AN ANNOTATED LIST of the Troglobltlc Anchlallne and FRESHWATER FAUNA of QUINTANA ROO
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AN ANNOTATED LIST OF THE TROGLOBlTlC ANCHlALlNE AND FRESHWATER FAUNA OF QUINTANA ROO - - Por Thomas M. lliffe In: Navarro, D. y E. Suarez-Morales (Eds.). Diversidad Biologica en la Reserva de la Biosfera de Sian Ka'an Quintana, Roo, Mexico. Vol. 11 CIQROISEDESOL (1 992) AN ANNOTATED LIST OF THE TROGLOBITIC ANCHIALINE AND FRESHWATER FAUNA OF QUINTANA ROO Thomas M. Iliffe Department of Marine Biology, Texas A & M University at Galveston, Galveston, Texas 775-53-1675, U.S.A. Twenty species of aquatic, troglobitic contain freshwater near the surface. (cave-limited) organisms are presently However, beneath an abrupt halocline, known from cenotes and caves in the the salinity increases to brackish or state of Quintana Roo, Mexico. These fully marine levels (14 to 35 ppt). include 18 species of crustaceans from Average salinity beneath the halocline 7 orders and 2 fish species from two in mainland caves is about 16 ppt orders. Twelve species inhabit (Coke, pers. comm.). The depth of the anchialine (coastal, brackish water) halocline and thus the thickness of the caves, while the remainder are limited freshwater lens increases with distance to subterranean freshwaters. Nearly all inland. Most of these caves are formed are of a marine origin. by preferential dissolution of limestone The word "cenote" is derived from at the mixing zone halocline between a Mayan term and denotes a fresh and brackish waters (Back et al., subterranean chamber that contains 1986). permanent water. Ha11 (1936) and Due to characteristic cave others have categorized cenotes as development at the halocline and the follows: (1) Jug-shaped cenotes have a orientation of cave passages along frac- small surface opening. (2) Vertical- tures, many anchialine caves extend for walled cenotes have a large surface considerable distances. With the opening. (3) Aguada-like cenotes development of advanced cave diving consist of large cavities with sloping methodology, it has become possible walls leading down to shallow pools. to explore and study these vast "Aguadas" are seasonal pools only underwater labyrinths. One of containing water during the wet season. Mexico's largest underwater cave (4) Cave-like cenotes contain a lateral systems is the Sistema Naranjal (the passage descending to a chamber Maya Blue-Najaron System) near containing water. Often, these pools are Tulum with 15,480 m of surveyed sumps with cave passages continuing passage (Coke, pers. comm.). Other underwater. Many cenotes and caves long underwater caves on the mainland near the Caribbean coast of Quintana include Nohoch Nah Chich (13,380 m), Roo contain anchialine pools. The term Cenote Sac ActGn (4,500 m), Cenote "anchialine" was coined by Holthuis Zapote (4,500 m), Cenote Ponderosa (1973) to denote, "pools with no (4,500 m) and Carwash Cenote (2,590 surface connection to the sea, m). Cueva Quebrada (7,600 m) and containing salt or brackish water, which Aerolito (6,100 m) are long anchialine fluctuates with the tides". Anchialine caves on the island of Cozumel. caves in Quintana Roo typically The limestone that makes up the TROGLOBITIC FAUNA: Thomas M. Iliffe Yucatan Peninsula ranges in age from document the rich anchialine fauna Miocene and Eocene in the interior to inhabiting coastal caves. The remipede Pleistocene and Holocene deposits near Speleonectes tulumensis, the osnacod the coast. Surface rocks along the east Danielopolina mexicana, the coast of Quintana Roo are composed thermosbaenacean Tulumella unidens of Pleistocene coral reef material dated and the amphipod Tuluweckelia cernua at 120,000 years (Back et al., 1986). are known only from anchialine caves Ha11 (1936) has examined the on the mainland near Tulum. The physical and chemical characteristics shrimp Agostocaris bozanici, of the more inland cenotes of Yucatan. Yagerocaris cozumel, Somersiella He found that water temperatures sterreri and Janicea anfiguensis have ranged from 21.9 to 28.5 "C with a mexican populations limited to caves mean of 25.4 "C. Three cenotes had in the island of Cozumel. Only the marked vertical gradients with warmer cirolanid isopod Bahalana rnayana has water at the surface an cooler water (by so far been obtained from both as much as 5 "C) at depth. Water in locations. several cenotes examined by Hall Reddell (1977) examined became anoxic and contained hydrogen distribution patterns of the troglobitic sulfide at depth, although the surface fauna in Yucatan. Four aquatic species waters were near saturation. Other -the mysid Antromysis cenotensis, the cenotes had relatively uniform oxygen shrimp Typhlafya mitchelli and the two concentrations at all depths. blind fish, Ogilbia pearsei and The first serious attempts to Ophisternon