'' BULLETIN DE L'INSTITUT RO YAL DES SCIENCES NAT URELLES DE BELGIQ UE, BULLETIN VAN HET KONINKLIJK BELGISCH lNSTITUUT VOO R NATUURWETENSCHAPPEN, BIOLOG fE , 76: 5-26, 2006 BIOLOG IE, 76: 5-26, 2006 The remarkable diversity of subterranean Cirolanidae (Crustacea: Isopoda) in the peri -Caribbean and Mexican Realm by Thomas M. ILIFFE & Lazare BOTOSANEANU We dedicate this paper to the memory of Jan H. Stock, indef atigable promoter of stygobiological research in the West Indies, almost ten years aft er his untimely death. Abstract With 42 ende mic spec ies (o ne of them with two subspecies) described to this day, most of th em belo nging to 11 ende mic genera, the peri-Caribbea n -Mex ican zone has definitely the richest fa un a of stygobitic (entirely subterranean-adapted) cirolanids of the world . After a succinct prese ntation of th e various habitats fo r stygobitic cirolanids in the zone, the species are grouped together (Table I) accordin g to the ir affili atio n to habitats of fres hwater or of marine wate r - water salinity being considered th e most reliable common denominator. A check-list in alphabeti cal order follows, including references, know n distribution , informati o n on habitats, etc. That this is a case of very hi gh biodi versity of stygobitic Ci ro lanidae res ults from comparison with the situati on in th e rest of the wo rld. Some aspects of morph ology in relation to sys te mati cs are reviewed (i. a. : evoluti ve significan ce of some morphological adaptations; monophyly of stygobitic genera). Attempts are made to reply to ques ti ons like: what can we kn ow about ancestors of stygobitic species? Can age of stygobitic taxa, or lineages, be appreciated with some degree of reliability? What can we guess about the roots of prese nt-day di versity (two types of habitat requ iring di ffe rent ex pl anati on fo r time and mode of groundw ater coloni zati o n being recog ni zed)? How deep are these roots? What is the legacy of Tethys? This case of hi gh biodi versity could be explained th rough the rich , turbule nt history of this vast and frag mented area; th rough the abundance and ex traordin ary di ve rsity of habitats propitious for stygoevolution; and thro ugh the (s upposed) abundance and hi gh diversity of pote ntial marin e ancestors. A final part of the paper is devoted to endangered or already destroyed habitats and fa un a, with a plea for protecti on. Key words: stygobitic C irol anidae, Caribbean-Mex ican area, subte rranean aqu atic habitats, biodi vers ity, stygoevo lution Resume Avec 42 especes ende miques ac tu elle me nt decrites (un e de celles ci avec deux sous-especes), appartenant pour Ia plupart a 11 ge nres e ndemiq ues, Ia zone Cara"ibe-Mex icaine possede sans doute Ia fa un e de cirolani des stygobies Ia plus ric he du mo nde. Apres une prese ntatio n succincte des divers habitats peupl es par des ciro lanides stygo bies dans Ia zo ne, les especes sont groupees (Tableau 1) d'apres ces habi tats: eau do uce o u eau marine- Ia sa linite de l' ea u etant conside ree com me denominate ur commun le plus digne de confiance. Une li ste des taxo ns e n ordre alphabetique presente les references, Ia distribu tion connue, des info rmati ons sur les habitats, etc. Une comparaiso n avec Ia situati o n pour le reste du globe montre qu ' il s ' agit ici d ' un cas de biodive rsite remarqu ablement grande. Sont passes en revue plusieurs aspects concern ant des relati ons e ntre morphologie et systematique (signi ficatio n evoluti ve de certaines adaptations morphologiques; monophylie des ge nres stygobies). On essaie de fo urnir des reponses a des questions comme: qu ' est-ce que peut etre connu sur les ancetres des especes stygobies? Est-ce que !'age des taxo ns (ou des lignees) stygobies peut etre apprecie avec un certain degre de certitude? Qu ' es t-ce qu ' on peut supposer concern ant les rac ines de Ia di versite actuelle (deux ty pes d ' habitat de mandant des explications di ffe rentes pour le te mps et Ia modalite de Ia colonisati on des eaux soute rraines etant distingues)? Combien profo ndes so nt ces rac ines? Que! est, dans ce cas co ncret, I' heritage de Ia Tethys? Ce cas de biodiversite pourrait etre ex plique par l'hi stoire ric he et mouve mentee de cette zone vaste et frag mentee; par l' abond ance et l'extraordinaire diversite des habitats propices a Ia stygoevolutio n; ainsi que par l'abond ance et Ia di versite (s upposees) des a ncetres marins potentiels. Une secti on fin ale du trava il es t consacree aux habitats et a Ia faun e menaces OU deja detruits. Mots-cJes: Cirolanidae stygobies, zone Cara1be-Mex icaine, habitats aquatiques so uterrains, biodiversite, stygoevolution. Introduction Although the first subterranean cirolanid isopod of the Caribbean -Mexican zone was di_scovered more than one century ago, it was only during the last half of the previous century that research on this group of stygobitic crustaceans has assumed large proportions, thanks to increas ing interest in speleology and stygobiology and to improving techniques for exploration and zoological sampling. This has led to discovery and description of a remarkable large number of species entirely bound to subterranean aquatic habitats of various kinds, making this zone the most extraordinary " hot spot" of the world for stygobitic Cirol anidae. The time seems to be ripe fo r attempting a sy nthetic presentation of know n fac ts, foc using on diversity and its poss ible explanation, in a naturalistic spirit, and limiting to a minimum reasonable speculati on not dictated by fashions. In order to avoid this paper taking exaggerated proportions, almost no reference to stygobionts other than Cirolanidae and to "generalized tracks" will be made. Also reference to publications not directly dea ling with Cirolanidae has been limited to some of II 6 T. M. ILIFFE & LAZARE BOTOSANEANU those shedding more light on aspects directly concerning the present paper. The stage The zone dealt with in this paper is called "Caribbean-Mexican basin" by ARGANO (1972), "The (sub) tropical Atlantic" by STOCK (1994), " the Central American/Caribbean intermediary areas" by BANARESCU ( 1995), or "Western Atlantic" by BRIGGS ( 1995) - who distinguishes here a "West Indian Province" and a "Caribbean Province". The wording used in the title is from Stygofauna Mundi (BOTOSANEANU, 1986). Stygobitic cirolanids of the peri-Caribbean - Mexican zone inhabit wholly or partially submerged caves and groundwater of varying salinity, all situated in karstic limestone. Due to locally characteristic differences in habitat type, each area within the zone will be considered separately. Generalized descriptions of habitat, geological setting, hydrology, water quality, and cirolanid stygofauna will be presented. Distribution of cirolanid species within the zone is illustrated in Fig. 1. Bermuda is a mid-Atlantic volcanic seamount, capped with Pleistocene aeolian and marine limestone. Karstic caves in Bermuda are concentrated in a small area of the island where the oldest limestone outcrops. Tidal , sea level pool s occur at entrances and in the dark interior of the caves. Although the upper (dry and shallow submerged) portions of Bermuda caves are dominated by collapse features, extensive horizontal passage development typically occurs at 18m de pth. The presence of large speleothems (stalactites, stalagmites, etc.) in all parts of the underwater caves indicates that the entire known portions were completely dry and air-filled for long times during glacial periods of lowered sea level. While surface waters in Bermuda's cave pool s range from slightly brack ish to nearly fully marine, deeper waters below about 35 m depths approach fully marine salinities. One stygobitic cirolanid, Arubalana arubaides, is known from Bermuda. The Bahamas archipelago consists of a series of shallow banks made up of a continuous, up to 5700 m thick, sequence of carbonates of shallow water origin which began depositing during the Jurassic. As the banks gradually subsided, carbonate