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1996 Iturralde-Vinent, M., G. Hubbell, R. Rojas, 1996. Catalogue of Cuban fossil Elasmobranchii (Paleocene to Pliocene) and paleogeographic implications of their Lower to Middle Miocene occurrence,

Boletín de la Sociedad Jamaicana de Geología, vol. 31, p. 7-21.

CATALOG OF CUBAN FOSSIL ELASMOBRANCHII (PALEOCENE--PLIOCENE) AND PALEOOCEANOGRAPHIC IMPLICATIONS OF THEIR LOWER--MIDDLE

MIOCENE OCCURRENCE

M. Iturralde-Vinent1 , G. Hubbell2 , R. Rojas1

1 Museo Nacional de Historia Natural, La Habana, Cuba

2 Jaws International, 150 Buttonwood Drive, Key Biscayne, FL 33149

Las Tablas 1,2, 4 y figura 3 están al final del texto.

Abstract

Although fossil Elasmobranchii have been found in Paleocene, Eocene, Miocene and Pliocene(?) rocks from Cuba, few papers are dedicated to their study and description. This paper includes a revised list of Cuban fossil Elasmobranchii (19 species from 10 genera) based on material preserved in several natural history collections of Cuba and reports in the literature.

Paleocene sharks (Carcharodon auriculatus, Striatolamia sp.) are known from localities in Pinar del Río, western Cuba. Middle to Late Eocene taxa (Carcharodon auriculatus, Striatolamia macrota, Isurus sp.) are known from localities in Havana city, Havana province, and the Sierra Maestra mountains northwest of Santiago de Cuba. Miocene sharks (Carcharhinus obscurus, C. perezi, Carcharias taurus, Carcharodon carcharias, C. megalodon, C. subauriculatus, Galeocerdo contortus, Hemipristis serra, Isurus hastalis, I. sp. cf. I. desori, Negaprion brevirostris, Sphyrna mokarran, S. prisca [?]) are well known from many quarries in western Cuba, especially from those in Havana and Matanzas provinces, but some taxa have been found in several sites throughout the country. Late Miocene-Pliocene(?) C. megalodon teeth are reported from quarries near Cárdenas in Matanzas province. Fossil remains of shark are mostly teeth, but some shark vertebrae are in the collections as well as reported in the literature. Several batoid remains have been found, including a dental plate of the Eocene (?) Aetobatus poeyi, as long as several unidentified tail spines and dental plates of two probably new Middle Miocene taxa, still under study.

The Elasmobranchii species found in Cuba are cosmopolitan, but, interestingly, C. megalodon and some other Lower-Middle Miocene species have been reported only from Havana and Matanzas provinces (western Cuba). This peculiar distribution is probably related to the existence of a Lower-Middle Miocene water channel accross western Cuba (Havana-Matanzas) that connected the Atlantic Ocean with the Caribbean Sea. At that time the Caribbean surface current probably joined the Florida-Gulf Stream currents across the present-day area of Havana-Matanzas, as several additional facts suggest.

Key words: sharks, Tertiary, Cuba, paleooceanography

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Resumen

Dientes fósiles de Elasmobranchios se encuentran en rocas del Paleoceno, Eoceno, Mioceno y Plioceno(?) de Cuba, pero muy pocas obras se han dedicado a su estudio y descripción. En el presente trabajo se ofrece una lista revisada de los Elasmobranchios fósiles cubanos (19 especies de 10 géneros), incluidas algunas especies que se reportan por primera vez para el territorio, a partir del material atesorado en las colecciones de historia natural de los museos cubanos y otras que aparecen en la literatura.

Tiburones del Paleoceno (Carcharodon auriculatus, Striatolamia sp.) se reportan de Pinar del Río en Cuba occidental. Los taxones del Eoceno Medio y Superior (Carcharodon auriculatus, Striatolamia macrota, Isurus sp.) se conocen de localidades en la ciudad de La Habana, la provincia de La Habana y en las montañas de la Sierra Maestra al NW de Santiago de Cuba. Los tiburones del Mioceno (Carcharhinus obscurus, C. perezii, Carcharias taurus, Carcharodon carcharias, C. megalodon, C. subauriculatus, Galeocerdo contortus, Hemipristis serra, Isurus hastalis, I. sp. cf I. desori, Negaprion brevirostris, Sphyrna mokarran, S. prisca [?]) se conocen bien de varias canteras situadas en el occidente de Cuba, especialmente en las provincias de La Habana y Matanzas, pero algunos han aparecido en otras localidades en el resto del país. Dientes de tiburón del Mioceno-Superior-Plioceno(?) (C. megalodon) se reportan de varias canteras cercanas a la ciudad de Cárdenas en Matanzas. Los restos fósiles de tiburón son principalmente dientes, pero en las colecciones hay algunas vertebras que han sido reportadas antes en la literatura. De Cuba se ha descrito el batoide del Eoceno(?) Aetobatus poeyi, y se encuentran, en lechos del Mioceno, espinas de la cola de batoides, así como dos placas dentales en proceso de identificación.

Las especies de tiburones encontradas en Cuba son cosmopolitas, pero es muy interesante que C. megalodon junto a otras especies del Mioceno Inferior-Medio se hayan encontrado solamente en Cuba occidental (La Habana y Matanzas). Esta distribución peculiar está relacionada probablemente con la existencia de un canal marino en Cuba occidental (Habana-Matanzas) durante el Mioceno Inferior a Medio, el cual comunicaba el Oceano Atlántico con el Mar Caribe.

INTRODUCTION

Fossil Elasmobranchii were reported in Cuba since the beginning of the XVIII century by different naturalist as Felipe Poey y Alloy, José Ramón de la Sagra, L.J.R. Agassiz, Justo Egozcue y Cia, and Manuel Fernández de Castro. This last author in 1873, in a paper describing a new fossil species of the Eocene(?) batoid (Aetobatis poeyi) 1, listed the following Cuban shark taxa: Carcharodon megalodon, C. heterodon, C. auriculatus, C. angustidens, Hemipristis serra, Sphirna prisca, Oxyrhina quadrans, O. hastalis, O. plicatilis, Odontapsis or Lamna

1 This species now is named Aetobatus poeyi (Fernández de Castro), Kruckow and Thies (1990). But the author's family name Fernández de Castro is incorrectly cited as "Castro" in this and other references.

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hopei, identified by Prof. D. Justo Egozcue y Cia2. In 1876 Fernández de Castro presented a catalog of Cuban fossils that included five genera and 13 species of sharks, but the names of the taxa were not listed. He also reported that C. megalodon teeth were found frequently west of Matanzas, and he identified H. serra as a typical Tertiary fossil from Cuba. The most important taxonomic paper about Cuban sharks was that of M. Sánchez Roig (1920) in which 12 species from six different genera were described and some were illustrated.

The senior author started this project in 1992 with the purpose of evaluating the biogeographic information that Cuban fossil Elasmobranchii might provide for use in tectonic and paleogeographic reconstruction of the Caribbean area. Following is an interesting result related to the Lower-Middle Miocene paleooceanography.

Abbreviations

MNHNH: Museo Nacional de Historia Natural de La Habana

MNHNH-P: Paleontological collections of Museo Nacional de Historia Natural de La Habana

IGP: Instituto de Geología y Paleontología de La Habana

CATALOG OF CUBAN FOSSIL ELASMOBRANCHII

The authors have examined the paleontological collections of Museo Nacional de Historia Natural in Havana, Museo Provincial "Ignacio Agramonte" in Camagüey, Museo Municipal de Cárdenas in Matanzas, and Instituto de Geología y Paleontología in Havana (where some of Sanchez Roig's materials are stored), and the private collection of J. L. Fernández Millera stored at the Instituto de Ecología y Sistemática in Havana. Unfortunately, some of these materials have very poor locality descriptions and age assignments are often questionable. For several examples, we were unable to identify the stratigraphic horizon with certainty. During the last four years, several paleontological sites yielding well localized material were established by the MNHNH staff, in collaboration with curators of local museums and paleontologists from the American Museum of Natural History (AMNH, New York).

