During the early 1930s, the region of Madruga in the Province of Mayabeque, western Cuba, became the center of attention to several North American petroleum geologists. Their initial surveys revealed the presence of Cretaceous and Cenozoic outcrops within a complex geological setting (DeGolyer, 1918DeGolyer, E. (1918). The geology of Cuban petroleum deposits. American Association of Petroleum Geologists Bulletin, 2, 133-167.; Wright and Sweet, 1924Wright, A. and Sweet, P.W.K. (1924). The Jurassic as a source of oil in western Cuba. Bulletin of the American Association of Petroleum Geologists, 8, 516-519.; Chawner, 1932Chawner, W.D. (1932). Geology and petroleum possibilities of northern Matanzas and Santa Clara Provinces. Oficina de Minerales, Inv. 2520, unpublished report.; Lewis, 1932Lewis, J.W. (1932). Geology of Cuba (with discussion by R. J. Metcalf). American Association of Petroleum Geologists Bulletin, 16, 533-555.; Palmer, 1932Palmer, R.H. (1932). Informe geológico del área de Madruga. Oficina Nacional de Recursos Minerales. Ministerio de Energía y Minas, La Habana.; Palmer, 1938Palmer, R.H. (1938). Field guide to geological excursion in Cuba. Secretaría de Agricultura.). In 1932, Chawner reported and mapped several fossil deposits within this geological complex, attracting the attention of Palmer, who revisited and resampled most of the outcrops available at the time (Palmer, 1948Palmer, R.H., 1948. List of Palmer Cuban Localities. Bulletins of American Paleontology, 31, 1-178.). Palmer’s compilation included over 3217 localities of which at least 110 are today within the region of Madruga, several of which are sites of geological and paleontological interest.
One of the most important Palmer’s fossiliferous locality from Madruga is “Cantera Jene 2 km. S. Aguacate” (Loc. 766 in Palmer, 1948: 321Palmer, R.H., 1948. List of Palmer Cuban Localities. Bulletins of American Paleontology, 31, 1-178.), but this reference is ambiguous because there are various quarries about 2 km south of Aguacate. One of them is approximately 100 m along the west side of the highway, another is about 550 m to the east, following an old road. Palmer’s locality likely denoted the first quarry, from which recently important fossil discoveries (e.g. shark teeth and turtles) have been made; most of which are as-yet-unpublished. Iturralde-Vinent and colleagues referred to this locality as “Havana Prov. Limestone quarry near the town of Aguacate” and briefly noted the occurrence of fossil sharks (Iturralde-Vinent et al., 1996Iturralde-Vinent, M., Hubbell, G. and Rojas, R. (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, 31, 7-21.). However, even though both quarries are richly fossiliferous, they are still poorly sampled and their fossil fauna remains unstudied.
Our short exploration of the Aguacate quarry revealed a rich mollusk fauna including chemosymbiotic bivalves of the family Lucinidae, as well as a few solitary corals. Because studies of Cuban Miocene bivalve faunas are rare (Cooke, 1919Cooke, C.W. (1919). Contributions to the geology and paleontology of the West Indies IV. Tertiary mollusks from the leeward islands and Cuba. Carnegie Institution of Washington Publications, 291, 103-156.; Franco, 1983Franco, G.L. (1983). Nuevos subespecies de Bivalvos del Neógeno de Cuba oriental, Contribucion a la geologia de Cuba Oriental. Editorial Científico-Técnica.; Kiel and Hansen, 2015Kiel, S. amd Hansen, B.T. 2015. Cenozoic methane-seep faunas of the Caribbean region. PLoS ONE, 10, e0140788.), we use this opportunity to provide an overview of the fossil fauna collected at the Aguacate quarry and put the fauna into a regional and paleoecological context.