deposition has kept pace, mai ntaining a shallow water environment. The banks are separated from one another by deep water channels reac hing 2,000 m de pths. Water-filled caves in the Bahamas are locally referred to as " blue holes". Inland blue holes occur in the interior of islands, while ocean blue holes are s ituated offshore, in shallow water. While many inland and ocean blue holes are karstic sinkhol es with no passage developme nt, others are parts of an extensive network fo rmed either by limestone di ssolution at the fresh/saltwate r inte1face (flank margin caves) or by slump faulting occurring along the platform marg in s (fault line caves). Ocean holes typica ll y have reversi ng, high velocity, tidal currents which flush the caves with seawater, while inland caves lack perceptible currents and have a fresh to brackish water le ns overl y ing fully marine waters. Stygobitic fa una of the Bahamas includes seven species of cirolanids from three gene ra, Bahalana, Ciralana (subgenu s Ciralana) and Exumalana. The northern coast of the Yucatan Peninsula is composed of Pleistocene and Holocene limestone, while Miocene to Eocene age rocks are exposed in the interior. Northern Yucatan is generally low, with little relief, and smface rivers or streams are absent. Water-filled caves are locally referred to as cenotes, a Spanish corruption of the Mayan word "dzonot". The most exte nsive underwater cave systems in the world are located along the eastern Caribbean coastline, with Systema Ox B-::1Ha at 130 km being the world's longest. In the interior of the Peninsula, a well defined semicircle of cenotes, mostly in the form of deep sinkholes, outlines the deeply buried rim of the 65 million year old Chicxulub impact crater. Groundwater throughout the Peninsula consists of a fresh water lens of varying thickness, floating on top of denser seawater. The depth of the halocline boundary between fresh and saltwater varies from 10 m depth along the coast to more than 60 m in the interior. The stygobitic fauna of Yucatan thus includes both freshwater and marine species. Seven species of stygobitic cirolanids from five genera, Metaciralana, Cirolana (subgenus Anapsilana), Creaseriella, Haptalana and Yucatalana , have been described from the Yucatan Penin sula (including Cozumel and Belize). The Greater Antilles consist of the four large islands of Pue rto Rico, Hispaniola (with Haiti and the Dominican Republic), Cuba, and Jamaica. They are part of a lower Cretaceous to Holocene island arc chain making up a submerged mountain range extending from Central America eastwards through the Caribbean. Each of the four islands is encircled by a coastal plain, backed on the north coast of Cuba, Jamaica, and Hispaniola by Pleistocene-raised shorelines that reach heights of 300 m. The extensive limestone outcrops on these islands have given rise to a tropical karst landscape, characterized by subterranean drainage, well developed cockpit and tower karst, and numerous caves including large river caves, deep sinkholes and long horizontal caves near the coast that formed under the influence of palaeo-sea levels. Two species each of stygobitic cirolanids are known from Cuba (Haptalana and Cirolana, subgenus Anapsilana), Jamaica (Aruba/ana and Ciralana, subgenus Anapsilana) and Hi spaniola (both Cirolana, subgenus Anapsilana). The Netherlands Antilles include the islands of Aruba, Bonai re and Cura<;ao _i n the southern Caribbean. The islands formed as a result of volcanic activity 90-95 million years ago, but half of the present day surface consists of Neogene or Quaternary limestone. Caves are generally small , but stygobitic fauna is also found in karstic springs, wells, anchialine pools and the marine interstitial. One stygobitic ciroland (Aruba/ana) is known from Aruba. The Cayman Islands are co mposed of two distinct limeston_e formations; dense and highly karstified Oligocene-Mioce ne limestone forms the central core of each island, and is surrounded by a coastal limestone terrace of Pleistocene age ca lled " ironshore". The island of Cayman Brae, due to its greater elevation , has more numerous and larger caves. One speci es of stygobiti c cirolanid ( Cirolana, subgenus An.apsilana) is known from Grand Cayman. One stygobitic cirolanid, Zulialana caalescens, is described from the Sierra de Perija in northwestern Venezuela. This regio n is a tropical rainforest with well developed karst in Cretaceous limestone. I I Caribbean stygobitic Cirolanidae The stygobitic cirolanid, Antrolana lira, is restricted to the Shenandoah Valley including caves in Augusta County, Virginia and Jefferson County, West Virginia. This isopod inhabits pools that intersect the groundwater table. Limestone in this area is Upper Cambrian in age. Caves and groundwater habitats in Central and West Texas including the Balcones Escarpment and Edwards Plateau are inhabited by two stygobitic cirolanids, Speocirolana hardeni and Cirolanides texensis texensis. The Edwards Aquifer which underlies this area has the highest diversity of stygobites in the United States (LONGLEY, 1986). The Edwards limestone which outcrops over most of this area was deposited in a shallow sea starting in the early Cretaceous. The receding shoreline passed through this area in the early Tertiary. The Sierra Madre Oriental of northeastern Mexico contains a wide variety of groundwater habitats (e.g., caves, thermal springs, wells, mines, and spring-fed pools) and an exceptionally diverse assortment of stygobitic cirolanids including 16 species from 4 genera (Cirolanides, Mexilana , Speocirolana and Sphaerolana). All of these systems contain freshwater, although water in thermal springs can be highly mineralized. During the Mesozoic, a thick layer of limestone was laid down in the shallow sea extending from south central Texas, across Mexico and Belize, and into Guatemala. In the late Cretaceous and early Tertiary, a series of mountainbuilding events known as the Laramide Orogeny uplifted, distorted and compressed these rocks creating the coastal ranges of the Sierra Madre Oriental. Water salinity apparently being the best possible "common denominator", we make an attempt to group together the species considered in this paper according to their affiliation to either freshwater or marine habitats. Needless to say, in rather many cases, it is difficult to appreciate correctly the situation, owing to problems with obtaining reliable data during difficult exploration, or brought about by sampling quite near Uust above or just below) haloclines. Moreover, some species are certainly more or less euryhaline. For a few species - included or not in Table 1 - the situation is at present rather problematic (Arubolana imula and parvioculata, Bahalana cardiopus, Speocirola.na thermydronis, Yuca.tala.na robustispina.: see explanatory notes under "The actors"). 7 Genus Antrolana BOWMAN, 1964 I. Antrolana lira BOWMAN, 1964 Fig. 7 COLLINS & HOLSINGER, 1981 ; HOLSINGER et a.l. , 1994; HOLSINGER, 2005 o,S? Phreatic water in ten caves in the Shenandoah Valley karst region of northwestern Virginia and eastern West Virginia, USA Freshwater Genus Arubolana BOTOSANEANU & STOCK, 1979 2. Arubolana aruboides (BOWMAN & ILIFFE, 1983) Originally described in Bermuda/ana n.g. ; transferred by NOTENBOOM, 1984, Bennudalana being synonymized with Arubola.na.. NOTENBOOM, 1984; BOTOSANEANU & ILIFFE, 1997 o,S? Bermuda, Hamilton Parish: Church Cave (Paynter's Vale Cave), Wonderland Cave (Whitby Cave, Fantasy Cave), Bitumen Cave, Wilkinson Quarry Cave (this last cave: new record) Salt water; salinity variable with depth but species found only in deeper, salt water 3. Arubola.na imula BOTOSANEANU & STOCK, 1979 Fig. 16 NOTENBOOM, 1984 o,S? Aruba by Lago Colony: Mangel Cora Tunnel Fully marine water(?), although in original description " . .. brackish (chlorinity 3600 mg/1)" 4. Arubola.na parvioculata. NOTENBOOM, 1984 o,S? The actors Here follows a concise presentation of all 42 described stygobitic species (one of them with two subspecies) of the peri-Caribbean and Mexican realm, in alphabetical order. For each species: publications, other than that containing the original description, bringing some relevant additional information ; male/female described; known distribution; habitats of freshwater/saline water; some additional information. To these 42 species, two more should be added: that, from Cuba, mentioned in the "Appendix to the check-list" and described as Bahalana bowman.i; and a new species of Bahala.na from Abaco, the Bahamas, described by BOTOSANEANU & JLJ FFE in a separate paper in this same volume of the Bulletin (pp. 27-31). Jamaica, Discovery Bay: interstitial in dry bed of Rio Secco near sea shore (possibly also in marine interstitial) Water salinity very different in various holes dug in sediment Microoculate Genus Baiza/ana CARPENTER, 1981 5. Bahalana. ca.icosa.na. BOTOSANEANU & ILIFFE, 2003a Plate I (a) o,S? North Caicos Island, Sandy Point: Cottage Pond (anchialine cave). Middle Caicos Island , Conch Bar: Conch Bar Cave Fully marine salinity '' 8 T. M. ILLFFE & LAZARE BOTOSANEANU -2 ) 30° 30° 37, 38, 39,40 30 20° ->g_) 41 31 29 28, 33 26, 27, 34, 35, 36 J2 ~9: 7 , ~\ ~6? 