'' BULLETIN DE L' INSTITUT RO YAL DES SCIENCES NAT URELLES DE BELGIQ UE, BIOLOG fE , 76: 5-26, 2006 BULLETIN VAN HET KO NINKLIJK BELGISC H lNSTITUUT VOO R NATUURWETENSCHAPPEN, 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, indefatigable promoter of stygobiological research in the West Indies, almost ten years after his untimely death.

Abstract globe montre qu ' il s'agit ici d'un cas de bi odiversite remarquable­ ment grande. Sont passes en revue plusieurs aspects concern ant des With 42 endemi c species (one of them with two subspecies) de­ relati ons entre morphologie et systematique (signi fication evolutive scribed to this day, most of th em belongin g to 11 endemi c genera, de certaines adaptations morphologiques; monophylie des genres the peri-Caribbean -Mexican zone has definitely the richest faun a stygobies). On essaie de fournir des reponses a des questions of stygobitic (entirely subterranean-adapted) cirolanids of the comme: qu 'est-ce que peut etre connu sur les ancetres des especes world. After a succin ct presentation of th e various habitats for stygobies? Est-ce que !'age des taxons (ou des li gnees) stygobies stygobitic cirolanids in the zone, the species are grouped together peut etre apprecie avec un certain degre de certitude? Qu 'est-ce (Table I) according to their affili ation to habitats of freshw ater or of qu ' on peut supposer concern ant les rac ines de Ia diversite actuelle marin e water - water salinity being considered th e most reli able (deux ty pes d' habitat demandant des explications differentes pour common denominator. A check-list in alphabeti cal order follows, le temps et Ia modalite de Ia coloni sati on des eaux souterraines etant including references, known di stribution, informati on on habitats, di stingues)? Combien profondes sont ces racines? Que! est, dans etc. That this is a case of very hi gh bi odi versity of stygobitic ce cas concret, I ' heritage de Ia Tethys? Ce cas de biodiversite pour­ Ci ro lanidae results from comparison with the situati on in th e rest of rait etre explique par l'histoire riche et mouvementee de cette zone the world. Some aspects of morphology in relation to systemati cs vaste et fragmentee; par l' abond ance et l'extraordinaire diversite are revi ewed (i. a. : evoluti ve signi fican ce of some morphological des habitats propi ces a Ia stygoevolution; ainsi que par l'abond ance adaptations; monophyly of stygobitic genera). Attempts are made to et Ia di versite (s upposees) des ancetres marins potentiels. Une sec­ reply to questi ons like: what can we kn ow about ancestors of ti on fin ale du travail est consacree aux habitats et a Ia faun e mena­ stygobitic species? Can age of stygobitic taxa, or lineages, be appre­ ces OU deja detruits. ciated with some degree of reliability? What can we guess about the Mots-cJes: Cirolanidae stygobies, zone Cara1be-Mexicaine, habi­ roots of present-day di versity (two types of habitat requiring di ffe r­ tats aquatiques souterrains, bi odiversite, stygoevolution. ent expl anati on for time and mode of groundwater coloni zati on be­ ing recogni 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, turbulent hi story of this vast and fragmented area; th rough Introduction the abundance and extraordinary di ve rsity of habitats propitious for stygoevolution; and through the (s upposed) abundance and hi gh di­ Although the first subterranean cirolanid isopod of the Carib­ versity of potential marine ancestors. A fin al part of the paper is bean -Mexican zone was di_scovered more than one century devoted to endangered or already destroyed habitats and faun a, with ago, it was onl y during the last half of the previous century a plea for protecti on. that research on this group of stygobitic crustaceans has as­ Key words: stygobitic Cirol anidae, Caribbean-Mexican area, sub­ sumed large proportions, thanks to increasing interest in spe­ te rranean aquatic habitats, bi odi versity, stygoevolution leology and stygobiology and to improving techniques for exploration and zoological sampling. This has led to discov­ Resume ery and description of a remarkable large number of species entirely bound to subterranean aquatic habitats of various Avec 42 especes endemiques actu ell ement decrites (un e de cell es ­ kinds, making this zone the most extraordinary "hot spot" of ci avec deux sous-especes), appartenant pour Ia plupart a 11 genres the world for stygobitic Cirolanidae. The time seems to be endemi ques, Ia zone Cara"ibe-Mex icaine possede sans doute Ia ripe fo r attempting a synthetic presentati on of known facts, fa un e de cirolani des stygobies Ia plus ric he du monde. Apres une focusing on diversity and its possible explanation, in a natu­ presentation succin cte des divers habitats peuples par des cirolanides stygobi es dans Ia zone, les especes sont groupees (Ta­ ralisti c spirit, and limiting to a minimum reasonable specula­ bleau 1) d'apres ces habi tats: eau douce ou eau marine- Ia salinite ti on not dictated by fashions. In order to avoid this paper tak­ de l'eau etant consideree com me denominateur commun le plus di­ ing exaggerated proportions, almost no reference to gne de confiance. Une li ste des taxons en ordre alphabetique pre­ stygobionts other than Cirolanidae and to "generalized sente les references, Ia distribution connue, des info rmati ons sur les tracks" will be made. Also reference to publications not di­ habitats, etc. Une comparaison avec Ia situati on pour le reste du rectly dealing with Cirolanidae has been limited to some of II