Here is reported for Cuba 19 species belonging to 10 different genera (table 1), most of them of worldwide distribution. Some of these species are illustrated in plate I. We identified the Cuban fossil material available in Cuban public and private collections, and the specimens were later sent for verification to Robert Purdy (Smithsonian Institution, Washington, D.C.). The identification of those specimens that were not available, was evaluated according to the illustrations provided in the paper, as in the example of Sánchez Roig (1920). The species listed without illustrations (i.e: Fernández de Castro 1873, 1876; Burchard, 1920; Sánchez Roig, 1928; Kruckow and Thies, 1990) were not accepted as a good record.

2 These fossils were exposed in the Universal Exhibition held in Paris, 1867.

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The fossil sharks reported by Fernández de Castro (1973, 1976) were not available, but some of the taxa have been reported later and there is fossil material available. Leriche (1938) revised the identifications of Sánchez Roig (1920) assuming that the age of the specimens was Miocene or Pliocene, so he misidentified several species (Table 2). Sánchez Roig's figured material probably were lost, but additional material collected and identified by him is preserved at the collection of IGP. Table 2 indicates our assignment of the specimens figured by Sánchez Roig (1920).

In 1928, Sánchez Roig published a catalog of the shark teeth stored in the collections of the Instituto Nacional de Investigaciones Científicas y Museo de Historia Natural in Havana. The catalog included specimens he identified as Lamna elegans, L. longidens, L. cuspidata, Isurus hastalis, Carcharodon auriculatus, C. lanciformis and C. megalodon. He only illustrated C. megalodon, and we could not validate his other identifications. In the same paper, Sánchez Roig corrected to Eocene his previous Miocene age assignment for the sequence of "Tejar Consuelo" (Sánchez Roig, 1920). But both Eocene and Miocene shark-bearing rocks crop out at the quarry (Bronnimann and Rigassi, 1963), as we will discuss later.

At the Instituto de Geología y Paleontología we examined shark teeth collected and identified by Sánchez Roig from "Tejar Consuelo" quarry in the city of Havana, but none of the specimens can be identified as those illustrated in his 1920 paper. In this collection his identification of Lamna longidens is a fragmentary tooth of Carcharodon sp. Several teeth labeled as "Middle Eocene" Isurus desori should be referred to Stratolamia macrota. An eroded tooth identified as Hemipristis serra has some similarities to Galeocerdo. Several teeth were correctly labeled as G. contortus and I. hastali.

From Charco Redondo Mine J. W. Gidley (in Burchard 1920) identified three species of sharks. He explained that Lamna cf. L. elegans teeth "are not typical L. elegans but seemed nearer this species than any other described. I. hastalis, or very near this species. Carcharodon auriculatus (?), the large serrate tooth is like this species only more slender than usual. These teeth seem to be of Cretaceous or Tertiary, probably Eocene age." We do not know the repository of these specimens, and they were not illustrated, so we can not verify Gidley's identifications. The collection of IGP contains specimens identified as I. desori and I. hastalis from the Charco Redondo Mines district (table 3). Most of these teeth can be referred to S. macrota, but some are definitely Isurus sp. Additionally, in the catallog of IGP it is recorded I. hastalis associated with Carcharodon sp. from an Eocene locality in central Cuba.

Kruckow and Thies (1990, Table 12) listed Aetobatus poeyi as Eocene, and the following Miocene species of Cuban Elasmobranchii (Eugomphodus acutissimus, E? sp., Procarcharodon megalodon, Carcharodon carcharias, Isurus desori, I. hastalis, Hemipristis serra, Galeocerdo aduncus, Sphirna sp.). This is the list of Leriche (1938) based on Sánchez Roig's (1920) descriptions and illustrations. Leriche assumed that all the specimens illustrated by Sánchez Roig from "Tejar Consuelo" quarry were Miocene or Pliocene in age, so he review Sánchez Roig's identifications with this opinion in mine. The authors have found that latest Eocene and Miocene fossils were actually recovered from "Tejar Consuelo" quarry, so the identifications by Leriche (1938) require further revision.

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Table 2 show the species identified and figured by Sánchez Roig (1920), the opinion of Leriche (1938) concerning these taxa, and that of the present authors taking into consideration not only the descriptions and excellent illustrations of Sánchez Roig, but also additional material from "Tejar Consuelo" quarry collected by Sánchez Roig. Representative specimens are illustrated in figure 1.

Figure 1. Photographs of some Cuban fossil shark teeth, actual size. For locality of specimen see table 3. A. Carcharhinus obscurus, MNHNH-P-3033. B. Carcharhinus perezi, MNHNH-P-2027. C. Carcharias taurus, MNHNH-P-2166. D. Carcharodon auriculatus, left to right: MNHNH-P-2043, 2049, 2048. E. Carcharodon subauriculatus, MNHNH-P-2142, 2150. F. Galeocerdo contortus, MNHNH-P-2216. G. Hemipristis serra, MNHNH-P-3036. H. Hemipristis serra, MNHNH-P-3034. I. Negaprion brevirostris, MNHNH-P-3037. J. Sphyrna mokarran, MNHNH-P-3032. K. Striatolamia macrota, MNHNH-P-2054-2057.

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Figure 2. Location map of main Cuban Elasmobranchii fossil localities. Distribution of Carcharodon

megalodon fossil sites and phosphatic rocks in Cuba.

A comprehensive catalog of Cuban fossil Elasmobranchii is presented in table 3. The following paragraphs comment on the localities and ages for Cuban fossil Elasmobranchii.

Paleocene

Paleocene shark teeth are found in the MNHNH-P collection, with additional material loaned from the private collection of J. M. Fernández Milera. The manganese-bearing matrix from all the specimens suggest localities in the Cordillera de Guaniguanico, Pinar del Río, where manganese ore deposits are concentrated in the deep-water limestones and shales of the Paleocene Ancón Formation. There are few possibilities for the younger formations to be fossiliferous. The Lower Eocene rocks on top of Ancón are devoid of macrofossils because represent clastic to chaotic synthrust sedimentation in the foldbelt. Middle Eocene and younger rocks are not known from Guaniguanico (Pszczolkowski 1978, Iturralde-Vinent 1994).

The actual paleontological sites are poorly identified. Some teeth found in the collection of MNHNH-P (table 3) are reported to be from Pico Chico, Sierra de San Andrés, about 6 km south of La Palma. Two manganese mines are known near this location. The site yield Striatolamia sp. and additional unidentified shark teeth. Several teeth loaned by J. M. Fernández Milera belong to a manganese deposit in "Alturas de Pizarras, Cordillera de Guaniguanico, Pinar del Río." Carcharodon landenensis is identified from his collection.

Another locality bearing shark teeth in Guaniguanico, poorly located as "near the small village of San Cayetano", was reported to Rafael Quiñones (MNHNH) by a group of young naturalists. They collected many specimens, but most of them were lost. A fragmentary tooth can be refered to C. auriculatus, and two fragmentary teeth as Striatolamia sp. All of them have manganese stain.

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Eocene

Eocene fossil shark localities are known in eastern Cuba (Charco Redondo Mines district), central Cuba (near Florida and near Cienfuegos), Havana Province (Nazareno), and Havana city ("Tejar Consuelo" quarry).

The manganese ores with fossil sharks in Charco Redondo Mines district are known to be associated with the Middle Eocene Charco Redondo limestones (Burchard 1920; Lewis and Straczek 1955). We identified C. auriculatus and S. macrota (tables 1) from a collection of several hundred teeth found embedded in the manganese ore (pyrolusite) of the Charco Redondo Mine. The collection is housed at Museo Provincial "Ignacio Agramonte" in Camagüey, but some representative specimens are deposited in the paleontological collections of MNHNH (table 3). According to Burchard (1920) the fossiliferous horizon is 1.8 to 3 m thick breccia with clays, sand, and boulders of oolitic limestone in manganese ore matrix with ..."a crust slightly richer in manganese oxide" with abundant shark teeth. The breccia is located within limestones and argillaceous limestones of the Charco Redondo Formation of early Middle Eocene age (Lewis and Straczek 1955). The shark species identified by J.W. Gidley (in Burchard 1920) were not illustrated so they can not be verified. From the Charco Redondo Mines district are stored shark teeth at the IGP collection (table 3) that the authors identified as C. auriculatus, S. macrota and Isurus sp. The specimens of Isurus sp. resemble I. hastalis, but are smaller and fragmentary.

From the area near Central (sugar mill) Argentina, in the Camagüey province, Late Oligocene I. hastalis and Carcharodon sp. are reported in the IGP catalog, but the specimens were lost. It is not known who identified the species. The senior author mapped the area of Central Argentina and found only Lower to Middle Eocene rocks.

Also at the IGP collection there is a tooth collected near Nazareno, a small town at the Havana Province, that can be identified as S. macrota (table 3). In this area the senior author mapped the Middle to Late Eocene sequence of the Nazareno Group (Albear and Iturralde-Vinent 1985).

The sharks teeth from "Tejar Consuelo" quarry unfortunately were mixed Eocene and Miocene specimens during the collection (Sánchez Roig, 1920; 1928). Eocene taxa can be easily identified, but the problem is that different Eocene units crop out at "Tejar Consuelo" quarry (Bronnimann and Rigassi, 1963): the Lower Eocene Capdevila Formation (sandstones and shales), the Middle Eocene Universidad Formation (deep-water radiolarian marls), and the Upper Eocene Consuelo Formation (marls with planktonic forams, gastropods and echinoids). At the same locality, the Lower Miocene Husillo limestones and the Middle Miocene Cojimar marly limestones and limestones crop out. No reworking is reported in this Miocene sequence (Bronnimann and Rigassi, 1963; Albear and Iturralde-Vinent, 1985). The source of the Eocene shark teeth can be either the Universidad or Consuelo Formations, as the Eocene teeth are embedded in marls (Sánchez Roig, 1920, Fig. 13). From this quarry fossil can be identified as latest the Middle to Late Eocene species: C. auriculatus (tables 2) and S. macrota (table 3). The specimens of S. macrota preserved at the IGP collection are embedded in yellow soft marls. The rock was sampled for age identification. According to Timothy Bralower (University of North Carolina), José Fernández and Silvia Blanco (Cuba Petroleum, CUPET), the samples include the following latest Eocene planktonic

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microfossils (nannofossils zones NP19-20): Chiloguembelina sp., Reticulofenestra hillae, R. umbilicus, Dictyococcoites scrippsae, D. bisectus, Sphenolithus predistentus, Calcidiscus protoannulus and the very rare Discoaster barbadiensis, Ericsonia subdisticha, E. formosa and Campylosphaera dela. Therefore, it is possible to refer C auriculatus and S. macrota from Sánchez Roig's materials to the Late Eocene Consuelo Formation.

The batoid Aetobatus poeyi was described by Fernández de Castro (1873) from a locality in the lands of Ingenio (sugar mill) Constancia (today Moncada) northwest of the city of Cienfuegos (Fig. 2). The Eocene age for this species assigned by Fernández de Castro (1873) is questionable. According to Fernández de Castro (1973) the fossil was collected in rocks that crop out in the lower reach of Damují river basin. He supposed that the underlying rocks are of Cretaceous age, and indicate that the overlain horizon resemble the C. megalodon -bearing rocks in other localities of the country. The age needs to be verified in the future.

Lower and Middle Miocene

Lower and Middle Miocene Elasmobranchii are common in Cuban natural history collections, reported mostly from localities in western Cuba (fig. 2).

A well-dated late Lower Miocene assemblage was found at Domo de Zaza in south-central Cuba. From this site were collected very few teeth of Carcharinus obscurus, Hemipristis serra, Negaprion brevirostris, Sphyrna mokarran and unidentified batoid tail spines (table 3). The site has been described in greater detail elsewhere (MacPhee and Iturralde-Vinent, 1994; MacPhee and others, in prep.). The section, about ten meters thick, is composed of interbedded shallow marine limestones and marls, alluvial sands and gravels, lagoonal clays and paleosols. The shark teeth and batoid teeth and tail spine are found in lag deposits, derived from the marine units. At this site fossil remains of bony fishes, crocodiles, sirenians (dugongs), turtles and land mammals (including a sloth, a primate and a rodent) were found. Fossils reworked from older horizons are not found.

At several quarries within the city of Havana Lower and Middle Miocene limestones and marls have yielded shark teeth (Sánchez Roig 1920, 1928; table 3). From Miocene (Cojímar Formation) rocks in "Tejar Consuelo" quarry we were able to validate as probably Miocene taxa: H. serra, C. megalodon, C. carcharias, I. hastalis, C. taurus, G. contortus and S. prisca (?).

Another Lower-Middle Miocene site is the Phosphorite Mine (Mederos 1987) west of Loma Candela, 6 km northwest of Güines, in the province of Havana (fig. 2). At this locality Carcharias taurus, C. subauriculatus, I. hastalis and I. sp. cf. I. desori (table 3) were identified from approximately 200 fragmentary teeth collected from a thin phosphorite bed at the base of the Middle Miocene limestone section (Cojímar Formation). These teeth are associated with fragmentary sirenian bones and a fragment of a batoid tail spine. The material lack evidence of reworking from older sources. The bed comprises rounded, non-fossiliferous, calcareous, phosphorite-rich pellets, a few centimeters in diameter, directly overlying Lower Eocene deep-water marls lacking macrofossils. There were no microfossils in the matrix of the phosphorite bed, but its stratigraphic position is almost certainly Lower-Middle Miocene (Mederos 1987).

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The senior author has contacted private collectors in Havana, and has learned that C. megalodon teeth are being found at Cantera del Husillo (type locality of the Lower Miocene Husillo Formation of Bronnimann and Rigassi, 1963) and other quarries in the city that exploit the same limestones. The IGP records contain an unverified report of C. megalodon from the type locality of the early Middle Miocene Cojímar Formation, in Cojímar, near Havana city (table 3).

At several localities in the city of Havana, and in the provinces of Havana and Matanzas (fig. 2) C. megalodon teeth have been recovered with those of Carcharhinus perezi, Hemipristis serra and Galeocerdo contortus (table 3). The locations of these fossil sites are too general to enable the identification of the horizons that could include the Lower Miocene Husillo limestones, the Middle Miocene Cojímar marls and limestones, or the Lower-Middle Miocene Güines limestones (Iturralde-Vinent, 1969; Albear and Iturralde-Vinent, 1985). Fossil remains of dugongs have been reported in the Lower-Middle Miocene limestones of the Güines Formation as well.

A fragmentary dental plate recovered from the Middle Miocene Cojímar Formation was identified by Sánches Roig (IGP collection, table 3) as Aetobatus poeyi . The identification of the species, collected from Cantera Cojímar in Havana, is highly questionable, in the present state of preparation of the specimen. Another very large and complete batoid dental plate embedded in Middle Miocene limestones of the Güines Formation was recently discovered in a cave located near Cueva de Paredones, south of Ceiba del Agua (table 3). Both specimens are currently under study.

Late Miocene-Pliocene(?)

Quarries near the city of Cárdenas produce Late Miocene-Pliocene(?) calcarenites of the Canímar Formation (Iturralde-Vinent, 1969) (fig. 2). The quarries at Loma Fines offer some of the best fossil-collecting sites in the region. From them several well-preserved teeth of C. megalodon recovered by J. M. Fernández Milera were identified by present authors and reside in the collections of Museo Municipal de Cárdenas (Matanzas) and in MNHNH (table 3). Workers at the Loma Fines quarries claim to have found an entire jaw of C. megalodon and a body impression, but only the teeth were recovered. Bony fish impressions are known from Loma Fines quarries. Many large teeth are owned by private collectors.

PALEOOCEANOGRAPHIC IMPLICATIONS OF LOWER-MIDDLE MIOCENE SHARKS BIOGEOGRAPHY

Lower to Middle Miocene Elasmobranchii have an interesting distribution in Cuba that deserves special consideration. Figure 1 shows that the gigantic C. megalodon has been reported only in western Cuba (Havana and Matanzas provinces). The senior author has conducted field work and paleontological prospection throughout the entire island without discovering further C. megalodon localities, although Lower and Middle Miocene rocks are well represented in the whole country (Iturralde-Vinent, 1969, 1994). The huge teeth of C. megalodon are very popular among collectors. They are often collected by workers at quarries, constructions sites and farm fields, but no reports of this species are known elsewhere in the country.

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Figure 3. Paleogeographic maps of the Caribbean indicating paleocurrents. A from Early Miocene to Early Middle Miocene, b. from latest Eocene to Early Oligocene. Modified from Emery and Uchupi (1972), Riggs (1984), Iturralde-Vinent (1969, 1988), Mederos (1987), Mederos and Franco 1992, Denny (1992), Denny and others (1994), MacPhee and Iturralde-Vinent (1994), Atlas Nacional de Cuba (1970) and data presented in this paper. Occurrences of C. carcharodon after Kruckow and Thies (1990).

A comparison (table 4) shows that different Lower and Middle Miocene shark assemblages occur in central (Domo de Zaza) and western Cuba (Havana-Matanzas), with only one species common to both areas (Fig. 2). The taxa found at Havana-Matanzas includes some sharks whose present-day representatives are found strictly in deep oceanic basins. Isurus species and C. carcharias live in those conditions today, and C. megalodon would probably have preferred the same habitat. In contrast, the Domo de Zaza assemblage includes species that prey in shallow, nearshore environments, such as Negaprion brevirostris. These distributions are based on collections of fossils representing five years detail work at Domo de Zaza, and more than one hundred years of collecting around Havana and Matanzas. Thus, the pattern of distribution of C. megalodon and associated species may not be an artifact, but reflects environmental and/or biological factors. Other occurrences of C. megalodon in the Caribbean basin and its surroundings (Kruckow and Thies (1990) and figure 2) suggest that the gigantic shark was

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preying across the Caribbean Sea, moving from and to- the Pacific and Atlantic Oceans.

Table 4. Lower--Middle Miocene sharks from localities in Havana and Matanzas provinces and central Cuba (= Domo de Zaza).

Taxon Domo de Zaza

Havana-Matanzas

Carcharhinus perezi X Carcharihnus obscurus X Carcharias taurus X Carcharodon carcharias X Carcharodon megalodon X Carcharodon subauriculatus X Galeocerdo contortus X Hemipristis serra X X Isurus hastalis X Negaprion brevirostris X Sphyrna mokarran X Sphyrna prisca (?) X

The known distribution pattern of C. megalodon and other sharks, as previously discussed, will be further explored in order to evaluate its possible implications.

Stratigraphic and paleogeographic research in Cuba demonstrate that during the Latest Eocene and untill early Middle Miocene most of present-day Cuba and its marine shelf were covered by deep-water basins separated by shallows seas and isolated lands (see paleogeographic maps in Iturralde-Vinent, 1969, 1988; MacPhee and Iturralde-Vinent 1994; MacPhee and others, in prep.; also Brezsnyánszky e Iturralde-Vinent 1978, 1985). Sea-water circulation accross present day Cuba most probably took place along two main deeper water channels, one in western Cuba (Havana-Matanzas Channel) and another in eastern Cuba, the northeast trending Cauto-Nipe Channel (figs. 12 - 14 of Iturralde-Vinent, 1988). The Havana-Matanzas Channel efficiently connected the Caribbean Sea with the Atlantic Ocean via the Strait of Florida; and the Cauto-Nipe Channel connected the Caribbean Sea with the Old Bahama Channel (Fig. 3).

During Middle Miocene an important tectonic event took place in Cuba and both deeper-water channels practically close due to shallowing of the sea and homogenization of the marine environment. Extensive carbonate platform deposition took place all along the Cuban territory (Iturralde-Vinent, 1969, 1978, 1988, 1994; MacPhee and Iturralde-Vinent 1994, MacPhee and others, in prep.). Before this event took place large parts of the present day Cuban territory were covered by shallow waters, with small islands, keys and locally shallows. Coastal plains, ponds estuarine and lagoonal environments are represented by clays, sandy clays, gravels, marls and a few limestones intercalations which yield a fossil biota typical of fresh, and brackish waters. This kind of sections are represented by the Paso Real (pars), Arabos, Lagunilla and Magantilla Formations. Local carbonate banks (biogenic limestones and calcarenites) are represented by the Paso Real (pars), Husillo and Güines (pars) Formations (Iturralde-Vinent 1969, 1978, 1988). There are two exceptions to these shallow-water depositional pattern. The Havana and Matanzas Channel that was a NNW-SSE trending broad marine basin were sedimentation of hemipelagic marls and limestones of the

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Jaruco and Cojímar Formations took place (Bronnimann and Rigassi, 1963; Iturralde-Vinent 1969; Albear and Iturralde-Vinent, 1985); and the Cauto-Nipe Channel were hemipelagic marls and limestones of the "Nipe Series" were deposited (Iturralde-Vinent, 1969; 1978). This paleogeographic framework drastically changed during the second half of the Middle Miocene, around the Globorotalia fohsi-Globorotalia mayeri zones, when ended the deposition of the Cojímar and equivalent deeper-water Formations and most of Cuba was covered by extensive carbonate platform environments. Later since the Late Miocene the whole territory emerged nearly as it is today, and no further marine channel evolved up to the present (Iturralde-Vinent, 1969, 1988).

Seismic stratigraphy studies carried out in the southern Florida Channel provide important information related to the paleogeographic framework described for Cuba. From Maastrichtian to the late Oligocene the west Florida ramp aggraded. There was little winnowing and no evidence of erosional surfaces. Within the late Oligocene-early Miocene period this aggradational ramp began to prograde from the east as illustrated by west dipping clinoforms. The Suwannee Strait closed at this time deflecting the Suwanee Current to the south thus forming the precursor to the modern Loop Current, but not as strong as the modern system (Hine 1993). During the Middle Miocene (12-15 M.y.), the Florida carbonate-ramp slope experienced an oceanographic event related to strong seafloor erosion that resulted in a major stratigraphic break (Mullins et al. 1987). Erosion took place on the Pourtales Terrace (Gomberg, 1974) and the Miami Terrace (Mullins and Newmann, 1979) along with a 50% decrease in the rate of sediment accumulation in the southern Straits of Florida (Austin and others, 1988). The modern Loop Current/Gulf Stream circulation was initiated and carbonate deposition changed drastically into a pelagic slope-front-fill system (Mullins et al. 1987). The usual explanation for this Middle Miocene event is that it was related to major tectonic changes within the Caribbean. This tectonic event is usually interpreted as the closure of the Panama Strait and collapse of the Nicaragua megabank, and therefore the Caribbean Current was feeded by the Guyana Current from the equatorial Atlantic and the speed of the flow increased in the Loop Current (Mullins et al. 1987, Hine 1993).

The paleogeographic and oceanographic change recorded in Cuba and the Straits of Florida within the Middle Miocene have counterparts in the Nicaragua Rise (Droxler and others, 1989), the Aves Ridge (MacPhee and Iturralde-Vinent 1995), and may be related to general tectonic deformation in the Caribbean (Mann and others, 1990, Iturralde-Vinent, 1994). The Late Tertiary paleogeographic reconstruction of the eastern Caribbean (Fox, 1971; Donnelly, 1989; MacPhee and Iturralde-Vinent, 1994, 1995) suggests that during the latest Eocene-early Miocene, the Aves Ridge existed as a N-S possitive barrier between South America and the Greater Antilles. This feature strongly reduced the sea-water circulation from the Equatorial Atlantic into the Caribbean. The Nicaragua megabank also represent a major obstacle for free circulation of the marine waters. At that time, the Caribbean Current possibly was fed only from the Pacific Ocean through the "Straits of Panama", resulting in a considerably lessened water flow. Subsidence of the Aves Ridge since the Miocene and up to the present (Fox et al. 1971), and the fragmentation of the Nicaragua rise (Droxler and others, 1989), gave rise to the continuity between the Guyana and the Caribbean Current and produced an increase of the water flow toward the west and northwest (Fig. 3).

12

Information concerning phosphatic deposits shade new light on this subject. Lower to Middle Miocene phosphatic rocks are known in different places in Havana-Matanzas and Banes (Cuba, fig. 2), the Florida peninsula (Scott 1988) and South Carolina (Riggs, 1984; Mederos, 1987), associated with shallow marine rocks fringing deep seaways (fig. 2). According to Mederos (1987) and Mederos and Franco (1992) the Caribbean Current was the source of the phosphate for the Cuban deposits, and speculated that the phosphatic sources were submarine volcanoes within the Caribbean sea floor, the Lesser Antilles and northern South America (see his figure 9). We agree that the Caribbean surface current was involve in the transportation of phosphates, but more probably as nekton and zoo-phytoplankton that, after dead, sank to the sea botton. The biomass in the warm Guyana/Caribbean Currents was probably high, enhanced by the organic input of the Orinoco, Amazon and other South American rivers. In Cuba, the deeper-water Jaruco and Cojímar Formations yield abundant benthic and nectic biotas, as long as a large number of planktic microfossils (Bronnimann and Rigassi, 1963; Iturralde-Vinent, 1969). The concentration of the Lower and Middle Miocene phosphates in the Havana-Matanzas and Banes areas of Cuba (fig. 2) is in agreement with the paleogeographic pattern generally recognized for the formations of these ores. It is, a shallow shelf with low detrital imput fringing a major seaway (Havana-Matanzas and Cauto-Nipe Channels). In this conditions, upwelling of cold waters into the shelf concentrate the phospates that often yield abundant fossil shark teeth (Riggs, 1984; Mederos and Franco, 1992).

Previous data provide the background necessary to explain the Lower--Middle Miocene biogeography of C. megalodon. The Late Tertiary paleogeography of the Caribbean as discussed previously suggests that during the Latest Eocene-Oligocene there existed several barriers-the Aves Ridge and the Nicaragua megabank-that impede active deep and surface water circulation within the Caribbean Sea. That pattern would explain why the Loop/Gulf Stream Current flowed slowly through the Straits of Florida during late Eocene, Oligocene and early Miocene epochs, producing active progradation of the west Florida slope (Mullins et al. 1987, Denny, 1992; Hine 1993; Denny and others, 1994). Continual subsiding of the Aves Ridge (Holcombe and others, 1990) and fragmentation of the Nicaragua megabank (Droxler and others, 1989) during the Lower Miocene, modified this flow pattern. Therefore, the South Atlantic (= Guyana) current has been increasing the water flow rate into the Caribbean Sea, since the Miocene.

As a consequence of these events, the Caribbean Current heading northwest divided into two branches in the western Caribbean-one following the Yucatan Channel, the other flowing across the Havana-Matanzas Channel (Figure 2). The branch following toward the Yucatan Channel join the Loop Current and produced the Early Miocene progradation from the east in the west Florida slope. The other branch turned due north just southeast of the Isle of Youth (south of central Cuba), then flowed across the Havana-Matanzas Channel to join the Florida-Gulf Stream flow. Under such conditions, Lower Miocene and early Middle Miocene sharks swam between the Atlantic shores of North America and the Caribbean Sea via the Havana-Matanzas Channel, that offered a favourable environment rich in prey.

About late Middle Miocene, the Havana-Matanzas Channel closed in response to extensive shallowing of marine environments and the uplifting of some parts of the Cuban territory, confining the Caribbean Current to the Yucatan Channel, as it

13

is confined today. This concentration of flow could possibly explain the strong erosion that took place during the Middle Miocene in the southern Straits of Florida (Mullins et al., 1987; Austin and others, 1988; Denny and others, 1994). The timing of this event can be coincidental with the change from deep to shallow water environment in western Cuba, a process that took place within the Globorotalia foshi s.l. and the Globorotalia mayeri zones (age of the top of the Cojímar Formation; Iturralde-Vinent 1969).

Conclusions

1. It is presented a revised list of Cuban fossil Elasmobranchii (19 species from 10 genera) based on material preserved in several natural history collections of Cuba and reports in the literature.

2. These Cuban taxa were recovered from Paleocene, Middle Eocene, Late Eocene, Lower-Middle Miocene and Late Miocene-Pliocene rocks.

3. Lower to Middle Miocene Elasmobranchii known from western Cuba (Havana-Matanzas) are different from those found elsewhere in the territory. The Havana-Matanzas sharks include deep-water dwelling species as Isurus, C. calcharias and C. megalodon.

4. This biogeographic pattern in Elasmobranchii, as long as stratigraphic and paleogeographic research demonstrate that from Late Eocene untill early Middle Miocene there was a deep-water channel in western Cuba (Havana-Matanzas Channel). The Caribbean Current flowed along this channel since Late Oligocene untill Middle Miocene, but this water gate closed within the Globorotalia fohsi s.l.-Globorotalia mayeri zones.

5. The closure of the Havana-Matanzas Channel was probably coincidental with important tectonic and oceanographic changes that took place elsewhere in the Caribbean and the west Florida slope.

Acknowledgments

The authors wish to thank RARE Center for Tropical Conservation for partially supporting this research through its Caribbean Scientific Initiative. Alfonso Silva Lee (Museo Nacional de Historia Natural, La Habana) kindly photographed the specimens. We thank Clare Flemming (AMNH) for copy-editing and proofreading. Drawings of shark teeth were produced by Stephen Díaz Franco (MNHNH) and computer illustrations by the senior author. The authors gratefully acknowledge the suggestions offered by J. D. Steward (NHM, Los Angeles County), Marcelo de Carvalho (AMNH, New York), Richard T. Buffler (UTIG, Austin) and Albert Hine (CMR, University of South Florida). We specially thank Robert Purdy (Smithsonian Institution, Washington, D. C.) for his revision of our identifications and his very careful critical review of the first draft of the manuscript. José Fernández and Silvia Blanco (Cuba Petroleum, La Habana) and Tim Bralower (University of North Carolina, Chapel Hill) identified the microfossils from two samples. The authors are grateful to curators of the Instituto de Geología y Paleontología and the natural history collections in Cárdenas, Camagüey and other cities of Cuba for permission to study their respective fossil collections, to J. M. Fernández Milera, who loaned his private collection and provided important information and to J. A. Genaro who provided some specimens and important locality information.

References

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14

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Albear, J. and Iturralde-Vinent, M., 1985. Estratigrafía de las provincias de La Habana. in: Contribución a la Geología de las provincias de La Habana y Ciudad de La Habana, Editorial Científico-Técnica, La Habana, pp. 12--54.

Bresznyánszky, K. and Iturralde-Vinent, M., 1978. Paleogeografía del Paleógeno de Cuba oriental: Geologie en Mijnbow 57 (2): 123-133.

Bresznyánszky, K. and Iturralde-Vinent, M., 1985. Paleogeografía del Paleógeno de La Habana: Contribución a la Geología de las provincias de La Habana y Ciudad de La Habana, Editorial Científico-Técnica, La Habana, pp. 100--115.

Bronnimann, P. and Rigassi, D., 1963. Contribution to the geology and paleontology of the area of the city of La Habana, Cuba, and its surroundings. Eclog. Geol. Helvetiae 56 (1): 193-480.

Burchard, E. F., 1920. Manganese ore deposits of Cuba. Am. Inst. Min., Eng. and Trans. 63: 51-104.

Cappetta, H. 1987. Condrichthyes II, Mesozoic and cenozoic Elasmobranchii. Handbook of Paleoichthyology 3B, Gustav Fischer Verlag, Sttugart, 1-193.

Denny, W. A. III, 1992. Seismic stratigraphy and geologic history of Mid-Cretaceous trough Cenozoic rocks, Southern Straits of Florida: Master's Thesis, University of Texas at Austin, 1-160.

Denny, W. A. III, Buffler R. T. and Austin, J. A., 1994. Seismic stratigraphy and geologic history of Mid-Cretaceous through Cenozoic rocks, Southern Straits of Florida. American Association Petroleum Geologists Bull. 78 (3): 461-487.

Donnelly, T. W., 1989. History of marine barriers and terrestrial connections: Caribbean paleogeographic inference from pelagic sediment analysis. In: C. A. Woods (ed.), Biogeography of the West Indies. pp. 103-118. Gainesville, Fl.: Sandhill Crane Press.

Droxler, A., Staples, S.A., Rosencrantz, E., Buffler, R.T., and Baker, P.A., 1989. Neogene tectonic desintegration of a carbonate Eocene-Early Miocene megabank along the northern Nicaragua Rise, Caribbean Sea: Geology of the Caribbean, Third Annual Meeting, St. Croix, U. S. Virgin Islands, 1989, Proceedings.

Emery, K. O. and Uchupi, E., 1972. Western North Atlantic Ocean: Topography, rocks, structure, water, life, and sediments. American Association Petroleum Geologists Mem. 17: 532 p.

Fernández de Castro, M., 1873. Aetobatis poeyi: Anales de la Soc. Española de Historia Natural II, Madrid.

Fernández de Castro, M., 1876. Catálogo y cuadro sinóptico de los fósiles cubanos. Anales de la Academia de Ciencias de la Habana , XIII: 319-330.

Fox, P.J., E. Schreiber, and B. C. Heezen, 1971. The geology of the Caribbean crust: Tertiary sediments, granitic and basic rocks from the Aves Ridge. Tectonophysics 12: 89-109.

Gomberg, D., 1974. Geology of the Pourtales Terrace. Florida Science v. 37, suppl. 1: 15 p.

Holcombe, T., Ladd, J.W., Westbrook, G., Terence Algar, N., Bowland, C.L., 1990. Caribbean marine geology; ridge and basins of the plate interior, in Dengo, G. and Case J.E., eds. The Caribbean region: The Geology of North America, Volume H: Boulder, Colorado, Geological Society of America, p.231-260

Iturralde-Vinent, M., 1969. Principal characteristics of Cuban Neogene stratigraphy. American Association Petroleum Geologists Bull. 53 (9): 1938-1955.

Iturralde-Vinent, M. 1978. Los movimientos tectónicos de la etapa de desarrollo platafórmico de Cuba: Geologie en Mijnbow 57 (2): 205-212 Holand [also 1979: Geotektonics (4): 63-76, Moscow (English version)].

Iturralde-Vinent, M., 1988. Naturaleza Geológica de Cuba. Editorial Científico-Técnica, La Habana, 246 pp.

Iturralde-Vinent, M., 1994. Cuban Geology: A new plate tectonic synthesis. Journal of Petroleum Geology, 17 (1): 39-71.

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Kruckow, T. and Thies, D., 1990. Die neoselachier der Paläokaribik (Pisces: Elasmobranchii): Courier Forsch.-Inst. Senckenberg: 119: 102 p.

Leriche, M. 1938. Contribution à l'étude des poissons fossiles des pays riverains de la Méditerrané américaine (Venezuela, Trinité, Antilles, Mexique): Mémoires de la Societé Paleontologique Suisse LXI:

Lewis, G. E. and Straczek, J. A., 1955. Geology of south-central Oriente Province, Cuba. U.S. Geol. Survey Bull. 975-D: 171-336.

MacPhee, R. and Iturralde-Vinent, M., 1994. First Tertiary Land Mammal from Greater Antilles: An Early Miocene Sloth (Xenarthra, Megalonychidae) From Cuba. Amer. Mus. Novitates, 3094: 1-13, New York.

MacPhee, R. and Iturralde-Vinent, M., 1995. Origin of the Greater Antillean Land mammal Fauna, 1: New tertiary Fossils from Cuba and Puerto Rico. Amer. Mus. Novitates, 3141: 1-31, New York.

MacPhee, R., Iturralde-Vinent, M. and Gaffney, E., in prep. Domo de Zaza, a Lower Miocene land-mammal locality in south-central Cuba: Summary of Geological and Paleontological investigations. Am. Mus. Novitates.

Mann, P., Schubert, C. and Burke, K., 1990. Review of Caribbean neotectonics, in Dengo, G. and Case J.E., eds. The Caribbean region: The Geology of North America, Volume H: Boulder, Colorado, Geological Society of America, p. 307-338.

Mederos, P., 1987. Condiciones paleogeográficas de formación de fosforitas en la región Güines-Pipián. Serie Geológica del CIDP (2): 68-83.

Mederos, P., Franco, G., 1992. Influencia de las paleocorrientes en las acumulaciones de fosfato del Mioceno Inferior-Medio de Cuba: Rev. Tecnológica XXII (2): 13-20, Havana.

Mullins, H. T. and Newmann, A. C., 1979. Geology of the Miami Terrace and its paleooceanographic implications. Marine Geology, 30: 205-232.

Mullins, H.T., Gardulski, A.F., S.W. Wise and J. Applegate, 1987. Middle Miocene oceanographic event in the eastern Gulf of México: Implications for seismic stratigraphic succession and Loop Current/Gulf Stream circulation: GSA Bull. 98: 702-713.

Purdy, R. 1987. Fossil shark teeth (Poster). National Museum of Natural History, Smithsonian Institution.

Pszczolkowski, A. 1978. Geosynclinal sequences of the Cordillera de Guaniguanico in western Cuba; their lithostratigraphy, facies development and paleogeography: Acta Geol. Polonica 28 (1): 1-96. (Revised Spanish version 1987 in Contribución a la geología de la provincia de Pinar del Río, Acad. Ciencias de Cuba, Edit. Científico-Técnica, La Habana, 253 p.).

Riggs, S., 1984. Paleooceanographic model of Neogene phosphorite deposition, U.S. Atlantic Continental Margin: Science: 223-231.

Sánchez Roig, M., 1920. Escuálidos del Mioceno y Plioceno de La Habana. Boletín de Minas, Dirección de Montes y Minas 6: 1-16.

Sánchez Roig, M., 1928. Memoria del Instituto Nacional de Investigaciones Científicas y Museo de Historia Natural, Imprenta Montalvo y Cárdenas, La Habana, 220 p.

Scott, T.M. 1988. The lithostratigraphy of the Hawthorn Group (Miocene) of Florida. Florida Geol. Survey Bull. 59: 1-148.

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Table 1. List of Cuban shark taxa identified by the authors, and a guide to the plates.

TAXON

PALEO-

CENE

(?)

MIDDLE

-LATE

EOCENE

LOWER-

MIDDLE

MIOCENE

LATE

MIOCENE-

PLIOCENE

Plate I:

figure

Carcharhinus obscurus X 1

Carcharhinus perezi X 2

Carcharias taurus X 3

*Carcharodon auriculatus X 4

Carcharodon landenensis X -

Carcharodon carcharias X -

Carcharodon megalodon X X -

Carcharodon subauriculatus X 5

Galeocerdo contortus X 6

Hemipristis serra X 7, 8

Isurus sp. X -

Isurus hastalis X -

Isurus sp. cf. I. desori X -

Negaprion brevirostris X 9

Sphyrna mokarran X 10

Sphyrna prisca (?) X (**)

Striatolamia macrota X 11

Striatolamia sp. X -

Myliobatidae X

Aetobatus poeyi X(?) -

(*) Some authors as Cappetta (1987) place the species megalodon, auriculatus and subauriculatus in the genus Carcharocles, but Kruchow and Thies (1990) place the species megalodon and auriculatus in the genus Procarcharodon. (**) This species was identified from an illustration by Sánchez Roig (1920), but they must be verified with actual material in the future.

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Table 2. Cuban shark taxa identified by Sánchez Roig (1920), and by Leriche (1938), based on the Sanchez Roig's illustrations and descriptions. Leriche assumed that all specimens were Miocene or Pliocene in age. Present authors' identifications based on the 1920 illustrations and descriptions, plus examination of some additional specimens collected by Sánchez Roig at the same localities. The age of some specimens was found to be Latest Eocene (see text for details).

Shark taxon

sensu Sánchez Roig

Figures in Sánchez

Roig (1920)

Shark taxon

sensu Leriche

Shark taxon

sensu present authors

Lamna cuspidata 1, 2 Hemipristis serra Hemipristis serra

Lamna elegans 3, 4 Odontapsis acutissima Carcharias taurus

Lamna contortidens 5 Odontapsis acutissima Carcharias taurus

Lamna longidens -

Isurus hastalis 6, 7 Oxyrhina hastalis Isurus hastalis

Isurus desori * 8, 9 Oxyrhina desori Stratolamia macrota *

Isurus minutus 10 Odontapsis (?) sp. Isurus sp. (juvenil?)

Carcharodon auriculatus * 11, 12, 13 Carcharodon sp. Carcharodon auriculatus *

Carcharodon megalodon 14-17, 21 Carcharodon megalodon Carcharodon megalodon

Carcharodon lanciformis 18, 19 Carcharodon carcharias Carcharodon carcharias

Galeocerdo contortus 20 Galeocerdo aduncus Galeocerdo contortus

Sphyrna prisca 22 Sphirna sp. Sphyrna prisca (?)

(*) These specimens are embedded in Latest Eocene marls

18

Table 3. Catalog of Cuban fossil sharks in the paleontological collections of Museo Nacional de Historia Natural in Havana and the Institute of Geology and Paleontology. ## Repository unknow. JMFM Private collection of J. M. Fernández Milera. ID = Identification by (1) present authors, (2) Sánchez Roig (1920) revised by the authors, and (3) by J. W. Gidley in Burchard (1920)

# AGE TAXON LOCALITY ID OBSERVATIONS

JMFM Late Miocene- Pliocene

Carcharodon megalodon Matanzas Prov., Municipio Cárdenas, Loma de Fines, Cueva Armada, 2 km from Cárdenas

1 Collected by L. M. Fernández Milera.


Carcharodon megalodon Matanzas Prov., Municipio Cárdenas, Loma de Fines, 3 1/2 km from Cárdenas a Coliseo



Carcharodon megalodon Matanzas Prov., Municipio Cárdenas, Loma de Fines, Limestone quarry Larrauri, 2 1/2 km from Cárdenas to Coliseo.



Carcharodon megalodon Matanzas Prov., Municipio Cárdenas, Loma de Fines, Limestone quarry "Garriga," 3 km de Cárdenas


MNHNH -P2020

Late Neogene Carcharodon megalodon Matanzas Prov., Municipio Carlos Rojas, Limestone quarry

1 Additional materialdeposited at the Museo Municipal de Cárdenas, Matanzas.

MNHNH -P2074

Late Neogene Carcharodon megalodon Matanzas Prov., Municipio Cárdenas 1

MNHNH -P2021

Lower-Middle Miocene

Carcharodon megalodon (?) fragmentary tooth

Havana Prov., Municipio Madruga 1

MNHNH -P2024

Miocene Carcharodon megalodon La Posa (No further information available)

1 Collected by Miguel L. Jaume.

MNHNH -P2082


Carcharodon megalodon (?) fragmentary tooth

Havana Prov., Tunel Calle 88a and 61, Marianao

1 Collected by Ricardo Lozano.

MNHNH -P2131--

2138


Carcharodon megalodon City of Havana, Excavación en Calle 222, near Cimex. Escuela Politécnica Química de Alimentación "Ejército Rebelde"

1

MNHNH -P2224


Carcharodon megalodon Havana Prov., Municipio San José de las Lajas, Limestone quarry Camoa y Somorostro

1 Collected by Stephen Díaz.

19

MNHNH -P2226


Carcharodon megalodon Havana Prov. , Municipio Tapaste, Limestone quarry near the town of Aguacate

1 Collected by J. González Perez

## Lower-MiddleMiocene

Carcharodon megalodon Havana Prov., Municipio Plaza, Canteras Sixto Abreus, Costado south of Cementerio de Colón, Vedado-Habana

2 Collected by M. Sánchez Roig.

## MiddleMiocene

Carcharodon megalodon City of Havana, Limestone quarry "Cojímar," Reparto Cojímar

From a fossil catalog at IGP.

## LowerMiocene

Carcharodon megalodon City of Havana, Limestone quarry "Husillo"

Reported and identified by Giraldo Alayón (MNHNH).


Carcharodon megalodon Havana Prov., Municipio San Antonio de los Baños

2


Carcharodon megalodon Havana Prov., Municipio Arroyo Naranjo, town of Arroyo Naranjo

2


Carcharodon megalodon Havana Prov., Municipio Arroyo, locality of San Jose de las Lajas, (Finca Cervantes)

2 Collected by M. Sánchez Roig.

IGP Lower-MiddleMiocene

Carcharodon megalodon Havana Prov., Municipio Artemisa, quarry near Pijirigua

1

IGP Lower-MiddleMiocene

Carcharodon megalodon Havana Prov., Municipio Bejucal, quarry near Cacahual

1

IGP Miocene Carcharodon megalodon Matanzas Prov., Municipio Carlos Rojas, quarry near Carlos Rojas town

1

## Miocene Carcharodon megalodon City of Havana, Municipio Cerro, Ferrocarril de Marianao, Estación La Ceiba, Puentes Grandes

1 Collected by M. Sánchez Roig and reported in his 1920 paper.

MNHNH -P2142,

2150

Lower Miocene

Carcharodon subauriculatus Havana Prov., Municipio Guines, Phosphorite mine, 6 km NW of Guines

1 Collected by J. A. Genaro, R. Rojas, M. Iturralde-Vinent, Stephen Díaz, et al.

MNHNH -P2027


Carcharhinus perezi Havana Prov., Municipio Caimito, south of Ceiba del Agua. Secondary deposit in Cueva Paredones

1 Collected by M. Iturralde-Vinent .

MNHNH -P2219,

3033

Lower Miocene

Carcharhinus obscurus S. Spíritus Prov., Municipio La Sierpe, Canal Presa Zaza, Domo de Zaza

1 Collected by R. MacPhee, M. Smith, R. Rojas, M. Iturralde-Vinent.

20

MNHNH -P2165-

2166

Lower Miocene

Carcharias taurus Havana Prov., Municipio Guines, Phosphorite mine, 6 km NW of Guines


MNHNH -P2139


Hemipristis serra Matanzas Prov., Municipio Bolondrón, secondary deposit in Cueva El Pichi

1 Collected by Stephen Díaz.

MNHNH -P2227


Hemipristis serra Havana Prov. Limestone quarry near the town of Aguacate

1 Collected by J. González Perez

MNHNH -P2221, 3034, 3036

Lower Miocene

Hemipristis serra S. Spíritus Prov., Municipio La Sierpe, Canal Presa Zaza, Domo de Zaza


MNHNH -P2214,

2215

Miocene (?) Hemipristis serra City of Havana, Municipio Boyeros, Alturas del Cacahual. Secondary deposit in Cueva de la Diaclasa

1 Collected by Osvaldo Jiménez.

MNHNH -P3037

Lower Miocene

Negaprion brevirostris S. Spíritus Prov., Municipio La Sierpe, Canal Presa Zaza, Domo de Zaza


MNHNH -P3032

Lower Miocene

Sphyrna mokarran S. Spíritus Prov., Municipio La Sierpe, Canal Presa Zaza, Domo de Zaza


## Miocene (?) Shark teeth City of Havana, Municipio Vento (Finca Francisco de Barcos)

Collected by M. Sánchez Roig.. Identified as Lamna (?) sp.

MNHNH -P2169 -2170, 2175


Isurus hastalis Havana Prov., Municipio Guines, Phosphorite mine, 6 km NW of Guines


MNHNH -P2185


Isurus sp. cf. I. desori fragmentary tooth

Havana Prov., Municipio Guines, Phosphorite mine, 6 km NW of Guines


IGP-P- 2348- 2354


Isurus hastalis 1

MNHNH -P2216

Miocene (?) Galeocerdo contortus City of Havana, Municipio Boyeros, Alturas del Cacahual. Secondary deposit in Cueva de la Diaclasa


21

IGP Miocene Galeocerdo contortus City of Havana, Municipio Cerro, quarry "Tejar Consuelo" in Ciénaga

1 Collected and identified by M. Sánchez Roig, who reported these teeth as Middle Eocene.

IGP (68a)

Miocene (?) Galeocerdo (?) sp. fragmentary tooth

City of Havana, Municipio Cerro, quarry "Tejar Consuelo" in Ciénaga

1 Collected by M. Sánchez Roig, who identified this tooth as Middle Eocene Hemipristis serra.

## Miocene Carcharias taurus Carcharodon megalodon Carcharodon carcharias Galeocerdo contortus Isurus hastalis Hemipristis serra Carcharodon sp. Sphyrna prisca (?)


2 Collected by M. Sánchez Roig and reported in his 1920 paper.

MNHNH-P

Lower Miocene

Batoid tail spine S. Spíritus Prov., Municipio La Sierpe, Canal Presa Zaza, Domo de Zaza


MNHNH-P

Lower Miocene

Batoid tail spine Havana Prov., Municipio Guines, Phosphorite mine, 6 km NW of Guines


MNHNH-P

Middle Miocene

Myliobatidae Lower dental plate

Havana Prov., Municipio Caimito, south of Ceiba del Agua, inside Cueva del Bicho

1 Collected by Grupo Espeleológico Cayoguasal

MNHNH-P2218

Miocene (?) Unidentified fragmentary teeth City of Havana, Municipio Boyeros, Alturas del Cacahual. Secondary deposit in Cueva de la Diaclasa


IGP MiddleMiocene

Myliobatidae Fragmentary lower dental plate

City of Havana, Municipio de Cojímar, Cojimar quarry. Type locality of Cojímar Formation

1 Collected by J.T. Sierra. Identified by M. Sánchez Roig as Aëtobatis poeyi

MNHNH unknown Unidentified fragmentary teeth Havana Prov., Municipio Bejucal, Lecho del Arroyo Govea (Santa Rosa)


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IGP Eocene (?) Shark teeth Camagüey Prov., Central (sugar mill) Florida (today = Argentina)

From a fossil catalog in IGP. This specimen is reported as Late Oligocene, but geological maps indicate Eocene outcrops in the area. The species were identified as Isurus hastalis and Carcharodon sp.

## Latest Eocene Carcharodon auriculatus Striatolamia macrota


2 Collected by M. Sánchez Roig and reported in his 1920 paper

IGP (L 508)

Latest Eocene Striatolamia macrota City of Havana, Municipio Cerro, quarry "Tejar Consuelo" in Ciénaga

1 Collected by M. Sánchez Roig, who identified the (3) teeth as I. hastalis. A. de la Torre change the identification to I. desiori. Specimen embedded in yellow soft marls of Latest Eocene age

IGP Middle-LateEocene

Striatolamia macrota Havana Prov., near the small town of Nazareno

1 Identified by A. de la Torre as Isurus

MNHNH -P2025, 2026, 2043, 2047-2049, 2052-2053. 2075-2076

Middle Eocene

Carcharodon auriculatus Granma Prov., Charco Redondo Mines District

1 Additional material deposited at Museo Provincial "Ignacio Agramontes" Camagüey Province. Collected by Juan Melo. Collected inside the mine.

MNHNH -P2054-2063, 2070-2072, 2078-2081

Middle Eocene

Striatolamia macrota Granma Prov., Charco Redondo Mines District

1 Additional materialdeposited at Museo Provincial "Ignacio Agramontes" Camagüey Province. Collected inside the mine by Juan Melo

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## MiddleEocene

Carcharodon auriculatus (?)

Granma Prov., Charco Redondo Mines District, Charco Redondo Mine located 12.8 km SE of Santa Riuta railroad station.

3 Collected by E. F. Burchard. He also reported Isurus hastalis (or very near to this one) and Lamna cf. L. elegans

IGP- 3299

Middle Eocene

Striatolamia cf. S. macrota (fragmentary tooth )

Granma Prov., Manganese Mine, Charco Redondo Mines District

1 Collected by Pallarés. Identified by A. de la Torre as I. desori

IGP MiddleEocene

Striatolamia macrota Granma Prov., Charco Redondo Mines district, Taratana Mine, Santa Rita, Bayamo.

1 Identified by A. de la Torre as Lamna cf. L. elegans

IGP (L-509)

Middle Eocene

Isurus sp. Granma Prov., Manganese Mine, Charco Redondo Mines District

1 Collected by Ernesto Segeth. Identified by A. de la Torre as I. hastalis

## Eocene (?) Aetobatus poeyi (isolated tooth)

Cienfuegos Prov., lands of Ingenio Constancia (today =Moncada) near the city of Cienfuegos.

1 Collected and identified by Fernández de Castro (1873). The age must ve verified in the future

MNHNH-P2228

Paleocene Striatolamia sp. (fragmentary tooth )

Pinar del Río Prov., Cordillera de Guaniguanico, north of San Cayetano (teeth with manganese stain).

1 Donated by Rafael Quiñones

MNHNH-P2229

Paleocene Carcharodon landenensis. (fragmentary tooth )

Pinar del Río Prov., Cordillera de Guaniguanico, north of San Cayetano (teeth with manganese stain).

1 Donated by Rafael Quiñones

JMFM Paleocene Carcharodon landenensis. (fragmentary teeth )

Pinar del Río Prov., Cordillera de Guaniguanico, Alturas de Pizarras (teeth with manganese stain).

1 Collected by Federico Gómez de la Masa. Collection of J. M. Fernández Milera.

MNHNH -P2028, 2029, 2032-2033, 2035

Paleocene Striatolamia sp. (fragmentary teeth )

Pinar del Río Prov., Finca Montalvo, Sierra de San Andrés (Pico Chico). 1 1/2 Legua from "La Palma" (Manganese mine).


MNHNH -P2031, 2034, 2036, 2038-2041

Paleocene Unidentified fragmentary shark teeth

Pinar del Río Prov., Finca Montalvo, Sierra de San Andrés (Pico chico). 1 1/2 Legua from "La Palma" (Manganese mine).


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catalog of cuban fossil elasmobranchii (paleocene … · catalogue of cuban fossil elasmobranchii...

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