Location and General Description
Distributed on a narrow latitudinal stripe this mangrove ecoregion’s mangrove woodland fringe almost 1/4 of the Venezuelan coastline, on the northernmost extreme of South America stretching from Laguna de Cocinetas in the west, to the western edge of the Orinoco Delta system in the far east. They cover nearly 5,900 km2, thus ranking as one of the largest mangrove ecoregions in South America (Conde & Alarcón 1993; Conde & Carmona-Suárez 2002). The main mangrove stand located on the Atlantic Ocean coast, between the Gulf of Paria and the Orinoco Delta, annual rainfall exceeds 2,000 mm, freshwater input is steady and geomorphic conditions are regarded as optimal. These pristine, structurally complex and extensive mangrove forests reach heights of 35-40 m and form almost continuous belts, extending through about 400 km of coastline. The rest of the ecoregion is located on a semi-desert belt along the Caribbean Coast of Venezuela, mostly in the Gulf of Venezuela and on the central-western and central-eastern coasts (MARNR 1986). Small stands are also found offshore on many of the islands.
Most of the coastal Venezuelan mangrove ecoregion stands are small and only a few surpass 40 km2. These are found in the Gulf of Paria, San Juan River, Limón River- San Carlos Island, Los Olivitos, Cuare-Morrocoy and in Laguna de Tacarigua. In the insular domain, smaller although substantial mangrove tracts are found in Los Roques and Las Aves Archipelagos, La Orchila Island and in Margarita Island.
Red mangrove (Rhizophora mangle), black mangrove (Avicennia germinans) and white mangrove (Laguncularia racemosa) are the most common species of mangroves in Venezuela. A fourth species, the buttonwood mangrove (Conocarpus erectus) is regarded as an "associate" rather than a "true" mangrove; it appears in mangrove stands located in arid zones. Some species are less abundant, geographically constrained or their presence is still to be unquestionably confirmed are R. harrisonii, R. racemosa, A. schaueriana and Pelliciera rhizophorae. Although locally called mangroves, Thespesia populnea, Calliandra riparia and Coccoloba uvifera are not mangroves in the strict sense. The chewstick (Symphonia globulifera), a tree whose leaves resemble those of the genus Rhizophora, was photo-interpreted as a mangrove, leading to an almost three-fold overestimation of mangrove coverage in a survey undertaken some 15 years ago, in this ecoregion (Conde & Alarcón 1993).
In general, the coastal climate is dry tropical for most of the Caribbean strip, with average temperatures around 27.1°C to 30.6 °C. Scant precipitation is distributed in two distinct periods. In many localities rainfall rarely exceeds 200 mm each year, although in a few cases it has reached 800 to 1000 mm due to local climatic variations. In the Atlantic facade, the climate is more humid and precipitation usually exceeds 1600 mm each year and monthly average temperatures oscillate between 25 and 27 °C. The Venezuelan Coast is influenced by the action of the trade winds from the east-northeast and northeast-southwest directions. In some coastal zones winds can reach very high speeds, such as in Laguna de Cocinetas, where speeds vary from 50 to 65 km/h, inducing processes of accelerated sedimentation (Conde & Alarcón 1993). It has been postulated that mangrove development and structural complexity are closely related to this climatic gradient (Pannier 1986).
In the continental domain of Venezuela, mangroves are confined to two geomorphologic units: coastal lagoons and alluvial plains. In turn, in the alluvial plains three categories can be distinguished: 1) those with sedimentary marine deposition, as is the case of Los Olivitos Swamp, Morrocoy Bay, and Gulf of Paria; 2) Deltaic swamps with marine-fluvial deposition, as in the San Juan River and the Gulf of Paria; 3) Swamp deltas with predominantly alluvial sedimentary processes, as in the Orinoco Delta. The characteristic deltaic plains of the oriental sector have induced a major diversity of landscapes, due to the interaction of microtopographic and pedological factors, mediated by local characteristics (Pannier 1986; Conde & Alarcón 1993).