6 T. M. ILIFFE & LAZARE BOTOSANEANU

those shedding more light on aspects directly concerning the Pleistocene and Holocene limestone, while Miocene to present paper. 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 The stage to as cenotes, a Spanish corruption of the Mayan word "dzonot". The most extensive underwater cave systems in The zone dealt with in this paper is called "Caribbean-Mexi­ the world are located along the eastern Caribbean coastline, can basin" by ARGANO (1972), "The (sub) tropical Atlantic" with Systema Ox B-:: 1Ha at 130 km being the world's longest. by STOCK (1994), "the Central American/Caribbean interme­ In the interior of the Peninsula, a well defined semicircle of diary areas" by BANARESCU ( 1995), or "Western Atlantic" by cenotes, mostly in the form of deep sinkholes, outlines the BRIGGS ( 1995) - who distinguishes here a "West Indian deeply buried rim of the 65 million year old Chicxulub im­ Province" and a "Caribbean Province". The wording used in pact crater. Groundwater throughout the Peninsula consists the title is from Stygofauna Mundi (BOTOSANEANU, 1986). of a fresh water lens of varying thickness, floating on top of Stygobitic cirolanids of the peri-Caribbean - Mexican zone denser seawater. The depth of the halocline boundary be­ inhabit wholly or partially submerged caves and tween fresh and saltwater varies from 10 m depth along the groundwater of varying salinity, all situated in karstic lime­ coast to more than 60 m in the interior. The stygobitic fauna stone. Due to locally characteristic differences in habitat of Yucatan thus includes both freshwater and marine species. type, each area within the zone will be considered separately. Seven species of stygobitic cirolanids from five genera, Generalized descriptions of habitat, geological setting, hy­ Metaciralana, Cirolana (subgenus Anapsilana), drology, water quality, and cirolanid stygofauna will be pre­ Creaseriella, Haptalana and Yu catalana, have been de­ sented. Distribution of cirolanid species within the zone is scribed from the Yucatan Peninsula (including Cozumel and illustrated in Fig. 1. Belize). Bermuda is a mid-Atlantic volcanic seamount, capped with The Greater Antilles consist of the four large islands of Pleistocene aeolian and marine limestone. Karstic caves in Puerto Rico, Hispaniola (with Haiti and the Dominican Re­ Bermuda are concentrated in a small area of the island where public), Cuba, and Jamaica. They are part of a lower Creta­ the oldest limestone outcrops. Tidal, sea level pools occur at ceous to Holocene island arc chain making up a submerged entrances and in the dark interior of the caves. Although the mountain range extending from Central America eastwards upper (dry and shallow submerged) portions of Bermuda through the Caribbean. Each of the four islands is encircled caves are dominated by collapse features, extensive horizon­ by a coastal plain, backed on the north coast of Cuba, Ja­ tal passage development typically occurs at 18m depth. The maica, and Hispaniola by Pleistocene-raised shorelines that presence of large speleothems (stalactites, stalagmites, etc.) reach heights of 300 m. The extensive limestone outcrops on in all parts of the underwater caves indicates that the entire these islands have given rise to a tropical karst landscape, known portions were completely dry and air-filled for long characterized by subterranean drainage, well developed times during glacial periods of lowered sea level. While sur­ cockpit and tower karst, and numerous caves including large face waters in Bermuda's cave pools range from slightly river caves, deep sinkholes and long horizontal caves near brackish to nearly fully marine, deeper waters below about 3- the coast that formed under the influence of palaeo-sea lev­ 5 m depths approach fully marine salinities. One stygobitic els. Two species each of stygobitic cirolanids are known cirolanid, Arubalana arubaides, is known from Bermuda. from Cuba (Haptalana and Cirolana, subgenus Anapsilana), The Bahamas archipelago consists of a series of shallow Jamaica (Aruba/ana and Ciralana, subgenus Anapsilana) banks made up of a continuous, up to 5700 m thick, sequence and Hispaniola (both Cirolana, subgenus Anapsilana). of carbonates of shallow water origin which began deposit­ The Netherlands Antilles include the islands of Aruba, ing during the Jurassic. As the banks gradually subsided, car­ Bonai re and Cura<;ao _i n the southern Caribbean. The islands bonate deposition has kept pace, mai ntaining a shallow water formed as a result of volcanic activity 90-95 million years environment. The banks are separated from one another by ago, but half of the present day surface consists of Neogene deep water channels reaching 2,000 m depths. Water-filled or Quaternary limestone. Caves are generally small, but caves in the Bahamas are locally referred to as "blue holes". stygobitic fauna is also found in karstic springs, wells, Inland blue holes occur in the interior of islands, while ocean anchialine pools and the marine interstitial. One stygobitic blue holes are situated offshore, in shallow water. While ciroland (Aruba/ana) is known from Aruba. many inland and ocean blue holes are karstic sinkholes with The Cayman Islands are composed of two distinct limeston_e no passage development, others are parts of an extensive net­ formations; dense and highly karstified Oligocene-Miocene work formed either by limestone di ssolution at the fresh/salt­ limestone forms the central core of each island, and is sur­ water inte1face (flank margin caves) or by slump faulting oc­ rounded by a coastal limestone terrace of Pleistocene age curring along the platform margin s (fault line caves). Ocean called "ironshore". The island of Cayman Brae, due to its holes typicall y have reversi ng, high velocity, tidal currents greater elevation, has more numerous and larger caves. One which flush the caves with seawater, while inland caves lack species of stygobitic cirolanid ( Cirolana, subgenus perceptible currents and have a fresh to brackish water lens An.apsilana) is known from Grand Cayman. overl ying fully marine waters. Stygobitic fa una of the Baha­ One stygobitic cirolanid, Zulialana caalescens, is described mas includes seven species of cirolanids from three genera, from the Sierra de Perija in northwestern Venezuela. This re­ Bahalana, Ciralana (subgenu s Ciralana) and Exumalana. gion is a tropical rainforest with well developed karst in Cre­ The northern coast of the Yucatan Peninsula is composed of taceous limestone. I I Caribbean stygobitic Cirolanidae 7

The stygobitic cirolanid, Antrolana lira, is restricted to the Genus Antrolana BOWMAN, 1964 Shenandoah Valley including caves in Augusta County, Vir­ ginia and Jefferson County, West Virginia. This isopod in­ I. Antrolana lira habits pools that intersect the groundwater table. Limestone BOWMAN, 1964 in this area is Upper Cambrian in age. Fig. 7 Caves and groundwater habitats in Central and West Texas COLLINS & HOLSINGER, 1981 ; HOLSINGER et a.l. , 1994; including the Balcones Escarpment and Edwards Plateau are HOLSINGER, 2005 inhabited by two stygobitic cirolanids, Speocirolana hardeni o,S? and Cirolanides texensis texensis. The Edwards Aquifer Phreatic water in ten caves in the Shenandoah Valley karst which underlies this area has the highest diversity of region of northwestern Virginia and eastern West Virginia, stygobites in the United States (LONGLEY, 1986). The USA Edwards limestone which outcrops over most of this area Freshwater was deposited in a shallow sea starting in the early Creta­ ceous. The receding shoreline passed through this area in the early Tertiary. Genus Arubolana BOTOSANEANU & STOCK, 1979 The Sierra Madre Oriental of northeastern Mexico contains a wide variety of groundwater habitats (e.g., caves, thermal 2. Arubolana aruboides springs, wells, mines, and spring-fed pools) and an excep­ (BOWMAN & ILIFFE, 1983) tionally diverse assortment of stygobitic cirolanids including 16 species from 4 genera (Cirolanides, Mexilana, Originally described in Bermuda/ana n.g. ; transferred by NOTENBOOM, 1984, Bennudalana being synonymized with Speocirolana and Sphaerolana). All of these systems contain freshwater, although water in thermal springs can be highly Arubola.na.. mineralized. During the Mesozoic, a thick layer of limestone NOTENBOOM, 1984; BOTOSANEANU & ILI FFE, 1997 was laid down in the shallow sea extending from south cen­ o,S? tral Texas, across Mexico and Belize, and into Guatemala. In Bermuda, Hamilton Parish: Church Cave (Paynter's Vale the late Cretaceous and early Tertiary, a series of mountain­ Cave), Wonderland Cave (Whitby Cave, Fantasy Cave), Bi­ building events known as the Laramide Orogeny uplifted, tumen Cave, Wilkinson Quarry Cave (this last cave: new record) distorted and compressed these rocks creating the coastal ranges of the Sierra Madre Oriental. Salt water; salinity variable with depth but species found Water salinity apparently being the best possible "common only in deeper, salt water denominator", we make an attempt to group together the spe­ cies considered in this paper according to their affiliation to 3. Arubola.na imula either freshwater or marine habitats. Needless to say, in BOTOSANEANU & STOCK, 1979 rather many cases, it is difficult to appreciate correctly the Fig. 16 situation, owing to problems with obtaining reliable data dur­ ing difficult exploration, or brought about by sampling quite NOTENBOOM, 1984 near Uust above or just below) haloclines. Moreover, some o,S? species are certainly more or less euryhaline. For a few spe­ Aruba by Lago Colony: Mangel Cora Tunnel cies - included or not in Table 1 - the situation is at present Fully marine water(?), although in original description " . .. rather problematic (Arubolana imula and parvioculata, brackish (chlorinity 3600 mg/1)" Bahalana cardiopus, Speocirola.na thermydronis, Yuca.ta­ la.na robustispina.: see explanatory notes under "The ac­ 4. Arubola.na parvioculata. tors"). NOTENBOOM, 1984 o,S? The actors Jamaica, Discovery Bay: interstitial in dry bed of Rio Secco near sea shore (possibly also in marine interstitial) Here follows a concise presentation of all 42 described Water salinity very different in various holes dug in sediment stygobitic species (one of them with two subspecies) of the Microoculate peri-Caribbean and Mexican realm, in alphabetical order. For each species: publications, other than that containing the Genus Baiza/ana CARPENTER, 1981 original description, bringing some relevant additional infor­ mation; male/female described; known distribution; habitats 5. Bahalana. ca.icosa.na. of freshwater/saline water; some additional information. To BOTOSANEANU & ILI FFE, 2003a these 42 species, two more should be added: that, from Cuba, Pl ate I (a) mentioned in the "Appendix to the check-list" and described as Bahalana bowman.i; and a new species of Bahala.na from o,S? Abaco, the Bahamas, described by BOTOSANEANU & JLJ FFE North Caicos Island, Sandy Point: Cottage Pond (anchialine in a separate paper in this same volume of the Bulletin (pp. cave). Middle Caicos Island , Conch Bar: Conch Bar Cave 27-31). Fully marine salinity '' 8 T. M. ILLFFE & LAZARE BOTOS ANEANU