The material described here was collected during two visits to the Aguacate quarry (coordinates: 22°58'N, 81°49'W) (Fig. 1) in 2018 and 2020. The material is deposited and available at the local Madruga Museum with the catalog numbers MPAL-1050 to 1093, and at the Swedish Museum of Natural History in Stockholm (NRM).
Traditionally, the rocks of the Aguacate quarry have been informally referred to the Güines Formation (early-middle Miocene), but geological maps and lithology indicate that they belong to the Cojímar Formation (upper Lower Miocene-lower Middle Miocene). The Cojímar Formation (named by Palmer (1934)Palmer, R.H. (1934). The geology of Habana, Cuba, and vicinity. The Journal of Geology, 42, 123-145.) lies concordantly over the Jaruco or Husillo formations on basin areas, and transitions in facies both laterally and vertically to the Güines Formation. In Madruga, the Cojímar Formation occurs roughly to the north of Sierra del Grillo and mainly includes calcareous and argillaceous marls and gray chalky limestones. In the Aguacate area, the formation crops out as limestones, similar to those of the Güines Formation, but with marls and intercalated limestones typical of Cojímar lithology and fauna (Albear and Iturralde-Vinent, 1985Albear, J.F. and Iturralde-Vinent, M., (1985). Estratigrafía de las provincias de La Habana. In: Iturralde-Vinent, M. (Ed.), Contribución a la geología de las provincias de La Habana y Ciudad de La Habana. Editorial Científico-Técnica.). The thickness of the Cojímar Formation is usually less than 390 m as estimated from drill cores (Albear and Iturralde-Vinent, 1985Albear, J.F. and Iturralde-Vinent, M., (1985). Estratigrafía de las provincias de La Habana. In: Iturralde-Vinent, M. (Ed.), Contribución a la geología de las provincias de La Habana y Ciudad de La Habana. Editorial Científico-Técnica.; Pardo, 2009Pardo, G. (2009). Post-middle Eocene stratigraphy, in G. Pardo, The geology of Cuba. AAPG Studies in Geology Series, 58, 277-293.).
The Aguacate quarry exposes thickly bedded limestone with a fine to medium texture (calcarenite) intercalated with marls (Fig. 2). The exposed surface is altered and covered with a light gray patina. The marls are usually more fossiliferous than the limestone. Along the northern portion of the quarry, there are Miocene outcrops with fissures filled with rendzina-like red paleosol.
The fossil fauna of the Cojímar Formation is diverse but has not been evaluated in detail (Albear and Iturralde-Vinent, 1985Albear, J.F. and Iturralde-Vinent, M., (1985). Estratigrafía de las provincias de La Habana. In: Iturralde-Vinent, M. (Ed.), Contribución a la geología de las provincias de La Habana y Ciudad de La Habana. Editorial Científico-Técnica.). So far, some of the benthic microfossils have been described by Palmer (1940-1941)Palmer, D.K. (1940-1941). Foraminifera of the upper Oligocene Cojimar formation of Cuba. Parts 1-5. Memorias de la Sociedad Cubana de Historia Natural,14, 15. and the planktonic by Brönnimann and Rigassi (1963)Brönnimann, P. and Rigassi, D. (1963). Contribution to the geology and paleontology of the area of the city of La Habana, Cuba, and its surroundings. Eclogae Geologicae Helvetiae, 56, 193-480., and Iturralde-Vinent (1969)Iturralde-Vinent, M. (1969). Principal characteristics of Cuban Neogene stratigraphy. American Association of Petroleum Geologists Bulletin, 53, 1938-1955.. Typically, it includes the following: Orbulina universa, O. suturalis, Trilobatus immaturus, T. sacculifer, T. trilobus, T. quadrilobatus, Globigerinoides ruber, Dentoglobigerina altispira, Globoquadrina dehiscens, Sphaeroidinella dehiscens, Globorotalia praemenardii, G. archeomenardii, Fohsella praefohsi, F. peripheroacuta, F. peripheroronda, Globigerinella obesa, G. siphonifera (=G. aequilateralis), and ‘Nummulites’ cojimarensis (Albear and Iturralde-Vinent, 1985Albear, J.F. and Iturralde-Vinent, M., (1985). Estratigrafía de las provincias de La Habana. In: Iturralde-Vinent, M. (Ed.), Contribución a la geología de las provincias de La Habana y Ciudad de La Habana. Editorial Científico-Técnica.; Pardo, 2009Pardo, G. (2009). Post-middle Eocene stratigraphy, in G. Pardo, The geology of Cuba. AAPG Studies in Geology Series, 58, 277-293.). Recently, thin sections from samples of the quarry confirm the occurrence of Orbulina universa (Gútierrez-Domech, R., pers. comm., 2021). This thanatocoenosis indicates a Middle Miocene age around the Langhian-Serravallian transition [M7-M8, Fohsella peripheroacuta and F. praefohsi zones of Wade et al. (2011)Wade, B.S., Pearson, P.N., Berggren, W.A. and Pälike, H. (2011). Review and revision of Cenozoic tropical planktonic foraminiferal biostratigraphy and calibration to the geomagnetic polarity and astronomical time scale. Earth-Science Reviews, 104, 111-142.] for these deposits. These rocks were deposited in bathyal to neritic conditions in water less than 300 m (1000 ft) deep (Albear and Iturralde-Vinent, 1985Albear, J.F. and Iturralde-Vinent, M., (1985). Estratigrafía de las provincias de La Habana. In: Iturralde-Vinent, M. (Ed.), Contribución a la geología de las provincias de La Habana y Ciudad de La Habana. Editorial Científico-Técnica.; Pardo, 2009Pardo, G. (2009). Post-middle Eocene stratigraphy, in G. Pardo, The geology of Cuba. AAPG Studies in Geology Series, 58, 277-293.).
The fauna of the Aguacate quarry is preserved largely as internal or external molds, which makes it difficult to assign precise taxonomic identifications. We found several specimens of solitary corals, likely belonging to a single species (“Caryophyllia”, Fig. 3A, B). Mollusks are common and diverse, with at least eight bivalve and six gastropod families being present. These include:
Nuculanidae. A single specimen showing the taxodont hinge (Fig. 3C).
Pectinidae. One specimen with sharp radial ribs and wide interspaces with fine commarginal lirae (Fig. 3D), resembling certain species of the genus Chlamys, and the species Amusium mimyum, described as having a ribbed umbonal area (Woodring, 1982: p. 587-588; pl. 122, fig. 2Woodring, W.P. (1982). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-F, 541-759.).
Glycymerididae. A few internal molds showing hinge teeth and a strongly ribbed inner ventral margin (Fig. 3E); similar species include Glycymeris tumulus from the Miocene of Virginia, USA (Gardner, 1943: pl. 1, fig. 6Gardner, J. (1943). Mollusca from the Miocene and Lower Pliocene of Virginia and North Carolina Part 1. Pelecypoda. U.S. Geological Survey Professional Paper, 199-A, 1-144.), Glycymeris duplinensis Dall (Gardner 1943: pl. 2, fig. 3Gardner, J. (1943). Mollusca from the Miocene and Lower Pliocene of Virginia and North Carolina Part 1. Pelecypoda. U.S. Geological Survey Professional Paper, 199-A, 1-144.) from Miocene and Pliocene of North Carolina, USA, and the Miocene to Recent Glycymeris pectinata (Gmelin) known from Central America and the Caribbean Sea, and extending as far north as North Carolina (Gardner, 1926: p. 38Gardner, J. (1926). The molluscan fauna of the Alum Bluff Group of Florida. U.S. Geological Survey Professional Paper, 142-A, 1-64.).
Thyasiridae. Several specimens of a subrectangular Thyasira reaching 12 mm in length, with sharp posterior ridge and groove, and a long, thin anterior adductor scar (Fig. 3F).
Lucinidae. There are two species: a single specimen probably belonging to Lepilucina aff. gratis (Olsson, 1964Olsson, A.A. (1964). Neogene mollusks from northwestern Ecuador. Paleontological Research Institution.) is preserved as internal and external mold (Fig. 3G, H). This species has previously been reported from the middle Miocene of Ecuador and Panama (Woodring, 1982Woodring, W.P. (1982). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-F, 541-759.). The other species (Figure 4) is the most common taxon at the Aquacate quarry; it is a very flat, medium-sized Anodontia (sensu lato) with an undulating pallial line and long, thin anterior adductor muscle scar, a narrow and edentulous hinge plate, and no lateral teeth. No external features are preserved. This species is very similar to the extant Caribbean type species Anodontia alba and the extant A. edentuloides (Verrill) from the Gulf of California (Taylor and Glover, 2005Taylor, J.D. and Glover, E.A. (2005). Cryptic diversity of chemosymbiotic bivalves: a systematic revision of worldwide Anodontia (Mollusca: Bivalvia: Lucinidae). Systematics and Biodiversity, 3, 281-338.) except for its rather weak inflation, which is unusual for Anodontia. This species may represent a new taxon for Cuba, but we refrain from describing it until better specimen are available. Coeval species from Florida include ‘A.’ corpulenta, ‘A.’ santarosana, and ‘A.’ janus (cf. Bretsky, 1976Bretsky, S.S. (1976). Evolution and classification of the Lucinidae (Mollusca; Bivalvia). Palaeontographica Americana, 8, 219-337.), all of which are more inflated than the specimens from the Aguacate quarry. The latter species was also reported (as Pegophysema janus) from the Gatun Formation in Panama (Woodring, 1982, pl. 111, figs. 13, 15-17Woodring, W.P. (1982). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-F, 541-759.).
Veneridae.-Several specimens of general veneroid shape, reaching 30 mm in length and present as internal molds only, representing more than one species (Figs. 3I, J).
Tellinidae.-A small, elongate specimen of about 10 mm length (Fig. 3K); similar species are described as Tellina (Moerella) piesa and T. (M.) ctenota from the early and middle Miocene of the Alum Bluff Group in Florida (Gardner, 1928Gardner, J. (1928). The molluscan fauna of the Alum Bluff Group of Florida. Part V. Tellinacea, Solenacea, Mactracea, Myacea, Molluscoidea. U.S. Geological Survey Professional Paper, 142-E(I-IV), 185-240.), and as Tellina (Eurytellina) nitens Adams, from the Gatun Formation in Panama (Woodring 1982: pl. 117:16, 17Woodring, W.P. (1982). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-F, 541-759.).
Turritellidae.-One large external mold of about 15 mm diameter, showing strong spiral ribs and angular whorl profile (Fig. 3L).
Cerithiidae.-A few small external molds showing strong axial ribs crossed by six spiral cords (Fig. 3M).
Naticidae.-A few internal and external molds of moderately high-spired specimens having fine opisthocline ribs on the upper half of the whorls (Figs. 3N, O); species with a similar sculpture on the spira include Natica (Naticarius) stenopa (Woodring, 1957: pl. 20, figs. 4-6Woodring, W.P. (1957). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-A, 1-145.), from the middle Miocene Gatun Formation in Panama and several species assigned to Natica (Naticarius) and Stigmaulax from the Neogene of northern Ecuador (Olsson, 1964Olsson, A.A. (1964). Neogene mollusks from northwestern Ecuador. Paleontological Research Institution.).
Turridae.-A single internal mold showing a strongly constricted internal whorl (Fig. 3P) might belong to the Turridae.
Olividae.-A single external mold (Fig. 3Q).
Conidae.-A large internal mold having a moderately high spire (Fig. 3R).
Architectonicidae.-A large, fragmentary external mold (Fig. 3S) not unlike Architectonica nobilis Röding as illustrated from the middle Miocene Gatun Formation in Panama (Woodring, 1959: pl. 29: figs. 1-6, 10-12, 14-16Woodring, W.P. (1959). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-B, 147-239.).
The fauna of the Aguacate quarry consists of typical tropical, shallow marine corals and mollusks. The most common taxon is a very flat, medium-sized Anodontia (s.l.), which may likely represent a new taxon (Fig. 4). A lucinid with a similar outline, inflation, and edentulous hinge was reported as Myrteopsis? sp. from the early Miocene Cantera Portugalete in Cuba (Kiel and Hansen, 2015Kiel, S. amd Hansen, B.T. 2015. Cenozoic methane-seep faunas of the Caribbean region. PLoS ONE, 10, e0140788.); but it has a much shorter anterior adductor scar than the Anodontia (s.l.) specimens from the Aguacate quarry and shows weak but distinctive commarginal sculpture. The abundance of the lucinid bivalve Anodontia (s.l.) suggests that these deposits may be interpreted as an ancient seagrass meadow. Anodontia species are often associated with seagrass beds (Taylor and Glover, 2005Taylor, J.D. and Glover, E.A. (2005). Cryptic diversity of chemosymbiotic bivalves: a systematic revision of worldwide Anodontia (Mollusca: Bivalvia: Lucinidae). Systematics and Biodiversity, 3, 281-338.). For example, the extant A. alba, which resembles the Aguacate specimens regarding pallial line and adductor scar characters, was reported in very large numbers from a seagrass meadow in Biscayne Bay, Florida (Moore et al., 1968Moore, H.B., Davies, L.T., Fraser, T.H., Gore, R.H. and Lopez, N.R. (1968). Some biomass figures from a tidal flat in Biscayne Bay, Florida. Bulletin of Marine Science, 18, 261-279.) and other places in the Caribbean regions (Jackson, 1973Jackson, J.B.C. (1973). The ecology of molluscs of Thalassia communities, Jamaica, West Indies. 1. Distribution, environmental physiology, and ecology of common shallow-water species. Bulletin of Marine Science, 23, 313-350.). However, the co-occurring and not uncommon medium-sized thyasirid bivalve is rather unusual for seagrass meadows, as they tend to prefer deeper and more dysoxic environments (Dando et al., 2004Dando, P.R., Southward, A.J. and Southward, E.C. (2004). Rates of sediment sulphide oxidation by the bivalve mollusc Thyasira sarsi. Marine Ecology Progress Series, 280, 181-187.; Oliver, 2015Oliver, P.G. (2015). Deep-water Thyasiridae (Mollusca: Bivalvia) from the Oman Margin, Arabian Sea, new species and examples of endemism and cosmopolitanism. Zootaxa, 3995, 252-263.). Biogeographically, the fauna of the Aguacate quarry shows links to other areas within the Caribbean region, to Florida and the tropical/subtropical Atlantic coast of North America, and the Miocene of northern Ecuador (Gardner, 1943Gardner, J. (1943). Mollusca from the Miocene and Lower Pliocene of Virginia and North Carolina Part 1. Pelecypoda. U.S. Geological Survey Professional Paper, 199-A, 1-144.; Olsson, 1964Olsson, A.A. (1964). Neogene mollusks from northwestern Ecuador. Paleontological Research Institution.; Woodring, 1957Woodring, W.P. (1957). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-A, 1-145., 1959Woodring, W.P. (1959). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-B, 147-239., 1982Woodring, W.P. (1982). Geology and paleontology of Canal Zone and adjoining parts of Panama. U.S. Geological Survey Professional Paper, 306-F, 541-759.; Moore et al., 1968Moore, H.B., Davies, L.T., Fraser, T.H., Gore, R.H. and Lopez, N.R. (1968). Some biomass figures from a tidal flat in Biscayne Bay, Florida. Bulletin of Marine Science, 18, 261-279.; Jones and Portell, 1988).