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International Geology Review, Vol. 48, 2006, p. 791–827.Copyright © 2006 by V. H. Winston & Son, Inc. All rights reserved.

Meso-Cenozoic Caribbean Paleogeography: Implications for the Historical Biogeography of the Region

MANUEL A. ITURRALDE-VINENT1

Museo Nacional de Historia Natural, Obispo no. 61, Plaza de Armas, La Habana 10100, Cuba

Abstract

Since the latest Triassic, the Caribbean started to form as a system of rift valleys within west-central Pangea, later evolving into a mediterranean sea where distinct volcanic and non-volcanicislands evolved. Since its very early formation, this sea has been playing an important rolecontrolling the historical patterns of ocean water circulation, moderating the world climate, anddetermining the possibilities of biotic exchange of the surrounding terrestrial and marine ecosys-tems. The formation of a Mesozoic marine seaway between western Tethys and the eastern Pacific,across west-central Pangea, has been postulated for the Early Jurassic (Hettangian–Pliensbachian)according to biogeographic considerations, but supporting stratigraphic data are lacking. Probablysince the Bathonian but certainly since the Oxfordian, the stratigraphic record indicates that thisconnection was fully functional and the Circum-Tropical marine current was active. Overlanddispersal between western Laurasia (North America) and western Gondwana (South America) wasinterrupted in the Callovian when the continents were separated by a marine gap. Later, a connect-ing land bridge may have been present during the latest Campanian/Maastrichtian (~75–65 Ma),and since the Plio-Pleistocene (2.5–2.3 Ma). Evidence for a precursor bridge late in the MiddleMiocene is currently ambiguous. Since the formation of the first volcanic archipelago within theCaribbean realm at about the Jurassic–Cretaceous transition, volcanic islands, shallow banks, andridges have been present in the paleogeographic evolution of the area. However, these lands weregenerally ephemeral, and lasted just a few million years. Only after the Middle Eocene (<40 Ma)were permanent lands present within the Caribbean realm, providing substrates for the formationand development of the present terrestrial biota.

It is this independence of biological from geo-logical data that makes the comparison of thetwo so interesting because it is hard to imaginehow congruence between the two could be theresult of anything but a causal history inwhich geology acts as the independent vari-able providing opportunities for change in thedependent biological world.

— Donn E. Rosen (1985, p. 637)

Introduction

THE CARIBBEAN since its very early formation as asystem of latest Triassic–Jurassic rift valleys withinwest-central Pangea, up to its present mediterra-nean position between North, Central, and SouthAmerica and the Bahamas, has been playing animportant role controlling the historical patterns ofocean circulation, moderating world climate, and

determining the biotic exchange of surroundingterrestrial and marine ecosystems (Iturralde-Vinent,2003a).

The constantly changing geographic scenario ofthe Caribbean region provided either barriers orhighways for faunal exchange and evolution of ter-restrial and marine ecosystems. Consequently, inorder to understand the historical biogeography onemust take into account the paleogeographic history.The fact is that much of the geologic, tectonic, andeven paleogeographic literature of the Caribbeanwas written without the needs of biogeography inmind. Accordingly, biologists hoping to integrategeological information into their work, are usuallyfacing the problem of having to uncritically acceptsome paleogeographic maps that were not desig-nated to be used for biogeographic purpose (Itur-ralde-Vinent and MacPhee, 1999).

In previous papers (Iturralde-Vinent, 1982,2003a, 2003b; Iturralde-Vinent and MacPhee,1999; MacPhee and Iturralde-Vinent, 2000, 2005),general problems of the paleogeographic evolution1Email: [email protected]

7910020-6814/06/892/791-37 $25.00

792 MANUEL A. ITURRALDE-VINENT

of the Caribbean were evaluated. The main goal wasto identify the historical occurrence of land in theCaribbean and its surroundings in order to evaluatethe modern biogeographic hypothesis regarding theorigin of the past and present-day Antillean terres-trial biotas. Furthermore, utilizing almost the sameset of data, Iturralde-Vinent (2003a) generally sur-veyed the historical evolution of the Caribbean as amarine seaway. The historical biogeography of somemarine animals was also investigated in the light ofnew paleontological findings (González Ferrer andIturralde-Vinent, 2004; Myczyñski and Iturralde-Vinent, 2005; Gasparini and Iturralde-Vinent, inpress; Schweitzer et al., in press). The present paperis another step in the same direction, where theprevious results are compiled and updated withadditional information from some recent literature(Lawver et al., 1999; Mann, 1999; Damborenea,2000; Aberham, 2001; Bartolini et al., 2003;Prothero et al., 2003; Iturralde-Vinent and Lidiak,2006; etc.), as well as from new paleontological andstratigraphical information for the Mesozoic andCenozoic obtained during field work in Argentina,Jamaica, Haiti, Dominican Republic, Puerto Rico,and Cuba (De la Fuente and Iturralde-Vinent, 2001;Fernández and Iturralde-Vinent, 2000; Gaspariniand Iturralde-Vinent, 2001, in press; Iturralde-Vinent, 2001; 2003a, 2003b, 2003c; 2005;MacPhee et al., 2003; Myczyñski and Iturralde-Vinent, 2005, Schweitzer et al., in press).

Thus, the purpose of this paper is to present aseries of paleogeographic maps and to discuss someof their biogeographic implications for the origin ofCaribbean terrestrial and marine biota. The paleo-geographic reconstructions were designed for twopurposes: (1) the search for land, land bridge, andlandspan in the Caribbean; and (2) the evolution ofthe Caribbean as a marine pathway. These issueswill be analyzed within selected time intervals fromLatest Triassic to Recent. Moreover, the time inter-vals depicted in each map were chosen with abso-lute prejudice, to coincide with some importantbiogeographic events, so the paleogeographic mapscan be used to evaluate how the geographic historymay have influenced the biological events.

During this research I have honored the para-digm of Rosen (1985), quoted above, concerning thenecessary independent evaluation of the geologicand biologic data. Iturralde-Vinent and MacPhee(1999), MacPhee and Iturralde-Vinent (2000,2005), and Iturralde-Vinent (2003a, 2003c, 2005)found that in more than one example there are

conflicting biological, paleontological, and paleo-geographic data that were not resolved. The veryexistence of conflicting biological and geologicalinterpretations demonstrates that the present levelof scientific information for the Caribbean is stillinadequate, but also suggest that a multidisciplinaryapproach is needed to resolve these problems. Noisolated science or method has all the answers.2

Paleogeographic Maps, Method, and Data

The paleogeographical maps presented in thispaper have utilized the same general principlesexplained earlier (Iturralde-Vinent and MacPhee,1999), and include maps for the Jurassic, Creta-ceous, Eocene, Oligocene, Miocene, and (for Cuba)the Pliocene–late Pleistocene interval. The Trias-sic–Jurassic maps are of two categories: world maps(Fig. 1) and Caribbean maps (Fig. 2). The plate tec-tonic framework of these maps is from Lawver et al.(1999) and Marton and Buffler (1999), with minormodifications regarding the position of the Andeanand Piñon-Dagua (including Siquisique) terranes.The coastlines were first redrawn from Smith et al.(1994), but consequently updated in the areas of theEastern Pacific, Central Atlantic, the Gulf ofMexico, and the Caribbean and its surroundings(sensu Gradstein et al., 1990; Riccardi, 1991; Sal-vador, 1991; Pindell and Tabbutt, 1995; Randazzoand Jones, 1997; Iturralde-Vinent, 1998; Cordani etal., 2000). The data for the reconstruction of thepaleoenvironments are summarized in Figures 3 and4, and are from Iturralde-Vinent and MacPhee(1999), and Iturralde-Vinent (2003a, 2003c).

The paleogeographic maps for the Early andlatest Cretaceous and Lower Eocene (Fig. 5) wereconstructed taking into account the allochthonousmodel for the origin of the Caribbean plate, the datacompiled by Iturralde-Vinent and MacPhee (1999),some recent literature (Mann, 1999, Iturralde-Vinent and Lidiak, 2006), and field work in theGreater Antilles by the author. The latest Eoceneand younger maps (Figs. 6–8) are updated from Itur-ralde-Vinent and MacPhee (1999) to accomodatenew field data and recent literature (van Gestel etal., 1998, 1999; Mann, 1999; Iturralde-Vinent,

2In this paper, the geochronology is after Gradstein et al.(2004). Ma is adopted as an abbreviation for millions of years,and the tectonic term “terrane” is applied to identify alloch-thonous crustal elements (oceanic or continental in origin),which typically occur along plate boundaries.

CARIBBEAN PALEOGEOGRAPHY 793

2001; MacPhee et al. 2003) and inclusion of moredetailed information for Central America (Coatesand Obando, 1996; Denyer and Kussmaul, 2002).Furthermore, the most important paleogeographicevents are summarized in Figures 9 (marine envi-ronments) and 10 (land environments).

Caribbean Paleogeography and Some Biogeographic Implications

This section evaluates the paleogeographicscenarios of the Caribbean realm and its surround-ings from the latest Triassic to Recent, includingsome notes about paleoclimatology and paleocean-ography, and their bearing on the biogeography ofterrestrial and marine biotas. For the purpose of thisanalysis, the evolution of the Caribbean will bedivided into three stages: (1) latest Triassic andJurassic (origin of the Caribbean); (2) Cretaceous toLate Eocene (vanishing islands and land bridge inthe Caribbean), and (3) Latest Eocene to Recent(formation of the present-day Caribbean).

Latest Triassic and Jurassic:Origin of the Caribbean

At about the end of Triassic, the earth’s pre-Mesozoic continental crust was clustered as a singlesupercontinent, Pangea, a huge sialic mass that hadbeen assembled during the late Paleozoic. TheTethys Ocean extended eastward to Panthalassa, theprecursor of the Pacific Ocean, which surroundedPangea (Bullard et al., 1965). The break-up ofPangea started under such conditions as depicted inFigure 1A. The approximate westward flow of theCircum-Tropical marine current started probably inthe Bathonian, but certainly since the Oxfordian,and is suggested by physical paleoceanography(Parrish, 1992; Frakes et al., 1992), and the sus-pected migratory routes of marine invertebrates(Berggren and Hollister, 1974; Imlay, 1984; Boomerand Ballent, 1996; Damborenea, 2000) and marinevertebrates (Gasparini and Iturralde-Vinent, inpress).