-2

30° ) 30°

37, 38, 39,40 30 ->g_) 41 31 ~9: ~6? 29 J2 28, 33 7 , ~\ - 26, 27, 16, 20 )6- 5 34, 35, ...... ~ 36 20° 11 -- 20° 4 CJ..: -:_· ? '\ If>; ~ \)

km 1"""'1 1"""'1 0 400

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 (A napsilana) 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. Yu cata!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 (redrawn from BOTOSANEANU et al. 1998). Figure 3: S. pubens (redrawn from BOWMAN 1981). Figure 4: S. prima, male (redrawn from SCHOTIE 2002). Figure 5: S. pelaezi, female (redrawn from BOLIVAR y PIELTAIN 1950). Figure 6: S. zwnbadora, female (redrawn from BOTOSANEANU et a!. 1998) II Caribbean stygobitic Cirolanidae 11

9 7

\

Figure 7: An.trolana lira (redrawn f rom BOWMAN 1964). Figure 8: Yucatalana robustispina, female (original 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 LIFFE 2002). 11 12 T. M. ILIFFE & LAZARE BOTOSANEANU

12

13

Figure 12: Exumalana replans, female (redrawn from BOTOSANEANU & lLI FFE 2003). Figure 13: Zulialana coalescens (redrawn from B OTOSANEANU & VILORIA 1993). Figure14: Creaseriella anops, female (ori ginal drawing). Figure 15: Sphaerolana interstitia/is, male (original drawing). I I

Caribbean stygobitic Cirolanidae 13

-· ------

______...... - .... ,~ 17

18

Figure 16: Arubolana imula, male (redrawn from BOTOSANEANU & STOCK 1979). Figure 17: Cirolana (Anopsilana) radicicola, male (redrawn from NOTENBOOM 1981). Figure 18: Mexilana saluposi, female (redrawn from BOWMAN 1975). Figure 19: Cirolana (Cirolana) troglexuma, female (original drawing). 14 T. M. ILIFFE & LAZARE BOTOSANEANU

Closely related to B. geracei and B. cardiopus, not to B. 10. Cirolana (Anopsilana) acanrhura yagerae or B. exumina (NOTENBOOM, 198 1) Origi nall y described in Hairilana n.g. Transfen ed to 6. Bahalana cardiopus Anopsilana in BOTOSANEANU er al., 1986, Haiti/ana being NOTENBOOM, 1981 synonymized with Anopsilana in that publicati on. 9 BOTOSANEANU & ILI FFE, 1999 Haiti : well at Mari got, southern coast 'of Departement de 0,9 !'Ouest. Freshwater Bahamas. Mayaguana Island, Little Bay: Mount Mi sery Cave; AckJins Island : Duncan Pond Cave. Water salinity, according to NOTENBOOM, 1981, only ca. 10 11 . Cirolana (Anops ilana) crenata g/1 (but this is highly improbable) (BOWMAN & FR ANZ, 1982) Only very slight differences from B. geracei. Origi nally described in g. Anopsilana 0,9 7. Bahalana exwnina Grand Cayman Island, NW end: West Bay Cave. BOTOSANEANU & ILIFFE, 2002(a) Weakly brackish water Fig. II 0 12. Cirolana ( Anopsilana) cubensis Bahamas, Exumas, Great Guana Cay: Oven Rock Cave HAY, 1903 Fully marine salinity BOLiVAR Y PIELTAIN , 1950; RIOJ A, 1953, 1956 (i n this last Maybe not far from B. geracei and B. cardiopus publicati on as n. g. Troglocirolana) ; SILVA TABOADA, 1974. Transferred to Anopsilana in BOTOSANEANU et al., 1986, 8. Bahalana geracei Troglocirolana being synonymized with Anopsilana in that CARPENTER, 1981 publication. o,s> NOTENBOOM, 1981 Cuba. Pinar del Rio: Caverna en San Isidro; Habana: Cueva o,s> between Madruga and Aguacate, Cueva del Quintana!, Cueva de Ia Yagruma. Matanzas: Cueva de Suarez. Isla de Bahamas, San Salvador Island : Dixon Hill Lighthouse Cave Pinos: Cueva de los Camarones, Cueva de los Murcielagos. Fully marine salinity (but cave ca. 1 km di stant from ocean) Freshwater

9. Bahalana yagerae 13. Cirolana (Anopsilana) pleoscissa (CARPENTER, 1994) (BOTOSANEANU & ILIFFE, 1997) Fig. 10 Originally described in Jamaica/ana n.g. Transferred by Originally described in Dodecalana n.g.; transferred by BOTOSANEANU & lLIFFE, 2000, Jamaica/ana being BOTOSANEANU & ILIFFE, 1997, Dodecalana being synonymized with Cirolana (Anopsilana) in that publication. synonymized with Bahalana. 9 BOTOSA NEANU & Iliffe, ] 997, 2002a, 2003a Jamaica, Westmoreland ~arish: Revival Water Pump Cave 0,9 Freshwater Grand Bahama Island: Bahama Cement Cave, Old Freetown 14. Cirolana (Anopsilana) radicicola Cave, Lucayan Cavern; Sweetings Cay (adjacent to Grand (NOTENBOOM, 1981 ) Bahama Island): Janet Pyfrom's Blue Hole, Asgard Cave, Fig. 17 Virgo Blue Hole, Sagittarius Blue Hole, Lucy's Cave; Great Ex uma Island: Basil Minn's Blue Hole; Andros Island : Originally described in Haitilana n. g. Transferred in Conch Sound Blue Hole. BOTOSANEANU et al., 1986, Haitilana being sy nonymized Fully marine salinity with Anopsilana in that publicati on. Species distinct from all others by absence of pereiopods VII 0,9 in mature specimens. Haiti, southern peninsula, near Jeremie: "Source Debarasse", large karstic spring flowing from cave. Freshwater Genus Cirolana LEACH, 1818 (sg. Anopsilana PA ULIA N & 15. Cirolana ( Anopsilana) yucatana DELAMARE DEBOUTEV ILLE, 1956) BOTOSANEANU & ILI FFE, 2000 Most species in this subgenus are marine/epigean, some of 9 them stygophilic. Mexico, Yucatan, Mucuyche: Cenote Dzonotila. '' Caribbean stygobitic Cirolanidae 15