16, 20 )6- 5 ~ 11 .... . . 20° -4 CJ..: -:_· ? '\ If>; ~ \) km 1"""'1 0 1"""'1 400 Figure 1: Map of the peri-Caribbean- Mexican zone illustrating the distribution of endemic stygobitic Cirolanidae. Numbers refer to the species presented in that order under the section entitled "The actors": l. Antra/ana lira, 2. Aruba/ana arubaides, 3. Aruba/ana imula, 4. Aruba/ana parviaculata, 5. Bahalana caicasana, 6. Bahalana cardiapus, 7. Bahalana exumina, 8. Bahalana geracei, 9. Bahalana yagerae, LO. Ciralana (Anapsilana) acanthura, II. Ciralana (Anapsilana) crenata, 12. Ciralana (Anapsilana) cubensis, 13. Ciralana (Anapsilana) pleascissa , 14. Ciralana (Anapsilana) radicicala, 15. Ciralana (Anapsilana) yucatani:l, 16. Ciralana (Ciralana) traglexuma, 17. Ciralanides texensis texensis, 18. Ciralanides texensis rnexicensis , 19. Creaseriella anaps, 20. Exumalana replans , 21. Haptalana belizana, 22. Haptalana bawmani, 23 . Haptalana trichastama, 24. Hapta lana yunca, 25. Metaciralana mayana, 26. Mexilana salupasi, 27. Speaciralana bafivari, 28. Speaciralana disparicarnis, 29. Speaciralana endeca, 30. Speacirolanajustiura , 31. Speacirolana guerrai , 32. Speaciralana hardeni, 33. Speaciralana lapenita, 34. Speacirolana pelaezi, 35. Speacirolana prima, 36. Speaciralana pubens , 37. Speaciralana thermydranis , 38. Speaciralana zumbadara , 39. Sphaeralana affinis , 40. Sphaeralana interstitia/is, 41. Sphaerolana karenae, 42. Yucata!ana rabustispina, 43. Zulialana caalescens ,, Caribbean stygobitic Cirolanidae 9 Plate 1: Digital photographs of live stygobitic cirolanids: (a) Bahalana caicosana, (b) Cirolana (C.) trog lexuma, (c) Creaseriella anops, (d) Exumalana reptans, (e) Yucatalana robustispina and (f) Metacirolana mayana. '' 10 T. M. ILIFFE & LAZARE BOTOSANEANU 4 3 6 Figure 2: Speocirolana thermydronis, male (red rawn from BOTOSANEANU et al. 1998). Figure 3: S. pubens (redrawn from BOWMAN 198 1). Figure 4: S. prima, male (redrawn from SCHOTIE 2002). Figure 5 : S. pelaezi, fema le (redrawn from BOLIVAR y PI ELTA IN 1950). Figure 6: S. zwnbadora, female (redrawn fro m BOTOSANEANU et a!. 1998) II Caribbean stygobitic Cirolanidae 7 9 \ Figure 7: An.trolana lira (redrawn f rom BOWMAN 1964). Figure 8: Yucatalana robustispina, female (ori ginal drawing). Figure 9: Cirolanides texensis texensis (redrawn f rom BOWMAN 1964). Figure 10: Bahalana yagerae, male (redrawn from CARPENTER 1994). Figure 11: B. exumina, male (redrawn from BOTOSANEANU & l LI FFE 2002). 11 11 T. M. ILIFFE & LAZARE BOTOSANEANU 12 12 13 Figure 12: Exumalana replans, fema le (redrawn from BOTOS ANEANU & lLI FFE 2003). Figure 13: Zulialana coalescens (redrawn fro m & VILORI A 1993). Figure14: Creaseriella anops , fe male (ori ginal drawing). Figure 15: Sphaerolana interstitia/is, male (orig inal drawin g). B OTOSANEANU I I Caribbean stygobitic Cirolanidae -·---------________ .......... - .... 17 18 ,~ Figure 16: Arubolana imula, male (redrawn fro m BOTOSANEANU & STOCK 1979). Figure 17: Cirolana (Anopsilana) radicicola, male (redrawn from NOTENBOOM 1981). Figure 18: Mexilana saluposi, fema le (redrawn from BOWMAN 1975). Figure 19: Cirolana (Cirolana) troglexuma, female (original drawing). 13 14 T. M. ILIFFE & LAZARE BOTOSANEANU Closely related to B. geracei and B. cardiopus, not to B. yagerae or B. exumina 10. Cirolana (Anopsilana) acanrhura (NOTENBOOM, 198 1) 6. Bahalana cardiopus NOTENBOOM, 1981 Origi nall y described in Hairilana n.g. Transfen ed to Anopsilana in BOTOSANEANU er al., 1986, Haiti/ana being synonymized with Anopsilana in that publication. BOTOSANEANU & ILI FFE, 1999 0,9 Bahamas. Mayaguana Island, Little Bay: Mount Mi sery Cave; AckJins Island: Duncan Pond Cave. Water salinity, according to NOTENBOOM, 1981, only ca. 10 g/1 (but this is highly improbable) Only very slight differences from B. geracei. 9 Haiti : well at Marigot, southern coast 'of Departement de !' Ouest. Freshwater 11 . Cirolana (Anopsilana) crenata (BOWMAN & FRANZ, 1982) Origi nally described in g. Anopsilana 0,9 7. Bahalana exwnina BOTOSANEANU & ILIFFE, 2002(a) Fig. II Grand Cayman Island, NW end: West Bay Cave. Weak ly brackish water 12. Cirolana (Anopsilana) cubensis HAY, 1903 0 Bahamas, Exumas, Great Guana Cay: Oven Rock Cave Fully marine salinity Maybe not far from B. geracei and B. cardiopus 8. Bahalana geracei CARPENTER, 1981 BOLiVAR Y PIELTAIN, 1950; RIOJA, 1953, 1956 (i n this last publication as n. g. Troglocirolana) ; SILVA TABOADA, 1974. Transferred to Anopsilana in BOTOSANEANU et al., 1986, Troglocirolana being synonymized with Anopsilana in that publication. Bahamas, San Salvador Island : Dixon Hill Lighthouse Cave Fully marine salinity (but cave ca. 1 km di stant from ocean) Cuba. Pinar del Rio: Caverna en San Isidro ; Habana: Cueva between Madruga and Aguacate, Cueva del Quintana!, Cueva de Ia Yagruma. Matanzas: Cueva de Suarez. Isla de Pinos: Cueva de los Camarones, Cueva de los Murcielagos. Freshwater NOTENBOOM, 1981 o,s> o,s> 9. Bahalana yagerae (CARPENTER, 1994) Fig. 10 Originally described in Dodecalana n.g.; transferred by BOTOSANEANU & ILIFFE, 1997, Dodecalana being synonymized with Bahalana. BOTOSA NEANU & Iliffe, ] 997, 2002a, 2003a 0,9 Grand Bahama Island: Bahama Cement Cave, Old Freetown Cave, Lucayan Cavern; Sweetings Cay (adjacent to Grand Bahama Island): Janet Pyfrom's Blue Hole, Asgard Cave, Virgo Blue Hole, Sagittarius Blue Hole, Lucy's Cave; Great Ex uma Island: Basil Minn's Blue Hole ; Andros Island : Conch Sound Blue Hole. Fully marine salinity Species distinct from all others by absence of pereiopods VII in mature specimens. Genus Cirolana LEACH, 1818 (sg. Anopsilana PA ULIA N & DELAMARE DEBOUTEV ILLE, 1956) Most species in this subgenus are marine/epigean, some of them stygophilic. 13. Cirolana (Anopsilana) pleoscissa (BOTOSANEANU & ILIFFE, 1997) Originally described in Jamaica/ana n.g. Transferred by BOTOSANEANU & lLIFFE, 2000, Jamaica/ana being synonymized with Cirolana (Anopsilana) in that publication. 9 Jamaica, Westmoreland Freshwater ~arish: Revival Water Pump Cave 14. Cirolana (Anopsilana) radicicola (NOTENBOOM, 1981 ) Fig. 17 Originally described in Haitilana n. g. Transferred in BOTOSANEANU et al., 1986, Haitilana being sy nonymized with Anopsilana in that publication. 0,9 Haiti, southern peninsula, near Jerem ie: "Source Debarasse", large karstic spring flowing from cave. Freshwater 15. Cirolana (Anopsilana) yucatana BOTOSANEANU & ILI FFE, 2000 9 Mexico, Yucatan , Mucuyche: Cenote Dzonotila. '' Caribbean stygobitic Cirolanidae Freshwater Genus Cirolana LEACH, 181 8 (sg. Cirolana) Almost all described species are marine/epigean. 