The Intertidal Estuarine Communities of Lake Maracaibo, Venezuela
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THE INTERTIDAL ESTUARINE COMMUNITIES OF LAKE MARACAIBO, VENEZUELA GlLBERTO RODRIGUEZ Instituto Venezolano de Investigaciones Cientificas (IVIC), Caracas, Venezuela ABSTRACT The communities of rocky, sandy and muddy shores in the estuary are analyzed and the dominant species are recorded. The observed salinity ranges in mixohaline waters are combined with ranges observed in euhaline waters to give the salinity tolerances for the invertebrates in- habiting the estuary. The action of the substratum is discussed. At the end a summary of the physical conditions and a list of the plants and invertebrates are given. INTRODUCTION In a previous paper (Rodriguez, 1959) I have analyzed the structure of the intertidal communities of the tropical marine shores of Venezuela. In the present contribution an attempt is made to extend this analysis to the intertidal communities in mixohaline waters. For this purpose the estuary of Lake Maracaibo has been chosen. Lake Maracaibo lies in a geosynclinal basin of Miocene origin that once was a part of the extensive sea that covered the central portion of Venezuela. At present its area is 14,344 km2• Its maximal length, from Santa Rosa Point to Palmas Point, is over 155 km, and its maximal width, from El Guaco to Motatan del Lago, is 120 km. The greatest depth recorded is 34 m. This body of fresh water is in free communication with the marine waters of the Golfo de Venezuela through a narrow strait 39 km long and 6 km wide in the narrowest part. It has two mouths, one very shallow between the southern tip of Zapara Island and the east coast of the lake, the other between the west coast of Zapara Island and the southern tip of San Carlos Island. This mouth has been dredged to allow navigation to Port Maracaibo. The dredged channel has a least depth of 10 m. It is possible that the channel will allow a greater flux of sea water to enter the estuary and consequently in time conditions of life will be altered. The present survey was conducted at the time the channel was being dredged (July, 1956). Literature on the hydrography and biology of this estuary is very scanty, although this is the oldest and most important center of oil production in the country. In 1940, E. F. Robinson made a preliminary survey of the hydrography in connection with a dredging project (Robinson, 1947). A more detailed study was carried out by the Woods Hole Oceanographic Institution in 1953 and 1954 (Redfield, Ketchum and Bumpus, 1955). The Instituto Nacional de Canalizaciones has published a report on circulation and total discharge (Obadia, 1957). 198 Bulletin of Marine Science of the Gulf and Caribbean [13(2) Gessner has made studies on the hydrography and plankton of the Lake (Gessner, 1953; Gessner and Vareschi, 1956), and Ten Broek (195'0) has published a descriptive catalog of the molluscs. Crane (1943) has reported on the fiddler crabs of the genus Uca. ACKNOWLEDGMENTS Several persons and institutions cooperated with the author during the survey in 1956. The study was made while holding a postgraduate scholarship from the Venezuelan Association for the Advancement of Science. The lnstituto Nacional de Canalizaciones and especially lng. Arturo Obadia, provided all the field facilities. Dr. Th. van Andel, from the Compania Shell de Venezuela, made the granulometric analysis. Dr. Ten Broek assisted in the identification of molluscs. Dr. A. C. Redfield obtained permission from the Creole Petroleum Corporation to use the salinity data contained in his report. Drs. Hilary B. Moore and Gilbert L. Voss helped in many ways. To all these persons and institutions the author expresses his deepest gratitude. DEFINITION OF TERMS Description of communities.- The terminology used herein is as defined in a previous paper (Rodriguez, 1959), as follows: Zone: A band or region on the shore defined by physical conditions. Association: A community dominated by more than one species that creates special conditions for the secondary species different from the bare physical factors. Consocies: A community with one single dominant. Formation: An extensive climax community that covers large portions of a region. The word community is used in its broadest sense as a group of organ- isms. Belt is interchangeable with zone, but is used specially for bands of plants. Classification of waters according to salinity.-The classification of marine waters used here is in accord with the classification of salinity approved by the Symposium on the Classification of Brackish Waters, Venice, 1958, which is a modification of the Redeke- Valikangas system. Euhaline 40-30 %0 Mixohaline 30-0.5%0 Polyhaline 30-18 %0 Mesohaline 18-5 %0 Oligohaline 5-0.5%0 Limnetic < 0.5%0 1963] Rodriguez: Intertidal Estuarine Communities 199 Tidal Levels.