Freshwater Mexico, Yucatan Peninsula: numerous caves (some of them true cenotes) in the states Quintana Roo or Yucatan. To those Genus Cirolana LEACH, 181 8 in the various publications above mentioned should be added (sg. Cirolana) the following new records: from Quintana Roo - Cenote Kaape Ha (Del Mar), Systema Ox Bel Ha, Tulum; from Almost all described species are marine/epigean. Yucatan- Cenote San Miguel and Cenote Chean Kab, both at Huhi, and Cenote San Antonio at Homun. 16. Cirolana (Cirolana) troglexuma Freshwater BOTOSANEANU & ILIFFE, 1997 Fig. 19, Plate l(b) 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 20. Exumalana reptans new one (Basil Minn's Blue Hole); these records not pub­ BOTOSANEANU & ILIFFE, 2003(b) lished. Fig. 12, Plate l(d) S? Bahamas. Exuma Cays, Great Guana Cay: Oven Rock Cave; S? Great Exuma Island: Basil Minn 's Blue Hole; Cat Island: Big Bahamas, Exumas, Norman's Pond Cay: Norman's Pond Fountain Blue Hole (this last record not yet published). Cave ("inland blue hole"). Fully marine salinity Water fully marine

Additional note: in BOTOSANEANU & ILIFFE, 1997, two subgenera are recognized in g. Cirolana. Genus Haptolana BOWMAN, 1966

21. Haptolana belizana Genus Cirolanides BENEDICT, 1896 BOTOSANEANU & lLI FFE, 1997 17. Cirolanides texensis texensis 0 BENEDICT, 1896 Belize: Inland Blue Hole, Caves Branch. Fig. 9 Freshwater BOWMAN, 1964, 1972, ] 992; BOTOSANEANU & lLIFFE, 2002b (here, inter alia: updated distribution) 22. Haptolana bowmani o,S? BOTOSANEANU & ILIFFE, 1997 36 records (caves, phreatic waters, springs) all from Texas (counties Bexar, Burnet, Coma!, Crockett, Edwards, Hays, o,S? Kendall, Kerr, Medina, Real, Schleicher, Terrell, Uvalde, Val Mexico, Yucatan: Grutas de Tzab-Nah (Dzab Nah) 2 km S of Verde) all in the southern part of Edward's Plateau. Tecoh on road to Telchaquillo; immatures probably of the Freshwater same species from three other Yucatan localities: Cenote Kambul (published as Noc Ac), Cenote Mucuyche, Cenote Yuncu. 18. Cirolanides texensis mexicensis Freshwater, at least in type locality BOTOSANEANU & ILIFFE, 2002b BOTOSANEANU et al., 1998; BOTOSANEANU & ILIFFE 1999. o,S? 23. Haptolanrt trichostoma N. Mexico. Nuevo Leon, Lampazos: Cueva de El Tule, BOWMAN, 1966 Cueva de La Espantosa; Coahuila, west and north from SILVA TABOADA, 1974 Ciudad Acuna: S6tano de Amezcua. 0 Freshwater Cuba, Province Camaguey, Sierra de Cubita: Cueva Bonita, Cueva de Ia Lechuza. Genus Creaseriella RIOJA, 1953 Freshwater Considered by BOTOSANEA NU & ILIFFE, 2000 as not being 19. Creaseriella anops closely related to H. belizana, H. bowmani and H. yunca (CREASER, 1936) which could have a common marine ancestor. Fig. 14, Plate l(c) Originally described in Cirolana; transferred by RIOJA, 1953 to Creaseriella n.g. 24. Haptolana yunca BOTOSANEANU & ILIFFE, 2000 Creaser, 1938 ; BOLIVAR Y PI ELTAlN, 1950; RIOJA, 1953; REDDELL, 1977; PEREZ ARANDA, 1984; BOTOSANEANU & S? ILI FFE, 1997, 1999, 2002a; ESCOBAR et al. , 2002. Mexico, Yucatan, Yuncu: (deep) Cenote Sabakha. o,S? Caught in freshwater near halocline II 16 T. M. ILIFFE & LAZARE BOTOSANEANU

Genus Metacirolana NIERSTRASZ, 1931 28. Speocirolana disparicomis BOTOSANEANU & IUFFE, 1999 The bulk of species in this genus are marine/epigean. 3,9 Mexico, Tamaulipas, Jaumave: Guayatejo spring 25. Metacirolana mayana Freshwater (BOWMAN, 1987) Plate 1(f) 29. Speocirolana endeca Originally described as Bahalana; transferred by BOWMAN , 1981 BOTOS ANEANU & ILIFFE, 2002a. BOTOS ANEANU & lL!FFE, 1997, 1999, 2002a; CARPENTER, 3,9 1994. Mexico, Tamaulipas, NW of Ciudad Victoria: S6tano de las Calenturas (Yerbabuena); Cueva del Tecolote (Los San 3,9 Pedro). Mexico, Yucatan, Quintana Roo. Isla Cozumel: Cueva Probably freshwater Quebrada, Cenote Aero! ito, Cenote Tres Potrillos. Other lo­ ca]jties in Quintana Roo: Cenote Temple of Doom Cave (near Tulum), Cenote Crustacea, originally published as 30. Speocirolana just iura Aayin Aak (Puerto Morelos), Cenote 27 Steps (Akumal). BOTOS ANEANU & ILIFFE, 1999 One additional known locality in Quintana Roo has never c) been published: Cenote Chac Moo! (Puerto Aventuras). Mex ico, Nuevo Le6n, Potreritos: Sima Chupacab1e Fully marine water Freshwater

Genus Mexilana BOWMAN , 1975 31. Speocirolana guerrai CONTRERAS-BALDERAS & PUR ATA -VELARDE, 1982 26. Mexilana saluposi 3,9 BOWMAN, 1975 Mexico, Nuevo Leon, Linares: Cueva de Ia Chon-era Fig. 18 Freshwater 3,9 Considered as more closely related to S. pubens Mexico, San Luis Potosf, 4 km NW Micos: Cueva del Huisache 32. Speocirolana hardeni Probably freshwater BOWMAN , 1992 c)' 9 S. Texas: several wells, artesian wells, or springs in the COUI1- Genus Speocirolana BOLiVAR Y PIELTAIN, 1950 ties Bexar and Val Verde Origi nall y described as sg. of Cirolana LEACH. Elevated to Freshwater generi c rank by BOWM AN, 1964. Considered as most closely related to S. thermydronis

33 . Speocirolana lapenita 27. Speocirolana bolivari BOTOSANEANU & ILIFFE, 1999 (RIOJA , 1953) 3,9 Originally described in sg. Speocirolana of Cirolana. Mexico, Tamaulipas, Ciudad Victoria: Manantial La Penita REDDELL & MITCHELL 1971 a, b; BOWMAN 1981 ; BOTOSA­ Freshwater NEANU & ILIFFE, 1999; BOTOSANEANU et al., 1998. Apparently related to S. guerrai and S. pubens. 3,9 Mexico. Tamaulipas: Cueva (Grutas) de Quintero (Sierra de 34. Speocirolana pelaezi El Abra); "Bee Cave" (S ierra de Guatemala); Cueva del (BOLi VAR Y PI ELTA IN, 1950) Nacimiento del Rfo Frio (Municipio Gomez Farias, Ej ido El Fig. 5 Nacimiento); Ojo Encantado (Jaumave, Canon Trejo). San Luis Potosf: spring at La Laja. Nuevo Leon: Sima Originally described in sg. Speocirolana of Cirolana. Chupacable (Potreritos). RI OJA, 1953; REDDELL & MITCHELL, 1971 a, b; BOTOSA­ Freshwater NEANU et al., 1998; BOTOSANEANU & lLIFFE, 1999. Di sc ussion of variability, characters enab ling or not di stinc­ 3,9 ti on from the closely related and so metimes coexisting S. Mex ico. San Luis Potosf: Cueva de los Sabinos, Sotano de pelaezi, possibility of hybridi zation, relative abundance of Pichijumo, Sotano de las Piedras, S6tano del Tigre, Sotano these two spec ies: RIOJA , 1953 ; COLE & MrNCKLEY, 1966; de Ia Tinaja, S6tano del Arroyo, Cueva Chica, Cueva de Ia MINCKLEY & COLE, 1968; BOWMAN, 198 1; BOTOSANEANU Curva, Sotaruto de Montecillos. Tamauli pas, Sierra de El & lLI FFE, 1999; BOTOSANEANU et al., 1998. Abra: Cueva de Ia Florida, Cueva de El Pachon, Grutas Caribbean stygobitic Cirolanidae 17