Throughout their range, Venezuelan mangroves are associated with various types of vegetation, ranging from thorn scrub woodlands to coastal evergreen forests. These various associations host diverse faunal assemblages, which, in turn, might be a consequence of the variations of nearby vegetation and their proximity to other pristine ecosystems. Exhaustive lists of the species are meaningless, since they can vary substantially from forest to forest, however, mention of some species can help in sketching the importance of mangroves and their diversity. Some of the plant species, seem to be constantly associated with mangrove forests throughout their distribution range in the Neotropics. Among them, the fern (Acrostichum aureum) and the Malvaceae (Hibiscus tileaceus) are the most widespread. These species frequently form dense belts along the landward edge of mangroves, on more elevated sites and around dry and saline areas inside mangroves.
In the desert coastal areas of western Venezuela, mangrove forests are associated with xerophilous littoral scrubs, and halophilous and psammophilous littoral meadows. Xerophilous scrubs are characterized by individuals with heights from 0.5 to 5 m. Typical species are Acacia tortuosa, Bourreria cumanensis, Cercidium praecox, and Hibiscus tiliaceus. Columnar cactus can also be present. Halophilous meadows appear in brackish depressions of coastal areas, where Atriplex pentandra, Heterostachya ritteriana, and Sesuvium portulacastrum prevail. Psammophilous meadows settle on sandy dunes, normally not flooded by the sea and Egletes postrata, Ipomea pes-caprae, and Sporobolus virginicum are distinctive species (Conde & Alarcón 1993).
Mangrove forests located at the southwestern margin of Lake Maracaibo merge with tall (30-40 m) partially flooded ombrophilous evergreen forests, where Anacardium excelsum, Cariniana pyriformis, Ceiba pentandra, and Gustavia hexapetala predominate while the endemics Rhodospatha perezii and Spathiphyllum perezii are related to the Amazonian flora. Medina & Barboza (2000) have provided a detailed description of these mangroves and the associated vegetation.
Some mangrove stands are close to tropophilous semi-deciduous seasonal forests, characterized by one or two strata, and heights from 5 to 8 m, with emergent individuals up to 10-12 m. Representative species are Capparis coccolobifolia and C. tenuisiliqua, and the endemic Apoplanesia cryptantha. Mangroves also flourish near tropophilous basimontane deciduous forests, with heights from 10 to 15 m, and emergent trees up to 20 m. Bauhinia megalandra, Bourreria cumanensis, Calliandra caracasana, Erythrina poeppigiana, Hura crepitans, and Tabebuia billbergii are some of the more abundant species. In some localities, Capparis coccolobifolia, Diospyros inconstans, Jacquinia revoluta and Maytenus officinale predominate.
Since mangrove woodlands lie on the littoral zone, the mangrove fauna comprises elements from marine and terrestrial habitats. In effect, few species can be considered exclusive inhabitants of mangroves, although many are most commonly found associated with mangroves, and it is only in this sense that they can be called mangrove fauna (Lacerda et al. 2001). Most of the animals to be found in mangroves also appear elsewhere in other coastal ecosystems, and even in areas hundreds of kilometers from the coastal area, as in the case of the scarlet ibis (Eudocimus ruber) (Conde & Alarcón 1993), although some endemics have been recorded, such as the Trochilidae birds (Lepidopyga lilliae and Amazilia tzacatl) the latter being responsible for the pollination of Pelliciera in Colombia (Lacerda et al. 2001). The presence of a given species depends on a number of factors, such as rainfall patterns, tides and life cycle stage. Hence the difficulty of characterizing a "true" mangrove fauna. Whether transient or permanent, the mangrove fauna is large and diversified. Over 140 species of birds and 220 species of fish and hundreds of species of terrestrial and marine invertebrates can bring about high diversity assemblages along mudflats and other intertidal habitats.
