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By Tom Ilifje, Phd BIOLOGY OF UNDERWATER CAVES by Tom IlifJe, PhD Recent speleological investigations in coastal areas have confirmed that anthropogenic pollutants can have potentially devastating effects on the fresh groundwater lenses and adjacent open water marine ecosystems including coral reefs, mangroves and sea grass. Many species ofcave animals are listed as critically endangered. These animals are a barometer ofthe health ofthe environment and virtual missing links that help to explain the evolution oflife in the sea and the evolution oflife on earth. The subterranean aquatic environment consists of known and studied from freshwater caves, the discov­ interstitial (small, water-filled spaces between uncon­ ery of similar animals from marine caves is a recent solidated sediments) and cave (larger voids within event brought to light by cave diving explorations. bedrock, either formed by solution or volcanic action) Anchialine (tidal, water-filled voids near the coast) ecosystems. The water in underground environments caves typically possess a highly stratified water col­ can vary from completely fresh to fully marine sa­ umn with deeper, fully marine waters separated by linities. Such habitats are characteristically lightless, a halocline from overlying fresh or brackish water. environmentally stable and have limited input offood Cave diving technology has been an essential tool to due to the absence ofphotosynthetic plants and barri­ explore and study the deeper waters in such systems. ers to external input. Anchialine caves include the cenotes of Mexico's Ecologically, aquatic cave animals can be subdi­ Yucatan Peninsula, the blue holes ofthe Bahamas and vided into stygobites (cave-adapted species restricted Belize, as well as numerous limestone and volcanic to subterranean waters), stygophiles (species inhabit­ caves, mostly on islands, around the Caribbean, Med­ ing caves and completing their entire life cycle there, iterranean and Indo-Pacific. Seven of the ten longest obut which also occur in similar open water habitats) underwater caves in the world are anchialine caves stygoxenes (species common in caves, but which from the Caribbean coast ofYucatan. This anchialine must leave the cave to feed or reproduce), and acci­ cave habitat is characterized by the absence oflight, a dentals (species that wander or are washed into caves, salinity and temperature stratified water column, very but which cannot survive there for very long). The limited food resources, low levels of dissolved oxy­ prefix "stygo" refers to the subterranean River Styx gen and stable environmental conditions. which from Greek mythology circles through Hades Anchialine and freshwater stygobites are mostly or the underworld. Thus, stygobites are literally the crustaceans, and include several higher groups of "aquatic cave life", stygophiles the "aquatic cave lov­ crustaceans found only or primarily in subterranean ers", and stygoxenes the "aquatic cave guest~". habitats. Such animals include: , • Remipedes - primitive, anchialine, "living fos­ sil" crustaceans with highly segmented bodies, remi­ niscent of the segmented worms from which crusta­ .' ceans are thought to have evolved. Remipedes have paired hollow fangs for capturing prey and are among the top predators in this habitat. They are up to 4.5 cm in length, usually colorless and blind, with elongate, centipede-like bodies. Seventeen species ofremipedes inhabit fully marine, oxygen-deficient waters in caves Mictocaris halope, Bermuda. Photo: Tom llifJe in the Bahamas, Caicos Islands, Cuba, Yucatan Pen­ Although cave-adapted animals have long been insula, Canary Islands and Western Australia. 63 • Thermosbaenaceans - small (4 mm or less), • SpeJaeogriphaceans - small (less than 1 cm eyeless or eye-reduced, anchialine and freshwater long) freshwater crustaceans represented by spe­ crustaceans with a dorsal brood pouch in females. cies from caves in South Africa, Brazil and Western Their wide distribution in caves and thermal springs Australia. They are most closely related to the micta­ around the Mediterranean and Caribbean, as well as ceans. This widely separated distribution implies an Australia and Cambodia, suggests an origin along the early origin for the group, at least 200 million years coastline of the Tethys Sea, a shallow sea separating before present in the Tethys Sea. the continents during the early Mesozoic some 200 • Bochusaceans - small (1.2-1.6 mm), semi-trans­ million years ago. They include at least 34 species. parent and eyeless crustaceans that include two an­ chialine species from the Bahamas and Cayman Is­ lands and three deep-sea species. • Copepods - a large and diverse group, compris­ ing the most common animals in marine plankton. Platycopioid, misophrioid, cyclopoid, harapacticoid and epacteriscid calanoid copepods inhabit anchia­ line caves in tropical regions aroung the globe. They are small (typically 1-2 mm long) and have a short, cylindrical body with head and thorax fused Thermosbaenacean. Photo: Tom lliffe into a cephalothorax. Most are planktonic filter-feed­ • Mictaceans - small (3-3.5 mm), eyeless and de­ ers, but some such as the harpacticoids and cyclopoids pigmented, non-predatory crustaceans represented by are benthic. a single species in anchialine caves in Bermuda. 