-Two levels are considered as critical, Mean Low Water (MLW) and Mean High Water (MHW). These determine three main zones: Spray, from MHW up; Splash, from MHW down; and the Algal Belt to MLW. Also used are the words littoral for the region between MLW and MHW, supralittoral for the upper region and infralittoral for the lower region. STATIONS In order to obtain a representative sample of the intertidal fauna and flora of the estuary, eight conveniently spaced stations were established (Fig. 1). Capitan Chico Point (Station 1) is the southern limit of penetration for several estuarine species, and has an almost limnetic biota. The channels of San Carlos Island (Stations 2 and 3) are densely covered by mangrove undisturbed by human activities. Calvert (Station 8) represented a recently formed shore in process of colonization. Vigia Point (Station 4), Piedras Point (Station 5), the north coast of Zapara (Station 6, substation 4) and Toas Island were chosen as representatives of rocky shores. Zapara Island deserves special attention (Station 6). It is a narrow strip of sand dunes that forms a barrier to the entrance of marine waters into the estuary (Fig. 5). Its north coast is in contact with high salinity water (23%0) while its southern coast is in contact with low salinity water (6-8%0)' The island has been strongly altered by human action; mangroves have partially disappeared and new shores have been formed with sand from the dredgings. A breakwater has been laid on the north coast to prevent deposition of sand in the channel by the clockwise current in Golfo de Venezuela. LIST OF STATIONS, LAKE MARACAIBO, JULY, 1956 Station 1. Capitan Chico Point, 7.5 km north of Maracaibo. Station 2. Juan Zenon Channel, San Carlos Island. Station 3. Mouth of Juan Zenon Channel, San Carlos Island. Station 4. Vigia Point, north of Palmas Point. Station 5. Piedras Point, 5 km south of Altagracia Port. Station 6. Zapara Island. Substation 1. Entrance to the breakwater. Substation 2. Small bay at the west of the island. Substation 3. Shore on the southern coast of the island. Substation 4. Shore on the northern coast of the island. Station 7. Toas Island, shore on the west coast of the island, near the quarry. Station 8. Calvert, small peninsula, south of San Carlos Island. 200 Bulletin of Marine Science of the Gulf and Caribbean [13(2) o , 2 3 .( 5 I<MS .'::~:;~:~::,/':~1; I I I " I '.: !:. DEPTH CONTOURS 2m. 6m. ''1 8·m. MARACAIBO 1\1 FIGURE 1. Map of the northern portion of Lake Maracaibo, showing position of stations. 1963] Rodriguez: Intertidal Estuarine Communities 201 ECOLOGICAL SURVEY OF SANDY AND ~UDDY SHORES Station 1 (Fig. 2) is a small peninsula 1 Ian long and 0.5 km wide. The coastline is covered by Rhizophora, which is sometimes replaced by a tall grass, Phragmitis communis, that fills the same ecological role as does mangrove. The roots of Rhizophora and the stems of Phragmitis were covered at ~HW by a zone of Balanus improvisus. This substratum was sometimes shared by a red alga, Caloglossa leprieurii. The roots and stems are often dotted with oil, which does not seem to affect the survival of either the cirripede or the alga. Above the balanoid zone there was an abundant community of an estuarine gastropod, Neritina reclivata. The branches on the upper part were shared by three crabs, Rhithropanopeus harrisii, Metasesarma rubripes and Aratus pisoni. The supralittoral zone of the sandy beach had a crowded community of Vca murificenta, with the entrance to the burrows surrounded by circular walls. Below ~LW there was an extensive formation of Ruppia maritima. In tropical estuaries this seagrass fills the same role as Thalassia in marine bottoms. In places not covered by Ruppia there was a species of pelecypod, Polymesoda arctata, buried in the sand and leaving only the apex of the shell exposed to the water. This apex is used as a hard substratum for attachment by filamentous green algae. Numerous speci- mens of the blue crab, Callinectes sapidus, were found swimming among the Ruppia branches. In a small bay south of Capitan Chico Point there was a dense consocies of a fresh water submerged plant, Ceratophyllum demersum. Among its branches there was a small species of fresh water gastropod, Potamopyrgus sp. Floating in the water there were abundant specimens of Eichornia crassipes, the water hyacinth. According to Villalobos (personal com- munication), the roots of this plant in ~exico are the habitat of small atyid crustaceans, as well as other forms, but we were not able to detect them. The day we made our collections we obtained values of 1.32%0 for the salinity at low tide. Grain size analysis for the substratum of Ruppia, Uca murificenta and Ceratophyllum demersum respectively, are presented in Table III. The structure of the communities at this Station could be summarized as follows: A. Infralittoral, exposed places 1.