Printer Friendly

Structure and composition of the polychaete community from Bahia San Quintin, Pacific Coast of Baja California, Mexico.

Abstract.--The diversity patterns of the polychaete fauna from a Pacific coastal lagoon were described. Polychaetes were collected in 1995 and 1998. This lagoon is formed by 2 arms: the western arm named Bahia Falsa and the eastern arm named Bahia San Quintin. 46 stations were sampled with a geological box corer. A total of 3,275 polychaetes, 28 families, 56 genera, and 104 species were identified in 1995, and 3,168 polychaetes were collected in 1998, 21 families, 39 genera and 65 species. From all the macrofauna collected in both surveys, polychaetes represented 45.2%. From the species collected, 55% correspond to new records for the area. Families Dorvilleidae, Polynoidae, Oweniidae, Scalibregmatidae, Sternapsidae and Sigalionidae present in 1995, were not in 1998 survey. The stations with higher abundances (> 100 specimens/0.02 [m.sup.2]) were located on the southern half of Bahia San Quintin. Species richness and diversity were also higher in San Quintin Bay. From the 30 families previously reported for San Quintin lagoon, 23 have been collected and 6 families were added: Ampharetidae, Oweniidae, Scalibregmatidae, Sternapsidae, Dorvilleidae and Sigalionidae. Families not found in both surveys were: Paraonidae, Magelonidae, Apistobranchidae, Sphaerodoridae, Trichobranchidae, Chrysopetalidae and Arenicolidae.

Results showed slightly lower redox potential values (-336 to +187 mV), slightly higher sediment temperatures (19.8[degrees]-22.1[degrees]C) and organic matter contents (0.3-4.1%) in 1998.

From 1995 to 1998 a change in the composition and structure of the polychaete communities was noted; species richness diminished from 104 to 65 species. The trophic complexity changed with an increase of deposit-feeders, the abundance of other trophic categories decreased, indicating a loss of complexity. Significant changes in the abundance of some families were detected, some increased their abundances: Spionidae from 17% to 48%, Orbiniidae from 4% to 13%; other families decreased in terms of abundance and number of species: Lumbrinereidae from 11% to 1.4%, Nereididae from 9% to 1% and Sabellidae from 14% to 5%. These modifications altered the composition and structure of the polychaete communities in this lagoon. Increased anthropogenic disturbance (oyster culture, agriculture) and environmental variability due to the ENSO 97-98 may have affected recruitment and survival of some polychaete species.


Some lagoons, located along the Pacific coast of Mexico, present ideal hydrological and sedimentary characteristics which make them potential sites for aquaculture. The San Quintin complex is one of these coastal lagoons that favor the development of bivalve aquaculture. It is considered ecologically important because it is a nursery area for several fish species, a resting site for migrating birds which have lost most of their resting and feeding areas in the United States, and its high productivity and diversity, in part due to upwellings which supply nutrients periodically. It is environmentally important to obtain baseline scientific data that help understand how benthic communities function and how they change during different climatological conditions.

The hydrology of San Quintin lagoon has been studied (Alvarez-Borrego & Chee-Barragan 1976; Alvarez-Borrego et al. 1975; del Valle & Cabrera-Muro 1981 a, b; Farfan & Alvarez-Borrego 1983). In contrast, there is a lack of information on macrofauna. One of the most neglected, major groups of marine invertebrates may be the polychaetous annelids that could be useful as indicators of varying degrees of marine pollution (Tsutsumi 1990; Pocklington & Wells 1992). Only three polychaete surveys were found on the literature: Reish (1963) 90 stations sampled in 1960 in Bahfa San Quintfn (BSQ eastern arm), Calderon-Aguilera & Jorajuria-Corbo (1986) 11 stations sampled in 1981-82, 8 in BSQ and 3 in Bahia Falsa (BF western arm); and Diaz-Castaneda & Rodriguez Villanueva (1998) 39 stations sampled in December 1992, 13 in BF and 26 in BSQ.

Coastal marine benthic communities are threatened by human activities, and the present rate of habitat degradation is alarming. Given that only a small fraction of the benthic organisms that reside on or are buried in sediments have been described, it is likely that species are being lost without ecologists knowing they existed (Snelgrove 1999). Polychaetes constitute an important macrofaunal group in this lagoon comprising about 70% of the benthic biomass and individuals (Barnard 1970; Calderon-Aguilera & Jorajuria-Corbo 1986). More than 1,450 polychaete species are known from Mexico (Salazar-Vallejo et al. 1989; Diaz-Castaneda & Rodriguez-Villanueva 1998). Polychaetes are a significant component of all marine ecosysytems, they dominate soft-bottoms communities in terms of numbers of species and individuals. These annelid worms are important in food webs and in energy transference, both as predators and as important prey items for other animals, including crustaceans, fish and wading birds (Knox 1960). They present different feeding modes (carnivores, herbivores, omnivores, deposit-feeders, symbiotic chemoautotrophic bacteria), this plasticity could be the reason of their success in many environments (Beesley et al. 2000). Many species are important bioturbators of sediment and facilitate the incorporation of organic matter into sediments. Polychaetes show a spectacular diversity of reproductive and developmental modes which allow them to live in different environments (Wilson 1991; Giangrande 1997). Because of their cosmopolitan distribution, polychaetes can be used as indicators of pollution and the "state of health" of a benthic community (Pearson & Rosenberg 1978; Reish 1980; Bellan et al. 1988; Pocklington & Wells 1992; Lardicci & Rossi 1998).

From a management perspective, they are useful organisms for identifying problem sites and for the assessment of the severity of the problem. They respond to disturbance induced by different kinds of pollution, by exhibiting quantitative changes in assemblage distribution. Polychaetes can also be used as indicators of recovery of benthic environments from perturbations since in many cases they are major elements of the recolonization process (Diaz-Castaneda et al. 1989; Diaz-Castaneda & Almeda-Jauregui 1999).

In spite of their importance in benthic communities few faunal studies have occurred in Mexico, in part because of identification problems due to lack of proper identification keys as well as the low number of polychaetologists (Salazar-Vallejo et al. 1989; Pocklington & Wells 1992).