(Cueva) de Quintero; Tamaulipas, Sierra de Guatemala: Cuatro Cienegas; Cueva La Zumbadora (Municipio La Ma­ Cueva de Ia Mina; also in Tamaulipas are: Cueva del drid). Nuevo Leon: fl ooded mine near Cueva de Ia Boca Nacimiento del Rfo Frfo (Municipio Gomez Farias, Ejido El (Villa Santiago) Nacimiento) and Ojo Encantado (Jaumave, Canon Trejo). Freshwater Freshwater For various observations: see last paragraph of text for S. 40. Sphaerolana interstitialis bolivari COLE & M!NCKLEY, 1970 Fig. 15 35. Speocirolana prima COLE, 1984; BOTOSANEANU et al., 1998 SCHOTIE, 2002 o ,

39. Sphaerolana affinis Appendix to the check-list COLE & M !NCKLEY, 1970 COLE 1984; BOTOSAN EANU et al., 1998. In ORTIZ et al. (1997), the blind and depigmented ci.rolanicl o,

water). This species is certrunly not a Bahalana, lacking the lana RACOVITZA - Turcolana ARGANO & PESCE - Maroco­ distinctive characters of this genus. Moreover, information lana BOULANOUAR et al. There are about 20 species de­ on some relevant morphological detruls is lacking from its scribed in this complex (possibly not all will prove to be description and illustration. Being unable to decide about the valid, whereas additional species will be described); together genus to which it belongs, we could not place it in the check­ with fi ve species described in Sphaeromides DOLLFUS (one list. In the same publication, Anopsilana magna n. sp. is de­ with one subspecies, one with two), one in Faucheria scribed from a cave with fully marine water near the locality DOLLFUS & VIRE, one in Kens/eya J?RUCE & HERRANDO­ of "Bahalana bowmani". Although depigmented, this spe­ PEREZ, and one in Metacirolana, this would give a total of cies has very large facetted eyes; being quite probably not a slightly more than L.5 endemic species in seven genera. There stygobiont, it will not be taken into consideration in this pa­ is, too, an ecological difference between the stygobitic per. cirolanid fauna of the two zones: genuine anchialine species The case of the extremely poorly described and unrecogniz­ are very poorly represented in the Mediterranean - possibly as able Coni/era stygia PACKARD, 1894 from Monterrey, result of the hypersalinity/drying-up crisis during the late Nuevo Leon, Mexico, has been discussed in several publica­ Miocene (Messinian) having forced ancestors of the tions; it is considered as probably being a species of stygobitic taxa directly into near-littoral subterranean fresh­ Speocirolana. water habitats (STOCK, 1981 ). The contrast becomes very impressive when we reach non­ *** Mediterranean Africa (one species of Cirolana (Anopsilana) known from Madagascar; two of Skotobaena FERRARA & Several cases of coexistence of two species at one site are MONOD and one of Haptolana known from the Horn of Af­ documented at present: Bahalana exumina and Cirolan.a (C.) rica), to the enormous mass of Pacific islands (3 species of troglexuma in Oven Rock Cave; B. yagerae and C. (C.) Cirolana (Anopsilana) described from Sulawesi, the Philip­ troglexuma in Basil Minn 's Blue Hole; Speocirolana bolivari pines, and Palau Islands), and to Australia (only one species and S. pelaezi in Cueva de Quintero, Cueva del Nacimiento of Haptolana described; but - information in litt. from N.L. de Rfo Frfo, and Ojo Encantado; S. bolivari and S.fustiura in BRUCE - two or possibly three more cirolanids recently dis­ Sima Chupacable; S. zumbadora and Sphaerolana affinis in covered from the extraordinarily productive "calcretes" of Cueva La Zumbadora; Sphaerolana affinis and S. Northwestern Australia). interstitia/is in springs and "pozos" near Cuatro Cienegas. Genera Annina BUDDE-LUND and Sahara/ana MONOD, Moreover, there is very probably coexistence of Cirolanides whose species cannot be considered as clearly stygobitic, texensis texensis and of Speocirolana hardeni at some Texan have been omitted from these considerations. In our opinion, sites. Likewise, coexistence will possibly be discovered also the future will not bring drastic changes in the situation as for Speocirolana thermydronis and · Sphaerolana ( affin.is? here sketched. interstitia/is?) at some sites in the neighbourhood of Cuatro Cienegas. Some aspects of morphology in relation to systematics A uniquely high biodiversity of stygobitic Cirolanidae The vast array of morphological (and other) adaptations of Cirolanidae to hypogean life has been summarized by The peri-Caribbean and Mexican Realm is the most extraor­ BOTOSANEANU (2001). Only two facts of paramount impor­ dinary area of the world in this respect, with 42 endemic spe­ tance in the evolution of stygobiont Cirolanidae will be cies (one of them with two subspecies) described to this day, evoked here. First: the highly diversified pereiopodal mor­ most of them belonging to 11 genera endemic for the area - phology with more or less strongly raptorial structures affect­ some of these morphologically very remarkable. Excep­ ing various groups of pereiopods, is, possibly, grosso modo, tional cases of specific diversification are offered by a result of partly abandoning feeding by scavenging (raptorial Speocirolana, Bahalana, or Cirolana ( Anopsilana), and pereiopods are an exception in epigean-marine cirolanids) for those of Sphaerolana, Haptolana, or Arubolana are, too, an acti vely prey-grasping mode of life. Second: the occur­ highly interesting. The tropics of the Western Atlantic have rence in possibly phylogenetically widely distant taxa, of been for stygobitic Cirolanidae a centre of strong evolution­ highly modified tailfans and especially uropod , having com­ ary radiation (B RIGGS, 1995: 238-242 discusses the contrast pletely lost their natatory fu nction (a result of abandoning with the comparati vely poor faunal area represented by the active swimming for creeping in smaller bodies of water). It tropics of the Eastern Atlantic). Undoubtedly, more discov­ may be suspected that drastic remoulding of the tailfan has eries can be expected, especially in Mexico, and maybe also had a mechanical impact on the pleonal type of segmentation in the practically unexplored smaller Central American COUI1- - considered by BOWMAN (1975) as being of prime impor­ tries, when their anchialine habitats are explored by diving. tance for grouping cirolanid genera, as well as for realization There is a clear contrast even with the zone of the globe sec­ of a morphology making more or less complete rolling into a ond in biodi versity: the European Mediterranean sensu lato. ball possible. We believe that characters resulting from these It is not possible to offer really exact figures of stygobitic major evolutionary events and shared by congeneric species species for this zone, because there are still some sub judice can be considered as synapomorphies; whereas much caution taxonomk problems with the generic complex Typhlociro- is recommended when making use of them for deciding about I I

Caribbean stygobitic Cirolanidae 19

kinship between genera (BOTOSANEANU & VILORIA, lifeline carrying ... genetic reinforcement for a thriving tropi­ 1993:169). As aptly worded by CAVALLI-SFORZA et al. cal community across some 1200 km of ocean"). (1994:372) "Natural selection causes convergence, or diver­ All this does not mean that generic attribution of some spe­ gence in ways and directions that have nothing to do with cies will not be questioned in the future, as it has been in the coancestry". past for other species. We can only hope that authors at­ Characters from almost all parts of the cirolanid body have tempting revisions in the future will take into account the el­ been successfully used in systematics - a partial exception ementary fact, often neglected by cladists, that subterranean being the mouthparts. Different morphological characters life has generated characteristics strongly blurring the image may have quite different evolutive (phylogenetic) strength of phylogenetic relationships. and significance- a field for future research in cirolanid sys­ tematics. For the time being, experience shows that, besides characters mentioned above, good tools for systematics are those offered by: rostrum-lamina frontalis-clypeolabrum; The questions pleopodal morphology and setation; the penes.