16. Cirolana (Cirolana) troglexuma BOTOSANEANU & ILIFFE, 1997 Fig. 19, Plate l (b) 15 Mexico, Yucatan Peninsula: numerous caves (some of them true cenotes) in the states Quintana Roo or Yucatan. To those in the various publications above mentioned should be added the following new records: from Quintana Roo - Cenote Kaape Ha (Del Mar), Systema Ox Bel Ha, Tulum; from Yucatan- Cenote San Miguel and Cenote Chean Kab, both at Huhi, and Cenote San Antonio at Homun. Freshwater Genus Exumalana BOTOSANEANU & lLIFFE, 2003(b) BOTOSANEANU & lLIFFE, 1999, 2003b. The species has been rediscovered (2003, 2004) both in the type locality and in a new one (Basil Minn 's Blue Hole); these records not published. S? Bahamas. Exuma Cays, Great Guana Cay: Oven Rock Cave; Great Exuma Island: Basil Minn 's Blue Hole; Cat Island: Big Fountain Blue Hole (this last record not yet published). Fully marine salinity S? Bahamas, Exumas, Norman's Pond Cay: Norman's Pond Cave ("inland blue hole"). Water fully marine Additional note: in BOTOSANEANU & ILIFFE, 1997, two subgenera are recognized in g. Cirolana. Genus Haptolana BOWMAN, 1966 Genus Cirolanides BENEDICT, 1896 21. Haptolana belizana BOTOSANEANU & lLIFFE, 1997 17. Cirolanides texensis texensis BENEDICT, 1896 Fig. 9 20. Exumalana reptans BOTOSANEANU & ILIFFE, 2003(b) Fig. 12, Plate l (d) 0 Belize: Inland Blue Hole, Caves Branch. Freshwater BOWMAN, 1964, 1972, ] 992; BOTOSANEANU & lLIFFE, 2002b (here, inter alia: updated distribution) 22. Haptolana bowmani BOTOSANEANU & ILIFFE, 1997 o,S? 36 records (caves, phreatic waters, springs) all from Texas (counties Bexar, Burnet, Coma!, Crockett, Edwards, Hays, Kendall, Kerr, Medina, Real, Schleicher, Terrell, Uvalde, Val Verde) all in the southern part of Edward's Plateau. Freshwater 18. Cirolanides texensis mexicensis BOTOSANEANU & ILIFFE, 2002b o,S? Mexico, Yucatan: Grutas de Tzab-Nah (Dzab Nah) 2 km S of Tecoh on road to Telchaquillo; immatures probably of the same species from three other Yucatan localities: Cenote Kambul (published as Noc Ac), Cenote Mucuyche, Cenote Yuncu. Freshwater, at least in type locality BOTOSANEANU et al., 1998; BOTOSANEANU & ILIFFE 1999. o,S? N. Mexico. Nuevo Leon, Lampazos: Cueva de El Tule, Cueva de La Espantosa; Coahuila, west and north from Ciudad Acuna: S6tano de Amezcua. Freshwater Genus Creaseriella RIOJA, 1953 19. Creaseriella anops (CREASER, 1936) Fig. 14, Plate l(c) Originally described in Cirolana; transferred by RIOJA, 1953 to Creaseriella n.g. Creaser, 1938 ; BOLIVAR Y PI ELTAlN, 1950; RIOJA, 1953; REDDELL, 1977; PEREZ ARANDA, 1984; BOTOSANEANU & ILIFFE, 1997, 1999, 2002a; ESCOBAR et al. , 2002. o,S? 23. Haptolanrt trichostoma BOWMAN, 1966 SILVA TABOADA, 1974 0 Cuba, Province Camaguey, Sierra de Cubita: Cueva Bonita, Cueva de Ia Lechuza. Freshwater Cons idered by BOTOSANEANU & ILIFFE, 2000 as not being closely related to H. belizana, H. bowmani and H. yunca which could have a common marine ancestor. 24. Haptolana yunca BOTOSANEANU & ILIFFE, 2000 S? Mexico, Yucatan, Yuncu: (deep) Cenote Sabakha. Caught in freshwater near halocline II 16 T. M. ILIFFE & LAZARE BOTOSANEANU 28. Speocirolana disparicomis BOTOSANEANU & IUFFE, 1999 Genus Metacirolana NIERSTRASZ, 1931 The bulk of species in this genus are marine/epigean. 3,9 25. Metacirolana mayana (BOWMAN, 1987) Plate 1(f) Originally described as Bahalana; transferred by BOTOSANEANU & ILIFFE, 2002a. BOTOSANEANU & lL!FFE, 1997, 1999, 2002a; CARPENTER, 1994. 3,9 Mexico, Yucatan, Quintana Roo. Isla Cozumel: Cueva Quebrada, Cenote Aero! ito, Cenote Tres Potrillos. Other loca]jties in Quintana Roo: Cenote Temple of Doom Cave (near Tulum), Cenote Crustacea, originally published as Aayin Aak (Puerto Morelos), Cenote 27 Steps (Akumal). One additional known locality in Quintana Roo has never been published: Cenote Chac Moo! (Puerto Aventuras). Fully marine water Genus Mexilana BOWMAN, 1975 26. Mexilana saluposi BOWMAN, 1975 Fig. 18 3,9 Mexico, Tamaulipas, Jaumave: Guayatejo spring Freshwater 29. Speocirolana endeca BOWMAN, 1981 3,9 Mexico, Tamaulipas, NW of Ciudad Victoria: S6tano de las Calenturas (Yerbabuena); Cueva del Tecolote (Los San Pedro). Probably freshwater 30. Speocirolana just iura BOTOSANEANU & ILIFFE, 1999 c) Mex ico, Nuevo Le6n, Potreritos: Sima Chupacab1e Freshwater 31. Speocirolana guerrai CONTRERAS-BALDERAS & PURATA-VELARDE, 1982 3,9 Mex ico, Nuevo Leon, Linares: Cueva de Ia Chon-era Freshwater Considered as more closely related to S. pubens Mexico, San Luis Potosf, 4 km NW Micos: Cueva del Hui sache Probably freshwater 32. Speocirolana hardeni BOWMAN, 1992 c)' 9 Genu s Speocirolana BOLiVAR Y PIELTAIN, 1950 Origi nally described as sg. of Cirolana LEACH. Elevated to generi c rank by BOWM AN, 1964. S. Texas: several wells, artesian wells, or springs in the COUI1ties Bexar and Val Verde Freshwater Considered as most closely related to S. thermydronis 33 . Speocirolana lapenita BOTOSANEANU & ILIFFE, 1999 27. Speocirolana bolivari (RIOJA , 1953) Originally described in sg. Speocirolana of Cirolana. REDDELL & MITCHELL 1971a,b; BOWMAN 1981 ; BOTOSANEANU & ILIFFE, 1999; BOTOSANEANU et al., 1998. 3,9 Mexico. Tamaulipas: Cueva (Grutas) de Quintero (Sierra de El Abra); "Bee Cave" (S ierra de Guatemala); Cueva del Nacimiento del Rfo Frio (Municipio Gomez Farias, Ej ido El Nacimiento) ; Ojo Encantado (Jaumave, Canon Trejo). San Luis Potosf: spring at La Laja. Nuevo Leon: Sima Chupacable (Potreritos). Freshwater Disc ussion of variability, characters enab ling or not di stinction from the closely related and so metimes coexisting S. pelaezi, possibility of hybridi zation, relative abundance of these two spec ies: RIOJA , 1953 ; COLE & MrNCKLEY, 1966; MINCKLEY & COLE, 1968; BOWMAN, 198 1; BOTOSANEANU & lLI FFE, 1999; BOTOSANEANU et al., 1998. 3,9 Mex ico, Tamaulipas, Ciudad Victoria: Manantial La Penita Freshwater Apparently related to S. guerrai and S. pubens. 34. Speocirolana pelaezi (BOLiVAR Y PIELTAIN, 1950) Fig. 5 Originally described in sg. Speocirolana of Cirolana. RIOJA, 1953; REDDELL & MITCHELL, 1971 a, b; BOTOSANEANU et al., 1998; BOTOSANEANU & lLIFFE, 1999. 3,9 Mex ico. San Luis Potosf: Cueva de los Sabinos, Sotano de Pichijumo, Sotano de las Piedras, S6tano del Tigre, Sotano de Ia Tinaja, S6tano del Arroyo, Cueva Chica, Cueva de Ia Curva, Sotaruto de Montecillos. Tamauli pas, Sierra de El Abra: Cueva de Ia Florida, Cueva de El Pachon, Grutas Caribbean stygobitic Cirolanidae (Cueva) de Quintero; Tamaulipas, Sierra de Guatemala: Cueva de Ia Mina; also in Tamaulipas are: Cueva del Nacimiento de l Rfo Frfo (Municipio Gomez Farias, Ejido El Nacimiento) and Ojo Encantado (Jaumave, Canon Trejo). Freshwater For various observations: see last paragraph of text for S. bolivari 35. Speocirolana prima SCHOTIE, 2002 Fig. 4 o,<i' Mexico, Tamaulipas: Nacimiento de R fo Mante just south of Ciudad Mante Freshwater " .. . most like S. pubens ... " 36. Speocirolana pubens BOWMAN, 1981 Fig. 3 o,<i' Cuatro Cienegas; Cueva La Zumbadora (Municipio La Madrid). Nuevo Leon: flooded mine near Cueva de Ia Boca (Villa Santiago) Freshwater 40. Sphaerolana interstitialis COLE & M!NCKLEY, 1970 Fig. 15 COLE, 1984; BOTOSANEANU et al., 1998 o , <i' Mexico. Coahuila: small springs and " pozos" near Cuatro Cienegas. Tamaulipas: Manantial de San Rafael de los Castro (Municipio Ciudad Mante). Freshwater 41. Sphaerolana karenae RODRIGUEZ-ALMARAZ & BOWMAN, 1995 0 Mex ico, Nuevo Leon: small spring adjacent to Pilon River, Montemorelos; mine near Cueva de Ia Boca, (Villa) Santiago Freshwater Mexico. San Luis Potosf, San Nicolas de los Montes: Cueva de Ia Bonita; Tamaulipas, 14 km of Ocampo: Cueva de l Ojo de Agua de Manantiales Freshwater Considered as more closely related to S. guerrai 37. Speocirolana thermydronis COLE & MINCKLEY, 1966 Fig. 2 MlNCKLEY & COLE, 1968; COLE, 1984; BOWMAN 1992; BOTOSANEANU et al., 1998. o,<i' Mexico, Coahuila: several " pozos" (thermal springs and associated habitats - marshes) SW of C uatro Cienegas, for instance Pozos de Ia Becerra and El Majarral; El Potrero 1 and E l Potrero 2 (Municipio Melchor Muzquiz). Thermal , possibly rather highly mineralized water Considered as closely related to S. hardeni. 