From an ecological perspective, mangroves might be very important, since in barren areas that otherwise are deprived of any vegetation, such as coastal strips in desertic areas, mangrove woodlands can offer exceptional opportunities for resting, feeding, sheltering and nursing to faunal elements. Due to the accelerated destruction of inland forests, mangrove stands have become important sanctuaries and also could function as stepping-stones in the migratory routes of various species, that other wise would be confined to small vegetational patches and thus threatened to extinction. For instance, small populations of the vulnerable American crocodile, Crocodylus acutus, inhabit mangrove swamps, which have become their main remaining shelters in Venezuela.
The inventory of animals that live in or are transiently associated with Venezuelan mangroves is far from being over. Typical species are considered the yellow warbler (Dendroica petechia), the bicolored conebill (Conirostrum bicolor), the clapper rail (Rallus longirostris), the great-tailed grackle (Cassidix mexicanus), the spotted tody-flycatcher (Todirostrum maculatum), the rufous crab-hawk (Buteogallus aequinoctialis), the crab-eating raccoon (Procyon cancrivorus), the American crocodile (Crocodylus acutus) and the arboreal snake (Corallus hortulanus). Yet, some of these species have also been reported in other habitats; for instance, P. cancrivorus is common in savanna forests, far from mangrove stands, and D. petechia can be found in grasslands and rain forests. Some invertebrates appear closely related to mangroves and could be considered typical; for example, the mangrove tree crab (Aratus pisonii) and the crabs (Goniopsis cruentata and Ucides cordatus), the bivalve (Crassostrea rhizophorae) and many sponges.
The avifauna of Venezuelan mangroves has been reasonably inventoried through surveys in seven mangrove stands. A total of 141 species have been tallied, including resident, opportunistic and winter migrant species (MARNR, 1986). The highest number in Venezuela (80 species) was totaled in the mangroves of the Orinoco Delta of the Guianan mangroves ecoregion, while only 26 species have been recorded in the xeric mangroves of Laguna de Cocinetas. Only four of these species are shared by all these localities: the common egret (Casmerodius albus), the black vulture (Coragyps atratus), the brown pelican (Pelecanus occidentalis) and the scarlet ibis (Eudocimus ruber). The latter, an endangered species in several countries is quite common in Venezuelan mangroves. The Caribbean flamingo (Phoenicopterus ruber) inhabits many mangrove swamps of Venezuela, where it can reach high numbers. In 1990, 15,000 individuals were estimated in Los Olivitos Swamp, one of only four Caribbean localities where this bird nests. The mangrove-bordered Laguna de Tiraya (Paraguaná Peninsula, State of Falcón) is one of the feeding grounds for large flocks of flamingos, which nest on the islands north of Venezuela.
The endemic plain-flanked rail (EN) (Rallus wetmorei) is restricted to brackish lagoons and mangroves along a small stretch of this ecoregion on Venezuela's north coast (Stattersfield 1998). Other endangered or vulnerable birds that have been observed in mangroves of this ecoregion are the long-winged harrier (Circus buffoni), the dark-billed cuckoo (Coccizus melacoriphus), the boat-billed heron (Cochlearius cochlearius), the striped-backed bittern (Ixobrychus involucris), the masked duck (Oxyura dominica), the red-capped cardinal (Paroaria gularis migrogenis) and the osprey (Pandion haliaetus), a boreal winter visitor.
Herons, egrets, terns and gulls are very abundant in Venezuelan mangroves. Also, the magnificent frigatebird (Fregata magnificens), the roseate spoonbill (Ajaia ajaja), the anhinga (Anhinga anhinga), and the jabiru (Jabiru mycteria) are common. Winter visitors are comprised of the sandpipers (Calidris mauri and Micropalama himantopus), the blue-winged teal (Anas discors) and several Nearctic limicolaes. Non-aquatic species can be occasionally observed in mangroves including the orange-winged parrot (Amazona amazonica), the yellow-headed parrot (A. ochrocephala), as well as dense populations of macaws such as Ara chloroptera and A. severa.
