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The Remarkable Diversity of Subterranean Cirolanidae (Crustacea: Isopoda) in the Peri -Caribbean and Mexican Realm

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The Remarkable Diversity of Subterranean Cirolanidae (Crustacea: Isopoda) in the Peri -Caribbean and Mexican Realm '' 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.
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