What can we know about ancestors of stygobitic species? About possible monophyly of some stygobitic genera There is a consensus between various authors that all stygobitic Cirolanidae in the area are of marine origin. There This is a study of biodiversity, and by no means one of has been some discussion around Antra/ana lira, not ques­ cladistics. Nevertheless, here are succinct remarks on the tioning its marine origin but considering it as an exceptional possible monophylg of some genera represented in the area. case: the only taxon found in an area unexposed to marine There were difficulties with Anopsilana (discussion of vari­ water since the Paleozoic (detailed discussion in BOWMAN, ous aspects: MONOD, 1976; BOWMAN & FRANZ, 1982; 1964; see also COLLINS & HOLSINGER, 1981 ). However, re­ BRUCE, 1981, 1992; BRUSCA etal., 1995; BOTOSANEANU & cently published evidence (KOENEMANN & HOLSINGER, ILIFFE, 1997) considered as morphologically not sharply dis­ 2001: fig. 26) shows that during middle to late Cretaceous, tinct from Cirolana (a problem having received a provisional the localities of A. lira in Virginia were flooded by a conti­ solution by distinction of two subgenera: BOTOSANEANU & nental marine embayment: which confirms the marine origin ILTFFE, 1997) or as a polyphyletic taxon- conclusion drawn, of this species. The alternati ve "shallow water ancestors" probably, from the geographical distribution, because we versus "deep sea ancestors" has been discussed in several have not seen morphological arguments. In our opinion publications not specifically devoted to Cirolanidae (i.e.: these difficulties have been exaggerated; we can very well lLIFFE et al., 1983; HART et al., 1985; MANNING et al., 1986; imagine the origin of various members of Anopsilana across STOCK, 1986a; ANKER & ILIFFE, 2000). Origin from taxa of the globe either from various marine-epigean species of shallow marine water seems clearly more plausible if the an­ Cirolana (which, of course, would displace the question to: is oxia crisis with its catastrophic consequences in the Atlantic Cirolana monophyletic, or not?); or, in some cases, even waters deeper than 200 mat the Mesozoic/Tertiary boundary, polytopically from various populations of one epigean as well as at the Oligocene/Miocene boundary, are taken into Cirolana - which would be a case similar to those of consideration. Nevertheless, the problem cannot be consid­ Astyana.x fasciatus/Anoptichthys in Central Mexico (i.a. : ered as definitely settled. That the immediate ancestors were WILKENS, 1981) or of Asellus aquaticus with its various animals well preadapted to hypogean life seems to be a "cave forms". strongly founded opinion (importance of preadaptation in the Despite impressive diversity of habitus in various species, case of Cirolanidae has been stressed by BOTOSANEANU, Speocirolana also is very probably monophyletic, 2001). monophyly being supported by a good number of shared Very little concrete evidence can be gathered, without falling morphological details (for instance: BOTOSANEANU et al., into wild speculation, about the actual ancestors. It should be 1998) and also by the remarkable compact distribution of kept in mind that "the subterranean often convergent evolu­ most species in northeastern Mexico (SCHOTIE, 2002: fi g. tion of characters might obscure the phylogenetic relation­ 4), a fine example of speciation by geographic isolation. ships between marine epigean and stygobiont taxa" Shared characters for various species of Haptolana are sum­ (GALASS I, 1997); and the possibility of speciation, in some marized in BR UCE & HUMPHR EYS, 1993 and in cases, from already stygobitic ancestors (HOLSINGER, 2004) BOTOSANEANU & ILTFFE (1 997, 2000), and at least some of should not be excluded. There is more solid evidence for the them are probably synapomorphic. species of sg. Anopsilana, very probably derived from ma­ Bahalana is an example of a monophyletic genus rine-epigean Cirolana spp., and for the subterranean (BOTOSANEANU & ILI FFE, 2003a) not needing much discus­ Metacirolana, for which descent from congeneric marine/ sion; and a fine example of speciation by insular isolation. epigean species is quite plausible (according to KENSLEY & As to Arubolana, despite the remarkable disjunct distribution SCHOTIE, 1989, from epigean marine waters of the Cari b­ of its species, there can be no doubt about monophyly (Bow­ bean, not including the Gulf of Mexico, two species of MAN & IL!FFE, 1983; see also B RIGGS, 1995:238 for discus­ Anopsilana, five of Cirolana, and four of Metacirolana are sion of the role played by the Gulf Stream as "Bermuda's known). 20 T. M. IUFFE & LAZARE BOTOSANEANU

