sign in sign up
<<
Home , Blindness in animals, Cave, Cenote, Mafia Island, Procaris, Speleology

AN ANNOTATED LIST OF THE TROGLOBlTlC ANCHlALlNE AND FRESHWATER OF

- - 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, . Vol. 11 CIQROISEDESOL (1 992) AN ANNOTATED LIST OF THE TROGLOBITIC ANCHIALINE AND FRESHWATER FAUNA OF QUINTANA ROO

Thomas M. Iliffe

Department of , Texas A & M University at Galveston, Galveston, Texas 775-53-1675, U.S.A.

Twenty species of aquatic, troglobitic contain freshwater near the surface. (-limited) organisms are presently However, beneath an abrupt , known from and in the the increases to brackish or state of Quintana Roo, Mexico. These fully marine levels (14 to 35 ppt). include 18 species of from Average salinity beneath the halocline 7 orders and 2 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 increases with distance to subterranean freshwaters. Nearly all inland. Most of these caves are formed are of a marine origin. by preferential dissolution of The word "" 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 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 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 ". Anchialine caves on the island of . 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 and 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 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 Agostocaris bozanici, of the more inland cenotes of Yucatan. Yagerocaris cozumel, Somersiella He found that water 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 shrimp Typhlafya mitchelli and the two at all depths. , Ogilbia pearsei and The first serious attempts to 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 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 and Characteristic features: Elongate, reduction in distant populations of these slender remipede without pigment or fish, as as in the shrimp Creaseria . Small cephalic shield; trunk morleyi, suggested to Wilkens, a segments increasing with age to a simultaneous start for cavernicole maximum observed number of 36 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 are moderately active, Class Ostracoda good swimmers. When collected and Order maintained in aquaria, the animals take Suborder 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 with spines on carapace. mouthparts would seem to imply a Size: Length and height of carapace: carnivorous mode of feeding. 0.95 mm by 0.74 mm (A-l? female). Indeed ...speleonectids (have been Habitat: Anchialine observed to feed) on Typhlarya garciai, Number of species in genus: Seven a caridean commonly associated with species, six of which are troglobitic and West Indian nectiopodans. The prey one from the deep sea. was grasped in the flexed mouthparts Genus range: , Bahamas, Jamai- and pressed tightly to the mouth. When ca (Kornicker & Iliffe, in press), feeding was completed, an empty Mexico (Quintana Roo), Canary cuticle was set afloat. Islands, Galapagos- all anchialine; Life history: According to Schram South Atlantic near the Equator at a (1986: 40), nothing is currently known depth of 3459 m. concerning nectiopodan breeding Species range: Known only from Maya habits nor details of development. Blue Cenote, located near Tulum, Several of the known species have been Quintana Roo. found in association with juveniles. In Ecology: Two specimens were general form they resemble the adults, collected from euhaline waters below but they are smaller, lack gut the halocline at 16 m depth. diverticula and gonopores, and only Life history: Not known. All instars have from 12 to 17 segments. are swimmers, probably living close to Closest related species: S. lucayensis or in the substrate (Kornicker, pers. from the Bahama Islands. comm.). Evolutionary origins: Remipedes are Closest related species: D. rnexicana the most primitive of known differs from its congeners by having a crustaceans. The combination of a large carapace with abundant surface spines number of primitive characters in the (Kornicker & Iliffe, 1989).D. mexica- animals is reminiscent of the ancestral na shares several characters in common . The nearest relative to with D. carolinae and may be more living remipedes is a closely related to this deep sea species species, Tesnusocaris goldichi, known than are any of the other cave species only from the record (Schram, (Kornicker, pers. comm.). D. mexica- 1983, 1986; Yager, 1986). na also shows relationships with D. sox (from the Galapagos) and with D. elizabethae (from Jamaica). TROGLOBITIC FAUNA: Thomas M. Iliffe 201