The crab-eating raccoon (Procyon cancrivorus) is one of the most common terrestrial mammals sighted in these mangroves. Visitors from the surroundings ecoregions include the crab-eating fox (Cerdocyon thous) and the cottontail rabbit (Sylvilagus floridanus). The spotted paca (Agouti paca), the kinkajou (Potos flavus), and the Orinoco agouti (Dasyprocta guamara) are also common. Among larger mammals, the jaguar (Panthera onca), the South American tapir (Tapirus terrestris), the giant anteater (Myrmecophaga tridactyla), and the ocelot (Felis pardalis) have been sighted, although their abundance is unknown. The red howler monkey (Alouatta seniculus) and capuchin monkey (Cebus sp.) have been also observed. In the waterways, aquatic mammals include the West Indian manatee (Trichechus manatus), the river dolphin (Sotalia fluviatilis), the Amazon River dolphin (Ina geoffrensis) and the river otter (Lutra sp.). Several of these species are included on IUCN status protection lists.
The American crocodile (Crocodylus acutus) inhabits many mangrove swamps. In the San Juan River, the spectacled caiman (Caiman crocodylus), an endangered species, is common. Intruding marine species, such as the endangered Atlantic green turtle (Chelonia mydas), are also sighted.
Many commercial fishes, which sustain subsistence and artisan fisheries, are associated to Venezuelan mangroves, including catfishes (Arius herzbergii, Cathorops spixii); snooks (Centropomus undecimalis, C. ensiferus); mullets (Mugil curema, M. liza); and mojarras (Diapterus plumieri, D. rhombeus, Gerres cinereus). Crustaceans, such as shrimp Penaeus spp. and swimming crab Callinectes spp., can also be very important in mangrove-based fisheries for humans and as a base for the food chain of mangrove ecosystems.
In some localities of the Venezuelan coast where waters are crystal-clear, a variegate community of sponges (33 species), tunicates (12), bivalves, and algae, can be found adhered to the submerged roots of R. mangle (Sutherland 1980; Díaz et al. 1985, 1992; Orihuela et al. 1991). The mangrove oyster (Crassostrea rhizophorae) used to be one of the most common species in this community and a high-ranked staple for subsistence fishermen, but nowadays is almost totally extinct due to overexploitation (Rodríguez & Rojas-Suárez 1995). This fragile ecosystem can be wiped out due to the re-suspension of sediments (Orihuela et al. 1991) or a combination of low temperatures and simultaneous plunges of salinity (Laboy et al. 2001).
Although mangrove woodlands have not been systematically monitored, some evidences suggest dramatic losses during the last decades. In the Vegetation Map of Venezuela, published in 1960 (Hueck 1960), mangroves are reported for most of Lake Maracaibo perimeter, whereas their present distribution is more restricted. The current cover of mangroves in this area has been estimated in 1/10 (Conde & Alarcón 1993) of that reported by Hueck (1960). In other ecoregions the reduction is also remarkable. In 1926, Henri Pittier described several mangrove forests where nowadays only small patches are left. Similarly, in Adícora (State of Falcón), Cumaná (Sucre), and at Píritu and Unare (Anzoategui), as well as along the coast of the State of Carabobo, where historical records afford evidence of extensive mangrove forests (Pittier 1926; Esteves 1980), they have disappeared almost entirely or only small patches are left. The causes of such deforestation is unknown; however, it is probable that in some of those areas semi-industrial exploitation was carried out, or that small-scale extraction levels, to meet local needs, were constant for decades. However, in a few localities, an inverse process has been observed, such as in Caño Mánamo (Delta Amacuro), where mangroves have expanded at a rate of 6 - 7 ha/year since 1965, when a cofferdam was built, leading to the salinization of the formerly limnetic waters (Colonnello & Medina 1998).
Mangroves have been specifically protected since 1974 when a Presidential Decree was passed (Decreto Presidencial N° 110; Gaceta Oficial de la República de Venezuela N° 30.408). This decree banned the direct destruction of mangroves and those activities that could threaten them, including discharging of wasted waters, dredging of marine bottoms and dumping. However, since 1991 a new Presidential Decree allows the option of intervention in those coastal localities regarded as economically depressed, to allow some destructive activities —that is, almost the entire Venezuelan coast (Decreto Presidencial N° 1843; Gaceta Oficial de la República de Venezuela N° 34.819). In this second decree the possibility of intervention of mangroves, with the administrative authorization of the MARNR as the only requisite, is open and the need for an Environmental Impact Assessment (EIA) is discretionary.