Latest Triassic–Early Jurassic. During the latestTriassic–Early Jurassic (Fig. 1A), Pangea repre-sented a major barrier for the dispersion of marinebiota, as it blocked equatorial dispersion. The non-pelagic marine biota was forced to surround Pangeain order to migrate along the continental shelf, butthis possibility was controled by climatic zones(Parrish, 1992). Land biota, on the other hand,would disperse across Pangea; however, climatic

zones, mountain ranges, and the occurrence of adeveloping network of rift basins and valleys (simi-lar to present-day East African rift valleys) wouldprobably create some restrictions and/or preferredpathways for the dispersal. A major branch of theserift basins extended along the present continentalmargin of North America into the Gulf of Mexicoand the Mexican terranes, representing the earlysuture between Gondwana and Laurasia (Figs. 1,2A, and 2B; Bullard et al., 1965; Klitgord et al.,1988; Gradstein et al., 1990; Pindell and Tabbutt,1995). Therefore, the latest Triassic–Jurassic riftbasins should not be thought of as the Caribbeanbasin or the Gulf of Mexico per se, but as precursorslocated within west-central Pangea (Iturralde-Vinent and MacPhee, 1999, Iturralde-Vinent,2003c).

Intracontinental extension persisted into theEarly Jurassic, widening the rift systems with largeaquatic basins—mostly as lakes and rivers (Fig. 1).The strata filling these rift basins are usuallydescribed as redbeds, including paleosols, alluvial,and lake sediments (Salvador, 1987, 1991; Poag andValentine, 1988; Gradstein et al., 1990; Milani andTomas Filho, 2000).

During the Jurassic, the rift system locatedwithin the present coastal areas of North Americaaborted, and the break-up of Pangea shifted to a newrift system represented today by the Atlantic mid-ocean ridge and its extinct Caribbean branch.Terrestrial sediments in these new basins generallypredate the deposition of evaporitic strata in theareas that later evolved into marine basins (Evans,1978). This process is evident from the North toCentral Atlantic, where marine inundation and saltdeposition extended southward since Hettangiantime (Poag and Valentine, 1988; Gradstein et al.,1990). Within the Florida-Bahamas block, whichlater played the role of gatekeeper between theAtlantic Ocean and the Caribbean Sea, seismicsections are interpreted as representing the two riftbasin systems. One rift basin is epicontinental andfilled with clastic sediments (Sheridan et al., 1988:their Fig. 7). The only well in the Bahamas inter-secting this section was the Great Isaac, whichrecovered redbeds of Late Jurassic (probably Call-ovian) age above basement rocks (Meyerhoff andHatten, 1974; Jacobs, 1977). Another rift basin islocated in the transition between the continental andoceanic crust; its sedimentary filling is interpretedas strata of Late Jurassic and younger limestones,dolomites, and evaporites (Sheridan et al., 1988:


FIG. 1. Jurassic world paleogeographic maps after Iturralde-Vinent (2003c). This scenario suggests that dispersionof marine animals across central Pangea was only possible along the rift valley system during sea-level highstands. Sincethe Oxfordian, the Caribbean opened as a marine corridor (white arrows) and interrupted latitudinal migratory routes ofterrestrial animals.


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their Fig. 7). This interpretation is confirmed bystratigraphic sections in the southern margin of theBahamas platform that crop out in north-centralCuba, where Oxfordian? through Early Cretaceousdolomite, limestone, anhidrite, gypsite and haliteoccur (Punta Alegre, Cayo Coco, and Perros Forma-tions; Meyerhoff and Hatten, 1968, 1974; Iturralde-Vinent, 1994, 1998). These salt deposits usuallyhave been correlated with the Gulf of Mexico’sWerner-Louan salts (Meyerhoff and Hatten, 1974),but this interpretation is challenged because theybelong to different basins separated by a basementhigh (the Florida-Yucatan ridge; Figs. 2B and 2C).Iturralde-Vinent (2003c) proposed that the Baha-mian—North Cuba evaporites are indeed an exten-sion of the evaporite systems developed during theopening of the Atlantic Ocean, which is youngersouthward (Evans, 1978; Gradstein et al., 1990). Onthe African side of the Atlantic, between the Dema-rara Plateau and the Cape Verde Islands, the oldermarine rocks are of Early Cretaceous age (Hayes, etal., 1972; Jones et al., 1995).

Therefore, the presence of marine environments,even those of short duration, within the latest Trias-sic and Lower Jurassic sections of the south CentralAtlantic, Florida-Bahamas, the Gulf of Mexico,northern South America, or the Caribbean realm

cannot be conclusively demonstrated (Iturralde-Vinent, 2003c). Only in Mexico are some marineintercalations of Sinemurian–early Pliensbachianage in the Triassic throughout Lower Jurassic terres-trial deposits of the Huayacocotla Formation present(Figs. 3 and 4; López Ramos, 1975); however, thesemarine incursions seem to be due to transgressionsfrom the Pacific Ocean (Salvador, 1987, 1991).Some of the old continental margin sections of theCaribbean may have been dragged into subductionzones and lost, but this does not seem to be the casefor Florida-Bahamas (Sheridan et al., 1988), norfor the southwestern terranes of Cuba (Pinos,Escambray, and Guaniguanico; Fig. 3), becausethey yield Jurassic and probably older rocks (Fig. 4;Millán and Somin, 1981; Somin and Millán, 1981;Pszczólkowski, 1999; Iturralde-Vinent, 1994;Pindell, 1994).

The stratigraphic data previously mentioned(Iturralde-Vinent, 2003c; Figs. 3 and 4) conflictwith the biogeographic thesis which holds that sincethe Hettangian or Pliensbachian, the “HispanicCorridor” (Smith, 1983) was active as a marine routefor exchange between west Tethyan and easternPacific biotas (Gasparini, 1978, 1992; Westermann,1981, 1992; Hillebrandt, 1981; Imlay, 1984; Bartoket al., 1985; Sandoval and Westermann, 1986;

FIG. 3. Present-day location map of the continents, blocks, and tectonostratigraphic terranes that were active duringthe early opening of the Caribbean (latest Triassic–Late Jurassic) after Iturralde-Vinent (2003c). Patterns representdiferent tectonic units. Numbers on the map show the general location of the columnar sections in Figure 4.


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Hillebrandt et al., 1992; Gasparini and Fernández,1996; Damborenea, 2000; Aberham, 2001). Thispaleontological evidence leads to the propositionthat some brief marine connections took placeduring sea-level highs, allowing some shallowmarine biota to migrate across Pangea (Dambore-nea, 2000). The sea-level curve is slightly support-ive for this point of view, because since the latestTriassic until the Middle Jurassic (Bajocian), therelative position of the sea level was generally low.However, a series of high stands may have producedpartial inundation of the lower part of the reliefalong the latest Triassic–Middle Jurassic rift basinsystems located within North America, in lesserdegree along the suture of the developing NorthAtlantic Ocean (Figs. 1, 2, and 4). Although puremarine sediments of this age have not been reportedin these rift basins, they contain lagoonal depositsand local evaporites yielding fish remains includingElasmobranchia of probably marine origin (vanHouten, 1962; Smith and Robison, 1988; Poag andValentine, 1988). This evidence opens the possibil-ity that some short-time marine incursions, althoughnot properly documented in these basins, may havetaken place. Thus, the documented migration of theTethyan marine biota probably reflected short-terminfluxes of taxa from Tethys to the Eastern Pacificand vice versa, and probably took place as a resultof periodic flooding of west-central Pangea duringsea-level high stands (Parrish, 1992). In any event,this paleogeographic scenario cannot be termed a“corridor,” because it was actually a “filter” forthe dispersion of the marine biota (Gasparini andIturralde-Vinent, in press).

Another possibility, contrary to the biogeo-graphic hypothesis, may be that the marine biota didnot disperse across west-central Pangea during thelatest Triassic–Middle Jurassic times, but aroundthe supercontinent, following the southern seaway orthe Viking corridor, or by both routes (Fig. 1). Suchdispersion may have been theoretically possiblewhen polar water was not so cold, during the warmclimatic period that expanded since Permian anduntil mid-Jurassic time (Frakes et al., 1992). How-ever, the fossil record does not support such a possi-bility (Westermann, 1992; Boomer and Ballent,1996; Damborenea, 2000; Aberham, 2001).

Middle Jurassic. A widening Pangean rift systemdeveloped due to extensional stress and develop-ment of intracontinental siliciclastic basins. How-ever, the Middle Jurassic separation betweenLaurasia and Gondwana was in progress along the

mid Atlantic, Gulf of Mexico, and the intra-Carib-bean rift system (Bullard et al., 1965; Klitgord et al.,1988; Gradstein et al., 1990; Bartok, 1993; Pindelland Tabbutt, 1995; Milani and Tomas Filho, 2000).

The Jurassic paleogeographic evolution of theGulf of Mexico has been documented by severalauthors, but there is no clear agreement. Accordingto Stephan et al. (1989) and Smith et al. (1994), theGulf was a marine tongue of the Pacific that occu-pied the Mexican terranes and the western Gulf ofMexico by the Sinemurian, and later in the Batho-nian expanded as far as Florida. This interpretationcontradicts concrete seismic and stratigraphic datawhich suggest that the Gulf of Mexico was not amarine basin until the late Bathonian-Callovian(Figs. 3 and 4; Meyerhoff and Hatten, 1974; LópezRamos, 1975; Sheridan et al., 1988; Salvador, 1991;Buffler and Thomas, 1994; Pindell, 1994; Martonand Buffler, 1994, 1999). By Callovian time, theGulf had developed an oceanic crust and hyper-saline environments that covered larger areas (Figs.1 and 2; Salvador, 1987, 1991, Winker and Buffler,1988; Sawyer et al., 1991). This basin was separatedfrom the early Caribbean by a long peninsularprojection of the North American continent (a landspan, sensu Iturralde-Vinent and MacPhee, 1999)which embraced the Maya Block (Yucatan),the southeastern Gulf of Mexico, and Florida. ThisFlorida-Yucatan emerged ridge (Figs. 1 and 2) musthave had a peculiar land biota that has not yet beenproperly investigated due to limited outcrops(López-Ramos, 1975; Viniegra-O., 1981). In theGuaniguanico terrane, Oxfordian fossils of terres-trial origin have been found, including plants, ptero-saurs (Colbert, 1969; Gasparini et al., 2004), anddinosaurs (De la Torre y Callejas, 1949; Gaspariniand Iturralde-Vinent, in press), probably represent-ing part of the Florida–Yucatan ridge biota.

With regard to the Caribbean area, Middle Juras-sic marine rocks of Bajocian-Bathonian age havebeen reported from the Guaniguanico terrane ofwestern Cuba and in the Siquisique basalts of Vene-zuela (Fig. 2B; Bartok et al., 1985; Bartok, 1993).Inasmuch as the Siquisique basalts are allochtho-nous (Fig. 1; Aleman and Ramos, 2000), they arenot a sure indication of the occurrence of marineenvironments within the Caribbean. In the Guan-iguanico terrane (Fig. 3), the Lower–Middle andUpper Jurassic San Cayetano Formation is generallyinterpreted as having been deposited in a continen-tal coastal plain, with intermingled terrestrial, allu-vial, lagoonal and shallow marine beds (Haczewski,


1976). However, Lower Jurassic strata have neverbeen identified with confidence in the San CayetanoFormation (Pszczólkowski, 1978, 1999), and thoseof Middle Jurassic age are represented by: (1)?Lower–Middle Jurassic black shales with thecoastal plant Piazopteris branneri (Areces-Malléa,1990), 2) Bajocian? marine sandstone with mollusksincluding Trigonia (Vaughonia) (Krömmelbein,1956; Pszczólkowski, 1978), and (3) Bajocian–Bathonian marine shales with palinomorphs anddinoflagelates (Dueñas Jiménez and Linares, 2001).Jurassic siliciclastic and carbonate rocks with vari-ous degrees of metamorphism have also beenreported from the Pinos and Escambray terranes inCuba (Figs. 3 and 4), but the scarce marine fossilsfound in these pre-Oxfordian sections are so poorlypreserved that precise age assignment remainsproblematic (Millán, 1981; Millán and Somin, 1981,Millán and Myczyñski, 1979; Somin and Millán,1981).

Another problem related to the southwesternCuban terranes (Fig. 3; Pinos, Guaniguanico, andEscambray) is their original position, which hasbeen largely debated in recent years. Severalsources of independent data yield the conclusionthat these terranes were formed in the Caribbeanborderland of the Maya Block (Fig. 2B; for a discus-sion see Iturralde-Vinent, 1994, 1996, 1998;Bralower and Iturralde-Vinent, 1997; Hudson et al.,1999; Pszczólkowski, 1999; Pszczólkowski andMyczyñski, 2003; Pessagno et al., 1999; Schaff-hauser et al., 2003; Pindell et al., 2006).

Within Florida and the Bahamas, the occurrenceof Middle Jurassic marine sediments is not welldocumented (Fig. 4; Meyerhoff and Hatten, 1974;Randazzo and Jones, 1997), so again the possibleexistence of a marine pathway across the Florida-Bahamas area is problematic because of the lack ofstratigraphic control. Middle Jurassic marine rocksare not reported from the area of the DemararaPlateau–Cape Verde Islands (Hayes, et al., 1972;Jones et al., 1995). Within northern South America,some Lower to Middle Jurassic, partially marinesections have been reported (Tinacoa and MacoitaFormations), but the age assignment remains contro-versial (González de Juana et al., 1980; Maze, 1984;Macsotay and Peraza, 1997).

Under the circumstances, the hypothesis thatduring Middle Jurassic time the Caribbean was afully functional marine seaway or “corridor” and theCircum-Tropical marine current was active (Grad-stein et al., 1990) remains problematic. The occur-

rence of marine sediments of this age in thesouthwestern terranes of Cuban can be explained asa reaction to the crustal extension between SouthAmerica and the Maya Block (Lawver et al., 1999),producing a marine basin that opened into thePacific. The possibility that this early Caribbeanintercontinental embayment communicated with theCentral Atlantic since the Bajocian can be acceptednow only as a working hypothesis (Figs. 1 and 2), butrequires further confirmation in the stratigraphicrecord of the circum-Caribbean region.

Late Jurassic. During the Late Jurassic, the gapbetween North America and Gondwana widened,and true marine basins with ocean crust were devel-oping both within the Caribbean and the Gulf ofMexico (Figs. 1 and 2D; Iturralde-Vinent, 2003c).The Gulf of Mexico was an independent marinetongue of the Pacific Ocean until the latest Jurassic(Kimmeridgian–Tithonian), when finally communi-cation between the Caribbean and the Atlantic wasdeveloped (Salvador, 1991; Marton and Buffler,1999). The Caribbean Seaway (now a true corridorfor marine biota) opened wide, allowing pelagicmarine biota exchange between western Tethys andthe eastern Pacific realms (Figs. 1 and 2; Gaspariniand Iturralde-Vinent, in press). The Circum-Tropi-cal marine current flowed across the Caribbean sea-way (Berggren and Hollister, 1974; Parrish, 1992).

The evolution of the Gulf of Mexico in the LateJurassic is fairly well understood (Salvador, 1991;Buffler and Thomas, 1994; Pindell, 1994; Martonand Buffler, 1999). Until Callovian it was arestricted saline basin, but in the Oxfordian, ageneralized marine transgression from the Pacificcovered wide areas with shallow carbonate andshale deposits of the Smackover, Zuloaga, andrelated formations (Salvador, 1991; Marton andBuffler, 1999). About Kimmeridgian–Tithoniantime, the southwestern Gulf was drowned by shallowmarine environments, and the Gulf of Mexicobecame a new corridor for the marine biota (addedto the Caribbean seaway) fully connecting the Atlan-tic with the Pacific Ocean. This event probably pro-duced a subdivision of the Circum-Tropical marinecurrent into two branches (Berggren and Hollister,1974). The opening of the Gulf of Mexico as a newseaway into the Caribbean triggered some changesin the composition of the marine biota of NorthAmerica (Westermann, 1992; Kriwet, 2001).Another implication of this event was the isolationwithin the Maya Block of the terrestrial biota thateventually inhabited the Florida-Yucatan emerged


ridge (Figs. 1 and 4). The widening gap betweenNorth America and Gondwana limited the possibil-ity of direct overland dispersal, between the landbiotas of these continental areas. Probably since theBajocian, but surely since the Oxfordian, such apossibility came to a close (Figs. 1 and 2; Iturralde-Vinent and MacPhee, 1999).

Within the Caribbean, Upper Jurassic marinerocks are well developed (Figs. 1, 2, and 4). In theCuban southwestern terranes (Guaniguanico andEscambray) the Callovian–Oxfordian was a time oftransition between siliciclastic and carbonatemarine deposition (Pszczólkowski, 1978; 1999;Somin and Millán, 1981). The mid-late Oxfordianage Jagua Formation of western Cuba yields a richfossil assemblage that includes terrestrial (plants,pterosaurs, and dinosaurs), coastal (plants, reptiles,fish, invertebrates), and open marine species(reptiles, fish, and invertebrates) (De la Torre yCallejas, 1949; Colbert, 1969; Pszczólkowski, 1978;Iturralde-Vinent and Norell, 1996; Fernández andIturralde-Vinent, 2000; De la Fuente and Iturralde-Vinent, 2001; Gasparini and Iturralde-Vinent,2001; in press; Kriwet, 2001). By the end ofthe Jurassic, this off-Yucatan continental margin(Guaniguanico terrane) developed into deepermarine environments (Pszczólkowski, 1978, 1999;Pszczólkowski and Myczyñski, 2003; Sánchez-Barreda, 1990; Schaffhauser, et al., 2003) andreached its maximum width probably in the UpperCretaceous (Pindell, 1994).

In the Bahamas, Upper Jurassic marine rockshave been intersected by exploratory wells and cropout in northern Cuba (Khudoley and Meyerhoff,1971; Meyerhoff and Hatten, 1968, 1974; Sheridanet al., 1988). These sections represent a passivemargin domain, with restricted shallow carbonateand evaporitic facies within the Bahamas, andsouthward, development of continental slope depos-its with deep-water limestone since the Kimmerid-gian (Meyerhoff and Hatten, 1968, 1974; Iturralde-Vinent, 1994, 1998). During the Late Jurassic,major Atlantic-Caribbean marine water circulationmay have taken place eastward of the Bahamas; theStrait of Florida opened as a deep marine channelonly by Early Cretaceous (Fig. 2C and 2D; Bufflerand Hurst, 1995).

In the northern continental margin of SouthAmerica, the paleogeographic scenario was differ-ent. Uppermost Triassic–Jurassic strata are gener-ally represented by redbeds with paleosols, andalluvial and lake deposits, which yield fossils of

terrestrial plants as well as those of fresh waterinvertebrates (Fig. 4; González de Juana et al.,1980; Maze, 1984). Some Upper Jurassic marinebeds occur within the terrestrial sections, mostlyrepresented by intercalations of limestone, clastic-carbonate rocks, and shales, which are morecommon and thicker toward the continental edge.These marine rocks contain Late Jurassic fish(Lepidotus and Elasmobranchia), mid-Late Jurassiccorals (Aplophyllia), and Kimmeridgian throughLower Cretaceous ammonites (González de Juana etal., 1980; Maze, 1984; Macsotay and Peraza, 1997;see Figs. 9 and 10 for a summary of the main Juras-sic paleogeographic events previously described).

Cretaceous to Late Eocene: Vanishing islandsand land bridge in the Caribbean

The paleogeography of this time interval is illus-trated by three maps for the Early Cretaceous (~125Ma), Latest Cretaceous (~70 Ma), and basal EarlyEocene (~55 Ma) (Fig. 5). These maps have beendesigned to represent intervals of “land maxima,” orin other words, the periods of maximum land expo-sure and interconectiveness. As a tectonic frame-work for these maps, the allochthonous Caribbeanplate model (Pindell 1994; Pindell et al., 2006), theposition of the continental terranes (Lawver et al.,1999), and the location of the volcanic arc terranescombining Lawver et al. (1999), Pindell et al.(2006), and Iturralde-Vinent (1998) were chosen.Regarding the occurrence of land, shallow seas,and deep-marine environs within these tectonicterranes, they have been depicted utilizing the data-base and method described by Iturralde-Vinent andMacPhee (1999), with new data obtained from fieldwork and some recent literature.

Cretaceous. During the Cretaceous, the paleogeo-graphic scenario of the Caribbean and Gulf ofMexico underwent a series of important modifica-tions (Figs. 5A, 5B, 9, and 10). In general, the Gulfof Mexico had achieved its present structuraldimensions, inasmuch as little crustal expansiontook place after the Berriasian, when the MayaBlock reached its present-day position relative toNorth America (Marton and Buffler, 1999). Themain paleogeographic changes in the Gulf of Mexicohad to do with the position of the coastline and themarine sedimentary facies (McFarland and Menes,1991). Moreover, due to the existence of a wideinterior sea in North America, periodically the Gulfof Mexico served as a gateway between the North


American epicontinental sea and the Caribbean(Fig. 5B).

With the exception of the Barremian, when it waspartially exposed, the Maya Block generally evolvedas an isolated shallow carbonate platform (Viniegra-O., 1981). Another large platform evolved above theYucatan-Bahamas Block, partially exposed in theBarremian. This platform subdivided into smallerunits after the Aptian and up to the Present (Fig. 5B;Khudoley and Meyerhoff, 1971; Salvador, 1991;Buffler and Hurst, 1995; Randazzo, 1997). Thenorthern South American margin became a silici-clastic coastal plain eventually inundated by epi-continental marine environments, and the coastlinegenerally extended far into the continent (Figs. 5A,5B; Pindell and Tabbutt, 1995; Cordani et al.,2000). The North Atlantic was already a wideoceanic basin, but communication with the SouthAtlantic was limited until the Aptian-Albian, whena marine seaway between Africa and South Americadefinitely opened (Bullard et al., 1965; Berggremand Hollister, 1974, Jones et al., 1995; Riccardi,1991).

Regarding the Caribbean, the maximum separa-tion between the Maya Block and South Americawas reached during the latest Cretaceous (Lawver etal., 1999; Pindell et al., 2006), and the paleogeo-graphic realm underwent continuous modification ofboth marine and terrestrial landscapes. Thesechanging scenarios were due to the onset and evolu-tion of volcanic and nonvolcanic islands, non-volca-nic ridges, rises, basins, and trenches. Two mainvolcanic archipelagos were active, one well withinthe Pacific Ocean (now the basement of CentralAmerica), and another generally located in the oce-anic gap between North and South America (now thebasement of the Greater Antilles, Lesser Antilles,and Aves Ridge) (Figs. 5A and 5B). For reference,they will be called the Central American volcanicarc system and the Antillean volcanic arc system,respectively.

Cretaceous marine fossil invertebrates stronglysuggest wide interrelationships of the westernTethys, North and Central Atlantic, Caribbean, andeastern Pacific biotas, implying the existence ofmarine currents connecting these water basins(Scott, 1984; Sohl and Kollmann, 1985; Rojas et al.,1995; Buitrón-Sánchez and Gómez-Espinosa, 2003;Pszczólkowski and Myczyñski, 2003). And as wasdiscussed by Iturralde-Vinent (1982, 2003a) andIturralde-Vinent and MacPhee (1999), during thistime interval no “permanent” landmasses existed in

the Caribbean that lasted until the present, a featurethat unfortunately is never evident when paleogeo-graphic maps are inspected.3 The fact is that, whenthe Cretaceous to Late Eocene stratigraphic recordof the Caribbean is investigated for either thepresent-day subaereal or submarine areas, very fewindications of land occurrence are found, and noneof those indications implies long-lasting or continu-ous emergence since the Mesozoic into the Cenozoic(Fig. 5; Iturralde-Vinent and MacPhee, 1999; Itur-ralde-Vinent, 2003a, MacPhee and Iturralde-Vinent, 2005; Myczyñski and Iturralde-Vinent,2005). This point of view was challenged by Hedges(2001), but addressed by MacPhee and Iturralde-Vinent (2005).

An intercontinental land bridge has been pro-posed to account for the late Campanian–Paleoceneexchange of land tetrapods between North and SouthAmerica (Fig. 5B; Gayet et al., 1992; Iturralde-Vinent and MacPhee, 1999), based on paleontologicground (Lucas and Alvarado, 1994; Gayet, 2001).According to Iturralde-Vinent and MacPhee (1999),during the latest Campanian–Maastrichtian“substantial subaereal exposure existed along thepartially extinct Cretaceous volcanic arc and adja-cent continental margins, as indicated by evidenceof deformation, angular unconformity, hiatuses,deep-seated erosion, mountain-building, and terres-trial sedimentation (including conglomerate andpaleosol development)” (Khudoley and Meyerhoff,1971; Mattson 1984; Maurrasse 1990; Iturralde-Vinent, 1994). One problem for this land bridge isthe observation by Gayet (2001) that the exchangeof land vertebrates was particularly active duringthe Maastrichtian and Paleocene, a time when“… transgressive late Maastrichtian marine sedi-ments are recorded in the [Antillean] Cretaceousvolcanic arc as well as North and South America”(Iturralde-Vinent and MacPhee, 1999). In recentyears, some stratigraphic sections previously datedas latest Maastrichtian in western and central Cuba(Cacarajícara, Peñalver, and related units) havebeen redefined as Cretaceous–Tertiary boundaryin age, and the previously underlying siliciclasticsections (implying land occurrence within theAntillean volcanic arc system) extended in age up tothe end of the Cretaceous (Tada et al., 2003).

3To avoid this kind of confusion, Figures 9 and 10 summarizethe presence of land environments in another way.


FIG. 5. Paleogeographic maps of the Caribbean during (A) Early Cretaceous, (B) latest Cretaceous, and (C) EarlyEocene; after Iturralde-Vinent (2004). This paleogeographic scenario illustrates the formation of the Caribbean plate inthe Pacific realm and its posterior insertion within the inter-American gap replacing the older proto-Caribbean crust.Volcanic arc systems occupied the leading and trailing edges of the plate. These volcanic arcs supported non-permanentislands that were drowned and new ones exposed repeatedly, while the arc migrated laterally. In these conditions, sub-sistence of terrestrial biota is not expected within the Caribbean. A suspected land bridge or stepping stones scenarioallowed biotic exchange between the continents near the end of the Cretaceous.


In the Dominican Republic, on the other hand,an extensive survey of the uppermost Cretaceous–Paleocene? Don Juan conglomerates has confirmedthat the section is a complex of terrestrial red beds,laterally merging into shallow-marine rocks (Bour-don, 1985; Lewis et al., 2002; Iturralde-Vinent,unpubl. observations, 2003). Therefore, the latestCretaceous land bridge may have been operationalsometime between 75 and 65 Ma, represented by asystem of islands and shallows, existing when exten-sive uplift took place along the old Antillean Creta-ceous volcanic arc system (Fig. 5B). Under thisscenario, the tetrapod exchange between North andSouth America across the Antillean volcanic arcsystem may have taken place during some sea-leveldrop, probably utilizing a combination of islandstepping stones and sweepstakes dispersal mecha-nisms, while this arc collided with the Maya Blockand South America near the end of the Cretaceous(Iturralde-Vinent et al., 2006; Pindell et al., 2006).

The possibility of other ephemeral land bridgesor stepping-stone islands along the Antillean volca-nic archipelago during the Aptian–Albian andSantonian–Campanian was also evaluated by Itur-

ralde-Vinent and MacPhee (1999), but they con-cluded that there is insufficient evidence to proposeactual land connections between North and SouthAmerica during these periods. Within the aforemen-tioned time intervals, many indications of localuplift are known within the volcanic archipelago(Maurrasse, 1990; Iturralde-Vinent and MacPhee,1999); therefore, we cannot exclude the possibilityof terrestrial faunal exchange between closelylocated islands, due to the operation of the sweep-stakes or stepping-stone scenarios (McKenna,1973); however, these mechanisms have not beendocumented in the Caribbean realm, other thanthose discussed in this paper. In this regard, Horne(1994) reported a mid-Cretaceous Ornithopoddinosaur femur from central Honduras (part of theChortis Block), suggesting that northern CentralAmerican terranes were physically connected toNorth America at about 85–95 Ma. This is correct,but the geographic feature does not represent a landbridge, but a landspan (continent-to-island connec-tion). In fact, neither stratigraphic nor paleontologicdata suggest that those north Central Americanlands extended to South America (Gayet, 2001)

FIG. 5. Continued (see facing page).


across the Central American or Antillean volcanicarc systems (Fig. 5; Denyer and Kussmaul, 2002;Iturralde-Vinent, 2005).

Hedges (2001) presented a diferent hypothesis,suggesting that “… the proto-Antillean island arcmore or less connected North and South Americaduring the late Cretaceous (~100 to 70 Ma).” Butthere is no fossil record supporting this 30 millionyear long–lasting connection between North andSouth America across either the Antillean or theCentral American volcanic arc systems (Lucas andAlvarado, 1994; Iturralde-Vinent and MacPhee,1999; Gayet, 2001, Iturralde-Vinent, 2005). Theproblem is that Hedges understood an “island arc”as an emergent ridge, but such an implication is notnecessary true, as was discussed in detail by Itur-ralde-Vinent and MacPhee (1999). A volcanicisland arc is a geologic unit with a dynamic relief ofbasins and highs, that may or not be temporallyuplifted (as a submarine ridge, a chain of islands andshallows, or an emerged ridge).

The impact of a large extraterrestrial bolide withthe earth at the Yucatan Peninsula at 65 Ma cer-tainly played an important role in the biogeographicevolution of the Caribbean region (Crother andGuyer, 1996). Although an extensive literature isdedicated to the analysis of the global consequencesof this event, the following is a brief summary toevaluate the local effects according to the latestknowledge acquired by a Cuban-Japanese researchteam (Tada et al., 2003, and references within). Inthe western Caribbean basin, the earthquake trig-gered by the impact at Chicxulub induced extensivecollapse along the edge of the carbonate platforms ofYucatan, Florida, the Bahamas, and the Cuban seg-ment of the Antillean volcanic arc system. Platformmargins were strongly transformed, and the localislands and continental seashores were engulfed bylarge tsunami waves produced by the collapse of theplatforms. Huge amounts of sediments were deliv-ered to the sea in a few hours, particularly in thewestern part of the Caribbean (Tada et al., 2003). Asa consequence, one can hypothesize that any low-land within the Caribbean and its margins musthave been strongly transformed and deprived ofmajor life forms. The shallow seas were probablydeeply eroded and buried by thick layers of clasticsediment, the benthic and nectonic biota beingeliminated. In the deep basins located in the north-western Caribbean and in the Gulf of Mexico, thebenthic, nectonic, and planktonic sea life also mayhave been terminated or severely damaged. This is

another reason to adopt the postulate that no Creta-ceous terrestrial life form in the Caribbean realmsurvived into the Tertiary, much less into the Present(Iturralde-Vinent, 1982; Iturralde-Vinent andMacPhee, 1999). If, eventually, some pre–latestEocene Caribbean terrestrial taxon proves to bepresent today on some island as postulated utilizingmolecular clocks (Hedges et al., 1992; Hedges,2001; Dávalos, 2004), this occurrence may havebeen due to the operation of the “Noah’s arc” mech-anism of dispersion described by McKenna (1973)for other regions, but yet awaiting concrete demon-stration in the Caribbean as discussed by Iturralde-Vinent and MacPhee (1999) and MacPhee and Itur-ralde-Vinent (2000, 2005).

In his critical observations of the paleogeo-graphic model developed by Iturralde-Vinent andMacPhee (1999), Hedges (2001) concluded that thegeological history of the Caribbean “… region is notknown in enough detail to support such specula-tion.” Mainly because “… other authors haveclaimed the opposite.” Then, Hedges (2001) sug-gested that at least Puerto Rico, but probably otherAntillean islands, have been emergent since theAlbian based largely on work by Donnelly (1992).However, more detailed investigations compiled byIturralde-Vinent and MacPhee (1999, Tables 1–3and Appendices) has allowed a new interpretation ofDonnelly’s (1992) data. Moreover, the presentauthor has visited all the localities referred to byDonnelly (1992) and several more in Cuba, Jamaica,Hispaniola, and Puerto Rico, and obtained samplesthat have been examined in order to verify age andpaleoenvironment. As early as the Neocomian orearly Aptian (~140 Ma) volcanic islands in theAntillean volcanic arc system were emergent (LosRanchos and equivalent Formation of DominicanRepublic; Smiley, 2002), but these islands weresubsequently submerged in the Albian (~110 Ma;Río Hatillo and equivalent formations in the Domin-ican Republic; Myczyñski and Iturralde-Vinent,2005). Furthermore, marine strata of Albian,Cenomanian, Turonian, Coniancian, Santonian,Campanian, Maastrichtian, Paleocene, and Eoceneage are well developed across the Antillean volcanicarc system in present-day Puerto Rico, Hispaniola,Cuba, and Jamaica, arguing for widespread repeatedsubmergence (Maurrasse, 1990; Rojas et al., 1997;Lidiak and Larue, 1998; Iturralde-Vinent andMacPhee, 1999; Lewis et al., 2002; Myczyñski andIturralde-Vinent, 2005, Schweitzer et al., in press).


Therefore, the present Antillean islands arenewly formed post–Middle Eocene geographic enti-ties (<40 Ma; Iturralde-Vinent, 1982; Iturralde-Vinent and MacPhee, 1999). These entities arecomposed of allochthonous older geologic units,amalgamated in the basement of the islands. There-fore, the paleogeography of those units greatlydiffers in all aspects from the present-day islands(Pindell et al., 2006). For example, Puerto Rico orHispaniola, as geographic entities, have no meaningin the Cretaceous or Paleocene. Their basementsconsist of a complex allochthonous assemblage ofMesozoic rocks of deep oceanic and crustal origin(Bermeja and Duarte complexes) and deformedallochthonous Cretaceous to Middle Eocene igneousand sedimentary rocks of island-arc and crustal ori-gin (Dengo and Case, 1990; Donovan and Jackson,1994; Mann, 1999; Bartolini et al., 2003; Iturralde-Vinent and Lidiak, 2006). The island’s basement, asan assemblage of all these geologic units, is ayounger, post–Middle Eocene entity, with in situlatest Eocene to Recent sedimentary rocks of marineand terrestrial origin (Iturralde-Vinent andMacPhee, 1999).

Early Tertiary. The Paleocene through MiddleEocene paleogeographic history is representedherein by a map for the basal Early Eocene (~ 55Ma; Fig. 5C). During this time interval, Yucatan,Florida, and the Bahamas evolved as shallow car-bonate platforms facing deep-water environments.Within the Caribbean, the evolution of two mainvolcanic archipelagos took place, one facing thePacific Ocean (Central American volcanic arcsystem); the other, in the Caribbean interior, movedeastward on the leading edge of the Caribbean plate(Antillean volcanic arc system; Fig. 5C). The preciseposition of the Antillean volcanic arc system duringthe Eocene has been distinctly depicted by differentauthors (e.g., Bartolini et al., 2003; Iturralde-Vinentand Lidiak, 2006), but the scenario presented in thispaper was selected to be sufficiently general so asnot to contradict substantially any allochthonousmodel of the Caribbean plate.

Challenging this reconstruction, Hedges (2001)stated that “The Bahamas platform has remained arelatively stable carbonate block for most of theCenozoic. However, the compressional forces of thecollision with the proto-Antilles during early Ceno-zoic may have caused uplift along the margin of theBahamas platform. Because the platform alreadywas near sea level, any uplift would have exposeddry land for colonization by terrestrial organisms.”

These conclusions are not correct for several rea-sons. First, the Bahamas platform, as it is today, haslittle to do with the “Bahamas platform” during theMesozoic and Early Tertiary, when it was muchlarger, with a different shape, and a distinct paleo-geography (Fig. 5C). Second, it is not accurate to saythat the “proto-Antilles” (which may imply actualislands) collided with the Bahamas platform,because the collision was between the Antillean vol-canic arc system (the leading edge of the Caribbeanplate) and the Bahamas borderland (the so calledPlacetas, Camajuaní, Remedios, and Cayo Cocobelts of northern Cuba, sensu Meyerhoff and Hatten,1974; Pushcharovsky, 1988). Third, this collisionhas been documented by Bralower and Iturralde-Vinent (1997) and Iturralde-Vinent (1994, 1996,1998) to show that it produced a local Paleoceneuplift (forebulge) in parts of the Bahamas borderland(the so-called Remedios and Cayo Coco belts) but itwas totally drowned in the Paleocene–Early Eoceneto become a deep water foreland basin. When thiscollision ended in the Middle to Late Eocene, thewestern part of the older Bahamas block remainedattached to the leading edge of the Caribbean plate(northern Cuba), and a marine transgressionsubmerged large parts of present Cuban area andfurther north to create the Canal Viejo de Bahamaschannel. Subsequently, since the Miocene, theBahamas assumed their present general shape (Fig.6; Khudoley and Meyerhoff, 1971; Meyerhoff andHatten, 1974; Sheridan et al., 1988). On the otherhand, there is no paleontologic record of terrestrialfossils or sediments within the Bahamas Cretaceousand Early Tertiary stratigraphic sections, whosesouthern margin extensively crops out in northernCuba (Meyerhoff and Hatten, 1974; Pushcharovsky,1988), so the statement by Hedges remains fullyunsupported and speculative.

Regarding the Cretaceous and Paleogene Anti-llean volcanic arc system, there are clear indicationsof land, shallows, and deep-water environmentsassociated with the volcanic ridge, but no evidenceof continuous land occurrence in time or space(Maurrasse, 1990; Lidiak and Larue, 1998; Itur-ralde-Vinent and MacPhee, 1999; Bartolini et al.,2003).

The Caribbean paleogeographic scenario dis-cussed above for the Cretaceous to Late Eocene iscompatible with the occurrence of Early EoceneNorth American land mammals (Hyrachyus sp.) inJamaica, found in association with fresh waterand marine vertebrates (Domning et al., 1997). As


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outlined by Iturralde-Vinent and MacPhee (1999),this is an example of the “Viking funeral ship” sce-nario of McKenna (1973). Recognizance of theEocene localities with land vertebrates in Jamaica(accomplished by the author during year 2000) sup-port this scenario. Early Eocene vertebrates occur inthe Guy Hill Formation, which represent a coastalsiliciclastic depocenter, filled with arcosic sand-stones and gravels intercalated with clay, silt, lignif-erous sands and clays, and isolated beds of marlylimestones. This unit was deposited within interme-diate fluvio-marine environments all along the west-central part of present-day Jamaica (Western Jamai-can block, sensu Iturralde-Vinent and MacPhee,1999). By the Early Eocene, western Jamaica wasprobably located in the latitude of Central America,in contact with the Chortis Block–Nicaragua Rise(Fig. 5C; Pindell, 1994; Domning et al., 1997).These facts imply that the sedimentary basin wherethe Guy Hill Formation was deposited was muchlarger than present-day Jamaica, and the sedimentsource must have been a continental terrane,because the clastic sediments are well sorted, andderived from sialic basement, implying long-dis-tance transport. Therefore, the source area wasprobably basement rocks of the Chortis Block/Nica-ragua Rise, as part of northern Central America. Inthis paleogeographic scenario, as suggested byDomning et al. (1997), the North American terres-trial biota populated the “Jamaican/Central Ameri-can territory” when it was a landspan (peninsularprojection) of North America (Iturralde-Vinent,2005; Fig. 5C).

This inclusion of Jamaica in a peninsular projec-tion of North America into Central America contra-dicts the statement of Hedges (2001) that the “…recent discovery of ungulate (rhynocerontoid) andiguanid lizard fossils from the Eocene (~50 Ma) ofJamaica… indicate that a diverse biota may haveexisted on some Caribbean islands in the earlyCenozoic.” But Jamaican terranes were detachedfrom Central America only after the Middle Eocene,due to the displacement of the Caribbean plate(Pindell, 1994), and much later uplifted as theCaribbean island of Jamaica near its presentgeographic position during the Neogene (Donovanand Jackson, 1994).

The existence of evanescent (non-permanent)islands, ridges, rises, and shallow banks in the Car-ibbean during the Cretaceous through Late Eocenecan be also visualized as non-permanent barriersagainst the free flow of the Circum-Tropical marine

current across the Caribbean Seaway (Figs. 5, 9, and10; Iturralde-Vinent, 2003a). This current wascertainly a major route for biotic dispersion, as sug-gested by the presence of several groups of marineanimals (Berggrem and Hollister, 1974; Smith,1984; Scott, 1984; Sohl and Kollman, 1985;Ricardi, 1991; Rojas et al., 1995; Kriwet, 2001;Buitrón-Sánchez and Gómez-Espinosa, 2003; Sch-weitzer et al., in press). This marine current wasinterrupted only between 75 and 65 Ma for no morethan 3–5 m.y., when the latest Campanian–Maas-trichtian suspected land bridge was probably con-structed along the Antillean volcanic arc system(Figs. 5B and 10; Iturralde-Vinent and MacPhee,1999). This brief shutdown of the Circum-Tropicalmarine current must have produced a noticeableeffect in the world climate and temporarily inter-rupted the marine biotic exchange along the Carib-bean seaway.

Around the Middle to Late Eocene, a profoundmodification of the Caribbean tectonic regime tookplace (Mattson, 1984; Iturralde-Vinent andMacPhee, 1999). A general uplift (Pyrenean orog-eny) is recorded in every marine topographic high orland area, both in the Caribbean realm and in itssurrounding marine and continental areas (Protheroet al., 2003). In the Caribbean, sedimentation ofterrestrial conglomerates was a widespread event,the so called “conglomerate event” of Iturralde-Vinent and MacPhee (1999). In concert with thesedevelopments, the paleogeographic regime under-went important changes that defined a new stage inthe evolution of the region (Iturralde-Vinent, 1994,1998; Iturralde-Vinent and Gahagan, 2002).

Latest Eocene to Recent: Formation of thepresent-day Caribbean

The latest Eocene to Recent stage of the Carib-bean paleogeographic evolution has been evaluatedin great detail (Iturralde-Vinent and MacPhee,1999; Iturralde-Vinent, 2001; MacPhee and Itur-ralde-Vinent, 2005). I have generally concludedthat uplift of the core of the present Antilles startedafter the Middle Eocene (~40 Ma). The Eocene–Oli-gocene transition ( ~35-33 Ma) was the peak of thisgeneral uplift (Pyrenean orogeny) when the amountof land in the Caribbean should have been at amaximum (Fig. 6). The Late Oligocene was a time ofhigh sea level, and therefore of minimum exposure(and probably minimal interconnectedness) of emer-gent areas (Fig. 7). Since the early Middle Miocene,further isolation of land areas took place as a conse-


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quence of active tectonic disruption of the northernand southern Caribbean plate boundaries. In thecase of the Greater Antilles, this resulted in the tec-tonic subdivision of previously existing landmasses(Fig. 8). This subdivision was very significant bio-geographically as it probably resulted in island-island vicariance of sloths (White and MacPhee,2001) and other terrestrial organisms (MacPhee andIturralde-Vinent, 1994, 1995, 2000).

Late Eocene–Oligocene transition. The paleogeo-graphic scenario for the Eocene–Oligocene transi-tion (zones P16 to P18 of Berggren et al., 1995) wascharacterized by the formation of a large peninsularextension of South America (Gaarlandia landspan)that reached as far as central Cuba. The western-most Cuban archipelago was isolated from nearbylands by the Havana-Matanzas and Yucatan chan-nels (Iturralde-Vinent, 1969, 1972; Iturralde-Vinentet al., 1996). The Central American volcanic arcsystem was partially uplifted, but the land areasremained generally isolated from nearby continents(Coates and Obando, 1996; Denyer and Kussmaul,2002; Iturralde-Vinent, 2005). Other small islandsand shallow banks were present in different parts ofthe Caribbean (Fig. 6). Under these conditions,Gaarlandia interrupted the Circum-Tropical marinecurrent, but limited flow took place between thePacific, Caribbean, and North Atlantic via the Straitof Florida (Fig. 6; Mullins and Neumann, 1979; Itur-ralde-Vinent and MacPhee, 1999). For a short timeperiod, < 3 million years, overland dispersal fromnorthern South America into Gaarlandia occurredfor various elements of the land biota (Borhidi,1985; MacPhee and Iturralde-Vinent, 1994, 1995,2000, 2005; Iturralde-Vinent and MacPhee, 1996,1999). The Caribbean waters may have cooled dueto the combined action of the California currentdrifting southward (Duque Caro, 1990; Droxler etal., 1998) and the general cooling of the worldclimate (Wright and Miller, 1993; Prothero et al.,2003).

Oligocene. Since the Early Oligocene (~32–30Ma), but certainly by Late Oligocene (zones P21b-P22 of Berggren et al., 1995), many Caribbean low-lands were inundated, and Gaarlandia was subdi-vided into distinct archipelagos (Fig. 7). TheCircum-Tropical marine current was re-established,producing another change in climate due to theinflux of warm Atlantic waters into the Caribbean(Duque Caro, 1990; Droxler et al., 1998; see alsoWright and Miller, 1993). The terrestrial organismsthat colonized Gaarlandia in the Eocene-Oligocene

transition became isolated from the continent, andrepresents the earliest true Greater Antillean biota.Elements of this biota with a clear South Americansignature have been found in Lower Oligocene unitsof Puerto Rico; in the Miocene of Puerto Rico, His-paniola, and Cuba; and in younger deposits of manyAntillean islands (Monroe, 1980; Borhidi, 1985;Graham, 1986, 1990; MacPhee and Wyss, 1990;MacPhee and Iturralde-Vinent, 1994, 1995, 2000;MacPhee and Grimaldi, 1996; MacPhee et al.,2003; Iturralde-Vinent and MacPhee, 1996, 1999;White and MacPhee, 2001). In the Caribbean, thefirst coral reef communities appeared in the Oli-gocene, and evolved to importance during the Neo-gene and Holocene (Frost et al., 1983; Budd et al.,1996; González Ferrer and Iturralde-Vinent, 2004).

Miocene and Pliocene. During the Miocene andPliocene, the Caribbean plate continued its east-ward displacement (Pindell, 1994; Iturralde-Vinentand Gahagan, 2002). The Greater Antillean tectonicterranes were transported, deformed, and skewedalong the contact between the Caribbean and NorthAmerican plates, creating deep marine straits thatcompleted the isolation of the major islands as inde-pendent geographic entities (Fig. 8; Iturralde-Vinent and Gahagan, 2002). The Circum-Tropicalmarine current drifted more efficiently across theCaribbean Sea into the Pacific Ocean, with a branchthat fed the early Gulf Stream (Iturralde-Vinent etal., 1996; Iturralde-Vinent and MacPhee, 1999).

Regarding the climate, the oxygen isotope curvesrecord Middle Miocene (17 and 14 Ma) warm excur-sions (relative maxima) both in the Atlantic andPacific (Tsuchi, 1993; Wright and Miller, 1993).These events have been correlated in the Caribbeanwith the unusual production of resin by the amber-producer tree Hymenaea protera in Hispaniola andPuerto Rico (Iturralde-Vinent and Hartstein, 1999;Iturralde-Vinent, 2001), suggesting that in bothmarine and terrestrial environments an unusualwarm climatic maxima probably occurred. After thisevent, a general cooling trend is evident in theoxygen isotope signature (Tsuchi, 1993; Wright andMiller, 1993).

About Late Pliocene (nearly 2.5–2.3 Ma), butstarting as a shallow ridge as early as 3.0 Ma (Jack-son et al., 1996), the Isthmus of Panama closed andformed the Central American land bridge. Neverthe-less, this closure was not fully erected, becauseTonnoidean gastropod larvae were capable of cross-ing between the Caribbean and the Pacific duringinterglacial maxima during the Early Pleistocene


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(Beu, 2001). This ridge probably had an ephemeralprecursor back in the late Middle Miocene, creatingsome closely spaced shallows and islands in CentralAmerica, when some South American mammals dis-persed into North America (Webb, 1985; Lucas andAlvarado, 1994). The Pliocene was also a time ofgeneral uplift in many areas of the Caribbean, pro-ducing the backbone of today’s relief (Iturralde-Vinent, 1969, 1978, 2003b; Mann et al., 1990;Coates and Obando; 1996; van Gestel et al., 1998,1999). As this paleogeographic situation evolved,the Caribbean region looked more like that of today.The island’s shelf became defined, the major moun-tain ranges formed, but the topographically lowparts of the landscape were subjected to repeatedinundation and desiccation due to oscillatory sealevel rise and fall (Haq et al., 1987) and verticalmotion of the earth’s crust (Mann et al., 1990; Itur-ralde-Vinent, 1978, 1998, 2003b). These eventsproduced short time subdivisions or ephemeral con-nections between closely related topographic highsand small islands in the Caribbean, but never againbetween the major islands (Cayman, Bahamas,Cuba, Jamaica, Hispaniola, and Puerto Rico).Neither were there connections between the islandsand the nearby continents, because they were sepa-rated by deep water channels generally since theMiocene and Pliocene (Mann et al., 1990; Iturralde-Vinent and MacPhee, 1999).

Concerning island connectivity, research by vanGestel et al. (1998, 1999) challenged the widelyheld conclusion that the island of Puerto Rico hadbeen continuously uplifted since the latest Eocene(Meyerhoff, 1933; Monroe, 1980; Iturralde-Vinentand MacPhee, 1995, 1999). According to van Gestelet al. (1999), the Oligocene to Pliocene phase ofdevelopment of Puerto Rico “… started with aperiod of non-deposition and erosion, resulting in aLatest Eocene to Oligocene unconformity”; butsince the Middle Oligocene the Puerto Rico and theVirgin Islands were covered by the sea (van Gestelet al., 1999, their Fig. 18). This statement wasaddressed in detail by MacPhee et al. (2003). First,during the Eocene–Oligocene transition, thick ter-restrial conglomerates, paleosol and alluvial sand-stone with terrestrial plant remains were deposited,clearly indicating erosion and non-marine deposi-tion (Monroe, 1980; MacPhee and Iturralde-Vinent,1995; Montgomery, 1998). Second, Oligocene andMiocene stratigraphic sections along the northernand southern coastal flanks of the island are differ-ent in many aspects, and normally contain clastic

material derived from the igneous-sedimentaryCretaceous–Eocene core of Puerto Rico (Monroe,1980; Iturralde-Vinent and Hartstein, 1998). Third,many findings of Oligocene and Miocene terrestrialfossil remains, undoubtedly indicating the existenceof land, have been reported from the island(Graham, 1986; MacPhee and Wiss, 1990; MacPheeand Iturralde-Vinent, 1995; Iturralde-Vinent andHarstein, 1998, Iturralde-Vinent, 2001; MacPhee etal., 2003).

On the other hand, the interpretation of severaloffshore seismic lines by van Gestel et al. (1998)provide important information concerning the inter-connectiveness of Puerto Rico with Hispaniola, theVirgin Islands, and Saba Bank. The Gulf line LS49(van Gestel et al. 1998, Fig. 12) strongly suggeststhat St. Croix was isolated from Saba Bank since theMiddle Miocene; Gulf lines LS50 to LS52 (vanGestel et al. 1998, their Fig. 11) also strongly sug-gest that the marine depression between Puerto Ricoand the easternmost Virgin Island was inundatedonly since the Pliocene. The interpretation of EW96-05 line 35 and UTIG north-south line VB by vanGestel et al. (1998, Figs. 14 and 15) accross theMona Passage between Puerto Rico and Hispaniola,on the other hand, suggest that the area was inun-dated since the Oligocene.

This interpretation contradicts Iturralde-Vinentand MacPhee (1999), who assumed the inundationto have taken place after the Miocene, mostlybecause the old pre-latest Eocene island cores areexposed or covered by transgresive “Pleistocenerocks” on both sides of the Mona passage. vanGestel et al.’s (1998) interpretation of the age of thesediments filling in the Mona Passage is based onthe assumption that they have “the same age as inthe Puerto Rico North Coast basin,” but they do nothave any direct stratigraphic control. It would bemore correct to correlate the sediments filling theMona Passage with those outcropping on both sidesof the passage (in easternmost Hispaniola and west-ernmost Puerto Rico). Those in westernmost PuertoRico are definitely Quaternary, but those on eastern-most Hispaniola are newly dated as Middle to LateMiocene in age. The age revision of the UpperTertiary rocks of Hispaniola conducted duringwinter 2003 by the author, included the recovery ofseveral samples from two quarries (one near Bassoraand the other east of La Romana), which yieldmicrofossils (Sorites marginalis (including speci-mens ~ 2 cm in diameter), Amphistegina spp.,Amphistegina angulata, Amphistegina cf. A. rotundata,


?Nummulites sp., Victoriella sp., Globigerinita sp.[rare], Globigerina sp. [rare], Acervulinidae, mol-lusks, echinoids, and red algae, identified by SilviaBlanco Bustamante). This fossil assemblage can bedated as Middle or Late Miocene, because of thepresence of Miocene taxa and the absent of EarlyMiocene species (as Lepidocyclina or Miogypsina).

On the other hand, fossils described by Kaye(1959) from the islands Mona and Monito suggest aLower or Middle Miocene age. In order to corrobo-rate this dating, paleontological samples kindlyprovided by Wilson Ramírez from the limestone anddolostones exposed in Mona island were investi-gated. These samples, also indentified by SilviaBlanco Bustamante, yield algae, spines of equino-derms, bryozoa and foraminifera (Nummulites sp.,Amphistegina angulata, Amphistegina cf. A. gibbosaand Orbulina? sp.) which can be dated as Middle orLate Miocene, also because the presence of Miocenetaxa and the absence of Early Miocene markers. Thefact that near Mona Island (within the Mona Pas-sage) the thickness of the oldest seismostratigraphicbeds wedge out, suggest that the older basementbelow Mona was exposed longer than the rest of thegraben, and the rocks of Mona are near the oldestbasement. Therefore, it is tentatively assumed here,following MacPhee et al. (2003), that the marineinundation of the Mona Passage and isolationbetween Puerto Rico and Hispaniola may havestarted around the mid-Oligocene (~30 Ma) but alsoin the Early Miocene, because Miocene rocks ofMona and Dominican Republic reflect shallow-marine environments.

Hedges (2001) argue against the Iturralde-Vinent and MacPhee (1999) paleogeographic recon-struction of Puerto Rico, because “Larue (1994)noted that shallow-water limestone facies are foundin north- and south-central Puerto Rico, suggestingthat the Central Block may have been a topographichigh in the Eocene.” Although this evidence doesnot contradict the interpretation that Iturralde-Vinent and MacPhee (1999) presented for PuertoRico, the same Larue et al. (1998) concluded thatthe core of the island was inundated in the Paleo-cene-Eocene, and later inversion and uplift tookplace between the Late Eocene and MiddleOligocene. Furthermore, geological prospecting ofthe island by the author (1999–2003) revealed thepresence of Paleocene to Middle Eocene rocks ofdeep-marine origin in the northern, western, andcentral areas of Puerto Rico, which suggests that thedeformed Antilles Cretaceous volcanic arc core was

covered by the sea during the Early Tertiary (seealso Larue et al., 1998; Montgomery, 1998). Later, inthe Late Eocene, the presence of a regional uncon-formity and terrestrial deposits in the island indi-cates that it was uplifted as part of Gaarlandia(Larue et al., 1998; Iturralde-Vinent and MacPhee,1995, 1999). Under this scenario, the occurrence ofOligocene and Miocene shallow-water marine rocksin both northern and southern Puerto Rico, with ter-restrially derived clastic material (Monroe, 1980,see also a full evaluation by MacPhee et al., 2003),as well as offshore in the North Basin of Puerto Rico(Larue et al., 1998), indicates that not only thecentral block, but the entire axial part of the islandwas uplifted since the latest Eocene as depicted byMeyerhoff (1933).

Hedges (2001) further disputed the authorsinterpretation of the “Aves ridge.” According toHedges (2001) … “the difference between an islandchain and a continuous land bridge is fundamentalfor biogeography …”, but “… geological support fora continuous land bridge vs. a chain of islands doesnot exist.” However, Iturralde-Vinent and MacPhee(1999) compiled a set of data that show that AvesRidge was a topographic high during the Eocene–Oligocene transition, with important possibilities ofbeing completely uplifted for a short period, but donot rule out the possibility that it may has been aclose chain of islands separated by a shallow shelf.Any of these arrangements provided a route forterrestrial dispersion of land taxa as confirmed bythe fossil record (see Table 1 of MacPhee et al.,2003; MacPhee and Iturralde-Vinent, 2005). On theother hand, this paleogeographic scenario explainswhy Aves Ridge operated as a filter and not as a fullcorridor for the dispersion of the land biota(MacPhee and Iturralde-Vinent, 2000).

Among the problematic aspects of the paleogeo-graphical reconstruction presented here (Figs. 6, 7and 8), pending further evaluation, is the subdivi-sion of Jamaica into two main independent terranes,the close positioning of the southwestern peninsulaof Hispaniola and the Blue Mountains Block ofJamaica in the Oligocene, as well as the inferredpermanent exposure of parts of eastern Jamaica(Blue Mountains Block) as early as 33–35 Ma (Itur-ralde-Vinent and MacPhee, 1999). For a summary ofthese relationships, see Figures 9 and 10.

Quaternary. The Pleistocene and Holocenepaleogeography of the Caribbean have not been fullydeveloped up to the present. Nevertheless, recentlyIturralde-Vinent (2003b) produced a set of three


paleogeographic maps for Cuba (Fig. 11), includingthe Pliocene–Lower Pleistocene, the Late Pleis-tocene (~125,000–120,000 a), and the latest Pleis-tocene (~25,000–20,000 a). These maps suggestthat the present shape of the island was attained8,000–6,000 years ago, that about 25,000 years ago

the entire present shelf was uplifted, and that about125,000 years ago only isolated islands occurredwithin the territory of present-day Cuba. Thepresent-day Island of Youth (Isle of Pines) and west-ern Cuba were connected at least between 125,000and 8,000 years ago, providing a route for animal

FIG. 9. Schematic illustration of the Jurassic to Recent evolution of water circulation across west-central Pangea/Caribbean Seaway, updated after Iturralde-Vinent (2003a) with new information provided in this paper and newchronology.


migration between these lands. Also I concludedthat today’s small islands and keys within the Cubanshelf are very young features, probably younger than25,000 years (Iturralde-Vinent, 2003b).

Concerning the marine biota, about 20,000 to25,000 years ago, the emergence of the whole Cubanshelf must have eliminated all marine life, sothe present-day coralline and related communitiesmust be younger (González Ferrer and Iturralde-Vinent, 2004). In relation to the climate, it was sug-gested that periodic stages of cool, dry weather coin-cided with every glacial period, while warm, wet

stages attended interglacial periods. Under thiscircumstance, the terrestrial biota populating Plio-Pleistocene Cuban territory experienced severalstages (glacial maxima) of high possibilities ofexchange and interaction (full connectivity withinthe territory), intercalated with stages of isolationand concentration within the highlands (interglacialmaxima), concurrent with climatic modifications(Fig. 11). These events may have shaped the evolu-tion of the biota in the Quaternary, and probablyaccount for extinctions and development of localendemics. The same events may have operated in

FIG. 10. Jurassic to Recent evolution of land occurrences and terrestrial interconnections within the Caribbean,modified and updated from Figure 12 of Iturralde-Vinent and MacPhee (1999), with additional data provided in thispaper and new chronology. Vertical patterns are lands, and horizontal shades imply non-permanent connections as landbridge and landspan. Large horizontal arrows suggest possible overland migratory routes.


FIG. 11. Late Pliocene to latest Pleistocene paleogeographic evolution of Cuba after Iturralde-Vinent (2003b). Thesemaps illustrate a typical scenario for conditions of glacial and interglacial maxima that repeatedly took place since thePliocene. Glacial maxima allowed extensive dispersion of terrestrial organisms, whereas interglacial maxima favoredisolation of the biota in high lands (transient island conditions). The 25–20,000 a glacial maxima produced the totalexposure of the present Cuban shelf, destroying all shallow-marine ecosystems. Consequently the present marine popu-lations of the Cuban shelf are younger. Since the beginning of the Holocene, the outline of the present land areas havebeen shaped and reshaped constantly as a consequence of sea level rise and fall and neotectonic movements.


the entire Caribbean area, but we cannot prove it atpresent.

The Late Eocene to Recent paleoceanographicmodel for the Caribbean, developed by Iturralde-Vinent (2003a), takes into account the importance oftopographic factors in the reconstruction of the maintrend of surface marine currents (Fig. 12). However,Hedges (2001) postulated that the present pattern ofsurface marine currents in the Caribbean was simi-lar during the entire Cenozoic. His position ismostly based on the belief that the Coriolis effect isthe only driving force for surface marine currents.This simplistic interpretation is flawed, as demon-strated by Iturralde-Vinent and MacPhee (1999),MacPhee and Iturralde-Vinent (2000, 2005), andIturralde-Vinent (2003a).

Conclusions

The conclusions of this paper are meant to beindependent of any biogeographical bias. They showthat we have advanced in our comprehension of theCaribbean paleogeography and paleoceanography,but at the same time suggest how far are we yet froma complete understanding of some important detailsof the area’s geological history.

1. Caribbean seaway. The history of the Carib-bean as a seaway is synthesized in Figure 9. Themarine connection between western Tethys and theeastern Pacific across west-central Pangea, accord-ing to biogeographic interpretations, is postulated tohave existed since the Early Jurassic (Hettangian–Pliensbachian) according to paleontological find-ings (Berggren and Hollister, 1974; Imlay, 1984;Boomer and Ballent, 1996; Damborenea, 2000;Gasparini and Iturralde-Vinent, in press), but thestratigraphic basis for this date is lacking. Probablysince Bathonian (~165 Ma), but certainly sinceOxfordian (~159 Ma), the stratigraphic and paleon-tologic record indicates that this connection wasfully functional as a corridor for marine biotas, andthe Circum-Tropical marine current established(Fig. 9).

2. Intercontinental connections. The history ofterrestrial connections within the Caribbean realmis summarized in Figure 10. The last time that over-land dispersal between western Laurasia (NorthAmerica) and western Gondwana (South America)had some probability of occurrence was during theMiddle Jurassic, because since the Oxfordian thecontinents were separated by a marine gap (Fig. 1).Later, in the latest Campanian/Maastrichtian (~75–

65 Ma) a landbridge or closely located islands andshallows may have been present along the axis of theAntillean volcanic arc system connecting North andSouth America (Fig. 5A). During that time, the Anti-llean volcanic arc system was located in the presentpossition of Central America. On the other hand, thepresent-day bridge (Panamanian Isthmus) wascompleted only in the Plio-Pleistocene as part of theCentral American volcanic arc system (2.5–2.3 Ma).Evidence for a precursor bridge late in the MiddleMiocene (~9 Ma; Fig. 10) remains ambiguous at thistime, because it is based only on paleontologicevidences (Webb, 1985).

3. Availability of lands in the Caribbean. The his-tory of land occurrence in the Caribbean realm issummarized in Figure 10. Since the formation of thefirst volcanic archipelago within the Caribbeanplate about the time of the Jurassic–Cretaceoustransition, volcanic islands, shallow banks, andridges have been always present in the paleogeo-graphic scenario of the region. But these lands weregenerally ephemeral, because they lasted just a fewmillion of years before being drowned (Iturralde-Vinent and MacPhee, 1999). Also, the tectonicterranes where these lands evolved were not locatedin the same paleogeographic position as today,because of subsequent lateral movements of thetectonic plates (Figs. 7, 8, and 9). Only after theMiddle Eocene (~40 Ma) have permanent landsbeen available within the Caribbean geographicsetting (Fig. 10). Gaarlandia emerged later at about35–33 Ma, forming a long ridge (landspan) thatbriefly connected northern South America with cen-tral Cuba (Fig. 6). Gaarlandia was partially drownedsince ca. 32–30 Ma (Fig. 7), and diverse groupsof islands developed and ultimately became thepresent day lands and shallow banks of theCaribbean.

4. Mesozoic and Early Cenozoic Caribbean biotas.Fossils remains of terrestrial fauna and flora areknown since the Lower Jurassic sedimentary sec-tions of western Pangea around the future Caribbeanrealm. From the Middle Jurassic (Bajocian) onward,the occurrence of marine fossils has been recordedin the Caribbean region (Guaniguanico, Escambray,and Pinos terranes of western Cuba), but theyrepresent biotic elements of intercontinental seachannels within Pangea (Fig. 2C). Terrestrial islandbiota are also recorded from the Cretaceous andthroughout the Eocene within the Antillean volcanicarc terranes, chiefly represented by plants andinvertebrates. Eocene vertebrates are reported from


FIG. 12. Caribbean Late Eocene to Recent paleoceanographic model after Iturralde-Vinent (2003a). This scenariotakes into account the constantly changing terrestrial and submarine relief of the region, as well as general patterns ofocean water circulation. This can be utilized as a tool to evaluate any model of Late Tertiary overwater dispersal.


western Jamaica, but this paleobiota belonged origi-nally to northern Central America (Fig. 5C). Never-theless, probably none of these pre–latest Eoceneterrestrial clades survived into the present-day biotaof the Caribbean islands, inasmuch as no paleonto-logic record exists to prove it. Cretaceous andEocene terrestrial lineages are assumed to havebeen rooted in the Antilles, solely based in molecu-lar clocks used to date phylogenetic divergence(Hedges, 2001; Dávalos, 2004; Roca et al., 2004);but the method has been challenged by severalauthors (Crother, 2002; Crother and Guyer, 1996;

Iturralde-Vinent and MacPhee, 1999; MacPhee andIturralde-Vinent, 2000, 2003, 2005).

5. The bolide that impacted the earth in Yucatan(Chicxulub) at 65 Ma induced unusual catastrophicevents that affected the Caribbean geographic andbiological environment. Both marine and terrestrialbiotas sustained a high level of mortality, probablylarger than in other areas of the planet. Conse-quently, the Caribbean biota were severely damagedand probably none of the local species, especiallyterrestrial ones, survived into the Tertiary. Thisimplies that the window of opportunity for coloniza-

FIG. 13. Caribbean lands chorogram for the Middle Eocene to Recent interval. Small black ovals represent points ofconvergence or divergence of land units. The triangular gray pattern associated with land divergence or convergencesuggests possible land connectivity during sea level drops. Gaarlandia is depicted as a large black oval. NA and SAindicate North and South America, respectively. Black blocks mean land, either island (vertically short) or continent(vertically long). Horizontal grey strips depict extensive land exposure and potential interconnectivity during tectonicuplift or glacial maxima.


tion of the Caribbean sea and lands can be limited tothe Cenozoic (Crother and Guyer, 1996; Iturralde-Vinent and MacPhee, 1999; MacPhee and Iturralde-Vinent, 2005).

6. Origins of the present-day land biota. Thepresent-day terrestrial biota of the CaribbeanIslands originated after the Middle Eocene (<40Ma), when permanent land masses became avail-able. These lands (peninsular projections of the con-tinents, archipelagos, isolated islands, and keys),underwent extensive modifications in terms of relief,extension, and geographic position. Periodicclimatic changes also took place, specially since thePliocene. Nevertheless, the lands were able toaccommodate a rich biota that evolved into today’sisland ecosystems. In order to characterize landoccurrence and land connectivity since the LowerOligocene within the Caribbean, Dávalos (2004,Fig. 1) produced a geological area chorogram mostlybased on Iturralde-Vinent and MacPhee (1999).Here I present a different version of the geologicalarea chorogram, extended backward to the MiddleEocene, based on all the plate tectonic and strati-graphic evidence presented in this paper (Fig. 13).This figure can be used to test biogeographicscenarios.

Acknowledgments

The author wishes to thank Ross D. E. MacPhee(American Museum of Natural History, New York)and Zulma Gasparini (Natural History Museum ofthe University of La Plata) for sharing relevant pale-ontological data with the author, and for extensivediscussions about the historical biogeography of theCaribbean land fauna and marine reptiles respec-tively. Silvia Blanco Bustamante (PetroleumResearch Center, Cuba) and Consuelo Díaz Otero(Institute of Geology and Paleontology, Cuba) identi-fied microfossils of key samples for this study. Rein-aldo Rojas, Stephen Díaz, William Suárez (MuseoNacional de Historia Natural, Cuba) and JuanitoGallardo (Museo de Viñales, Cuba), provided greatsupport during field work. Lisa Gahagan and LarryLawver (University of Texas at Austin), providedaccess to the program PLATES for the plate tectonicreconstruction of the Caribbean. Support for thefield and laboratory research for this paper inArgentina and the Greater Antilles was provided bygrants 6001/97, 6009/97, and 6984/1 from theNational Geographic Society, the American Museumof Natural History, the Institute for Geophysics and

the Institute for Latin American Studies of the Uni-versity of Texas at Austin, the Spanish DGES-MECproject BTE2002-01011 (University of Granada,Spain), the Museo Nacional de Historia Natural deCuba, and other sources. Field reconnaissance workcarried out in Central America, and some locationsin the Lesser and Greater Antilles, was also pro-vided by the UNESCO/IUGS International GeologicCorrelation Program for the years 2000–2004.

Note

While this paper was in press, a meeting washeld in Punta Cana, Dominican Republic, July 5–8,2006, dedicated to “Biological Diversification onthe West Indies Archipelago” [www.lacertilia.com/WIndies/]. During this event, among other issues,the paleogeographic background for biologicalcolonization, diversification, extinction, and perma-nence of terrestrial life in the islands was evaluated.Of particular interest to evolutionary biologistswere: (1) land permanence, and (2) the erection anddisappearance of terrestrial connections, bothbetween continent and islands and between islands.

Regarding land permanence, geologists arguethat island landscapes in the past may have been oftwo general kinds: (a) islands with mountains andplains (as Cuba or Puerto Rico today); and (b) shal-lows and low sandy keys with mangrove vegetation(as some Bahamian keys today). These two geo-graphic scenarios imposed different constraints tospecies survival, as the first type of islands may lastlonger, geologically speaking, whereas the secondtype may be ephemeral, subject to reduction ordisappearance by the combined action of tectonicsubsidence, strong hurricanes, tsunamis, and sealevel rise.

Regarding connections between lands, thoseevents may occur due to local or general tectonicuplift (as at the end of the Eocene in the Caribbean)and/or sea level decline. The interruption of landconnections can take place in two steps. Initially amarine transgression may override the connector,producing shallow seas and ephemeral sandy keysand shallows, as occur today between Cuba and theIsland of Youth. Eventually, tectonic forces mayproduce a break-up of the shelf and erect a deepchannel, widening and complicating the gapbetween lands, as the present-day WindwardPassage produced by the Cayman trench system.Mona Passage was presented as an example ofinitial drowning starting in the Upper Oligocene


or the Lower Miocene, later separated by a deepchannel since the Pliocene. Biologically speaking, ashallow-water passage with keys may eventuallyallow overland dispersal of terrestrial organisms, butcan behave more as a filter than as a corridor. TheAves Ridge was presented as an example of a con-necting feature between South America and thenucleus of the Greater Antilles during the Eoceneand Oligocene transition, represented by smallislands, sandy keys, and shallows that may be com-pletely subaerial for a short period of time.

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