Danielopol (1990) concluded that D. Undetermined thermosbaenaceans have mexicana is more primitive than other been reported from Cueva Quebrada on congeners mainly because of the Cozumel (Bowman, 1987). presence of a Bellonci organ. Ecology: Three specimens were Evolutionary origins: Of the 5 genera collected from near the halocline at - comprising the family 10 to -18 m depth. above and Thaumatocyprididae, 2 are known only below the halocline at -14 m were 1.5 from (from the and and 35 ppt, respectively. Upper and Lower ), 2 known Life history: Not known. The 29 only from the deep sea, while the genus specimens of the two Bahamian species Danielopolina is the only member of reported by Yager (1987) are all the family inhabiting anchialine caves. females. While Iliffe et al. (1984) have Closest related species: Both T. postulated a deep sea origin for unidens and T. grandis from the Danielopolina, Danielopol (1990) Bahamas share a long and multi- believes anchialine Danielopolina segmented first antennae, although the species were derived from a shallow latter species is considerably larger. water ancestor. Evolutionary origins: The distribution of thermosbaenaceans with hypogean species in the West Indies, Yucatan, Texas, the Canary Islands, the Class Mediterranean region, Somalia and Subclass Eumalacostraca Cambodia is taken to indicate a Superorder Pancarida Tethyan origin associated with the Order Thermosbaenacea breakup of Pangea in post-Jurassic time Family Halosbaenidae (Schram, 1986; Cals and Monod, Subfamily Halosbaeninae 1988). Tulumella unidens Bowman & Iliffe, 1988 Characteristic features: Unpigmented, blind mysid with small Superorder Peracarida but non-functional eyestalks. Order Mysidacea Size: Length to 3 mm. Suborder Mysida Habitat: Anchialine. Family Mysidae Sumber of species in genus: Three, Antromysis (Antromysis) cenotensis all anchialine. Creaser, 1936 Genus range: Bahamas (Grand Characteristic features: Small, blind Bahama, Abaco and South Andros and unpigmented mysid. Islands), Mexico (Quintana Roo). Size: 3-4 mm in length from tip of Species range: Know from Najaron rostrum to end of telson. Cenote (Bowman and Iliffe, 1988), Habitat: Freshwater. Temple of Doom Cenote and Carwash Number of species in genus: Nine. Cenote (Holsinger, 1990). Genus range: Four troglobitic species 202 TROGLOBITIC FAUNA: Thomas M. Iliffe from Mexico (Yucatan, Quintana Roo outer rami of third uropod. and Oaxaca), Cuba and Jamaica make Size: Up to 10 mm. up the subgenus Antromysis. Marine Habitat: Typically anchialine. members of two other subgenera are Number of species in genus: Seven, found in the Bahamas, , all but one anchialine. Caribbean coast of Colombia, Costa Genus range: Bahamas (Grand Rica, Surinam and the Keys Bahama and Abaco Islands), Caicos (Bowman, 1977). Islands (Providenciales), Haiti and Species range: Widespread in caves in Mexico (Quintana Roo). Yucatan and Quintana Roo. Collection Species range: Bahadzia n.sp. A is sites in Quintana Roo include Actun know only from Xcan-ha Cenote on Ha, Cenote de San Martin, Cenote de Cozumel, while Bahadria n.sp. B Las Ruinas, Cenote de Santo Domin- inhabits one cave on the mainland of go, Cenote de Juan Coh, and Temple Yucatan and two on the island of of Doom Cenote (Reddell, 1977; Cozumel (Holsinger, in press) Bowman, 1977; Bowman, 1987). Ecology: Bahadzia is usually found in Ecology: Enormous numbers of A. mesohaline to euhalink waters below cenotensis are present swimming about the halocline in anchialine caves. in the water column of many cave Life history: Not known. pools. Closest related species: These two Life history: Not known. new species appear to be sibling taxa Closest related species: A. cubanica that evolved from a common ancestor (Cuba) and A. peckorurn (Holsinger, in ms.). (Jamaica). Evolutionary origins: The genus Evolutionary origins: All members of Bahadzia is most closely allied the subgenus Antromysis phylogenetically with Ma~.aweckelia lack eyes and inhabit caves. Their and Tuluweckelia from the Yucatan distribution in the Yucatan, Cuba and Peninsula (Holsinger, in press). Species Jamaica suggests descent from a B is thought to be close to the putative preadapted marine ancestor that ancestral species which independently colonized coastal caves in the northern and simultaneously colonized caves on Caribbean. both Cozumel and the mainland (Holsinger, in press). Species A, inhabiting an adjacent but apparently separate system on Cozumel, Order Amphipoda has diverged to a greater degree. Suborder Gammaridea Family Hadziidae Mayaweckelia cenoticola Holsinger, Bahadzia n.spp. A & B Holsinger, in 1977 press Characteristic features: Small to Characteristic features: Amphipod medium-sized amphipod lacking eyes lacking eyes and pigment; and pigment. Characterized by second segment on Size: Males to 4.0 mm; females to 6.0 TROGLOBITIC FAUNA: Thomas M. Iliffe 203 mm. Habitat: Anchialine. Habitat: Freshwater. Number of species in genus: One. Number of species in genus: Two, Species range: Known only from six both uoglobitic caves in Quintana Roo. Collection sites Genus range: Mexico (, include Temple of Doom Cenote, Cue- Yucatan and Quintana Roo). va de la Calavera, Carwash Cenote, Species range: Known from 11 caves Maya Blue Cenote, Cenote in Yucatan, Campeche and Quintana and Najaron Cenote, all near Tulum. Roo (Holsinger, 1990). Collection sites Ecology: Found in fresh to weakly in Quintana Roo include five caves - brackish waters of anchialine caves Cenote de San Martin, Cueva de near the northeastern coast of Quinta- Tancah, Carwash Cenote, Cenote de na Roo. Las Ruinas, Cenote de Santo Domingo Life history: Highly disproportionate (Reddell, 1977; Holsinger, 1990). sex ratio favoring females. Of the 56 Ecology: M. cenoticola inhabits specimens reported by Holsinger predominantly fresh or, at most, weakly (1990), 71% were female, 7% male and brackish cave waters. In several caves, 21% juveniles. Some larger females it was found in small, partially isolated had setose brood plates but none was pools (Holsinger, 1977). ovigerous. Life history: A strongly skewed sexual Closest related taxa: Tuluweckelia ratio favoring females may be present belongs to the weckeliid group of for this species. Hadziidae that consists of marine relict Closest related species: M. species inhabiting subterranean yucatanensis, known from a single cave freshwaters of southcentral Texas, in the state of Campeche. northern Mexico, Yucatan, Cuba, Puer- Evolutionary origins: Mayaweckelia to Rico, and Haiti (Holsinger, 1990). has taxonomic affinities with marine Evolutionary origins: According to forms. It inhabits caves situated Holsinger (1990), the marine ancestors between the Pliocene and Pleistocene of Tuluweckelia colonized caves during shorelines. Holsinger (1977, 1989) a Pleistocene regression of higher sea concludes from this that Mayaweckelia levels. Evidence for this is its originated from a marine ancestor distribution in partly brackish cave "stranded" by retreating sea levels waters near the coast. during the late Tertiary or early Quaternary.

Tuluweckelia cernua Holsinger, 1990 Order Characteristic features: Amphipod Suborder Flabellifera without eyes and pigment. Family Cirolanidae Anterior body region bent downward Bahalana mayana Bowman, 1987 at a sharp angle. Characteristic features: Medium- Size: Males to 5.0 mm; females to 8.0 sized cirolanid isopod without eyes and mm. pigment. 204 TROGLOBITIC FAUNA: Thomas M. Iliffe

Size: Males to 10.0 mm; females to 8.5 Yucatan and Quintana Roo. Collection mm. sites in Quintana Roo include Temple Habitat: Anchialine. of the Doom Cenote, Cenote de San Number of species in genus: Three, Martin, Cueva Ajin and Cenote Tos all anchialine Virlol (Reddell, 1977; Wilkens, 1982; Genus range: Bahamas (San Salvador Bowman, 1987). and Mayaguana Islands), and Mexico Ecology: Numerous specimens (Quintana Roo). collected from freshwater cave pools Species range: Known only from two using traps baited with meat (Creaser, caves in Quintana Roo: Cueva Quebra- 1936, 1938). da on the Island of Cozumel, and Tem- Life history: Not known. ple of Doom Cenote, near Tulum. Closest related tam: Not known. Ecology: In Cueva Quebrada, Evolutionary origins: Derived specimens were collected from marine forms in early Pleistocene approximately 800 m from the entrance times (Wilkens, 1982). at a depth of about 5 m. Salinity at this point was 21 ppt and dissolved oxygen, Superorder Eucarida 3.0 ppm. Specimens in Temple of Order Doom Cave were collected in 12 to 18 Suborder Pleocyemata m depths, from water of 14 ppt salinity Infraorder Caridea (Bowman, 1987). Superfamily Procaridoidea Life history: Collected specimens Family Procarididae include 9 males, 6 females and 1 Procaris n. sp. Kensley, in prep. juvenile (Bowman, 1987). Characteristic features: Closest related species: The two Phyllobranchiate gills; maxillipeds and Bahamian species appear to be most pereiopods with strong exopods; none closely related while B. mayana differs of pereiopods chelate or subchelate; from both in a number of features. rostrum small and unarmed. Evolutionary origins: The members of Size: Similar to other congeners the genus appear to be derived from a where the carapace length reaches 9- marine ancestor ranging the area from 10 mm. the Bahamas to the Yucatan Peninsula Habitat: Anchialine. (Holsinger, 1989). Number of species in genus.- Four, all anchialine. Creaseriella anops (Creaser, 1936) Genus range: Bermuda, Ascension Characteristic features: Large Island, , Mexico (Quintana cirolanid isopod without eyes and Roo). pigment. Species range: Anchialine caves on Size: To 21.8 mm (Creaser, 1936). Cozumel. Habitat: Freshwater. Ecology: Abel and Felgenhauer Number of species in genus: One. (1985) have studied the ecology of the Species range: Known from numerous Ascension species. They report that cenotes and caves in the states of smaller individuals spend most of their TROGLOBITIC FAUNA: Thomas M. Iliffe 205

time in crevices, while larger Superfamily Atyoidea individuals are observed swimming in Family Atyidae open water. Examination of gut Typhlatya mitchelli Hobbs & Hobbs, contents reveals that they feed on both 1976 plant matter and crustaceans including Characteristic features: Translucent amphipods and atyid . to white or pigmented small shrimp. Life history: Little is known of either Pigment, which occurs only in the reproductive biology or ontogeny individuals from some localities, of Procaris (Schram, 1986). A sex appears grayish to black or brown ratio, determined by the presence of (Hobbs, 1979). Eyes lack facets and genital apertures, of eight females Lo pigment. Rostrum does not extend one male has been determined for the anteriorly beyond eyessize: Total Ascension species, P. ascensionis length to about 16 mm; carapace length (Felgenhauer et al., 1988). Although of females to 4.8 mm. more than 1,000 specimens of P. Habitat: Freshwater. ascensionis were observed in the field, Number of species in genus: Nine, all no ovigerous females were seen. A troglobilic. single female maintained in the Genus range: Ascension Island, laboratory bore approximately 60 Bermuda, West Indies (Barbuda, bright orange eggs on the endopods of Caicos Islands, Cuba, Dominican pleopods. The large size of these eggs Republic, Mona, Puerto Rico), Mexico (0.83-0.93 mm) suggests the existence (Yucatan, Campeche and Quintana of a zoeal larval stage (Felgenhauer et Roo) and Galapagos Islands. al., 1988). Species range: Known from 17 caves Closest related species: All known and cenotes in the states of Yucatan and species of Procaris are quite similar, Quintana Roo. Collection sites in Quin- showing few differentiating characters tana Roo include Cueva del Fermin and (Hart and Manning, 1986). Actin Ha (Hobbs, 1979). Evolutionary origins: Hart and Ecology: According to Hobbs Manning (1986:416) note that the (1979:622), similarities between species and their In all localities shrimps were living in highly anomalous distribution in lentic situations varying from very marine caves indicate an extremely small shallow (less than 0.3 m depth) slow rate of evolution. They suggest pools to extensive underground lakes that, "Procaris, or its predecessors, (greater than 2.0 m depth). Generally may, at one time, have been widely the pools were in total darkness, but T. distributed throughout the oceans, mitchelli occurs in entrance areas that surviving today only in cryptic habitats receive direct light from the surface. removed from some of the Substrates of the pools consisted of environmental necessitating guano, silt, organic debris, rocks and change." combinations thereof. Shrimp were found on the subsaate, "hanging" from submerged walls along the edges of 206 TROGLOBITIC FAUNA: Thomas M. Iliffe

lakes, free-swimming, and among den- of females to 5.2 mm (Hobbs and se, branched root systems hanging Hobbs, 1976). down from the roof of cenotes. Habitat: Freshwater. Life history: Males and ovigerous Species range: Known from 21 caves females are as yet unknown (Hobbs & and cenotes in the states of Yucatan, Hobbs, 1976). Campeche and Quintana Roo. Closest related species: T. mitchelli is Collection sites in Quintana Roo intermediate, both in morphology and include Cueva del Ferrnin, Pozo de San in geographic position, between the Martin, Cenote de Juan Coh, Cenote de Caribbean species with a short rostrum Santo Domingo, Cenote Tos Virlol and (from Cuba and Mona) and the Cenote Coop (Hobbs, 1979). seemingly disjunct species from the Ecology: According to Hobbs (1979), Galapagos (Hobbs & Hobbs, 1976). This species is found in lentic habitats Evolutionary origins: Of the nine ranging from small pools (less than 0.5 species in the genus, four (from the m in depth) to deep lakes (greater than Galapagos Islands, Bermuda, 8 m). These bodies of water are Ascension Island and the Caicos generally floored with silt or gua- Islands) inhabit brackish or marine no ... Typhlatya pearsei is generally waters, while the remainder are found found in total darkness but occasionally in freshwater habitats. According to individuals are seen in enuance areas Iliffe (1986:7), "species within the where pools receive either direct or genus appear to have evolved from an indirect light from the surface. open water marine ancestor in the Dissolved oxygen levels in pools Atlantic which spread westward containing this shrimp are as low as through the Caribbean into the Pacific 0.78 mg 1-I or about 10% saturation with prevailing currents before the (Hall, 1936). closure of the Panama land bridge." Life history: Of the 230 specimens Iliffe et al. (1983) suggested an origin examined by Hobbs (1979), 88% were of the genus on submerged and females, 5% males and 6% juveniles. emergent seamounts associated with Closest related species: T. pearsei the Mid-Atlantic Ridge during the belongs to a group in which the rosuum separation of the American and African extends beyond the eyes. This group continental masses. includes Typhlatya species from Bermuda, Ascension Island, Cuba and Typhlatya pearsei Creaser, 1936 Campeche (Mexico). Characteristic features: Small, light, Evolutionary origins: Both T. pearsei whitish shrimp, somewhat opaque but and T. mitchelli show identical degrees not transparent; eyes lack facets and of eye reduction suggesting that they pigment. started the cavemicolous evolution at Rostrum extends anteriorly well the same time (Wilkens, 1982). beyond eyes. Size: To about 17 mm in total length. Carapace length of males to 3.8 mm; TROGLOBITIC FAUNA: Thomas M. Iliffe 207

Superfamily Rhynchocinetoidea Superfamily Palaemonoidea Family Agostocarididae Family Palaemonidae Agostocaris bozanici Kensley, 1988 Creaseria morleyi (Creaser, 1936) Characteristic features: Small, Characteristic features: Larger, unpigmented shrimp with eyes not translucent to white shrimp. drfferentiated into and stalk. Eyes somewhat bullet-shaped but Integument with sparse scattering of lacking pigment. Carapace with two tiny black (Kensley, spines along anterior lateral margin. 1988). Size: To approximately 70 mm in to- Size: Carapace length to 8.0 mm tal length; carapace length in males to Habitat: Anchialine. 20.3 mm, in females to 29.0 mm Number of species in genus: Two, (Hobbs, 1979). both anchialine. Habitat: Freshwater. Genus range: Bahamas (Grand Number of species in genus: One. Bahama Island), Turks and Caicos Species range: Widely distributed in Islands (Providenciales), Mexico at least 32 caves and cenotes in (Quintana Roo) Yucatan, Carnpeche and Quintana Roo Species range: Known only from (Hobbs, 1979). Xcan-ha Cenote on the island of Collection sites in Quintana Roo Cozumel, Quintana Roo. include Cueva del Fermin, Pozo de San Ecology: Specimens were collected in Martin, Cenote de Juan Coh, Cenote de euhaline waters (salinity 34 ppt) at 18 Las Ruinas, Cenote de Santo Domin- to 31 m depths. The foregut of two go, Cenote de Tos Virlol, Cenote Ajin specimens were packed with the and Cueva Coop. filaments of the benthic Ecology: Acoording to Hobbs cyanobacterium Oscillaroria (1979:630), corallinae. Creaseria morleyi occurs in quiet water Life history: Of the 9 specimens habitats ranging from small, shallow collected, 4 were females and 5 were pools (less than 0.5 m deep) and natu- inmature. None of the females were ral well (5 m deep) to large, deep lakes ovigerous (Kensley, 1988). (greater than 3 m in depth). These Closest related species: A. williamsi bodies of water are floored by guano from Grand Bahama Island and Provi- deposits, rich organic silt, debris, and denciales. rocks. Shrimp were found in total Etolutionary origins: Affinities for darkness as well as in entrance areas this family, which contains only two receiving direct and indirect light. bqxcies, are unclear. The genus shares Creaser (1936) observed these shrimp some features with the Bresiliidae, a crawling over the bottom, as well as iarn~lyof shrimp inhabiting deep sea swimming in the water column. He h) drotherrnal vents (Hart & Manning, states that they are extremely sensitive 1986; Kensley, 1988). to vibrations in the water. Specimens were collected in a wire trap baited with meat. Examination of stomach 208 TROGLOBITIC FAUNA: Thomas M. Iliffe contents indicates that the species is Cozumel indicate an extended cannibalistic (Creaser, 1938). Holthuis planktonic larval life that would help (1977: 187) states that, "these animals to explain its wide distribution are very aggressive, if two or more are (Kensley, 1988). placed together in a smallish container Closest related taxa: Closest affinities they will attack and mutilate each are to Somersiella sterreri (Hart and other." Manning, 1981). Life history: Of the 92 specimens Evolutionary origins: Considering the examined by Hobbs (1979), 70% were open, nonanchialine of the type females, 27% males and 3% juveniles. locality - a seawall in Antigua, Closest related taxa: No known close planktonic larval dispersal by ocean relatives. currents may explain its widespread Evolutionary origins: This shrimp is distribution (Kensley, 1988). believed to be derived from a marine ancestor stranded by sea level Sornersiella sterreri Hart & Manning, regressions in the early Pleistocene 1981 (Wilkens, 1982). Characteristic features: Large, uniformly reddish shrimp with three prominent white spots dorsally, 1 middorsally on the sixth abdominal Superfamily Alpheoidea somite and one on each exopod of the Family Hippolytidae uropods. The cornea is black, while the Janicea antiguensis (Chace, 1972) antennae are reddish basally, becoming Characteristic features: Medium- white for most of their length. The sized, red shrimp with pigmented eyes walking legs and the body are red (Hart and multiarticulate walking legs. and Manning, 1981). Size: Carapace length to 10.0 mm. Size: Carapace length to 16.1 mm. Habitat: Anchialine. Habitat: Anchialine. Number of species in genus: One. Number of species in genus: One. Species range: Antigua (from seawall), Species range: Anchialine caves in Bermuda (from reef cave in 15 m Bermuda and Quintana Roo. Cueva depth), Grand Bahama Island Quebrada on Cozumel is the only (Bahamas) and Cozumel (Quintana Mexican cave where this species has Roo) (both latter locations from been found. anchialine caves). Ecology: Cueva Quebrada, which is The only Mexican record for this more than 2700 m long, connects with species is from Cueva Quebrada the sea through entrances on the west (Kensley, 1988). coast of Cozumel. Maximun depth in Ecology: Collected from marine waters the cave is only 12 m, while the salinity of an extensive anchialine cave is 21 ppt (Bowman, 1987). connecting directly with the sea. Life history: An ovigerous female Life history: Numerous very small from Cozumel carried an estimated eggs on the ovigerous females from 2,000 tiny eggs. The small size of the TROGLOBITIC FAUNA: Thomas M. Iliffe 209 eggs suggests an extended existence in reproductive phase (Kensley, 1988). the plankton and may explain its Closest related taxa: Bears a superfi- presence at Bermuda (Kensley, 1988). cial resemblance to the genus Closest related taxa: Closest affinities Calliasmata represented by one species are to Janicea antiguensis (Hart and in the Red Sea area and one from the Manning, 1981). Dominican Republic. Evolutionary origins: Iliffe et al. Evolutionary origins: Pterygostomian (1983) suggested that S. sterreri is a spine and configuration of telson have Tethyan relict. The presence of no equivalents among the hippolytid numerous small eggs may imply genera. planktonic dispersal by ocean currents.

Yagerocaris cozumel Kensley, 1988 Characteristic features: Small shrimp Phylum CHORDATA with weakly pigmented eyes. Carapace Class Teleostomi with strong pterygostomian spine. Order Ophidiiforme~ Size: Carapace length to 7.2 mm. Suborder Bythitoidei Habitat: Anchialine. Family Bythitidae Number of species in genus: One. Subfamily Bythitinae Species range: Known only from Ae- Tribbe Dinematichthyini rolito Cenote and Cueva Quebrada on Ogilbia pearsei (Hubbs, 1938) the island of Cozumel, Quintana Roo. Characteristic features: Clear white Ecology: Specimens were collected brotulid becoming pinkish along the from eurihaline waters (marine salinity) posterior margins. Total lack of eyes; at 9 to 12 m depths. The foregut of large and well developed sensory dissected specimen contained an cavities on head. unidentifiable brown finely macerated Size: To 90.5 mm. mush (Kensley, 1988). Habitat: Freshwater. Life history: Individual shrimp show Number of species in genus: Six, plus a combination of both male and female numerous undescribed ones (Cohen & characteristics, possibly indicating Nielsen, 1978). protandrous hermaphroditism. Five of Genus range: From tropical American nine specimens collected were reefs, freshwater caves in the Yucatan ovigerous. Protandrous and brackish waters in the Galapagos. hermaphroditism in cave species has a Probably occurring in the Indo-Pacific primary advantage correlated with as well (Cohen & Nielsen, 1978), larger size of breeding females. Species range: Yucatan and Quintana Secondarily, considering the somewhat Roo. Navarro-Mendoza (1988) reports restricted gene pool within caves, it 0.pearsei from four locations within would enhance opportunity for gamete Quintana Roo. exchange if two individuals Ecology: Although in Yucatan this fish encountering one another were both has been reported from small or hermaphroditic and in the appropriate freshwater pools, in Quintana Roo it 210 TROGLOBITIC FAUNA: Thomas M. Iliffe occurs in larger and more diverse ca- ves in Yucatan. Blind probably ves and cenotes (Navarro-Mendoza, belonging to this species have been 1988). In these caves, it occurs with seen in Act6n Ha and Cenote de Santo such predators as the American Domingo, Quintana Roo (Reddell, Anguilla rostrara and the 1977). Coke (pers. comm.) has Rhamdia guatemalensis. Wilkens observed large eels of this species to (1982) reported that in Yucatan 0. 60 cm in length from Carwash Cenote. pearsei is usually found associated with He has also noticed specimens in Sac the eel Ophisternon infernale. Actun, Najaron, and Maya Blue. Life history: The species is viviparous Ecology. 0.infernale inhabits lightless with a reproductive period occurring waters containing organic during the months of December to rich sediments where it constructs February (Navarro-Mendoza, 1988). mucus-lined (Navarro- The young, which range in size from Mendoza, 1988). Although blind and 22 to 34.5 mm, are independent and inhabiting a totally dark environment, actively swimming from the moment this eel shows a circadm cycle with of birth. The number of young ranges maximal feeding activity occurring from 2 to 11, depending upon the size during the night. Laboratory of the mother. experiments suggest a diminished Closest related species: Congeners photoreception capacity in that inhabiting coral reefs in the Atlantic. specimens held in aquaria avoid light Evolutionary origins: Derived from (Navarro-Mendoza, 1988). When marine forms (Wilkens, 1982). dissolved oxygen levels fall below 1.3 mglliter, these eels may stick their Order Synbranchiformes heads out of water and take in Ophisternon infernale (Hubbs, 1938) atmospheric air. Characteristic features: Synbranchid Life history: The reproductive biology eel with total lack of external eyes, of this species is under study by Nava- degenerate brown pigmentation and rro-Mendoza. enhanced development of dermal Closest related species: 0. organs on the head (Hubbs, 1938). aenigmaticum, a secondary freshwater Size: To 60 cm. fish living in Mexico and Cuba (Rosen Habitat: Freshwater. & Greenwood, 1976). Number of species in genus: Six Evolutionary origins: 0. infernale (Rosen & Greenwood, 1976). appears to be clustered in caves located Genus range: , Ceylon, along the old Pliocene shoreline Philippines, Northern , (Wilkens, 1982). Although other Western Australia, Portuguese Guinea, cavernicolous species have secondarily , Guatemala, Cuba and spread over northern Yucatan through Mexico (Campeche, Tabasco, Chiapas, channels in the subterranean aquifer, Oaxaca, Veracruz, Yucatan and Quin- both 0.infernale and Ogilbia pearsei tana Roo). seem to be primarily restricted to ca- Species range: Known from five ca- ves and cenotes near their original sites TROGLOBITIC FAUNA: Thomas M. l'liffe 211 of cave colonization (Wilkens, 1982). from Jamaica and Oaxaca, MCxico, and a redescription and new records for A. cenotensis. Association for Mexican Acknowledgements Cave Studies Bulletin, 6:27-38. Bowman, T.E. 1987. Bahalana A 1986 cave diving expedition to mayana, a new troglobitic cirolanid study and collect from the anchialine isopod from Cozumel Island and the caves of the Yucatan was sponsored by Yucatan Peninsula. Proceedings of the National Science Foundation grants Biological Society of Washington, BSR-8215672 and ,BSR-8417494. 100:659-663. During this expedition, cave diving and Bowman, T.E. and T.M. Iliffe. logistic assistance was provided by 1988. Tulumella unidens, a new genus James Coke, Dinah Drago, Juan JosC and species of thermosbaenacean Fucat, Michael Madden and Dr. John crustacean from the Yucatan Peninsula, Markham. The following individuals Mexico. Proceedings of the Biological generously provided information and Society of Washington, 101:221-226. reviewed portions of the manuscript: Cals, P. and T. Monod. 1988. Drs. Thomas E. Bowman, Horton ~volutionet biogbographie des Hobbs 111, John Holsinger, Brian Crustacbs ThermosbCnacCs. Comptes Kensley, Louis Kornicker, James Rendus de I'Acade'mie des Sciences, Reddell, and Jill Yager. Mr. James Paris, 307 (SCrie III):341-348. Coke supplied information on Chace, F.A., Jr. 1972. The shrimps the state of cave diving explorations of of the Smithsonian Bredin Caribbean the anchialine caves as well as perso- Expedilions with a summary of the nal observations on the cave fauna. West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology Bibliography 98:x + 179. Cohen, D.M. and J.G. Nielsen. Abele, L.G. and B.E. Felgenhauer. 1978. Guide to the identification of 1985. Observations on the ecology and genera of the fish order feeding behavior of the anchialine with a tentative classification of the shnmp Procaris ascensionis. Journal order. NOAA Technical Report NNFS of Crustacean Biology, 5: 15-24. Circular, 4 17: 1-72. Back, W., B.B. Henshaw, J.S. Creaser, E.P. 1936. Crustaceans Herman, and J.N. Van Driel. 1986. from Yucatan. Carnegie Institute of Differential dissolution of a Pleistocene Washington Publications, 457: 117- reef in the ground-water mixing zone 132. of coastal Yucatan, Mexico. , Creaser, E.P. 1938. Larger cave 14: 137-140. Crustacea of the Yucatan Peninsula. Bowman, T.E. 1977. A review of Carnegie Institute of Washington the genus Antromysis (Crustacea: Publications, 491: 159-164. .Mysidacea), including new species Danielopol, D.L. 1990. The origin 212 TROGLOBITIC FAUNA: Thomas M. Iliffe of anchialine cave fauna the "deep sea" Mexico. Association for Mexican Cave versus the "shallow water" hypothesis Studies Bulletin, 6: 15-25. tested against the empirical evidence Honsinger, J.R. 1989. Preliminary of the Thaumatocyprididae zoogeographic analysis of five groups (Ostracoda). Bijdragen tot de of crustaceans from anchialine caves in Dierkunde, 60: 137-143. the West Indian region. Proceedings of Felgenhauer, B.E., L.G. Abele and the Tenth International Congress of W. Kim. 1988. Reproductive , Budapest, Hungary, pp. 25- morphology of the anchialine shrimp 26. Procaris ascensionis (Decapoda: Holsinger, J.R. 1990. Tuluweckelia Procarididae). Journal of Crustacean cernua, a new genus and species of Biology, 8:333-339. stygobiont amphipod crustacean Hall, F.G. 1936. Physical and (Handziidae) from anchialine caves on chemical survey of cenotes of Yucatan. the Yucatan Peninsula of Mexico. Carnegie Institute of Washington Beaufortia, 41:97-107. Publications, 4575- 16. Holsinger, J. R. In press. Two new Hart, C.W., Jr. and R.B. Manning. species of the subterranean amphipod 1981. The cavernicolous caridean genus Bahadzia (Hadziidae) from the shrimps of Bermuda (Alpheidae, Yucatan Peninsula region of southern Hippolytidae, and Atyidae). Journal of Mexico, with an analysis of phylogeny Crustacean Biology, I :44 1-456. and biogeography. Stygologia. Hart, C.W., Jr. and R.B. Manning. Holthuis, L.B. 1973. Caridean 1986. Two new shrimps (Procaididae shrimps found in land-locked saltwater and Agostocarididae, new family) from pools at four Indo-West Pacific marine caves of the Western North localities (Sinai Peninsula, Funafuti Atlantic. Journal of Crustacean Atoll, Maui and Hawaii Islands), with Biology, 6408-4 16. the description of one new genus and Hobbs, H.H. 111. 1979. Additional four new species. Zoologische notes on cave shrimps (Crustacea: Verhandelingen, 128:1-48. Atyidae and Palaemonidae) from the Hubbs, C.L. 1938. from the Yucatan Peninsula, Mexico. caves of Yucatan. Carnegielnstitute of Proceedings of the Biological Society Washington Publications, 491:261- of Washington, 92:618-633. 295. Hobbs, H.H. I11 and H.H. Hobbs, Iliffe, T.M. 1986. The zonation Jr. 1976. On the troglobitic shrimps of model for the evolution of aguatic the Yucatan Peninsula, Mexico in anchialine waters. Stygologia, (Decapoda: Atyidae and Palaemoni- 2:2-9. dae). Smithsonian Contributions to Iliffe, T.M., C.W. Hart, Jr. and R.B. Zoology, 240: 1-23. Manning. 1983. Biogeography and the Holsinger, J.R. 1977. A new genus caves of Bermuda. Nature, 302: 141- and two new species of subterranean 142. amphipod crustaceans (Gammaridae Iliffe, T.M., H. Wilkens, J. Parzefall s.lat.) from the Yucatan Peninsula in and D. Williams. 1984. Marine TROGLOBITIC FAUNA: Thomas M. Iliffe 213 cave fauna: Composition, systematics of synbranchiform fishes. biogeography and origins. Science, Bulletin of the American Museum of 225:309-3 11. Natural History, 1575-69. Kensley, B. 1988. New species and Schrarn, F.R. 1983. Remipedia and records of cave shrimps from the crustacean phylogeny. Pp. 23-28 In: Yucatan Peninsula (Decapoda: F.R. Schram, ed., Crustacean Agostocarididae and Hippoly tidae). Phylogeny. A.A. Balkema, Rotterdam, Journal of Crustacean Biology, 8:688- xi + 372 pp. 699. Schram, F.R. 1986. Crustacea. Kornicker, L.S. and T.M. Iliffe. Oxford University Press, Oxford, xii + 1989. New Ostracoda (Halocyprida: 606 pp. Thaumatocyprididae and Halocypridi- Wilkens, H. 1982. Regressive dae) from anchialine caves in the evolution and phylogenetic age: The Bahamas, Palau and Mexico. history of colonization of freshwaters Smithsonian Contributions to Zoology, of Yucatan by fish and Crustacea. 470: 1-47. Association of Mexican Cave Studies Navarro-Mendoza, M. 1988. Inven- Bulletin, 8:237-243. tario ictico y estudios ecol6gicos pre- Yager, J. 1986. Remipedia. Pp. liminares en 10s cuerpos de agua 382-383 In: L. Botosaneanu, ed., continentales en la Reserva de la Mundi. E.J. Brill, Leiden, Biosfera de Sian Ka'an y keas circun- vi + 740 pp. vecinas en Quintana Roo, Mexico. Yager, J. 1987. Speleonecres Technical Report, U.S. Fish and tulumensis n. sp. (Crustacea, Wildlife Service. Remipedia) from two anchialine Pearse, A.S. (ed.) 1936. Fauna of cenotes of the Yucatan Peninsula, the caves of Yucatan. Carnegie Mexico. Stygologia, 3: 160-166. Institute of Washington Publications, 457: 1-304. Pearse, A.S. (ed) 1938a. Fauna of the caves of Yucatan. Carnegie Institute of Washington Publications, 491: 1-304. Pearse, A.S. 1938b. Fauna of the caves of Yucatan, Introduction. Carnegie Institute of Washington Publications, 491: 1- 17. Reddell, J. A. 1977. A preliminary survey of the caves of the Yucatan Peninsula. Association of Mexican Cave Studies Bulletin, 6:215-296. Rosen, D. E. and P. H. Greenwood. 1976. A fourth neouopical species of synbranchid eel and the phylogeny and 214 TROGLOBITIC FAUNA: Thomas M. Iliffe

Fig. 1: Map of Quintana Roo showing locations of caves where aquatic troglobites have been reported.

1) Cenote de Santo Domingo; 2) Cenote de Las Ruinas; 3) Cenote de Juan Coh; 4) Cenote Tos Virlol; 5) Cenote Naharon; 6) Maya Blue Cenote; 7) Carwash Cenote; 8) Cueva de Tancah; 9) Temple of Doom Cenote; 10) Cenote Sac Actlin; 11) Actcn Ha; 12) Cenote Mojara; 13) Cenote Ponderosa; 14) Cenote de San Martin and Pozo de San Martin; 15) Cueva del Fermin; 16) Cenote Ajin; 17) Cenote Coop; 18) Cueva Quebrada; 19) Xcan-ha Cenote;6 20) Aereolito (Modified from Reddell, 1977). TROGLOBITIC FAUNA: Thomas M. lliffe 215

Fig. 1