However, since Venezuela has an extensive system of protected areas, which embraces nearly 1/3 of its territory, many mangrove stands are within the limits of conservation units, which in some cases have the rank of national parks and natural monuments (Conde & Alarcón 1993). Among them, it is worthwhile mentioning Parque Nacional Morrocoy, P. N. Mochima, P. N. Laguna de Tacarigua, and P. N. Archipiélago de Los Roques.
Types and Severity of Threats
There is a protracted story of use and abuse of Venezuelan mangroves that at least dates back to the pre-Columbian times when mangroves where extracted extensively by the indigenous dwellers in Los Roques Archipelago, 100 km north of the Venezuelan mainland (Amend 1992; Conde & Alarcón 1993). The exploitation levels were so intense that mangroves disappeared from some islands. During the XIX Century's second half, the Curaçoleans also exploited Los Roques mangroves, mostly R. mangle, to be used as charcoal for the steamship's furnaces, and tannins from the bark. Charcoal industries were installed in several of these desolate islands (Amend 1992; Conde & Alarcón 1993). Recent examples of exploitation worth mentioning follow. Including the extraction of mangrove trees for use as lumber in the copper mines of Aroa (State of Falcón) by the English company Bolivar Mining Association in 1930. As well as heavy mangrove extraction at the southern zone of the Lake Maracaibo (State of Zulia), for exportation to Germany and the USA prior to the Second World War. Also deforestation occurred during the 50’s, due to the expansion of coconut plantations and oil exploitation, which affected the Lake Maracaibo mangroves. Finally cutting of mangroves, for production of firewood and charcoal, in the Limón River.
Cutting, farming and oil spills have provoked important mortality events in some mangrove forests in the same area of the ecoregion. Currently, threats vary from one mangrove forest to another, but, in general, the main impacts are furtive deforestation and more recently habitat conversion due to urban, tourist and demographic growth, a common trend in countries in the Caribbean basin (Ellison & Farnsworth 1996). Other activities, which could put extra pressure on mangroves, are the creation and expansion of saltworks and aquaculture ponds, and oil spills in the Orinoco Delta, where intense oil prospecting activity has been mounting during recent years, and although not in this ecoregion is not far along the coast from the Grotto de Paria (Conde & Alarcón 1993; Conde 2001). The establishment of shrimp farms to meet the consumer demand for luxury shrimp has been a driving factor of massive mangrove losses in several Latin American countries such as Ecuador but not in Venezuela where this industry is circumscribed to a few places.
In Laguna de Tacarigua, an estuarine coastal lagoon bordered by mangroves, sustained massive deforestation occurred during 1920, 1927-1931, and 1953-1957. Other substantial impacts are the detour of the Guapo River’s original course in 1964, that has brought about an increase in the erosion and sediment transported towards the lagoon, provoking progradations and, it is presumed, a decrease in fish catches. A compendium of the impacts on this lagoon includes accumulation of tannins due to changes in the hydrodynamic patterns; dredging with consequent modification of tidal intrusion patterns; furtive exploitation of crocodiles and turtles; and overfishing with illegal fishing gear. More recently, resort development has put new pressures on the mangroves. Exclusion of the western bar of the legal protection regime; probable use of defoliants; and the alleged scarce interest of governmental entities to enforce the pertinent laws must be added to the list. The pressures should be subdued, due to its current National Park status however, during the 80's in the neighboring areas, several resorts were built (Díaz & Zelwer 1985).
Justification of Ecoregion Delineation
Classification and linework for all mangrove ecoregions in Latin America and the Caribbean follow the results of a mangrove ecoregion workshop (1994) and subsequent report (Olson et al. 1996).
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Prepared by: Jesús Eloy Conde (Centro de Ecología, Instituto Venezolano de Investigaciones Científicas) & Clara Alarcón (PDVSA Intevep)
Reviewed by: Christine Burdette