El Nino is an important phenomenon throughout the world. Its effects on marine ecosysytems and organisms may go beyond temperature change. Invertebrates have complex life cycles in which certain life stages, and therefore the dynamics of entire populations, are at the mercy of various physical processes acting within the ocean-atmosphere system (Arntz & Tarazona 1990; Bakun 1996; Escribano et al. 2004). During El Nino Southern Oscillation (ENSO) 1997-1998, high temperatures and low nutrient concentrations resulted in widespread mortality of giant kelp forests (Macrocystis pyrifera) (Tegner & Dayton 1987) and other marine organisms in the region. Temperature anomalies greater than I[degrees]C persisted continuously for 8 months in the west coast of Baja California, in some cases anomalies attained +3[degrees]C (Dayton et al. 1992).

The purpose of this study is to describe the composition and structure of polychaete communities in San Quintin lagoon in 1995 and 1998 before and after the El Nino 1997-98.

Study Area

San Quintin complex is a slightly hypersaline, highly productive coastal lagoon located between 30[degrees]24'-30[degrees]30' N and 115[degrees]57'-116[degrees]01' W in the Pacific coast of Baja California (Fig. 1). This lagoon has an area of 42 [Km.sup.2] (4,200 hectares) and around 80% of it is covered by the eelgrass Zostera marina (Inclan-Rivadeneyra & Acosta-Ruiz 1988; Poumian-Tapia & Ibarra-Obando 1999). It has been exploited for many years (mariculture) but it can still be considered a relatively non disturbed area, although oyster culture is increasing. The region is arid, with a mean annual rainfall of about 150 mm. About 90 percent of the rainfall occurs between October and March.


Seagrass beds are important nursery areas for many species of fish and invertebrates, including several of economic importance (Stoner 1980 a, b; Orth & van Montfrans 1984, 1990). They also help to stabilize sediments thus reducing coastal erosion and are responsible for the composition and diversity of the seagrass infauna.

The lagoon has the shape of an inverted "Y", it consists of two sub-basins: BF (west) and BSQ (east). BF has an average depth of 4 m whereas BSQ has an average depth of 8 m. The bay has extensive intertidal and shallow subtidal shoals and channels up to about 10 m deep extending along the length of each basin. It has a permanent entrance and exchanges water with the coastal ocean. During low tides around 20% of the seafloor is exposed. An important aspect of the marine environment is the pattern of coastal upwelling, which is strongest between May and August (Aguirre-Munoz et al. 1999). The granulometric studies show that in shallow areas as well as to the north of both arms clay and silty-sand predominate, whereas near the mouth very fine sands are more abundant. The channel sediments are highly diverse, going from medium to fine sand and silt (Calderon-Aguilera 1992; Camacho-Ibar et al. 1997; Poumian-Tapia & Ibarra-Obando 1999). The lagoon margins present a typical saltmarsh flora with Spartina foliosa and Salicornia virginica and other vascular plants (Dawson 1962; Barnard 1970).

Material and Methods

Forty six stations were sampled in December 1995 and April 1998, including 15 stations in BF and 31 in BSQ (Fig. 1). Samples were collected using a box corer (16 cm internal diameter, 13 cm depth, sampling area of 0.02 [m.sup.2]). Temperature and redox potential were measured immediately after collection of each sample by probing 2-3 cm inside the sediments an electrode coupled to a field potentiometer and a thermometer. Sediments were sieved in the field using a 1.0 mm mesh size and retained material was fixed in 10% buffered formaldehyde. In the laboratory, samples were washed and transferred to 70% isopropanol. Different zoological groups and particularly polychaetes were then sorted and identified at species level whenever possible.

Organic matter (percent of dry weight) was evaluated by ignition loss (Byers et al. 1978). Statistical methods were used to describe the structure and organization of the polychaete communities within the bay. Shannon diversity index and Pielou equitability were calculated in order to study the structure and degree of organization of the communities (Shannon & Weaver 1963; Frontier 1985; Pielou 1977). Trophic groups were determined using Fauchald & Jumars (1979) and Rouse & Pleijel (2001).

Olmstead and Tukey's test (Sokal & Rohlf 1995) was applied to analyze spatial distribution of polychaetes. This technique plots the frequency of appearance in each site sampled expressed as percentage against the density of organisms for each species. A mean average was calculated for both axes, resulting in four quadrants: I Frequent and abundant species, II Non frequent and abundant species, Ill Non frequent and non abundant species and IV Frequent and non abundant species.

Stress predictability (Alcolado 1992) modeling was applied to establish the level of environmental stress existing in the bay. Environmental severity or stress was predicted based on values of diversity (H') and evenness (J'), coupled with redox potential values.

Ordination and classification methods were used to detect spatial patterns among the polychaete fauna. The relationship between sample stations is reflected by the position they display in factorial space; when the two stations were close to each other, they had more similar faunistic profiles (Frontier & Pichod-Viale 1993; Diaz-Castaneda et al. 1993). A factorial correspondence analysis was carried out on the faunistic data: abundance of species and 46 stations. Cluster analysis using Pearson and Bray-Curtis coefficients (Bray & Curtis 1957; Sokal & Rohlf 1995) was employed to evaluate the level of association of different stations and species. A non-metric multidimensional scaling (MDS) method was used for the community ordination (Program PRIMER 5.1.1 for windows) since this technique has demonstrated to be suitable for multiple ecological purposes (Clarke 1993; Clarke & Green 1988). The MDS is based on the calculation of similarity/ dissimilarity coefficients among samples, in this case, the similarity coefficient of Bray-Curtis. One data matrix was created for each sampling period using abundance per species. Data were treated using Primer Program 5.1.1 for windows and Statistica v. 5.0, after transformation to [log.sub.10] (X + 1) as suggested by Frontier (1983) and Legendre & Legendre (1984).


In 1995, the redox potential values (Eh) were negative in most of the stations. In the eastern arm they varied between -340 and + 162 mV; BF presented values between--320 to +161 mV. Sediment temperatures oscillated between 19.1 and 22.0[degrees]C. Organic matter values varied between 0.3 to 3.4% in BSQ and 0.1 to 3.1% in BE In 1998 only 43% of stations were measured for Eh and temperature.

In the eastern arm the Eh varied between -336 mV and + 154, while the western arm presented values between--308 and + 187 mV. Sediment temperatures were in the range 19.8 to 22.1[degrees]C, while the organic matter content ranged between 0.5 to 4.0% in BSQ and 0.3 to 4.1% in BF (Table 1). These results show slightly lower Eh values and slightly higher temperature and organic matter contents in 1998.