Can the age of stygobitic taxa and lineages be REDDELL, 1971. appreciated with some degree of reliability? • Mainland of Mexico: ARGANO, 1972; BOTOSANEANU et a!., 1998; COLE, 1984; COLE & MINCKLEY, 1966, 1970; This is a ticklish question. To correlate age with the geologi­ HENDRICKSON & KREJCA , 2000; VILLALOBOS eta!., 1999. cal age of the rock in whkh the habitat of the stygobiont is • Yucatan Peninsula: lLIFFE, 1992b, 1993; REDDELL, 1977; developed, would be simply absurd (MONOD, 1975). It SCHMITTER-SOTO et a!. , 2002; SUAREZ-MORALES et a!. , would be equally wrong to correlate it with the supposed 2004; WILKENS, 1982. time of emergence above sea level of its habitat, because it is • Bahamas, Turks and Caicos: ALVAREZ et a/., 2004; quite possible (and in some cases quite probable) that a long BOTOSANEANU & lLIFFE, 2003a,b; FOSSHAGEN & lLIFFE, period of stygoevolution below sea level has preceded this 2004a,b; KOENEMANN et al. , 2003, 2004; KORNICKER et emergence. Several authors have published ideas in this con­ al. , 2002. text (HART et al., 1985; STOCK, 1986a, 1994; ILIFFE, 1990; • Greater Antilles: BRIGGS, 1984; KORNICKER & lLIFFE, ANKER & lLIFFE, 2000; KORNICKER & ILIFFE, 2000; and in­ 1992; JAUME, 2001; JUARRERO & ORTIZ, 2000; STOCK , terestingly, some additional evidence seems to come from ] 977, 1981, 1986b. palaeontology: LOZOUET, 2004). This idea is aptly worded, • Aruba: BOTOSANEANU & STOCK, 1979. for instance, in STOCK (1994:109-110): " ... stygobionts did • Sierra de Perija (Venezuela): BOTOSANEANU & VJLORlA , not need subaerial edifices to evolve, their marine ancestors 1993. could have Jived on shallow submerged banks or sea mounts and then adapted to subaerial conditions after such banks arose above sea-level". 1\vo types of habitat requiring different explanations for Can the age of a taxon or lineage be correctly inferred from time and mode of groundwater colonization · morphology? If unconditionally answering with "yes", this would imply that we believe that rates of stygoevolution are Despite differences in interpretation of known facts by vari­ identical in all lineages - something which cannot be ac­ ous authors, most situations may be reduced to two main cepted. Nevertheless, a morphological diversification like types of habitat and groundwater colonization history. The that seen in Speocirolana could be evidence for long-lasting author having most consistently considered this problem, of­ evolution. Genera in the fauna of the zone, with morphology fering solid generalizations is STOCK (l986a, 1986b, 1994), most radically moulded by subterranean life, are and we shall mainly borrow ideas from these publications. Sphaerolana, Zulialana, Exumalana, and Yucatalana: are STOCK (1986a:929-930) criticizes the idea that "all stygo­ they all very old/ the oldest? In some cases possibly yes bionts, be they high-salinity or low-salinity ones, have sup­ (Sphaerolana, Zulialana), in some. others possibly not posedly evolved to an identical evolutionary scenario and (Exumalana, Yu catalana); if such speculation can be made, it time-scale". Zones of two distinct types can be distinguished is because morphological evidence seems to be supported by in the peri-Caribbean and Mexican Realm: evidence from habitat and distribution. At the opposite end a). Lower ("sinking", or "subsiding") zones more or less re­ of the spectrum we have the least radically transformed taxa: cently emerged form the sea, habitats with fully or nearly still macro-oculate like Anopsilana magna, or, for instance, fu lly marine water and still in (indirect) contact with the sea: micro-oculate like Arubolana parvioculata: in such cases Bahamas and Turks and Caicos, Cayman Islands, Yucatan­ and possibly also in that of Cirolana (Cirolana) troglexuma Belize-Cozumel, near-shore localities of some greater Anti­ there is serious evidence for relatively recent stygobitization. lles. For such zones a "passive" ("regression") model of colonization of relatively recent date is proposed, with shorter periods available for stygoevolution. What can we guess about the roots of pr·esent day b). Higher ("emerging", or "rising") zones, since long/very diversity? long time not flooded ·by the sea, habitats generally with freshwater (possibly sometimes slightly brackish), and with­ Various authors have expressed opinions (sometimes contra­ out contact with the sea: mainland of Mexico, Virginia, dictory- even in publications by the same author!) about time Texas, more or Jess inland localities on some Greater Anti­ of groundwater colonization in various parts of the peri-Car­ lles, Sierra de Perija. For such zones, an "active" model of ibbean/Mexican Realm. Here follows an incomplete list of colonization is proposed, having started long ago either by such publications. geotectonic positive movements or by eustatic regressive sea . • Concerning more than one geographic unit belonging to level movements. This has allowed deeper penetration of the our zone: ALVAREZ et al. , 2005; HART et al. , 1985; colonized territories (with, in some cases, impressive "climb­ HOLSI NGER, 2000; HUMPHREYS , 2000; lLIFFE, 1992a, ing" of stygobionts in the mountains: Zulialana coalescens 2000, 2005; MONOD, 1975; NOTENBOOM , 1984; SK ET, to 400 m. a.s.l. , Speocirolana thermydronis to about 600 m. , 1996, 2005; STOCK, 1986a, 1994. S. pubens to about 800 m., S. endeca to 1460 m., S.fustiura to • Bermuda: ANKER & lUFFE, 2000; BOWMAN & IUFFE, more than 1500 m. a.s.l.). 1983; BRlGGS, 1995; lLIFFE, 1994, 2003, 2004; lLIFFE et In other words: the most reliable element for inferring time al. , 1983; HART eta!., 1985; MANNING eta!. , 1986; SKET and mode of stygoevolution seems to be the transition from & lLI FFE, 1980; STERRER, 1998. marine to freshwater. Table 1 summarizes what we know • Texas: HEN DRI CKSO N & KR EJCA , 2000; HOLSINGER & about stygobitic species belonging to one of the two catego­ LONGLEY, 1980 ; LONGLEY, 1986, 2004; MITCHELL & ri es. The situation of Cirolana (A nopsi/ana) spp. 1s seem- ,,

Caribbean stygobitic Cirolanidae 2 1

Table I of cirolanid stygoevolution and for making from the Western Atlantic a centre of evolutionary radiation (in contrast with SPECIES BOUND TO SPECIES BOUND TO FULLY the faunistically much poorer Eastern Atlantic) and it seems FRESHWATER OR NEARLY OR NEARLY FULLY MARINE FRESHWATER WATER plausible that the Palaeocene (a period of djsappearance of Antra/ana lira Aruba/ana arubaides the epicontinental seas, a period of shallow, tropical seas, as Cira/ana (Anopsi/ana) acantlwra Aruba/ana imula (?) result of the fall of eustatic sea level at the end of the Creta­ Cira/ana (Anapsilana) crenata Baha/ana caicasana ceous) was a period of (generic?) diversificafion and estab­ An ap.~ilana) Ciralana ( cubensis Baha/ana exu111ina lishment of centres of endemism; and that the appearance of Cirolana (Anapsilana) pleascissa Baha/ana geracei Cirolana (Anopsilana) radicicala Bahalana yagerae most present day species occurred from the Miocene through Cirolana (Anapsilana) yucatana Ciralana (Ciralana) traglexu111a the Pleistocene. Ciralanides texensis texensis Exumalana replans Ciralanides texensis mexicensis Metaciro/ana 111ayana Creaseriel/a a naps "Baha/ana baw111ani" What is the legacy of Tethys? Haptalana belizana (~pecies inquirendo) Haptalana bawmani SPECIES COLLECTED FROM Haptalana tric/wstama WATERS OF VARIABLE OR The Tethys Sea, restricted prior to the Cretaceous to Mediter­ Haptalana yunca QUESTIONABLE SALINITY ranean Europe and the Indo-Pacific, spread westwards in the Mexilana sa/uposi Aruba/ana parviaculaw early Cretaceous across the developing North Atlantic to the Speaciralana balivari Bahalana cardiapus New World tropics, to form a circumtropic marine belt (ilis­ Speacira/ana disparicamis Speocirolana endeca rupted during the M iocene). A vast amount of evidence from Speaciralana Just iura practically all groups of stygofauna has accumulated, point­ Speaciralana guerrai ing to the importance of the Tethys in generating distri bu­ Speacirolana hardeni tional patterns indubitably inherited from this circumglobal Speaciralana /apenita Speacira/ana pela e~i sea, and we cannot here refer to the vast bibliography on the Speaciralana prima subject. Speaciralana pubens Concerning Cirolanidae we can, first, distinguish a fine case Speaciralana thermydranis of what could be called «a loud echo from Tethys»: that of Speaciralana zwnbadara Haptolana. From this genus, six species are described: H. Sphaerolana affinis Sphaeralana interstitia/is trichostoma from Cuba; H. bowmani, H. belizana, and H. Sphaera/ana karenae yunca from Yucatan; H. somata MESSANA & CHELAZZI, Zulialana caa/escens 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. ingly contradictory: origin probably not very old, but most Morphological evidence shows that there is close relation­ species living in freshwater: a hint to importance of speed of ship between the three Yucatan species, all well distinct from stygoevolution in different lineages. And, from all points of the Cuban H. trichostoma (BOTOSANEANU & IUFFE, 2000) view, the situation of Bermuda is a very special one. and that there is more simjlarity between the Yucatan species and those from Somalia and from Australia (BOTOSANEANU & lLI FFE, 1997). How deep are the roots? On the other hand, there are «muffled echoes» of Tethys ian descent which can be traced from relationships between It is true that the idea has been expressed (HART et al., 1985; stygobitic genera a). in the Western Atlantic and b). in the MANNING et al., 1986) that "marine caves of Bermuda and Mediterranean sensu lata. T,able 2 (data mainly from the Caribbean" could have been invaded as early as the BOTOSANEANU et al., 1986) summarizes what we can say at Jurassic by ancestors of stygobitic species, thus before the present about these relationships. It is certajn that these data opening of the Atlantic, on both sides of the forming ocean will be strongly refined in the future. (and STOCK, 1986a, agrees with this idea.) But it is likely that the most propitious time for the starting of a flourishing pe­ riod of stygoevolution in the area was the Cretaceous, a pe­ How could this high biodiversity be explained? riod coinciding with the opening of the Atlantic. As early as the Aptian time (ca 124 My ago) a Caribbean Province be­ 1). Through the rich, turbulent history of this vast and frag­ came distinct as result of the Atlantic Ocean spreading in mented area, with many phases of marine transgressions/re­ such a manner that the Caribbean shallow waters became gressions affecting its various parts, and with alternatingly geographically isolated, gene flow from the East being inter­ rising and subsiding zones either continental or never having rupted. But it is especially during the Late Cretaceous, ape­ been part of a continental block. Climatic changes, although riod vividly characterized by BRJGGS (1 995) as a momentous not particularly severe, have certai nly been very important; a period in the Earth's history when - under a greenhouse cli­ factor of paramount importance in the passive colonization mate - high sea level had resulted in the formation of exten­ of subterranean habitats by marine ancestors of stygobionts sive epicontinental seas, the epicontinental flooding allowing (see BOTOSANEAN U & HOLSING ER, 199 1) has been repre­ rivers to deposit sediments in the shallow sea waters. All this sented by the strong tropical storms sweeping the area. With suggests ideal conditions for development of the deep roots respect to all this, the contrast with, for instance, the eastern II

22 T. M. ILIFFE & LAZARE BOTOSANEANU

Table 2: Genera of stygobitic Cirolanidae in two geographically disjunct reational use. Bermuda caves inhabited by Arubolana groups with Tethysian ties. aruboides are threatened by groundwater pollution, quarry­ ing of limestone and development (lLIFFE et al., 1984; lLIFFE, West Atlantic Mediterranean (sensu Jato) 2003). Many cenotes along the Caribbean coastline of the "Cirolana-group" Yucatan Peninsula have been set up for use by recreational Creaseriella Haptalana Haptalana scuba divers. An epigean fish, Asf)anax mexicanus, which is normally restricted to surface pools• of the cenotes has "Sphaeromides-group" learned to follow divers into the caves, feeding off stygobitic Antra/ana Sphaeromides crustaceans illuminated by the diver's lights. Considering Ba/ralana "Typhlocirolana-complex": that some caves are visited by hundreds of divers each week, Cirolanides Typlr/ociro/ana Mexilana Turcalana the cave fauna of the entire area has been severely impacted Speociralana Maraca/ana and in places is locally extinct. So-called "deep well injec­ Metacirolana Metacirolana tion" (to 30 to 100 m depths) of partially treated sewage and "Faucheria-group" other waste waters going on in Bermuda and the Yucatan Pe­ Sphaerolana Faucheria ninsula, among other areas, is particularly troubling. Cave Kensleya Skatabaena and groundwater are naturally depleted in dissolved oxygen due to limited contact with the atmosphere, long residence N.B. The position of Aruba/ana, Yucatalana, Zulialana, and Exumalana in times of the water and absence of photosynthetic oxygen pro­ such a table is at present uncertai n. duction. Enrichment of subterranean waters with excess or­ ganic material in the waste water stimulates growth of bacte­ ria, consumption of the limited oxygen and results in anoxia, anaerobic hydrogen sulfide production and extermination of Atlantic is considerable. all aerobic organisms. 2). Through the abundance and extraordinary diversity of habitats propitious for stygoevolution (see "The stage" for detruls). This has too, certru nly been of paramount impor­ tance in determining the present-day diversity of Cirolanidae Acknowledgements and other stygobionts. 3). Through the supposed abundance and high diversity of L. BOTOSANEANU traveled to Galveston, Texas, for two potential marine ancestors. If we consider the present-day weeks of work with Th. M. ILI FFE, thanks to a grant from the fauna of marine-epigean Cirolanidae in the area (KENSLEY & Netherlands Organization for Scientific Research (NWO). SCHOTTE, 1989 enumerate only 24 species in five (four) gen­ Dr. N.L. BRUCE (National Institute of Water and Atmos­ pheric Research, New Zealand) kindly replied to several era for the Caribbean not including the Gulf of Mexico; from questions asked by the authors and also offered other useful an in litteris information from N.L. BRUCE, we learn that suggestions. In addition to a grant from the Biodiversity Sur­ from the whole western North Atlantic, 35 species in 16 gen­ veys and Inventories Program of the National Science Foun­ era are recorded at present) this does not seem particularl y dation (DEB-0315903) toT. ILI FFE, other support and assist­ rich (STOCK, 1994 observes that "the number of stygobiont ance with field studies has already been cited in resulting taxa is always much higher than the number of marine taxa of publications. the same genus"). Of course, our exact knowledge about the ancestors of stygobitic Cirolanidae is extremely limited; nev­ ertheless, there is evidence from palaeontology about the marine fauna of the area having been particularly rich during References the Cretaceous. ALVAREZ, F., ILIFFE, T.M. & VI LLALOBOS , J.L., 2005. New species of the genus Typhlatya (Decapoda: Atyidae) from anchialine caves Endangered or already destroyed habitats and fauna: in Mexico, the Bahamas, and Honduras. Journal of Crustacean Bi­ a plea for protection. ology, 25(1): 81- 94. ALVAREZ, F., VILLALOBOS, J.L. & ILIFFE, T.M., 2004. A new species At least three stygobitic cirolanids are currently on endan­ of Agostocaris (Caridea: Agostocarididae) from Acklins Island, Ba­ gered species lists. Arubolana aruboides and 24 other hamas. Proceedings of the Biological Society of Washington, stygobitic taxa from Bermuda are considered as "critically 11 7(3): 368- 376. endangered" on the IUCN Red List. Creaseriella anops ANK ER, A. & ILI FFE, T.M., 2000. Description of Bennudacaris from Yucatan is listed as threatened in the official Mexican harti, a new genus, and species (Crustacea: Decapoda: Alpheidae) endangered species list - Ia Norma Oficial Mexicana (NOM- from anchi aline caves of Bermuda. Proceedings of the Biological 059-ECOL-2001). Antrolana lira from Virginia is designated Society of Washington 11 3(3): 761-775. as threatened by the U.S. Fish and Wildlife Service. ARGANO, R., 1972. On a troglobitic Cyathura from subterranean Futhermore, a number of signjficant habitats are threatened. waters of Mexico. Academia Na-;.ionale dei Lincei an no 369: 23-34. Cuatro Cienegas, or "Four Marshes", located in Mexico's BANAR ESCU, P. , 1995. Zoogeography of Fresh Waters, Vol. 3: Dis­ Chihuahuan Desert has been set aside as a biological reserve, tribution and dispersal of freshwater ani mals in Africa, Pacific Ar­ but is still threatened by mining of gypsum and increased rec- eas and South America. AULA-Verlag, Wiesbaden. '' Caribbean stygobitic Cirolanidae 23

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