up as juveniles in the lagoon system, whose conservation is thus important not only for the sake of tourism or for intrinsic reasons, but also for the welfare of fishermen not yet assimilated by the tourism-oriented economy of the region.

 

Acknowledgements

We thank El Colegio de la Frontera Sur and the Universidad de Quintana Roo, who provided financial support, equipment and facilities to conduct this research; Roberto Herrera, Juan Pablo Ortega, Marco Antonio Jiménez, Ricardo Briceño, and Máximo Suárez, for their assistance in field work in Chacmochuch; Héctor Ventura for GIS assistance, and Cecilia Hernández for her translation support. We also thank two anonymous reviewers, and Sophie Calmé, who corrected the English. Othón Cervantes provided unpublished climatic data from CNA.

This work is part of a M. Sc. thesis by the first author.

 

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