The lists of species found in each survey are given in Table 2. In 1995, a total of 8,680 benthic organisms were collected, of which 38% were polychaetes, 36.5% were crustaceans and 27.4% were molluscs. The 3,275 polychaetes collected and identified belonged to 28 families, 56 genera and 104 species (Table 2).

The families best represented were Capitellidae (19%), Spionidae (17%), Sabellidae (14%), Lumbrinereidae (11%), Nereididae (9%), Cossuridae (8%) and Syllidae (8%). The 10 top dominant species were Prionospio heterobranchia (331), Chone infundiliformis (295), Mediomastus californiensis (264), Cossura candida (236), Scoletoma crassidentata (222), Exogone lourei (206), Capitella capitata (147), Armandia brevis (130), Neanthes arenaceodentata (121), Chone mollis (117). The first eight species constitute 55% of the total abundance, the first five have been reported as abundant in previous studies (Reish 1963; Calderon-Aguilera 1986; Diaz-Castaneda & Rodriguez-Villanueva 1998). Reish (1963) found six species that constituted the dominant bay species on the basis of number of specimens. These were, in decreasing order of importance, Prionospio malmgreni, Exogone verugera, Cossura candida, Capitia ambiseta, Scoloplos acmeceps and Fabricia limnicola. Calderon-Aguilera (1992) reported five numerically dominant species: Exogone occidentalis, Pseudipolydora kempi, Scoloplos acmeceps, Prionospio heterobranchia and Neanthes arenaceodentata.

In April 1998, a total of 5,584 benthic organisms were collected, of which 56.7% were polychaetes, 27.2% were crustaceans and 7.5% were molluscs. The 3,168 polychaetes identified, belonged to 21 families, 39 genera and 65 species (Table 2). The families best represented were Spionidae (47.6%), Capitellidae (12.3%), Syllidae (10.5%), Paraonidae (7%) and Orbiniidae (6.8%). The first six species constitute around 75% of the total abundance. The ten top dominant species were Prionospio heterobranchia (832 specimens), Polydora websteri (548), Scoloplos acmeceps (370), Exogone lourei (291), Mediomastus californiensis (273), Cirriformia spirabrancha (128), Capitella capitata (68), Chone mollis (62), Megalomma pigmentum (59) and Fabricinuda limnicola (50).

The following families present in 1995 were not found in the 1998 survey: Dorvilleidae, Polynoidae, Oweniidae, Scalibregmatidae, Sternapsidae and Sigalionidae. Some of these families have species that are carnivorous. The increase in temperature in 1998 is related to a diminution of prey items which in turn may have affected their abundances.

Olmstead & Tukey's graph is only presented at the family level (Fig. 2a), because there were too many species to produce a clear graph. In 1995 and 1998 the polychaete families were placed in three out of four possible categories: dominant, restricted and rare. In 1995, in quadrant I (frequent and abundant), 6 polychaete families were characterized as dominant. Spionidae, Nereididae, Sabellidae, Lumbrinereidae, Capitellidae and Syllidae families displayed high densities and wide distribution throughout the lagoon. The families Spionidae, Capitellidae, and Sabellidae presented the highest densities and combined accounted for 45% of the total abundance of polychaetes. Ten families restricted to certain areas of the lagoon were located in quadrant II (non-frequent and abundant) and corresponded to 32% of all families. Within quadrant III (non-frequent and non-abundant), 12 polychaete families were located, classified as rare or occasional. No families were located in quadrant IV corresponding to frequent and non-abundant families. Approximately 35% of species were located in quadrant I (36 species). In 1998 (Fig. 2b), in quadrant I, only 3 polychaete families were characterized as dominant. The families Spionidae, Orbiniidae and Capitellidae displayed high densities and


Although abundances were similar in 1995 and 1998, we noted a change in the composition and structure of the polychaete communities; species richness diminished signficantly from 104 to only 65, reflecting the diminution of diversity values and equitability of the communities. The trophic structure also changed as deposit-feeders increased their numbers, and the abundance of other trophic categories diminished, indicating a modification in community organization and loss of complexity.

Significant variations in the abundance of some important families were detected between 1995 and 1998; some increased their abundances: Spionidae from 17% to 48%, Orbiniidae from 4% to 13%. These families are essentially deposit-feeders although some species of spionids can also feed as filter-feeders. Other families decreased in terms of number of species: Lumbrinereidae from 11% (4 species) to 1.4% (1 species), Nereididae from 9% (5 species) to 1% (only 2 species) and Sabellidae from 14% to 5% (3 species). Lumbrinereids live in sand or mud, in plant roots or in algal hold-fasts, nereidids are omnivores and herbivores and sabellids are filter-feeders. These modifications changed the diversity and structure of polychaete communities in this lagoon.

Diversity values ranged between 1.18-4.04 in 1995, and 1.10-3.51 in 1998, with higher values mainly located in BSQ, especially the middle section (Fig. 3). Diversity in BF diminished in 1998, reaching a maximum of 2.71. In 1995, 55% of stations presented equitability (Pielou index) values higher than 0.800, indicating a more or less even distribution of polychaetes at the species level. Whereas in 1998 only 35% of stations reached more than 0.800. BF presented in both surveys lower equitability and diversity values, probably this is related to the increasing oyster culture which seem to affect the benthos due to negative Eh values.

Values of species diversity (H') and evenness (J') for stations sampled were analyzed and placed into four "environments" (Fig. 4) as defined by the Stress-Predictability modeling (Alcolado 1992). We found seven times more stations classified in environment I (very favorable and stable) in 1995 compared to 1998.


In 1995 (Fig. 4a), environment I, which included 21 stations with the highest values of diversity (H') (2.52-4.04) and evenness (J') (0.740-0.910), was characterized as being very favorable and stable. Environment II was represented by 11 stations (10, 13, 21, 24, 26, 28, 32, 33, 34, 38, 42) located in the middle areas of BSQ and BE it was favorable and stable, H' values ranged between 2.68-3.62 and J' values between 0.694-0.910. Environment III (stations 2, 3, 4, 15, 37, H' 2.41-2.89, J' 0.653-0.849) was characterized as being constant, with a degree of environmental stress. Finally, environment IV which corresponded to stations 18, 35, 36 and 39, was moderately favorable, with unstable conditions and a certain degree of environmental stress. Environment IV station diversity values (H") oscillated between 1.17-2.83, evenness values (J") from 0.700 to 0.960, and were the lowest of this study.


Whereas in 1998 (Fig. 4b), environment I included only 3 stations (6, 15, 25) with the highest values of diversity (H') (3.24-3.52) and evenness (J') (0.777-0.878), was characterized as being very favorable and stable. Environment II was constituted by 10 stations (2, 3, 4, 10, 11, 12, 13, 14, 18, 21) situated mainly in the medium section of BSQ, it was favorable and stable, H' values ranged between 2.42-3.18 and J' values between 0.723-0.918. Environment III increased with respect to 1995 survey, it was formed by 12 stations located in both arms (5, 8, 9, 16, 19, 26, 27, 32, 35, 36, 39, 43), H' 2.16-2.51, J' 0.575-0.836) characterized as being constant, with a degree of environmental stress. Finally, environment IV which corresponded to stations 30, 33, 34, 38, 40, 41, 42 and 45, located in the middle of BF was moderately favorable, with unstable conditions and a certain degree of environmental stress. Environment IV station diversity values (H") oscillated between 1.56-2.16, evenness values (J") from 0.640 to 0.831, and were some of the lowest of this study. The number of stations in this last category almost doubled in relation to 1995.


In relation to trophic groups, polychaete species found in 1995 and 1998 were in decreasing order of importance: deposit-feeders, carnivores, filter-feeders and herbivores. Deposit-feeders increased their numbers in 1998. We also detected an important presence of carnivores, essentally syllids, which increased their abundance in 1998. Spionidae and Capitellidae which were among the most abundant families have species which are surface and subsurface deposit-feeders, as well as filter-feeders. The dominant species Prionospio heterobranchia can feed as a deposit-feeder or as a filter feeder, this capacity allows them to stay and even increase their abundances in disturbed conditions.

The families best represented in 1995, by decreasing order of importance were Capitellidae, Spionidae, Sabellidae, Lumbrinereidae and Nereididae, they changed in 1998: Spionidae, Orbiinidae, Capitellidae, Syllidae, Sabellidae.

Bray-Curtis coefficient of similarity was used to measure the level of association of samples. The dendrogram (Fig. 5a) generated with samples and replicates from 1995 revealed a clear separation between both arms, to the right of the graph are located BSQ samples and to the left BF samples (except stations 4 and 15). BF revealed four groups at 30% level of similarity: group I with 6 stations from the middle and mouth, group II formed by six stations from the southern section and mouth, group III constituted by five stations from the north and near the mouth and group IV formed by 6 stations from the western section. Groups III and IV are formed by stations from the northern section which is the least well flushed. At the left of the figure appear 9 stations connected to the other groups with low levels of similarity. At 42% similarity SQB samples are separated into three groupings: group V comprised 21 stations, it is subdivided in two groups, VA constituted by 13 stations located in the middle and south and VB formed by eight stations from the north, group VI gathered seven stations from the middle and south and group VII is constituted by 12 stations from the middle section of the bay.


The dendrogram (Fig. 5b) generated with samples from 1998 revealed also a separation between both arms, although it is less clear than in 1995. To the right of the graph are located mainly SQB samples and to the left most of BF samples. At 30% level of similarity which correspond to a faunistic dissimilarity for the bay we observe 3 groups: group I formed by 13 stations from middle SQB and east BE at a higher level of similarity it is subdivided in two groupings IA formed by stations from middle BF and south BSQ and IB which gather stations from the southern half of BSQ. Group II formed by four stations located near the mouth and group III constituted by 18 stations subdivided in three groupings which represent respectively north BF and middle BSQ, middle BSQ and north BSQ.


Factorial Correspondence Analysis (FCA) and non metric MDS were applied to a matrix of 104 species and 45 stations in 1995, and a matrix of 65 species and 42 stations in 1998. Data treated correspond to the polychaete species abundances. Some representative graphics were selected.

In 1995, the first two axis extracted 45.5% of total inertia. In the factorial plane 1-2 we observed that samples from each lagoon arm gathered, forming BF and BSQ two separate groups in the factorial space (Fig. 6a). Most of the stations of BSQ were located on the positive side of axis 2 whereas all stations of BF were situated in the negative side of this axis which indicate that polychaete communities inhabiting each lagoon arm are not exactly the same, even if the species list is similar, the proportion between species change. This can be explained by the fact that hydrological conditions are not the same in both arms, depth averages are 4 and 8 m in BF and BSQ respectively. Conditions in BF can favor the development of more sensible species, those that can not tolerate very low oxygen concentrations and/or very negative Eh values because in this arm the water exchange with the sea is faster (Diaz-Castaneda & Rodriguez-Villanueva 1998). However if oyster aquaculture increases, it may cause a negative effect in the water and sediment quality, affecting these species.


In 1998, the first two axes extracted 55.2% of total inertia. In the factorial plane 1-2 (Fig. 6b), BF stations appear together, forming two groups. One of the groups is located on the positive side of axis 2, the second group which correspond to the southern section is situated on the negative side of this axis, near the stations from north BSQ. BSQ stations form a group that extends along axis l, the stations from the north (negative side of axis 2), middle and southern sections differentiate.


Polychaete communities were also examined using non metric multidimensional scaling. This analysis also provided evidence of a separation between samples of the eastern and western arms of the bay. MDS analysis applied to data from 1995 (Fig. 7a) revealed a group of stations located in the center which correspond to BSQ; to the right are located stations from the northern area. Around these stations are located those from BE The stations near the mouth are surrounded by a dotted circle. Stress value (0.15) indicated that the configuration was a good representation of the faunistic similarities between stations. In 1998 (Fig. 7b) the separation between stations from BSQ and BF is also observed but not so clearly, stations from north BSQ are situated in the right and upper section. Most BSQ stations are located in the right side, while BF are mainly in the left. Stress value is 0.13. The mouth stations are essentially located in the same position than in 1995.


In conclusion, the study of polychaete communities in both arms of San Quintin lagoon showed in both surveys differences in species diversity, composition of species and abundances. Polychaete communities seemed to have balanced according to the physico-chemical characteristics prevailing in the area in 1995 and 1998. The proportion of families changed, some families disappeared completely in the 1998 survey: Dorvilleidae, Polynoidae, Oweniidae, Scalibregmatidae, Sternapsidae, Sigalionidae. Seven families found in 1992 (Diaz-Castaneda & Rodriguez-Villanueva 1998) were not collected in 1995 or 1998: Paraonidae, Magelonidae, Apistobranchidae, Sphaerodoridae, Trichobranchidae, Chrysopetalidae and Arenicolidae.