infernale- are widespread investigate the cave fauna of the over the Coastal Plain. The cirolanid Yucatan Peninsula were carried out in isopod, Creaseriella anops, is known 1932 and '36 by expeditions from the from both the Coastal Plain and Sierra Carnegie Institution of Washington de Ticul, but has not been found in the (Pearse, 1936, 1938 a). A total of 306 Sierra de Bolonchtn. The amphipod species was collected from aquatic and Mayaweckelia cenoticola occurs in terrestrial caves during this study of both the Coastal Plain and the Sierra which 28 were considered as de Bolonchtn. Two shrimp, Typhlatya troglobites (Pearse, 1938 b). pearsei and Creaseria morleyi, are Between 1973 and '75, systematic found in the Coastal Plain, Sierra de studies of the cave fauna of entire Ticul and Sierra de BolonchCn. An Yucatan Peninsula were carried out by asellid isopod, Caecidotea sp., is biologists from Texas Tech University known only from the Northwestern (Reddell, 1977). These investigations Coastal Plain. Similarly, the amphipod increased the total number of species Mayaweckelia yucatanensis, and the reported from caves to 565 of which shrimp Typhlatya campecheae, are 11 were aquatic troglobites and 23 restricted to the Sierra de BolonchCn. terrestrial. Reddell (1977) concludes that at this Cave diving investigations starting time it is not possible to speculate in the mid 1980's have just begun to whether these distribution patterns TROGLOBITIC FAUNA: Thomas M. Iliffe 199 result from, "different times of 1989). Four genera (Janicea, Procaris, invasion, a greater degree of mobility Typhlatya and Somersiella) occur in of the species, or a slower evolutionary both Yucatan and Bermuda, while three rate". genera (Danielopclina, Typhlatya and Wilkens (1982) plotted the Ogilbia) cohabit Yucatan and dismbution of the aquatic troglobites Galapagos caves. in relation to the position of Pliocene The following list describes the and Pleistocene shorelines in Yucatan. known aquatic troglobites inhabiting He supposed that the transition of caves in Quintana Roo. Locations of marine species to freshwater caves in these caves are shown in Figure 1. Yucatan was dependent on the existence of coastal (anchialine) caves. Wilkens believed that the more stable Pliocene shoreline was the primary site Phylum CRUSTACEA of cave colonization. Indeed, most ca- Class Remipedia ves containing the cave fish Ogilbia Order Nectiopoda pearsei and Ophisternon infernale are Family Speleonectidae located near the ancient Pliocene coast. Speleonectes tulumensis Yager, 1987 The similar degree of eye and pigment Characteristic features: Elongate, reduction in distant populations of these slender remipede without pigment or fish, as well as in the shrimp Creaseria eyes. Small cephalic shield; trunk morleyi, suggested to Wilkens, a segments increasing with age to a simultaneous start for cavernicole maximum observed number of 36 evolution of each species. Since the eye (Yager, 1987). rudiments of Ophisternon infernale are Size: To 27.5 mm. less reduced than those of Ogilbia Habitat: Anchialine. pearsei and Creaseria morleyi, this Number of species in genus: Four, all former species probably had a later anchialine. beginning of regressive evolution. Genus range: Bahamas (Grand Wilkens interprets the present Bahamas and Abaco Island), Mexico widespread distribution of many (Quintana Roo), Canary Islands, Belize troglobites as a result of secondary (Yager, pers. comm.). dispersal through the subterranean Species range: Known from anchialine water system. caves in Belize and Quintana Roo. The aquatic troglobitic fauna of the Collection sites in Quintana Roo Yucatan Peninsula has taxonomic include Carwash and Najaron Cenotes. affinities to species inhabiting caves on Ecological notes: Schram (1986:40) various oceanic islands. At least 8 ge- has noted that remipedes in general, nera (Agostocaris, Bahadzia, ... live below a distinct halocline in Bahalana, Danielopolina, Janicea, brackish layers of waters generally Typhlatya, Tulumella and deep within the caves ...oxygen in the Speleonectes) are common to caves in remipede habitat is very low, around the Yucatan and Bahamas (Holsinger, 0.5 parts per billion-virtually anoxic. 200 TROGLOBITIC FAUNA: Thomas M. Iliffe Yet the animals are moderately active, Class Ostracoda good swimmers. When collected and Order Halocyprida maintained in aquaria, the animals take Suborder Halocypridina to ceaseless, rather frenetic swimming Superfamily Thaumatocypridoidea and literally burn themselves out with Family Thaumatocyprididae a few days. Danielopolina mexicana Kornicker & Swimming at any speed is achieved Iliffe, 1989 with regular metachronal beats. Characteristic features: Halocyprid The robust, prehensile to subchelate ostracod with spines on carapace. mouthparts would seem to imply a Size: