Physical geography
The region under consideration consists of two semi-enclosed basinsâthe Caribbean Sea and the Gulf of Mexicoâas well as the westernmost embayment of the Atlantic Ocean. Bounded to the west and south by Central and South America, the Caribbean is bounded to the north by the Greater Antilles (Cuba to Puerto Rico) and to the east by the great arc of small islands called the Lesser Antilles.
Figure 1 The many Exclusive Economic Zone (EEZ) and Fishery Zone (FZ) claims in the Caribbean region complicate regional conservation schemes.
Most of the Caribbean Sea is centered on the Caribbean Plate, a relatively small tectonic plate (section of Earthâs crust) that formed between 75 and 90 million years ago. The movements of this plate form subduction zones to the east and west (where one plate slides beneath another) and transform boundaries (where two tectonic plates slide past each other) to the north and south. These tectonic interactions have created the island chains of the Greater and Lesser Antilles as well as the mountainous perimeters across Central America and the southern Caribbean Sea. Most of the remainder of the region lies on the North American Plate (Pindell 1994, Meschede & Frisch 1998).
The Caribbean Sea is itself divided into four smaller basins separated by shallower ridges, while the Gulf of Mexico can be considered a separate fifth deep basin. Deep trenches are found to the north of Puerto Rico and to the south of Cuba and the Cayman Islands. The basins, as well as the deep trenches, are isolated from each other by shallow underwater sills and, although there is some water exchange, each may contain unique and important features (Tomczak & Godfrey 1994, Smith, Carpenter & Waller 2002). With the exception of the Bahamas Banks, the only extensive shallow areas are on the continental shelves, notably around the southern U.S. coast, the northern coast of the Yucatan Peninsula and the Nicaraguan Rise.
Surface water movement is driven by the North Equatorial Current and the North Brazil (or Guiana) Current. These combine to form the Antilles Current, which sweeps up the outer edge of the Lesser Antilles and the eastern side of the Bahamas. There is also considerable flow between the Lesser Antilles islands, and to a lesser degree through the channels of the eastern Greater Antilles, creating a broad westward-flowing Caribbean Current across the Caribbean Sea. This current is weak and there are surface eddies creating a relatively complex pattern of surface flow. Further west, the water flow is concentrated through the Yucatan Channel and becomes much stronger. Once in the Gulf of Mexico, this current flows northwest, then in a broad clockwise circle âthe Loop Currentâwhich flows back southward along Floridaâs west coast, shedding eddies westward into the Gulf before leaving the system east of Florida. The Florida Current, enhanced by flow from the Antilles Current, forms a broad stream of warm waterâthe Gulf Streamâwhich flows northeast across the Atlantic (Tomczak & Godfrey 1994, Longhurst 1998).
Rivers have a considerable influence in the southeast Caribbean, where outflows of the Amazon and Orinoco Rivers create slightly lower salinities and more suspended sediments over wide areas (Tomczak & Godfrey 1994).
Deep water flows into the Caribbean Sea through a limited number of deep passes: through the Jungfern Passage east of Puerto Rico into the Venezuela and Colombian Basins, through the Windward Passage east of Cuba into the Cayman and Yucatan Basins and through a number of eastern passages into the Aves and Grenada Basins. The deeper water areas have remarkably consistent temperatures. Little is known about the turnover of deep water in the Caribbean basins, but most estimates suggest periods of hundreds of years (Tomczak & Godfrey 1994, Tyler 2003).
Biodiversity
The Caribbean is perhaps best known for its tropical shallow marine ecosystems. Its coral reefs cover about 20,000 square kilometers (Spalding, Ravilious & Green 2001). This is only about 7% of the worldâs shallow reefs, but they are of enormous biological and human importance. Biologically they are unique, with very little overlap in species with other parts of the worldâexcept for Brazil and to a small degree West Africa, both of which have related assemblages but fewer species (CortĂŠs 2003). The Caribbean is also a center of diversity and endemism for some mangrove and seagrass species, although a higher proportion of these extends into the Eastern Pacific and across to West Africa. The Caribbean has roughly 22,000 square kilometers of mangrove (amended from Spalding, Blasco & Field 1997), which is about 12% of the global total. No accurate estimate of the extent of seagrass area is available, although the Caribbean is likely to be a highly significant region for these habitats (Onuf et al 2003, Creed, Phillips & Van Tussenbroek 2003). Unpublished estimates from work at the United Nations Environment Programme (UNEP) World Conservation Monitoring Centre suggest as much as 33,000 square kilometers of Caribbean seagrass beds (Spalding et al 2003), which would be about 18 to 19% of the global total.
An explanation for this diversity and endemism in the Caribbean is at least partly linked to the geological history (Table 1) of the region. During the Triassic, about 200 million years ago, all of the worldâs land areas were connected as one supercontinent known as Pangea. Pangea was then divided into northern and southern continents by seafloor spreading, and the growing gap between them was filled by a globe-encircling equatorial sea known as the Tethys (Wikipedia 2004). Without land masses to disrupt biological migration and genetic flow, much of the subtropical and tropical marine regions of the world shared a similar array of animals, referred to as a pan-Tethyan fauna, until the Miocene (Veron 1995, Saenger & Luker 1997). Since that time, the open-sea connection between the Caribbean and the Pacific became increasingly restricted as the Central American land bridge rose above sealevel, which finally separated the two bodies of water about three million years ago (GuzmĂĄn 2003). For the corals, and likely for many other species, considerable changes were then caused by the Pliocene/Pleistocene glaciation periods, including the extinction of many marine species. This was followed by significant biological âradiation eventsâ (the rapid evolution of new species to fill new habitats) in isolation from the rest of the worldâs tropical seas (Rosen 1984, Veron 1995), which led to the present levels of Caribbean coral diversity and endemism.
Table 1
The geologic time scale. Epochs are shown only for the more recent Cenozoic Periods. Many of the Era and Period boundaries are based on major episodes of extinction or other biological factors.
| Period | Epoch | Years ago |
| Cenozoic Era | | |
| | 11 thousand |
| Pleistocene (glacial) | 2 million |
| Tertiary | Pliocene | 7 million |
| Quaternary Holocene | Miocene | 23 million |
| Oligocene | 38 million |
| Eocene | 53 million |
| Paleocene 65 | million |
| Mesozoic Era | | |
| Cretaceous | | 135 million |
| Jurassic | Cretaceous | 135 million 195 million |
| Triassic | | 230 million |
| Paleozoic Era | | |
| Permian | | 280 million |
| Pennsylvanian (Upper Carboniferous) | 310 million |
| Mississippian (Lower Carboniferous) | 345 million |
| Devonian | | 395 million |
| Silurian | | 435 million |
| Ordovician | | 500 million |
| Cambrian | | 600 million |
| Precambrian Era | | 600 million to 4 billion |
Source: Modified from the American Heritage Dictionary 2000.
The boundaries and even the definitions of marine habitats other than coral reefs in the Caribbean remain poor. âProxyâ measures (scientifically assumed from the extension of known data) can be developed from knowledge of physical features such as surface circulation patterns, surface chlorophyll distributions or bathymetry. The location of upwelling areasâwhere deep, cold, nutrient-rich waters come to the surfaceâis extremely important for biodiversity and ocean productivity. Some sites are documented, such as the Yucatan Shelf edge (Merino 1997), the eastern coast of Venezuela and parts of the Gulf of Mexico (Longhurst 1998), but it may be possible to improve our understanding of the location, timing and frequency of these and other upwellings by looking at patterns of surface chlorophyll.
Dividing the Caribbean region into zones based on bathymetry (water depth) indicates that about 10% of the seafloor falls within areas less than 30 meters deepâwhere we find the highest light penetration and primary productivityâthe transfer of sunlight into food through photosynthesis. Waters on the continental shelf edges (to 200 meters depth) cover a similar area. Waters between 2,000 and 5,000 meter depths dominate the region (Figure 2). The basins and deep trenches are clearly separated below 4,000 meters depth, however. Existing studies suggest that levels of biodiversity below the continental shelf edge may be very high and that, even in the deep trenches where relatively fewer types of organisms exist, endemism is a key feature, with important examples of species unchanged since ancient times (Zezina 1997). The few studies undertaken in the Puerto Rico Trench suggest that it has lower than expected diversity, however (Vinogradova 1997, Tyler 2003).
Some of the most important features of the deep seafloor include rocky outcrops with no sediment cover, deep shelf bioherms (limestone rock structures derived from the shells of living organism) and hydrothermal vent areas where hot minerals spew up from cracks in the seafloor. There has been no systematic attempt to map these, but it might be possible to produce maps through a combined literature survey and expert interpretation of feat...