The eastern arm (SQB) presented in both surveys, higher polychaete densities and diversity values, probably the oyster aquaculture in Falsa Bay although not intensive has produced a certain impact in the benthic communities due to an excess of organic matter, resulting in negative Eh values and reduction of abundance and diversity. Comparing the polychaete composition and the values of some ecological indexes (Shannon Index, Pielou, Simpson dominance) between 1995 and 1998 we detected an important diminution of the species richness and diversity.

In BSQ several stations presented diversity values higher than 3.50, one station had more than 4.0; in BF only one station presented a higher value than 3.50, no stations had values higher than 4.0. Higher species richness values were located in BSQ (up to 26/station), indicating that this is an adequate environment for polychaetes development. It seems that the physico-chemical characteristics prevailing in each arm, as well as water exchange rates influence benthic communities development. The mathematical methods applied allowed a better global vision of the organization and structure of these populations.

Increased environmental variability due to the ENSO 97-98 probably may have affected recruitment, survival and reproductive patterns of polychaete species, modifying the composition and structure of polychaete communities between 1995 and 1998. In Chile, Escribano et al. (2004) found increased benthic bioturbation during the same period. Probably there was seagrass reduction (Zostera marina) in 1998 that could have indirectly affected zoobenthic recruitment, which is partially driven by the hydrodynamic environment modified by seagrass meadows, as has been observed by other authors (Bostrom & Bonsdorff 2000).

Finally, in San Quintin lagoon more than 50% of stations had relatively high values of diversity and evenness, indicative of healthy benthic communities. In general, the bay is characterized by high diversity values and approximately 70% of stations were favorable and constant environments for polychaetes, especially in the southern area of BF where the bivalve culture takes place and the middle section of BSQ.


We would like to thank E. Gutierrez for his help during field work in 1995. G. de la Selva and M. Necoechea for helping sort the macrofauna and C. Almeda for his help with statistical programs. J. Dominguez and F. Ponce helped with the preparation of figures. The comments of an anonymous reviewer were greatly appreciated.

Literature Cited

Aguirre-Munoz, A., R. Buddemeier, V. Camacho-Ibar, J. Carriquiry, S. Ibarra-Obando, B. Massey, S. Smith & E Wulff. 1999. Sustainable versus unsustainable resource utilization in an isolated coastal ecosystem. Regional Environmental Change. Editor-in-chief: W. Salomons. Springer-Verlag. Berlin, Germany.

Alcolado, M. P., 1992. Sobre la interpretacion del ambiente marino mediante el empleo de los indices de diversidad y equitatividad. Ciencias Biologicas 24, 124-127.

Alvarez-Borrego S., Ballesteros G. & A. Chee-Barragan. 1975. Estudio de algunas variables fisicoquimicas en Bahia de San Quintin, en verano otono e invierno. Ciencias Marinas, 2(2):1-9.

Alvarez-Borrego S. & A. Chee-Barragan. 1976. Distribucion superficial de fosfatos y silicatos en Bahia de San Quintin, B.C., Ciencias Marinas 3(1):51-61.

Arntz, W. E. & J. Tarazona. 1990. Effects of El Nino on benthos, fish and fisheries off the South American Pacific coast, In: Glynn, R W. (Ed.), Global Ecological Consequences of the 1982-83. El Nino-Southern Oscillation. Elsevier Oceanography Series, Amsterdam, pp. 323-360.

Bakun, A. 1996. Patterns in the Ocean. Ocean Processes and Marine Population Dynamics. CIBNOR, Baja California Sur. 323 pp.

Barnard J. L., 1970. Benthic Ecology of Bahia San Quintin, Baja California. Smithsonian Contr. Zool, 44:1-60.

Beesley, R, G. Ross & C. J. Glasby. 2000. Polychaetes and Allies: The Southern synthesis Fauna of Australia. Vol. 4A Polychaeta, Myzostomida, Pogonophora, Echiura, sipuncula. CSIRO Publishing: Melbourne, 465 pp.

Bellan G., G. Desrosiers & A. Wilsie (1988) Use of an Annelid Pollution Index for monitoring a moderately polluted littoral zone. Marine Pollution Bulletin 19:662-665.

Bostrom, C. & E. Bonsdorff. 2000. Zoobenthic community establishment and habitat complexity--the importance of seagrass shoot-density, morphology and physical disturbance for faunal recruitment. Mar. Ecol. Prog. Ser. 205:123-138.

Bray, J. R. & T. Curtis. 1957. An ordination of the upland forest communities in southern Wisconsin. Ecological Monographs 27, 325-349.

Byers, S. C., E. Mills & P. Stewart. 1978. A comparison of methods to determine organic carbon in marine sediments with suggestions for a standard method. Hidrobiologia 58: 43-47.

Calderon-Aguilera, L. 1992. Analysis of the benthic infauna from San Quintin, with emphasis on its use in impact assessment studies. Ciencias Marinas 18 (4): 27-46.

Calderon-Aguilera, L. & A. Jorajuria-Corbo. 1986. Nuevos registros de especies de poliquetos (Annelida: Polychaeta) para la Bahia de San Quintin, B.C:, Mexico. Ciencias Marinas 12 (3): 41-61.

Camacho-Ibar, V., J. Carriquiry & S. Smith. 1997. Bahia San Quintin, Baja California (a teaching example), p. 9-15. In S. Smith, S. Ibarra-Obando, R Boudreau and B. Camacho-Ibar (eds.),

Comparison of Carbon, Nitrogen and Phosphorus fluxes in Mexican Coastal lagoons. LOICZ Reports & Studies N[degrees] 10, Neth. Inst. Sea Res., Texel, Netherlands.

Clarke, K. 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18:117-143.

Clarke, K. & R. Green. 1988. Statistical design and analysis for a biological effects study. Mar. Ecol. Prog. Ser. 46: 213-226.

Dawson, E., 1962. Benthic marine exploration of Bahia de San Quintin, Baja California, 1960-61. Marine and Marsh Vegetation. Pacific Nat. 3(7): 275-280.