64 • Ostracods - small (approximately 0.5-2 mm), Caribbean, Europe, Galapagos, India, Indonesia, Ja­ benthic or planktonic bivalve crustaceans. Halocyprid pan, Malaysia, NOI1h and Central America and Poly­ ostracods include anchialine species with a distribu­ nesia. Cirolanid isopods have been found in freshwa­ tion and co-occurrence similar to that of remipedes. ter and anchialine caves clustered in Mexico and the More than 300 species of podocopid ostracods have Caribbean as well as in Europe and the Mediterra­ been found in springs, caves and anchialine habitats. nean. • Mysids - small (approximately 3-20 mm), • Amphipods - laterally compressed body (flat­ shrimp-like crustaceans including stygobitic species tened from side to side), occurring in freshwater and found in freshwater and anchialine habitats in Africa, marine habitats. Stygobitic representatives are present the Caribbean, Mediterranean, and India. Their distri­ in the bogidiellid, crangonyctid, hadziid and niphar­ bution suggests that they were stranded in caves by gid families of the amphipod suborder Gammaridea. lowering of sea levels in the Tethys and Mediterra­ They are very widely dispersed with large numbers nean. inhabiting caves in Central and Southern Europe, the Mediterranean, eastern and southern North America, and the Caribbean. • Decapods - possess five pairs of pereiopods or legs (hence the name Decapoda). Anomuran crabs (e.g., hermit, porcelain, mole and sand crabs) inhabit a freshwater cave in Brazil and an anchialine lava tube in the Canary Islands. Stygobitic crayfish are pres­ ent in caves in North America and Cuba. Brachyuran crabs (i.e., true crabs) are widely distributed from Stygiomysid. Photo: Tom Iliffe caves in the tropics and subtropics. Caridean shrimp include freshwater and anchialine representatives • Isopods - dorsoventral1y compressed body (flat­ from caves mostly in tropical latitudes. tened from top to bottom); occuring in terrestrial, freshwater and marine habitats. Stygobitic isopods • Fish - nearly 100 species of stygobitic fish are range from several millimeters to several centimeters presently described, primarily from tropical and sub­ in length. tropical regions. They occur principally in freshwater caves, but anchialine fish are present in Caribbean is­ lands and Yucatan. The principle stygobitic fish are cyprinids (carps and minnows), balitorids (river 10­ aches), and siluriform fishes (catfish). Six species of amblyopsid cave fish occur in the southern and east­ ern United States. Isopod. Photo: Tom llifJe Cavefish - Ogilbia persei. Photo: Tom llifJe Anthurid isopods occur in anchialine and freshwa­ ter caves in the Canary Islands, Caribbean and Indian • SaJamanders - Ten species of stygobitic sala­ Ocean islands, Mexico and South America. Asel10t manders are known, including Proteus, the 25-30 cm isopods inhabit anchialine and freshwater caves in the long world-famous blind salamander from caves near the Adriatic Sea, and nine species from the US (Tex- 65 as, southwestern Ozark Plateau, Tennessee, Alabama, subsequently were dispersed by plate tectonic rafting Georgia and Florida). as the continents separated and moved to their present Most ofthese animals have lost their eyes and pig­ positions. A number ofother anchialine animals show mentation in response to life in the constant and total close relationships with present deep-sea species im­ darkness of the cave. A number of less-obvious evo­ plying a possible deep-sea origin for cave fauna. Fi­ lutionary modifications have also occurred. In com­ nally, some cave animals are thought to have been parison to their surface relatives, cave animals tend to 'stranded' in their present locations by receding wa­ be longer-lived, produce fewer but larger eggs, have ters of the Tethys Sea. lower rates of metabolism, possess more abundant Unfortunately, many of these unique and fascinat­ tactile and chemo-receptors, have longer antennae ing animals are threatened with extinction due to the and long, thin, body forms. actions of man. In Bermuda alone, 25 species ofcave animals are internationally recognized as "critically endangered." This is the highest level of threat and roughly equates to a 50% chance of the species go­ ing extinct if nothing is
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