38. Speocirolana zumbadora BOTOSANEANU et al. , 1998 Fig. 6 <jl Mexico, Coahuila, M unicipio La Madrid: Cueva La Z umbado ra Freshwater Genus Sphaerolana COLE & MINCKLEY, 1970 39. Sphaerolana affinis COLE & M !NCKLEY, 1970 COLE 1984; BOTOSANEANU et al., 1998. o,<i' Mexico. Coahuil a: vario us " pozos" ("springfed wells") near 17 Genus Yucata/ana BOTOSANEAN U & ILIFFE, 1999 42. Yucatalana robustispina BOTOSANEANU & IUFFE, 1999 Fig. 8, Plate l (e) BOTOSANEANU & lLIFFE, 2000, 2002a o,<i' Mexico, Yucatan: Cenote Pabakal (Eknakan); Cenote Kankirixche, Cenote Kakuel , Cenote Chuih-Hol Dos (all three: Mucuyche); Cenote Xacha, Huhi, Cenote San Geronimo, Sotuta (these latte r two records not yet published). Probably caught both above or below the halocline Genus Zulialana BOTOSANEANU & VILORIA, 1993 43. Zulialana coalescens BOTOSANEANU & .VILORIA, 1993 Fig. 13 o,<i' NW Venezuela, Estado Zu lia, Sierra de Perija: Cueva de Toromo, near hacie nda Medellfn. A second population of this species has been recently discovered by Francesco F. Herrera (IVIC, Caracas) in a cave about 100 km distant fro m the type locality, but still in Sierra de Perija: Mara 2 Cave, Guasare river basin. Freshwater Appendix to the check-list In ORTIZ et al. (1997), the blind and depigme nted ci.rolanicl Bahalana bowmani is desc ribed fro m "Cue va de l Humo, Playa Giron , Provi ncia de Matanzas, C uba" (fully mar ine II 18 T. M. ILLFFE & LAZARE BOTOSANEANU water). This species is certrunly not a Bahalana, lacking the distinctive characters of this genus. Moreover, information on some relevant morphological detruls is lacking from its description and illustration. Being unable to decide about the genus to which it belongs, we could not place it in the checklist. In the same publication, Anopsilana magna n. sp. is described from a cave with fully marine water near the locality of "Bahalana bowmani". Although depigmented, this species has very large facetted eyes; being quite probably not a stygobiont, it will not be taken into consideration in this paper. The case of the extremely poorly described and unrecognizable Coni/era stygia PACKARD, 1894 from Monterrey, Nuevo Leon, Mexico, has been discussed in several publications; it is considered as probably being a species of Speocirolana. *** Several cases of coexistence of two species at one site are documented at present: Bahalana exumina and Cirolan.a (C.) troglexuma in Oven Rock Cave; B. yagerae and C. (C.) troglexuma in Basil Minn 's Blue Hole; Speocirolana bolivari and S. pelaezi in Cueva de Quintero, Cueva del Nacimiento de Rfo Frfo, and Ojo Encantado; S. bolivari and S.fustiura in Sima Chupacable; S. zumbadora and Sphaerolana affinis in Cueva La Zumbadora; Sphaerolana affinis and S. interstitia/is in springs and "pozos" near Cuatro Cienegas. Moreover, there is very probably coexistence of Cirolanides texensis texensis and of Speocirolana hardeni at some Texan sites. Likewise, coexistence will possibly be discovered also for Speocirolana thermydronis and · Sphaerolana (affin.is? interstitia/is?) at some sites in the neighbourhood of Cuatro Cienegas. lana RACOVITZA - Turcolana ARGANO & PESCE - Marocolana BOULANOUAR et al. There are about 20 species described in this complex (possibly not all will prove to be valid, whereas additional species will be described); together with fi ve species described in Sphaeromides DOLLFUS (one with one subspecies, one with two), one in Faucheria DOLLFUS & VIRE, one in Kens/eya J?RUCE & HERRANDOPEREZ, and one in Metacirolana, this would give a total of slightly more than L.5 endemic species in seven genera. There is, too, an ecological difference between the stygobitic cirolanid fauna of the two zones: genuine anchialine species are very poorly represented in the Mediterranean - possibly as result of the hypersalinity/drying-up crisis during the late Miocene (Messinian) having forced ancestors of the stygobitic taxa directly into near-littoral subterranean freshwater habitats (STOCK, 1981 ). The contrast becomes very impressive when we reach nonMediterranean Africa (one species of Cirolana (Anopsilana) known from Madagascar; two of Skotobaena FERRARA & MONOD and one of Haptolana known from the Horn of Africa), to the enormous mass of Pacific islands (3 species of Cirolana (Anopsilana) described from Sulawesi, the Philippines, and Palau Islands), and to Australia (only one species of Haptolana described; but - information in litt. from N.L. BRUCE - two or possibly three more cirolanids recently discovered from the extraordinarily productive "calcretes" of Northwestern Australia). Genera Annina BUDDE-LUND and Sahara/ana MONOD, whose species cannot be considered as clearly stygobitic, have been omitted from these considerations. In our opinion, the future will not bring drastic changes in the situation as here sketched. Some aspects of morphology in relation to systematics A uniquely high biodiversity of stygobitic Cirolanidae The peri-Caribbean and Mexican Realm is the most extraordinary area of the world in this respect, with 42 endemic species (one of them with two subspecies) described to this day, most of them belonging to 11 genera endemic for the area some of these morphologically very remarkable. Exceptional cases of specific diversification are offered by Speocirolana, Bahalana, or Cirolana (Anopsilana), and those of Sphaerolana, Haptolana, or Arubolana are, too, highly interesting. The tropics of the Western Atlantic have been for stygobitic Cirolanidae a centre of strong evo lutionary radiation (B RIGGS, 1995: 238-242 discusses the contrast with the comparatively poor faunal area represented by the tropics of the Eastern Atlantic). Undoubtedly, more discoveries can be expected , especially in Mexico, and maybe also in the practically unexplored smaller Central American COUI1tries, when their anchialine habitats are explored by diving. T here is a clear contrast even with the zone of the globe second in biodi versity: the European Mediterranean sensu lato. It is not possible to offer really exact f igures of stygobitic species for this zone, because there are still some sub j udice taxonomk problems with the generic complex Typhlociro- The vast array of morphological (and other) adaptations of Cirolanidae to hypogean life has been summarized by BOTOSANEANU (2001). Only two facts of paramount importance in the evolution of stygobiont Cirolanidae will be evoked here. First: the highly diversified pereiopodal morphology with more or less strongly raptorial structures affecting various groups of pereiopods, is, possibly, grosso modo, a result of partly abandoning feedi ng by scavenging (raptorial pereiopods are an exception in epigean-marine cirolanids) for an actively prey-grasping mode of life. Second: the occurrence in possibly phylogenetically widely distant taxa, of highly modified tailfans and especially uropod , having completely lost their natatory fu nction (a result of abandoning active sw imming for creeping in smaller bodies of water). It may be suspected that drastic remoulding of the tailfan has had a mechanical impact on the pleonal type of segmentation - considered by BOWMAN (1975) as being of prime importance for grouping cirolanid genera, as well as for realization of a morphology making more or less complete rolling into a ball possible. We believe that characters resulting from these major evolutionary events and shared by congeneric species can be considered as synapomorphies; whereas much caution is recommended when making use of them fo r deciding about I I Caribbean stygobitic Cirolanidae kinship between genera (BOTOSANEANU & VILORIA, 1993:169). As aptly worded by CAVALLI-SFORZA et al. (1994:372) "Natural selection causes convergence, or divergence in ways and directions that have nothing to do with coancestry". Characters from almost all parts of the cirolanid body have been successfully used in systematics - a partial exception being the mouthparts. Different morphological characters may have quite different evolutive (phylogenetic) strength and significance- a field for future research in cirolanid syste matics. For the time being, experience shows that, besides characters mentioned above, good tools for syste matics are those offered by: rostrum-lamina frontalis-clypeolabrum ; pleopodal morphology and setation ; the penes. 19 lifeline carrying . .. genetic reinforcement for a thriving tropical community across some 1200 km of ocean"). All this does not mean that generic attribution of some species will not be questioned in the future, as it has been in the past for other species. We can only hope that authors atte mpting revisions in the future will take into account the ele mentary fact, often neglected by cladists, that subterranean life has generated characteristics strongly blurring the image of phylogenetic relationships. The questions What can we know about ancestors of stygobitic species? About possible monophyly of some stygobitic genera This is a study of biodiversity, and by no means one of cladistics. Nevertheless, here are succinct remarks on the possible monophylg of some genera represented in the area. There were difficulties with Anopsilana (discussion of various aspects: MONOD, 1976; BOWMAN & FRANZ, 1982; BRUCE, 1981, 1992; BRUSCA etal., 1995; BOTOSANEANU & ILIFFE, 1997) considered as morphologically not sharply distinct from Cirolana (a problem having received a provisional solution by distinction of two subgenera: BOTOSANEANU & ILTFFE, 1997) or as a polyphyletic taxon- conclusion drawn, probably, from the geographical distribution, because we have not seen morphological arguments. In our opinion these difficulties have been exaggerated; we can very well imagine the origin of various members of Anopsilana across the globe either from various marine-epigean species of Cirolana (which, of course, would displace the question to: is Cirolana monophyletic, or not?) ; or, in some cases, even polytopically from various populations of one epigean Cirolana - which would be a case similar to those of Astyana.x fasciatus/Anoptichthys in Central Mexico (i.a. : WILKENS, 1981) or of Asellus aquaticus with its various "cave forms". Despite impressive diversity of habitus in various species, Speocirolana also is very probably monophyletic, monophyly being supported by a good number of shared morphological details (for instance: BOTOSANEANU et al., 1998) and also by the remarkable compact distribution of most species in northeastern Mexico (SCHOTIE, 2002: fig. 4), a fine example of speciation by geographic isolation. Shared characte rs for various species of Haptolana are summarized in BRUCE & HUMPHREYS, 1993 and in BOTOSANEANU & ILTFFE (1 997, 2000), and at least some of the m are probably synapomorphic. Bahalana is an example of a monophyletic genus (BOTOSANEANU & ILIFFE, 2003a) not needing much discussion; and a fine example of speciation by insular isolation. As to Arubolana, despite the re markable disjunct distribution of its species, there can be no doubt about monophyly (BowMAN & IL!FFE, 1983; see also B RIGGS, 1995 :238 for discussion of the role played by the Gu lf Stream as "Bermuda's There is a consensus between various authors that all stygobitic Cirolanidae in the area are of marine origin. There has been some discussion around Antra/ana lira, not questioning its marine origin but considering it as an exceptional case: the only taxon found in an area unexposed to marine water since the Paleozoic (detailed discussion in BOWMAN, 1964; see also COLLINS & HOLSINGER, 1981 ). However, recently published evide nce (KOENEMANN & HOLSINGER, 2001: fig . 26) shows that du ring middle to late Cretaceous, the localities of A. lira in Virginia were flooded by a continental marine e mbayment: which confirms the marine origin of this species. The alternative "shallow water ancestors" versus " deep sea ancestors" has been discussed in several publications not specifically devoted to Cirolanidae (i.e.: lLIFFE et al., 1983; HART et al., 1985; MANNING et al., 1986; STOCK, 1986a; ANKER & ILIFFE, 2000). Origin from taxa of shallow marine water seems clearly more plausible if the anoxia crisis with its catastrophic consequences in the Atlantic waters deeper than 200 mat the Mesozoic/Tertiary boundary, as well as at the Oligocene/Miocene boundary, are taken into consideration. Nevertheless, the proble m cannot be considered as definitely settled. That the immediate ancestors were animals well preadapted to hypogean life seems to be a strongly fo unded opinion (importance of preadaptation in the case of Cirolanidae has been stressed by BOTOSANEANU, 2001). Ve ry little concrete e vidence can be gathered, without falling into wild speculation, about the actual ancestors. It should be kept in mind that " the subte rranean ofte n convergent evolution of characters might obscure the phylogenetic relationships between marine epigean and stygobiont taxa" (GALASSI, 1997); and the possibility of speciation, in some cases, from already stygobitic ancestors (HOLSINGER, 2004) should not be excluded. The re is more solid evide nce fo r the species of sg. Anopsilana, very probably derived from marine-epigean Cirolana spp., and for the subterranean Metacirolana, fo r which descent from congeneric marine/ epigean species is qu ite plausible (according to KENSLEY & SCHOTIE, 1989, fro m epigean marine waters of the Cari bbean, not including the G ulf of Mexico, two species of Anopsilana, five of Cirolana, and four of Metacirolana are known). 20 T. M. IUFFE & LAZARE BOTOSANEANU Can the age of stygobitic taxa and lineages be appreciated with some degree of reliability? This is a ticklish question. To correlate age with the geological age of the rock in whkh the habitat of the stygobiont is developed , would be simply absurd (MONOD, 1975). It would be equally wrong to correlate it with the supposed time of emergence above sea level of its habitat, because it is quite possible (and in some cases quite probable) that a long period of stygoevolution below sea level has preceded this emergence. Several authors have published ideas in this context (HART et al., 1985; STOCK, 1986a, 1994; ILIFFE, 1990; ANKER & lLIFFE, 2000; KORNICKER & ILIFFE, 2000; and interestingly, some additional evidence seems to come from palaeontology: LOZOUET, 2004). This idea is aptly worded, for instance, in STOCK (1994:109-110): " ... stygobionts did not need subaerial edifices to evolve, their marine ancestors could have Jived on shallow submerged banks or sea mounts and then adapted to subaerial conditions after such banks arose above sea-level" . Can the age of a taxon or lineage be correctly inferred from morphology? If unconditionally answering with "yes", this would imply that we believe that rates of stygoevolution are identical in all lineages - something which cannot be accepted. Nevertheless, a morphological diversification like that seen in Speocirolana could be evidence for long-lasting evolution. Genera in the fauna of the zone, with morphology most radically moulded by subterranean life, are Sphaerolana, Zulialana, Exumalana , and Yucatalana: are they all very o ld/ the oldest? In some cases possibly yes (Sphaerolana , Zulialana), in some. others possibly not (Exumalana, Yucatalana); if such speculation can be made, it is because morphological evidence seems to be supported by evidence from habitat and distribution . At the opposite end of the spectrum we have the least radically transformed taxa: still macro-oculate like Anopsilana magna, or, for instance, micro-oculate like Arubolana parvioculata: in such cases and possibly also in that of Cirolana (Cirolana) troglexuma there is serious evidence for relatively recent stygobitization. What can we guess about the roots of pr·esent day diversity? Various authors have expressed opinions (sometimes contradictory- even in publications by the same author!) about time of g roundwater colonization in various parts of the peri-Caribbean/Mexican Realm . Here follows an incomplete list of such publications. • Concerning more than one geographic unit belonging to our zone: ALVAREZ et al. , 2005; HART et al. , 1985 ; HOLSI NGER, 2000 ; HUMPHREYS , 2000 ; lLIFFE, 1992a, 2000, 2005 ; MONOD, 1975 ; NOTENBOOM , 1984; SK ET, 1996, 2005 ; STOCK, 1986a, 1994. • Bermuda: ANKER & lUFFE, 2000; BOWMAN & IUFFE, 1983; BRlGGS, 1995 ; lLIFFE, 1994, 2003 , 2004; lLIFFE et al. , 1983 ; HART eta!., 1985; MANNING eta!. , 1986; SKET & lLI FFE, 1980; STERRER, 1998. • Texas: HEN DRICKSON & KREJCA , 2000 ; HOLSINGER & LONGLEY, 1980 ; LONGLEY, 1986, 2004; MITCHELL & REDDELL, 1971. • Mainland of Mexico: ARGANO, 1972; BOTOSANEANU et a!., 1998; COLE, 1984; COLE & MINCKLEY, 1966, 1970; HENDRICKSON & KREJCA , 2000; VILLALOBOS eta!., 1999. • Yucatan Peninsula: lLIFFE, 1992b, 1993; REDDELL, 1977; SCHMITTER-SOTO et a!. , 2002; SUAREZ-MORALES et a!. , 2004; WILKENS, 1982. • Bahamas, Turks and Caicos: ALVAREZ et a/., 2004; BOTOSANEANU & lLIFFE, 2003a,b; FOSSHAGEN & lLIFFE, 2004a,b; KOENEMANN et al. , 2003 , 2004; KORNICKER et al. , 2002. • Greater Antilles: BRIGGS, 1984; KORNICKER & lLIFFE, 1992; JAUME, 2001; JUARRERO & ORTIZ, 2000; STOCK, ] 977, 1981, 1986b. • Aruba: BOTOSANEANU & STOCK, 1979. • Sierra de Perija (Venezuela): BOTOSANEANU & VJLORlA , 1993. 1\vo types of habitat requiring different explanations for · time and mode of groundwater colonization Despite differences in interpretation of known facts by various authors, most situations may be reduced to two main types of habitat and groundwater colonization history. The author having most consistently considered this problem, offering solid generalizations is STOCK (l986a, 1986b, 1994), and we shall mainly borrow ideas from these publications. STOCK (1986a:929-930) criticizes the idea that "all stygobionts, be they high-salinity or low-salinity ones, have supposedly evolved to an identical evolutionary scenario and time-scale". Zones of two distinct types can be distinguished in the peri-Caribbean and Mexican Realm: a). Lower ("sinking", or "subsiding") zones more or less recently emerged form the sea, habitats with fully or nearly fu lly marine water and still in (indirect) contact with the sea: Bahamas and Turks and Caicos, Cayman Islands , YucatanBelize-Cozumel , near-shore localities of some greater Antilles. For such zones a "passive" (" regression") model of colonization of relatively recent date is proposed , with shorter periods available for stygoevolution. b). Higher ("emerging", or " rising") zones, since long/very long time not flooded ·by the sea, habitats generally with freshwater (possibly sometimes slightly brackish), and without contact with the sea: mainland of Mexico, Virginia, Texas , more or Jess inland localities on some Greater Antilles, Sierra de Perija. For such zones, an " active" model of colonization is proposed, having started long ago either by geotectonic positive movements or by eustatic regressive sea . level movements. This has allowed deeper penetration of the colonized territories (with , in some cases , impressive "climbing" of stygobionts in the mountains: Zulialana coalescens to 400 m. a.s.l. , Speocirolana thermydronis to about 600 m. , S. pubens to about 800 m., S. endeca to 1460 m., S.fustiura to more than 1500 m. a.s.l.) . In other words: the most reliable element for inferring time and mode of stygoevolution see ms to be the transition from marine to freshwater. Table 1 summarizes what we know about stygobitic species belonging to one of the two categori es. The situation of Cirolana (A nopsi/ana ) spp. 1s seem- ,, Caribbean stygobitic Cirolanidae Table I SPECIES BOUND TO FRESHWATER OR NEARLY FRESHWATER SPECIES BOUND TO FULLY OR NEARLY FULLY MARINE WATER Antra/ana lira Cira/ana (Anopsi/ana) acantlwra Cira/ana (Anapsilana) crenata Ciralana ( An a p.~ilan) cubensis Cirolana (Anapsilana) pleascissa Cirolana (Anopsilana) radicicala Cirolana (Anapsilana) yucatana Ciralanides texensis texensis Ciralanides texensis mexicensis Creaseriel/a a naps Haptalana belizana Haptalana bawmani Haptalana tric/w stama Haptalana yunca Mexilana sa/uposi Speaciralana balivari Speacira/ana disparicamis Speocirolana endeca Speaciralana Just iura Speaciralana guerrai Speacirolana hardeni Speaciralana /apenita Speacira/ana p e la e~i Speaciralana prima Speaciralana pubens Speaciralana thermydranis Speaciralana zwnbadara Sphaerolana affinis Sphaeralana interstitia/is Sphaera/ana karenae Zulialana caa/escens Aruba/ana arubaides Aruba/ana imula (?) Baha/ana caicasana Baha/ana exu111ina Baha/ana geracei Bahalana yagerae Ciralana (Ciralana) traglexu111a Exumalana replans Metaciro/ana 111ayana " Baha/ana baw111ani" (~ p ecis inquirendo) SPECIES COLLECTED FROM WATERS OF VARIABLE OR QUESTIONABLE SALINITY Aruba/ana parviaculaw Bahalana cardiapus ingly contradictory: origin probably not very old, but most species living in freshwater: a hint to importance of speed of stygoevolution in different lineages. And, from all points of view, the situation of Bermuda is a very special one. How deep are the roots? It is true that the idea has been expressed (HART et al., 1985; MANNING et al., 1986) that "marine caves of Bermuda and the Caribbean" could have been invaded as early as the Jurassic by ancestors of stygobitic species, thus before the opening of the Atlantic, on both sides of the forming ocean (and STOCK, 1986a, ag rees with this idea.) But it is like ly that the most propitious time for the starting of a flourishing period of stygoevolution in the area was the Cretaceous, a period coinciding with the opening of the Atlantic. As early as the Aptian time (ca 124 My ago) a Caribbean Province became distinct as result of the Atlantic Ocean spreading in such a manner that the Caribbean shallow waters became geographically isolated, gene flow from the East being interrupted. But it is especially during the Late Cretaceo us, aperiod vividly characterized by BRJGGS (1 995) as a mome ntous period in the Earth 's history whe n - under a greenho use climate - hig h sea leve l had resulted in the formation of extensive epicontine ntal seas, the epicontinental flooding allowing rivers to deposit sediments in the shallow sea waters. All this suggests ideal conditio ns for de ve lopme nt of the deep roots 21 of cirolanid stygoevolution and for making from the Western Atlantic a centre of evolutionary radiation (in contrast with the faunistically much poorer Eastern Atlantic) and it seems plausible that the Palaeocene (a period of djsappearance of the epicontinental seas, a period of shallow, tropical seas, as result of the fall of eustatic sea level at the e nd of the Cretaceous) was a period of (generic?) di versificafion and establishment of centres of ende mism; and that the appearance of most present day species occurred from the Miocene through the Pleistocene. What is the legacy of Tethys? The Tethys Sea, restricted prior to the Cretaceous to Medite rranean Europe and the Indo-Pacific, spread westwards in the early Cretaceous across the developing North Atlantic to the New World tropics, to form a circumtropic marine belt (ilisrupted during the M iocene). A vast amount of evidence from practically all groups of stygofauna has accumulated, pointing to the importance of the Tethys in generating distri butional patterns indubitably inherited from this circumglo bal sea, and we cannot here refer to the vast bibliography on the subject. Concerning Cirolanidae we can, first, distinguish a fine case of what could be called «a loud echo from Tethys»: that of Haptolana. From this genus, six species are described: H. trichostoma from Cuba; H. bowmani, H. belizana, and H. yunca from Yucatan; H. somata M ESSANA & CHELAZZI, 1984, from northern Somalia; and H. pholeta BRUCE & HUMPHREYS , 1993, from W. Australja (Barrow Island). This is an exemplary case of a Tethysian pattern of distribution. Morphological evidence shows that there is close relationship between the three Yucatan species, all well distinct fro m the Cuban H. trichostoma (BOTOSANEANU & IUFFE, 2000) and that there is more simjlarity between the Yucatan species and those fro m Somalia and from Australia (BOTOSANEANU & lLI FFE, 1997). O n the other hand, there are «muffled echoes» of Tethys ian descent which can be traced from re lationships between stygobitic genera a). in the Weste rn Atlantic and b). in the Mediterranean sensu lata. T,able 2 (data mainly from BOTOSANEANU et al., 1986) summarizes what we can say at present about these re lationships. It is certajn that these data will be strongly refined in the future. How could this high biodiversity be explained? 