Dayton, P., M. Tegner, P. Parnell & P. Edwards. 1992. Temporal and spatial patterns of disturbance and recovery in a kelp forest community. Ecol. Monogr. 62: 421-445.

del Valle & H. Cabrera-Muro. 1981a. Aplicacion de un modelo numerico unidimensional a Bahia San Quintin, Baja California. Ciencias Marinas. Vol. 7: 1-15.

del Valle & H. Cabrera-Muro. 1981b. Analisis estadistico de condiciones hidrodinamicas en la Bahia de San Quintin, Baja California. Ciencias Marinas. Vol. 7: 17-29.

Diaz-Castaneda, V., A. Richard & S. Frontier. 1989. Preliminary results on colonisation, recovery and succession in a polluted area of the southern North Sea. Proc. 22nd EMBS. Barcelona, Spain. Topics in Marine Biology. J. Ros Ed. Scient. Mar. Vol. 53 (2-3): 705-716.

Diaz-Castaneda, V., Frontier, S., V. Arenas. 1993. Experimental re-establishment of a soft-bottom community: utilization of multivariate analyses to characterize different benthic recruitments. Estuarine Coastal and Shelf Science 37, 387-402.

Diaz-Castaneda, V. & V. Rodriguez-Villanueva. 1998. Polychaete fauna from San Quintin Bay, Baja California, Mexico. Bull. Southern California Acad. Sci. 97: 9-32.

Diaz-Castaneda, V. & C. Almeda. 1999. Early benthic organism colonization on a Caribbean coral reef (Barbados, W.I.): a plate experimental approach. Marine Ecology. 20 (3-4): 197-220.

Diaz-Castaneda, V. & G. San Martin. 2001. Syllidae (Polychaeta) from San Quintin Bay, Baja California, Mexico, with the description of a new genus. Proc. Biol. Soc. Wash., 114: 708-719.

Farfan, C. & S. Alvarez-Borrego. 1983. Variability and fluxes of nitrogen and organic carbon at the mouth of a coastal lagoon. Estuarine, Coastal and Shelf Science, 17: 599-612.

Fauchald, K. & P. Jumars. 1979. The diets of worms: a study of polychaete feeding guilds. Ocean. Mar. Biol. Ann. Review 17, 193-284.

Frontier, S. 1983. Strategies d' echantillonage en Ecologie. Masson, Paris. 494 p.

Frontier, S. 1985. Diversity and structure in aquatic ecosystems. Ocean. Mar. Biol. Ann. Review 23: 253-312.

Frontier, S. & D. Pichod-Viale. 1993 Ecosystemes. Structure, fonctionnement et evolution. Ed. Masson, Paris. 447 p.

Giangrande, A. 1997. Polychaete reproduction patterns, life-cycles and life-histories: an overview. Ocean. Mar. Biol. Ann. Review 35: 323-386.

Inclan Rivadeneyra, R. & M. Acosta Ruiz. 1988. Analisis de la Comunidad Incrustante en las Balsas para el Ostion Japones Crassostrea gigas en Bahia San Quintin, Baja California, Mexico. Ciencias Marinas. 15(1): 21 38.

Knox, G. 1960. Littoral ecology and biogeography of the Southern Ocean. Proc. R. Soc. Lond. B 152: 577-625.

Lardicci, C. & F. Rossi. 1998. Detection of stress in macrozoobenthos: evaluation of some methods in a coastal Mediterranean lagoon. Marine Environmental Research 45: 367-386.

Legendre, L. & P. Legendre. 1984. Ecologie Numerique. La structure des donnees ecologiques. Masson, Paris & Presses Univ. Quebec, 254 pp.

Orth, R., K. Heck & J. van Montfrans. 1984. Faunal communities in seagrass beds: a review of the influence of plant structure and prey characteristics on predator-prey relationships. Estuaries 7: 339-350.

Orth, R. J. and J. van Montfrans. 1990. Utilization of marsh and seagrass habitats by early stages of Callinectes sapidus: a latitudinal perspective. Bull. Mar. Sci. 46(1): 126-144.

Pearson, T. H. & R. Rosenberg. 1978. Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. Oceanogr. Mar. Biol. Ann. Review 16:229-311.

Pielou, E. C., 1977. Mathematical Ecology. John Wiley and Sons. New York. 385 pp.

Pocklington, P. & P. G. Wells. 1992. Polychaetes. Key taxa for Marine Environmental Quality Monitoring. Marine Pollution Bulletin 24: 593-598.

Poumian-Tapia, M. & S. Ibarra-Obando. 1999. Demography and biomass of the seagrass Zostera marina in a Mexican coastal lagoon. Estuaries 22: 834-847.

Reish, D. 1959. New species of Spionidae (Annelida: Polychaeta) from southern California. Bull. Southern California Acad. Sci., 58: 11-16.

Reish, D., 1963. A quantitative study of the benthic polychaetous annelids of Bahia de San Quintin, Baja California. Pacific Naturalist, 3: 399-436.

Reish, D. 1980. Use of polychaetous annelids as test organisms for marine bioassay experiments. In: Buikema AL Jr & J Cairns Jr (eds) Aquatic invertebrate bioassays: 140-154. American Society for Testing and Materials, Special Technical Publication No. 715, Philadelphia, Pennsylvania.

Rouse, G. & F. Pleijel. 2001. Polychaetes. Oxford University Press. 354p.

Salazar-Vallejo, S., J. A. de Leon-Gonzalez & H. Salaices-Polanco. 1989. Poliquetps (Annelida: Polychaeta) de Mexico: Generalidades, Claves ilustradas para familias y generos, y Bibliografia-Lista de especies. Libros Univ. Auton. Baja Calif. Sur, La Pax, Mexico. 212 p.

Shannon, C. E. & W. Weaver. 1963. The mathematical theory of communication. Univ. Illinois Press, Urbana. 125 pp.

Snelgrove, P. V. 1999. Getting to the bottom of marine biodiversity: sedimentary habitats. Ocean bottoms are the most widespread habitat on earth and support high biodiversity and key ecosystem services. BioScience 49: 129-138.

Sokal, R., & F. J. Rohlf. 1995. Biometry. Freeman. New York. 887 pp.

Stoner, A. W. 1980a. Perception and choice of substartum by epifaunal amphipods associated with seagrasses. Mar. Ecol. Prog. Ser. 3:105-111.

Stoner, A. W., 1980b. The role of seagrass biomass in the organization of benthic macrofaunal assemblages. Bull. Mar. Sci., 30: 537-551.

Tegner, M. & P. Dayton. 1987. El Nino Effects on Southern California kelp forest communities. Monograph: Advances In Ecological Research. Editor: A. Macfadyen & E. Ford, Vol. 17, pp 243-279.

Tsutsumi, H. 1990. Population persistence of Capitella sp. (Polychaeta; Capitellidae) on a mud flat subject to environmental disturbance by organic enrichment. Mar. Ecol. Prog. Ser. 63: 147-156.

Wilson, W. H. 1991. Sexual reproductive modes in polychaetes: classification and diversity. Bull. Marine Science, 48: 500-516.

Victoria Diaz-Castaneda, (1)* A. de Leon Gonzalez, (2) and E. Solana Arellan (1)

(1) Departamento de Ecologia, CICESE, Km 107 Carr. Tijuana-Ensenada, Ap. Postal 2732, Ensenada, Baja California

(2) Laboratorio de Biosistematica, Facultad Ciencias Biologicas, UANL San Nicolas de Los Garza, Nuevo Leon C. P. 66451 Mexico

* Corresponding author. E-mail:

Accepted for publication 10 February 2005.
Table 1. Physico-chemical values of San Quintin lagoon sediments.

 1995 1998
Station Eh(mV) T[degrees]C % O.M. Eh(mV) T[degrees]C % O.M.

 2 3.40
 3 --340 21.8 2.03 --336 20.2 4.05
 4 155 21.6 1.68 --208 21.1 2.90
 5 --106 20.2 2.84
 7 --160 20.9 1.86 --175 21.4 2.04
 8 --196 20.8 1.38 --180 21.6 1.35
 9 --245 20.8 2.01 --253 20.9 2.14
10 130 21.2 2.19
11 --73 21.0 1.10
12 --102 20.3 1.03 --98 20.8 1.66
13 --155 20.2 0.84
14 --197 21.0 1.32 --144 21.2 1.48
16 --187 19.9 2.35
17 --167 20.0 1.83
18 --109 20.1 2.04 --120 21.4 2.40
19 --219 20.6 2.12
21 --177 1.15 --174 21.5 1.57
22 --82 19.1 1.50 --92 21.4 1.92
24 162 21.7 2.46
25 --112 21.1 1.27 --126 21.7 1.94
26 --95 21.6 0.80
28 --62 20.5 1.38 103 21.5 1.45
29 66 21.1 0.91
30 --43 21.3 0.30 154 19.8 0.50
31 --69 21.5 0.42
32 --92 21.4 1.30
33 161 21.7 0.22 187 21.3 0.55
34 --24 21.3 2.14
35 --128 21.1 2.45
36 --85 21.6 1.60 --106 21.8 1.66
38 --123 21.6 2.90 --152 21.8 1.87
39 --136 21.2 2.78
40 --49
41 33 21.3 2.90 --190 20.9 2.70
42 --176 21.5
44 --320 21.2 3.27 --277 21.8 2.95
45 --308 22.1 4.11
46 --249 21.4 3.30

Table 2. Polychaete species from San Quintin lagoon, Baja California.

Species 1995 1998

 Ampharete labrops Harman, 1961 x
 Ampharete sp x
 Amphicteis acutifrons Grube, 1850 x
 Amphicteis sp Grube, 1850 x

 Capitella capitata Fabricius, 1780 x x
 Mediomastus californiensis Hartman, 1944 x x
 Mediomastus sp x x
 Notomastus magnus Hartman, 1947 x
 Notomastus tennis Moore, 1909 x x
 Notomastus sp x

 Aphelochaeta marioni Saint-Joseph, 1894 x
 Aphelochaeta sp x
 Cirriformia spirabranchia Moore, 1904 x x
 Monticellina tesselata Harman, 1960 x
 Protocirrineris socialis Blake, 1996 x
 Protocirrineris sp x

 Cossura candida Hartman, 1955 x x
 Cossura sp A x x

 Dorvillea sp x

 Lysidice ninetta Verril, 1900 x
 Marphisa disjuncta Harman, 1961 x
 M. sanguehea Montagu, 1815 x x
 Morphysa sp x x

 Pherusa capulata Moore, 1909 x x
 Piromis arenosus Kinberg, 1867 x x
 Piromis sp x

 Glycera americana Leidy, 1855 x x
 G. tennis Hartman, 1944 x x

 Goniada brunnea Treadwell, 1906 x
 G. littorea Hartman, 1950 x x

 Podarkeopsis glabra Hartman, 1961 x x
 Podarke pugettenisis Johnson, 1901 x x

 Scoletoma crassidentata Fauchald, 1970 x
 S. erecta Moore, 1904 x
 S. monroi Fauchald, 1970 x
 S tetraura Schmarda 1860 x x

 Axiothella rubrocincta Johnson, 1901 x
 Axiothella sp Verril, 1900 x
 Clymenura gracilis Moore, 1923 x
 Euclymeninae sp A Ardwidsson, 1906 x
 Isocirrus longiceps Moore, 1923 x
 Maldane sp x

 Nephtys caecoides Hartman, 1938 x x
 Nephtys sp x

 Neanthes caudata delle Chiaje, 1828 x x
 Nereis latescens Chamberlin, 1919 x
 N. pelagica Linne, 1758 x
 Nereis sp x
 Platynereis bicanaliculata Baird, 1863 x
 P. marphysa x

 Arabella Tricolor Montagu, 1804 x x
 A. pectinata Fauchald, 1970 x
 Drilonereis falcata Moore, 1911 x
 D. longa Webster, 1879 x
 D. mexicana Fauchald, 1970 x
 Drilonereis sp x
 Notocirrus californierisis Hartman, 1944 x

 Kinbergonuphis sp x x

 Armandia bioculata Hartman, 1938 x
 A. brevis Moore, 1906 x x
 Ophelia pulchela Tebble, 1953 x
 Polyophthalmus picuts Dujardin, 1839 x

 Leitoscoloplos mexicanus Fauchald, 1972 x
 L. normalis Day, 1977 x
 Naineris grubei Gravier, 1908 x
 Phylo felix Kinberg, 1866 x
 P. ornatus Verril, 1873 x
 Scoloplos actneceps Chamberlain, 1919 x x
 S. armiger Muller, 1776 x
 S. ohlini Ehlers, 1901 x
 S. texana Maciolek & Holland, 1978 x

 Owenia collaris Hartman, 1955 x

 Eteone Pacifica Hartman, 1936 x
 Eteone sp x x
 Eulalia bilineata Johnston, 1840 x
 Eumida sp x

 Harmothoe imbricata Linne. 1767 x
 Harmothoe sp x

 Chone infundibuliformis Kruyer, 1856 x
 C. mollis Bush, 1904 x x
 Fabricinuda limnicola Hartman, 1951 x x
 Megalommra pigmentum Reish, 1963 x x

 Scalibregma sp x

 Sthenelais fusca Johnson, 1897 x

 Aporionospio pigmaeus Hartman, 1961 x
 Boccardiella hamata Webster, 1879 x x
 Microspio pigmentata Reish, 1959 x
 Minuspio cirrifera Wiren, 1883 x
 Polydora socialis Schmarda, 1861 x
 P. websteri Hartman. 1943 x
 Prionospio heterobranchia Reish, 1959 x x
 P. lighti Maciolek, 1985 x x
 Pseudopolydora pauchibranchiata Okuda, 1937 x
 Scolelepis squamata Muller, 1806 x
 Spiophanes bombyx Claparede, 1870 x
 S. duplex Chamberlain, 1919 x x
 S. missionensis Hartman, 1941 x
 Spio pacifica Blake & Kudenov, 1978 x
 Spio sp x

 Cicese sphaerosylliformis Diaz & San Martin, 2001 x x
 Eusyllis sp x
 Exogone lourei Berkeley & Berkeley, 1938 x x
 Grubeosyllis mediodentata Westheide, 1974 x x
 Pinrosyllis Sp x x
 Sphaerosyllis californiensis Hartman, 1966 x
 Syllis aciculata Treadwell, 1945 x
 S. gracillis Grube, 1840 x x
 S. heterochaeta Moore, 1909 x
 Syllis sp x


 Eupolymnia nebulosa Montagu, 1818 x
 Pista alata Moore, 1909 x x
 Pista sp x x
 Polycirrus sp x

Table 3. Polychaete species recorded in previous studies at San Quintin
lagoon, Baja California. Numbers correspond to the different taxa
recorded in the area.

Reish, 1963

1 Anaitides ca. multiseriata
2 Anaitides williamsi
3 Arabella iricolor
4 Arenicola cristata
5 Armandia bioculata
6 Axiothella rubrocincta
7 Brania clavata
8 Capitella capitata
9 Capitita ambiseta
10 Chone mollis
11 Chrysopetalum occidentals
12 Cirrifornia luxuriosa
13 Cirrifornia spirabrancha
14 Cossura candida
15 Dorvillea articulata
16 Eteone dilatae
17 Eteone pacifica
18 Eulalia bilineata
19 Exogone verugera
20 Fabricia limnicola
21 Glycera americana
22 Goniada brunnea
23 Hapioscoloplos elongatus
24 Lepidonotus caelorus
25 Lumbrineris erecta
26 Lumbrineris minima
27 Marphysa sanguinea
28 Megalomma pigmentum
29 Nephtys caecoides
30 Nereis caudata
31 Nerinidaes maculata
32 Notomastus magnus
33 Onuphis microcephala
34 Ophiodromus puggettensis
35 Pista alata
36 Platynereis bicanaliculata
37 Polydora uncata
38 Polyophthalmus pictus
39 Prionosopio malmgreni
40 Prionospio pygmaeus
41 Scoloplos (L) ohlini
42 Scoloplos acmeceps
43 Scyphoproctus oculatus
44 Sphaerodorum minutum
45 Spiophanes missionensis
46 Trypanosyllis gemmipara
47 Typosyllis variegata
48 Aedicira pacifica
49 Aricidea suecica
 Armandia bioculata
 Axiothella rubrocincta
 Brania clavata
 Chone mollis
 Cirrifornia luxuriosa
50 Cossura soyeri
 Eteone dilatae
 Eteone pacifica
 Exogone occidentalis
 Fabricia limnicola
 Kinbergonuphis microcephala
 Leitoscoloplos pugettensis
 Lepidonotus sguamatus
 Lumbrineris erecta
 Lumbrineris mimima
51 Magelona pitelkai
 Marphisa sanguinea
52 Mediomastus ambisetus
53 Mediomastus californiensis
 Megalomma pigmentum
 Neanthes arenaceodentata
54 Nephtys caecoides
55 Nephtys ferruginea
 Notomastus magnus
 Notomastus tenuis
 Pherusa capulata
56 Phylo felix
 Pista alata
 Platynereis bicanaliculata
 Polyophtalmus pictus
57 Prionospio cirrifera
58 Prionspio heterobranchia
59 Prionospio malmgreni
60 Prionospio newportensis
61 Pseudopolydora kempi
 Scolelepis maculata
 Scolopios acmeceps
 Scyphoproctus oculatus
 Spiophanes missionensis
62 Apistobranchus sp
 Arabella tricolor
63 Brada villosa
 Brania sp.
64 Cirrifornia cf. spirabrancha
 Chaetozone sp.
 Chone sp.
65 Clymenura gracilsis
 Cossura candida
66 Euchone sp.
67 Exogone dispar
68 Exogone lourei
69 Goniada maculata
 Kinbergonuphis cf microcephala
70 Leitoscoloplos mexicanus
71 Lysidice ninetta
72 Marphysa sanguinea
 Megalomma bioculatum
73 Monticellina tesselata
 Neanthes acuminata
 Nereis sp.
 Notomastus sp.
74 Pionosyllia sp.
75 Polydora socialis
76 Praxillela sp.
77 Prionospio multibranchiata
 Prionospio (Minuspio) cirrifera
 Prionospio heterobranchia
78 Scoletoma tetraura
79 Scoloplos rubra
80 Spio pettiboneae
81 Syllis (Syllis) gracilis
COPYRIGHT 2005 Southern California Academy of Sciences
No portion of this article can be reproduced without the express written permission from the copyright holder.
Copyright 2005 Gale, Cengage Learning. All rights reserved.

Article Details
Printer friendly Cite/link Email Feedback
Author:Diaz-Castaneda, Victoria; Gonzalez, A. de Leon; Arellano, E. Solana
Publication:Bulletin (Southern California Academy of Sciences)
Geographic Code:1MEX
Date:Aug 1, 2005
Previous Article:The reef fish assemblage of the outer Los Angeles Federal Breakwater, 2002-2003.
Next Article:Composition of the epifaunal community associated with the seagrass Zostera marina in San Quintin Bay, Baja California.

Terms of use | Privacy policy | Copyright © 2021 Farlex, Inc. | Feedback | For webmasters