1). Through the rich, turbule nt history of this vast and fragme nted area, with many phases of marine transgressio ns/regress ions affecting its various parts, and with alte rnatingly rising and subsiding zones eithe r continental or never having been part of a continental block. C limatic changes, altho ugh not particularly severe, have certai nly been very important; a factor of paramount importance in the passive colonization of subte rranean habitats by marine ancestors of stygobionts (see BOTOSANEANU & HOLSINGER, 199 1) has been represented by the strong tro pical storms sweeping the area. With respect to all thi s, the contrast with, for instance, the eastern II 22 T. M. ILIFFE & LAZARE BOTOSANEANU Table 2: Genera of stygobitic Cirolanidae in two geographically disjunct groups with Tethysian ties. Mediterranean (sensu Jato) West Atlantic "Cirolana-group" Creaseriella Haptalana Haptalana "Sphaeromides-group" Antra/ana Ba/ralana Cirolanides Mexilana Speociralana Metacirolana Sphaeromides "Typhlocirolana-complex": Typlr/ociro/ana Turcalana Maraca/ana Metacirolana "Faucheria-group" Sphaerolana Faucheria Kensleya Skatabaena N.B. The position of Aruba/ana, Yucatalana , Zulialana, and Exumalana in such a table is at present uncertain. Atlantic is considerable. 2). Through the abundance and extraordi nary diversity of habitats propitious for stygoevolution (see "The stage" for detruls). This has too, certru nly been of paramount importance in determining the present-day diversity of Cirolanidae and other stygobionts. 3). Through the supposed abundance and high diversity of potential marine ancestors. If we consider the present-day fauna of marine-epigean C irolanidae in the area (KENSLEY & SCHOTTE, 1989 enumerate only 24 species in five (four) genera for the Caribbean not including the Gulf of Mexico; from an in litteris information from N.L. BRUCE, we learn that from the whole western North Atlantic, 35 species in 16 genera are recorded at present) this does not seem particularl y rich (STOCK, 1994 observes that "the number of stygobiont taxa is always much higher than the number of marine taxa of the same genus"). Of course, our exact knowledge about the ancestors of stygobitic Cirolanidae is extremely limited; nevertheless, there is evidence from palaeontology about the marine fauna of the area having been particularly rich duri ng the Cretaceous. Endangered or already destroyed habitats and fauna: a plea for protection. At least three stygobitic ciro lanids are curre ntly on endangered species lists. Arubolana aruboides and 24 othe r stygobitic taxa from Be rmuda are considered as "critically endangered" on the IUC N Red List. Creaseriella anops from Yucatan is listed as threatened in the official Mexican e ndangered species list - Ia Norma Of icial Mexicana (NOM059-ECOL-2001 ). Antrolana lira from Virginia is designated as threatened by the U.S. Fish and Wildlife Service. Futhermore, a number of signjf icant habitats are threate ned . C uatro C ienegas, or "Four M arshes", located in Mexico 's C hihuahuan Desert has been set aside as a biological reserve , but is still threate ned by mining of gypsum and inc reased rec- reational use. Bermuda caves inhabited by Arubolana aruboides are threatened by groundwater pollution, quarrying of limestone and development (lLIFFE et al., 1984; lLIFFE, 2003). Many cenotes along the Caribbean coastline of the Yucatan Peninsula have been set up for use by recreational scuba divers. An epigean fish, Asf) anax mexicanus, which is normally restricted to surface pools• of the cenotes has learned to follow divers into the caves, feeding off stygobitic crustaceans illuminated by the diver 's lights. Considering that some caves are visited by hundreds of divers each week, the cave fauna of the entire area has been severely impacted and in places is locally extinct. So-called "deep well injection" (to 30 to 100 m depths) of partially treated sewage and other waste waters going on in Bermuda and the Yucatan Peninsula, among other areas, is particularly troubling. Cave and groundwater are naturally depleted in dissolved oxygen due to limited contact with the atmosphere, long residence times of the water and absence of photosynthetic oxygen production. Enrichment of subterranean waters with excess organic material in the waste water stimulates growth of bacteria, consumption of the limited oxygen and results in anoxia, anaerobic hydrogen sulfide production and extermination of all aerobic organisms. Acknowledgements L. BOTOSANEANU traveled to Galveston, Texas, for two weeks of work with Th. M. ILIFFE, thanks to a grant from the Netherlands Organization for Scientific Research (NWO). Dr. N.L. BRUCE (National Institute of Water and Atmospheric Research, New Zealand) kindly replied to several questions asked by the authors and also offered other useful suggestions. In addition to a grant from the Biodiversity Surveys and Inventories Program of the National Scie nce Foundation (DEB-03 15903) toT. ILIFFE, other support and assistance with field studies has already been cited in resulting publications. References ALVAREZ, F., ILIFFE, T.M. & VILLALOBOS, J.L., 2005. New species of the genus Typhlatya (Decapoda: Atyidae) from anchialine caves in Mexico, the Bahamas, and Honduras. Journal of Crustacean Biology, 25(1): 8 1- 94. ALVAREZ, F., VILLALOBOS, J.L. & ILIFFE, T.M., 2004. A new species of Agostocaris (Caridea: Agostocarididae) from Ac klins Island, Bahamas. Proceedings of the Biological Society of Washington, 11 7(3): 368- 376. ANKER, A. & ILIFFE, T.M., 2000. Description of Bennudacaris harti, a new genus, and species (Crustacea: Decapoda: Alpheidae) from anchialine caves of Bermuda. Proceedings of the Biological Society of Washington 11 3(3): 761-775. ARGANO, R., 1972. On a troglobitic Cyathura from subterranean waters of Mexico. Academia Na-;.ionale dei Lincei an no 369: 23-34. BANARESCU, P. , 1995. Zoogeography of Fresh Waters, Vol. 3: Distribution and dispersal of freshwater ani mals in Africa, Pacific Areas and South America. AULA-Verlag, Wiesbaden. '' Caribbean stygobitic Cirolanidae BENEDICT, J.E., 1896. 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Notes on the intraspecific variability of Cirolanides texensis Benedict, 1896 (lsopoda: Cirolanidae) from Texas and Mexico. Bulletin de l'lnstitut Royal des Sciences Naturelles de Belgique, Biologie, 72: 113-117. BRUCE, N.L., 1981. Cirolanidae (Crustacea: Isopoda) of Australia: Diagnoses of Cirolana Leach, Metacirolana Nierstrasz, Neocirolana Hale, Anopsilana Paulian & Debouteville, and three new genera - Natatolana, Politolana and Cartetolana. Australian Journal of Estuarine and Freshwater Research, 32: 945-966. BOTOSANEANU, L. & lLIFFE, T.M., 2003a. A new species of the stygobitic cirolanid genus Bahalana from the Caicos Islands in the Caribbean (Isopoda: Cirolanidae). Travaux du Museum National d 'Histoire Naturelle <<Grigore Antipa», 45: 83-93. BRUCE, N.L., 1992. Anopsilana barnardi, a new species of estuarine cirolanid crustacean isopod frGm tropical eastern Australia. Memoirs of the Queensland Museum, 32(1 ), 1-8. BOTOSANEANU, L. & lLIFFE, T.M. , 2003b. 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New species of troglobitic shrimps from Mexico, with the description of Troglomexicanus, new genus (Decapoda: Palaemonidae). Journal of Crustacean Biology, 19: lll-1 22. WILKENS , H., 1982. Regressive evolution and phylogenetic age: the history of colonization of freshwaters of Yucatan by fish and crustacea. Association of Mexican Cave Studies Bulletin 8 (and Texas Memorial Museum Bulletin 28): 237-243. Thomas M. lLIFFE Department of Marine Biology Texas A&M University at Galveston Galveston, TX 77553-1675, USA E-mail: iliffet@tamug.edu Lazare BOTOSANEANU Zoologisch Museum Universiteit van Amsterdam Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands