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732
J. Paleont., 75(3), 2001, pp. 732–742
Copyright q2001, The Paleontological Society
0022-3360/01/0075-732$03.00
AN EXCEPTIONAL COASTAL UPWELLING FISH ASSEMBLAGE IN THE
CARIBBEAN NEOGENE*
ORANGEL AGUILERA
AND
DIONE RODRIGUES DE AGUILERA
Universidad Nacional Experimental Francisco de Miranda,
Centro de Investigaciones en Ciencias Ba´sicas,
Complejo Docente Los Perozos, Carretera variante sur, Coro, Estado Falco´n. Venezuela,
,aguilero@unefm.edu.ve.
A
BSTRACT
—We report the discovery of an extremely rich, previously undescribed Caribbeanlate Miocene to early Plioceneichthyofauna
represented by one hundred forty species of elasmobranchs and teleosteans from the Cubagua Formation, northeastern Venezuela. The
fauna exhibits significant ecological differences compared with common neritic Caribbean Neogene assemblages. The bathymetric
distributions of taxa, based on living counterparts, ranges from 0 to 100 m depth. The exceptional co-occurrence of deep water
(epipelagic, mesopelagic and benthopelagic), and shallow water (neritic) taxa is best interpreted as the consequence of ocean upwelling
in the proximity to the deep-water Cariaco Trench. Patterns of predator and prey are established and corroborate upwelling. Special
remarks are made regarding previously unknown late Miocene to early Pliocene Caribbean ichthyofaunas, the absence or rarity of
reported fossil taxa in the Recent Caribbean fauna, and a paleo- upwelling indicator (Lampadena jacksoni new species).
INTRODUCTION
I
N TROPICAL
Middle America, the closure of the Isthmus of Pan-
ama (Coates, 1999; Coates et al., 1992; Coates and Obando,
1996) resulted in major geographic and environmental changes.
Principal freshwater drainage systems into the south Caribbean
changed course, including the Magdalena River (Kay et al., 1997),
the Orinoco River (Diaz de Gamero, 1996), and the Amazon-
Orinoco systems (Lundberg et al., 1998). This resulted in pro-
gressive oceanographic and environmental changes in the Carib-
bean during the Tertiary. The paleoecology, diversity, and com-
position of Caribbean Neogene fish faunal assemblages reveal the
prevalence of neritic environments in many sedimentary basins,
including Panama (Gillette, 1984), Panama and Costa Rica
(Aguilera and Aguilera, 1999; Collins et al., 1999), Trinidad
(Nolf, 1976), northwestern Venezuela (Nolf and Aguilera, 1998),
the Dominican Republic (Nolf and Stringer, 1992), Jamaica
(Stringer, 1998), and Cuba (Iturralde-Vinent et al., 1996).
The ichthyofaunas of the Cubagua Formation, treated herein,
are ecologically very different from those previously documented
Caribbean Neogene fish assemblages. The Cubagua fauna is a
spectacular coastal upwelling faunal association that existed dur-
ing the late Miocene to early Pliocene in northeastern Venezuela.
The association is characterized by the high abundance of me-
sopelagic, epipelagic, and benthopelagic elasmobranchs (sharks
and rays), teleosteans (bony fish), and the exceptional co-occur-
rence with shallow water elasmobranch and teleostean species. In
terms of species composition and abundance, the teleostean fossil
fauna is dominated by the most diverse groups of oceanic mid-
water fish, such as the myctophids (lantern fish). The elasmo-
branch fauna is dominated by the squaliformes (spiny dogfish)
that, as a group, have the broadest bathymetric and geographical
distribution of all sharks. For comparative purposes, we use the
fauna of the early Miocene Cantaure Formation of northwestern
Venezuela to represent a typical neritic environment. The Cantau-
re fauna is the best known Caribbean neritic fish assemblage (Nolf
and Aguilera, 1998) and starkly contrasts the Cubagna fauna. It
is dominated by shallow water sciaenids (croakers) and carchar-
hinid sharks (Fig. 1).
Today, the ecological environments in coastal northeastern Ven-
ezuela are controlled by the interplay among topographic setting,
oceanographic conditions, and seasonal variation in the flux of
*Panama Paleontology Project, Smithsonian Tropical Research Institute,
Panama.
nutrients from the Orinoco outflow, and variability of wind forces.
Together, these conditions produce the exceptional upwelling phe-
nomenon in the Caribbean Sea (Akl et al., 1997; Febres-Ortega
and Herrera, 1975; Gine´s, 1972; Herrera and Febres-Ortega, 1976;
Okuda, 1974; Varela et al., 1997), which may have been present
under similar circumstances during the late Miocene to early Pli-
ocene.
PHYSICAL AND GEOLOGICAL SETTING
,
STRATIGRAPHY AND
COLLECTIONS
The geologic unit treated in the present study is the late Mio-
cene to early Pliocene Cubagua Formation exposed along the Ar-
aya Peninsula, Cubagua Island, Margarita Island and the northern
Paria Peninsula (northeastern Venezuela) (Fig. 2). This northern
continental shelf of Venezuela represents the southeastern margin
of the Caribbean Sea and forms the southern prolongation of the
Lesser Antillian Arch (Sellier, 1974), which is clearly influenced
by the waters of the Atlantic Ocean (Okuda, 1974; Febres-Ortega
and Herrera, 1976). The coast is topographically complex and
forms the Cariaco Gulf, between the Araya Peninsula and the
continental coast. The Cubagua and Coche islands are located on
the continental platform to the north of the Araya Peninsula, sep-
arated by a body of water of approximately 42 m depth.Margarita
Island is located just to the north of both islands and is separated
from them by the Margarita Channel, which has a depth of ap-
proximately 30 m.
The Araya saddle (depth 300 to 450 m) and the Cubagua saddle
(depth 150 to 260 m) represent the submarine western prolon-
gation of the Araya Peninsula and the Cubagua Bank, and help
form the submarine valley located between the Araya Peninsula,
Cubagua Bank, and Margarita Island. This submarine valley has
the same orientation as the Cariaco Trench, which is a depressed
tectonic zone in the continental platform of northern Venezuela,
between the Unare Platform and the Tortuga Bank. It consists of
two small sub-basins, each reaching depths of about 1,400 m,
which are separated by a central saddle that shoals to a depth of
about 900 m. A distinctive characteristic of this trench is a zone
of anoxia below 330 m depth, a product of restricted water cir-
culation (Febres-Ortega and Herrera, 1975; Herrera and Febres-
Ortega, 1976).
According to the Le´xico Estratigra´fico de Venezuela (1997),
the holostratotype section of the late Miocene to early Pliocene
Cubagua Formation is located in the Can˜o´n de La Caldera (Cub-
agua Island) and is 70 m thick. The hypostratotype section is
located in the borehole of well Rio Caribe-1, the top of which is
733AGUILERA AND AGUILERA—A CARIBBEAN NEOGENE FISH ASSEMBLAGE
F
IGURE
1—Present-day bathymetric ranges of taxa represented in the ear-
ly Miocene Cantaure Formation (northwestern Venezuela). Teleostean
records following Nolf and Aguilera (1998), that for elasmobranchs are
original data. Interrupted lines indicate extinct species.
located at 649 m of depth sediment below sea level, with the
bottom at 2,708 m of depth. The Cubagua Formation sits con-
formably upon the Tres Puntas Formation and extends conform-
ably up into the overlying Cumana´ Formation.
The lower Cubagua Formation consists of gray shale with glau-
conite, abundant pyrite nodules, gray limonite, and some sandy
intercalation with fine clastics probably carried out by turbidity
currents. Occasionally clastic metamorphic and volcanic compo-
nents are present. The upper Cubagua Formation consists of bio-
clastic banks of reef, mollusk and bryozoans, quartz sandstone,
calcareous sandstone, bioclastic limestone, with interstratified ol-
ive gray shale, laminar glauconitic clay and gray limonite. The
sediments of the lower part appear to have been deposited in deep
water, and the upper part in shallow tropical water.
The Cubagua Formation is divided into four formal members:
the Cerro Verde Member, exposed on the Cubagua Island and
westernmost Araya Peninsula; the Cerro Negro Member, exposed
along the Western Araya Peninsula and Cubagua Island; and the
La Tejita and Las Hernandez members, exposed on Margarita
Island.
The oldest of these, the Cerro Verde Member (late Miocene to
early Pliocene), consists of conglomerate with sandy matrix, that
continues vertically into fossiliferous conglomerate sandstone,
bioclastic sandstone, and shale. This member has a thickness of
45 m at the type section. The Cerro Verde Member rests uncon-
formably upon the metamorphic rocks of the Late Jurassic to Ear-
ly Cretaceous Manicuare Formation. It passes transitionally up
into the overlying Pliocene Cerro Negro Member.
The Cerro Negro Member (Pliocene) consists of a 2 m thick
basal sandy marl, which is highly fossiliferous with Ostrea hai-
tensis and O. crassissima, followed by thin intercalations of sandy
lime, fossiliferous marl, and limestone marl. The total thickness
of the member is 22 m at the type locality. The Cerro Negro
Member sits conformably and transitionally on the Cerro Verde
Member. It appears to continue conformably up into the overlying
Pleistocene to Quaternary Barrigo´n Formation.
The La Tejita Member (late Miocene to early Pliocene) consists
of a 45 m thick basal conglomerate with quartz pebbles, schist
and igneous rocks, followed by gypsum clay, calcareous argilla-
ceous sandy and fossiliferous marl, which is highly fossiliferous
with O. crassissima. The La Tejita Member rests unconformably
upon metamorphic rocks and the Eocene Punta Carnero Group.
It passes unconformably up into the overlying Pliocene Manguillo
Formation.
The Las Hernandez Member (Early Pliocene) consists of un-
consolidated marl, clay, and lime. The thickness of this member
is unknown and is overlain by alluvium.
The sample area (Fig. 2) includes the following localities in
the Araya Peninsula: northwest of Cerro Barrigo´n, south of Cerro
El Macho, north of Cerro La Cruz, north of Cerro Cangrejero,
southwestern of Castillo de Araya, ENSAL Pyramid, south of
Araya Salina. The localities on the Cubagua Island are Can˜o´n de
La Caldera and Can˜o´n de Charagato, both in the northern part of
the island. The locality on Margarita Island was the SantiagoMar-
in˜o International Airport, close to Laguna de Las Marites.
The late Miocene to early Pliocene Cubagua Formation age is
based on local geological and stratigraphic relations (Bermudez,
1966; Gonzalez de Juana et al., 1980; Le´xico Estratigra´fico de
Venezuela, 1997; Sellier, 1959; Vignali, 1965). A preliminary bio-
stratigraphic age of 4.2 Ma (early Pliocene) is used throughout
this paper and is based on the median value of the age ranges of
planktonic foraminifera obtained from all sampled areas (L. S.
Collins, personal commun., 1998) except from the Margarita Is-
land locality: La Tejita and Las Hernandez members. The taxa
examined include Dentogloboquadrina altispira (Cushman and
Jarvis), last appearance datum (LAD 52.5 Ma), Globigerinoides
734 JOURNAL OF PALEONTOLOGY, V. 75, NO. 3, 2001
F
IGURE
3—Distribution of the collections, the total number of elasmo-
branchs and teleosts by faunule.
F
IGURE
4—Cumulative curve of the number of taxa (Elasmobranchii,
Teleostei and both) as a function of the number of collections.
F
IGURE
2—Location of the sampling sites in northeastern Venezuela. IN-
SETS OF FIGURE 2.–– Detailed map showing the sampling locations.
735AGUILERA AND AGUILERA—A CARIBBEAN NEOGENE FISH ASSEMBLAGE
F
IGURE
5—Present-day bathymetric ranges of taxa represented in the late Miocene to early Pliocene Cubagua Formation, northeastern Venezuela.
Interrupted lines indicate extinct species.
736 JOURNAL OF PALEONTOLOGY, V. 75, NO. 3, 2001
737AGUILERA AND AGUILERA—A CARIBBEAN NEOGENE FISH ASSEMBLAGE
←
F
IGURE
6—Common upwelling elasmobranchs species, late Miocene to early Pliocene Cubagua Formation, Cerro Verde Member, Venezuela. 1, 2,
Heptranchias perlo (Bonnaterre), labial and lingual view of lower tooth, UNEFM-PF-01, loc. OA-99–55, Cerro El Macho, Araya Peninsula. Scale
bar equal to 1 mm. 3, 4, Notorynchus sp., labial and lingual view of lower tooth, UNEFM-PF-02, loc. OA-99–55, Cerro El Macho, Araya Peninsula.
Scale bar equal to 1 mm. 5, Dalatias sp., labial view of lower tooth, UNEFM-PF-03, loc. PPP2556, Cerro Barrigo´n, Araya Peninsula. Scale bar
equal to 1 mm. 6, 7, Centrophorus sp., labial and lingual view of lower tooth, UNEFM-PF-04, loc. PPP3058, Cerro Barrigo´n, Araya Peninsula.
Scale bar equal to 0.5 mm. 8, Deania sp., lingual view of upper tooth, UNEFM-PF-05, loc. PPP2557, Cerro Barrigo´n, Araya Peninsula. Scale bar
equal to 0.5 mm. 9, Etmopterus sp., lingual view of lower tooth, UNEFM-PF-06, loc. PPP2565, Cerro Barrigo´n, Araya Peninsula. Scale bar equal
to 100 mm. 10, 11, Isistius aff. triangulus (Probst), labial and lingual view of lower tooth, UNEFM-PF-07, loc. PPP3055, Cerro Barrigo´n, Araya
Peninsula. Scale bar equal to 0.5 mm. 12, Trigonognathus aff. kabeyai Mochizuki and Ohe, lateral view of anterior tooth, UNEFM-PF-08, loc.
PPP2555, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 1 mm. 13, 14, Pristiophorus sp., dorsal and anterior view of rostral tooth, UNEFM-
PF-09, loc. PPP3054, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 1 mm. 15, 16, Squatina aff. dumerili LeSeur, labial and lateral view of
lateral tooth, UNEFM-PF-010, loc. PPP2563, Cerro El Macho, Araya Peninsula. Scale bar equal to 1 mm. 17, Heterodontus sp., lingual view of
anterior tooth, UNEFM-PF-011, loc. PPP3058, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 18, 19, Odontaspis ferox (Risso),
labial and lingual view of anterior tooth, UNEFM-012, loc. PPP2563, Cerro El Macho, Araya Peninsula. Scale bar equal to 10 mm. 20, 21,
Pseudocarcharias aff. kamoharai (Matsubara), labial and lingual view of anterior tooth, UNEFM-PF-013, loc. PPP2556, Cerro Barrigo´n, Araya
Peninsula. Scale bar equal to 5 mm. 22, 23, Alopias aff. superciliosus (Lowe), labial and lingual view of upper lateral tooth, UNEFM-PF-014, loc.
OA-99–54, Cerro El Macho, Araya Peninsula. Scale bar equal to 5 mm. 24, 25, Isurus sp., labial and lingual view of anterior tooth, UNEFM-PF-
015, loc. OA-99–54, Cerro El Macho, Araya Peninsula. Scale bar equal to 10 mm. 26, Mustelus sp., occlusal view of lateral tooth, UNEFM-PF-
016, loc. PPP2563, Cerro El Macho. Scale bar equal to 0.1 mm. 27, 28, Raja sp., lateral and lingual view of lateral tooth, UNEFM-PF-017, loc.
PPP2554, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm.
obliquus Bolli (LAD 51.8 Ma), G. obliquus Bolli var. extremus
Bolli and Bermudez (LAD 51.8 Ma), Globorotalia crassaformis
(Galloway and Wissler) first appearence datum (FAD 54.3 Ma),
G. margaritae Bolli and Bermudez (FAD 55.6 Ma, LAD 53.4
Ma), G. miocenica Palmer (FAD 53.4 Ma, LAD 52.2 Ma), G.
pseudomiocenica Bolli and Bermudez (LAD 52.2 Ma), Neob-
loloquadrina acostaensis (Blow) (LAD 51.8 Ma), N. dutertrei
(d’Orbigny) (FAD 53.0 Ma, LAD 51.8 Ma) and Orbulina
universa d’Orbigny. These preliminary results are consistent with
the earlier age estimates reported by Bermudez (1966).
In several cases, the precise stratigraphic placement of samples
in relation to unit boundaries is not known, however, studies of
the physical stratigraphy are in progress (A. Coates, personal
commun., 1999). We have followed the collectionsand processing
methods used in the Panama Paleontology Project (see Jackson
et al., 1999). Samples were processed using 2 mm and 500 micron
sieves. Teeth and otoliths were picked at 6 to 103magnification
using a stereomicroscope. All teeth and otoliths were identified to
species. The basic references use to identify the Elasmobranch
teeth and Teleostean otolith are Cappetta (1987) and Nolf (1985)
respectively. We have posted digital images of otoliths and the
diagnostic character used for identification on the NMITA website
(nmita.geology.uiowa.edu).
A total of 38 collections were grouped into nine ‘‘faunules’’ in
an attempt to lessen the effects of sample size on the analysis of
patterns and trends of faunal assemblages. The groupings were
made on the basis of location, while taking into consideration the
continental and insular outcrop of the Cubagua Formation. Each
faunule corresponds to a single fossiliferous stratigraphic horizon
at a single outcrop. Due to pervasive bioturbation at the great
majority of sites, bedding could not usually be observed, there-
fore, packages of lithologically identical sediment, typically
amounting to a few meters of section, were treated as a single
horizon. Nearby but physically separate outcrops of the same
stratigraphic horizon were treated as separate faunules for pur-
poses of replication in the analyses (e.g., faunule 7: Can˜o´n de La
Caldera, and faunule 8: Can˜o´n de Charagato, both on Cubagua
Island).
All of the samples from the late Miocene to early Pliocene
Cubagua Formation were used to make one estimate of paleo-
bathymetry, using both elasmobranch and teleostean data follow-
ing the methods described in Aguilera and Aguilera (1999). The
method is based on the assumption that the taxa encounteredlived
together in the same environment represented by the sedimentary
horizons sampled. All of the identifiable teeth and otoliths from
the samples were identified and depth ranges were assigned to
each taxon based on the known depths of living counterparts.
COMPOSITION OF THE FAUNA
Sixty-three families, 115 genera, and 140 species in the 38
collections (Appendix 1) represent the late Miocene to early Pli-
ocene Cubagua Formation ichthyofauna. One hundred or more
specimens in the 38 collections comprise only 16 out of the total
152 taxa. These include 69 percent Myctophidae with other minor
groups of less than five percent in abundance (e.g., Gobiidae,
Sternoptychidae, and Merluccidae). The presence and abundance
of Myctophidae (including six genera and 11 species) in almost
47 percent of the collections, can be attributed to upwelling con-
ditions.
Taxonomic diversity in these nine faunules ranges from a min-
imum of three to a maximum of 125 taxa, and abundance ranges
from three to 1,989 specimens (medians: 33 taxa, 2,327 speci-
mens per faunule). Elasmobranch diversity varies a few fold but
teleostean diversity varies enormously (three to 55 fold). This
variation is not due to differences in numbers of samples (Fig. 3).
Elasmobranchs are clearly well sampled, as demonstrated by the
leveling off of the sampling effort curve (Fig. 4), but teleosts
appear to be less thoroughly sampled, with diversity still increas-
ing with numbers of collections.
The bathymetric distribution based on living representative
counterparts is 0 to 100 m (Fig. 5). However, this ichthyofauna
is characterized mostly by the co-occurrence of oceanic and ne-
ritic fish.
Faunules 1, 2, and 3 (Appendix 1) at the Araya Peninsula show
very low diversity and abundance, and an analysis of the faunal
associations was not possible.
Faunule 4 at the Cerro Barrigo´n (Araya Peninsula) is the most
diverse and abundant faunule from the Cubagua Formation. The
sampled horizon corresponds to the calcareous fine sandstone and
shale levels, presumably from the middle part of the Cerro Verde
Member. The assemblage is characterized by the co-occurrence
of epipelagic, mesopelagic and benthopelagic taxa together with
neritic taxa. This mixture of faunas is strongly indicative of up-
welling conditions close to the Cariaco Trench.
Faunule 5 at the Castillo de Araya (Araya Peninsula) is very
poorly fossiliferous and is not considered in the present analysis.
Faunule 6 at the Cerro El Macho (Araya Peninsula) is signifi-
cantly lower in diversity and abundance compared to Faunule 4.
738 JOURNAL OF PALEONTOLOGY, V. 75, NO. 3, 2001
F
IGURE
7—Common upwelling teleostean species, late Miocene to early Pliocene Cubagua Formation, Cerro Verde Member, Venezuela. 1, 2, Diaphus
dumerili (Bleeker), right otolith (inner view), UNEFM-PF-018, UNEFM-PF-019, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal
to 0.5 mm. 3, 4, Diaphus splendidus (Brauer), left otolith (inner view), UNEFM-PF-020, UNEFM-PF-021, loc. PPP2553, Cerro Barrigo´n, Araya
Peninsula. Scale bar equal to 1 mm. 5, 6, Diaphus sp. 1, left otolith (inner view), UNEFM-PF-022, UNEFM-PF-023, loc. PPP3055, CerroBarrigo´n,
Araya Peninsula. Scale bar equal to 0.5 mm. 7, 8, Diaphus sp. 2, left and right otolith respectively (inner view), UNEFM-PF-024, UNEFM-PF-
025, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 9, Diaphus sp. 3, right otolith (inner view), UNEFM-PF-026,
loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 10, Electrona rissoi (Cocco), left otolith (inner view), UNEFM-PF-
027, loc. PPP2572, Can˜o´n de Charagato, Cubagua Island. Scale bar equal to 1 mm. 11, Hygophum macrochir (Gunther), left otolith (inner view),
UNEFM-PF-028, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 12, Lampanyctus cupriarius (Taaning), right otolith
(inner view), UNEFM-PF-029, loc. PPP2553, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 13, 14, Lampanyctus aff. latesulcatus
Nolf and Stringer, left and right otolith respectively (inner view), UNEFM-PF-030, UNEFM-PF-031, loc. PPP3055, Cerro Barrigo´n,
739AGUILERA AND AGUILERA—A CARIBBEAN NEOGENE FISH ASSEMBLAGE
←
Araya Peninsula. Scale bar equal to 0.5 mm; 15–21, Lampadena jacksoni n. sp., left otolith (inner view), UNEFM-PF-032 to UNEFM-PF-038,
loc. PPP3055, Cerro Barrigo´n, Araya Peninsula (Holotype UNEFM-PF-035), Scale bar equal to 0.5 mm. 22, Symbolophorus sp., left otolith (inner
view), UNEFM-PF-039, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 1 mm. 23, Coelorinchus aff. coelorinchus Risso, right
otolith (inner view), UNEFM-PF-040, loc. PPP2556, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 24, Merluccius sp., right otolith
(inner view), UNEFM-PF-041, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm. 25, Steindachneria argentea Goode
and Bean, left otolith (inner view), UNEFM-PF-042, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 1 mm. 26, Bregmaceros
aff. cantori Miliken and Houde, otolith (inner view), UNEFM-PF-043, loc. PPP3055, Cerro Barrigo´n, Araya Peninsula. Scale bar equal to 0.5 mm.
The ichthyofauna is characterized primarily by the presence of
elasmobranchs and a relative scarcity of teleosteans, and the pres-
ence of deep-water shark species suggests an external neritic en-
vironment. The horizon sampled corresponds to calcareous sand-
stone and marl, apparently from the upper part of the Cerro Verde
Member of the Cubagua Formation.
Faunule 7 at Can˜o´n de La Caldera and Faunule 8 at Can˜o´n de
Charagato (Cubagua Island) have similar fish assemblages which
are moderately abundant and characterized by a typical neritic
ichthyofauna. The horizon sampled corresponds to calcareous
sandstone and marl, probably from the middle to upper part of
the Cerro Verde Member of the Cubagua Formation.
Faunule 9 from the La Tejita Member exposed at the Margarita
International Airport (Margarita Island) has low abundance; how-
ever, the presence of elasmobranch species suggests an inner ne-
ritic environment.
Many distinctive circumstances produce the general paleoeco-
logical conditions of this remarkable coastal upwelling (Araya
Peninsula horizon) and inner neritic (Cubagua and Margarita Is-
lands horizon) assemblage fauna in the late Miocene to early Pli-
ocene Cubagua Formation. Additional evidence supporting up-
welling can be found by examining the ecological associations of
the common living fish, particularly those relating to diet and
feeding habits.
Ebert et al. (1992) clearly showed that the dominant fish prey
in the diets of fifteen species of spiny dogfish sharks, Centropho-
rus (collected between 380 to 800 m of depth), Deania (400 to
800 m of depth), Etmopterus (450 to 925 m of depth) and Squalus
(50 to 550 m of depth) was mainly myctophids and merlucciids.
Species such as, Diaphus sp., Epigonus sp., Merluccius sp., Coe-
lorinchus sp., Maurolicus muelleri (Gmelin), and Lepidopus cau-
datus (Euphrasea) are present in the squalid diet.
These predatory bathyal sharks and the mesopelagic prey re-
lationship from the west coast of South Africa, can be used in the
analysis of the fossil Cubagua Formation fish assemblages (up-
welling fauna), because both groups (predators and prey) are very
well represented in the collections (Appendix 1).
Other living epibenthic and benthic predators in the northeast-
ern Atlantic, such as Mustelus sp., Squatina sp., and Raja sp.,
feed primarily on crustaceans, but also on fish such as ammody-
tids, callyonimids, gadoids, gobiids, and pleuronectiforms (Ellis
et al., 1996), which also are present in the fossil associations.
In contrast, the diet of neritic predators such as the bonnethead
shark, Sphyrna tiburo (Linnaeus), from southwest Florida (Cortes
et al., 1996) is dominated by crustaceans, principally shallow wa-
ter blue crabs (Callinectes sapidus Rathbun). The diet of the lem-
on shark, Negaprion brevirostris (Poey), collected from the Ba-
hamas, is dominated by inner neritic fish, like sparids, lutjanids
and gerreids (Cortes and Gruber, 1990). In addition, the diet of
the southern stingray, Dasyatis americana Hildebrand and
Schroeder, is dominated by crustaceans, mainly decapods and por-
tunids (Gillian and Sullivan, 1993).
Coastal upwelling is the principle oceanographic phenomenon
in the Caribbean associated with higher biological productivity.
Upwelling of colder water, less than 248C and rich in nutrients,
stimulates phytoplankton growth which, in turn, supports the sec-
ondary productivity and fisheries in the region. Upwelling is the
product of the seasonal winds along the northeastern Venezuelan
coast during February to April, when maximal wind speeds reach
5 m/s (Herrera and Febres-Ortega, 1975), and chlorophyll con-
centration reaches a maximal value in the same period (Varela et
al., 1997). Seasonal productivity increases due to upwelling over
much of the northeastern Venezuelancoast, and produces the most
important fishery activity in the country (Gimenez et al., 1993).
Evidence that higher biological productivity also occurred in
this area during the geologic past is found in the Neogene sedi-
mentary basin along the northeastern Venezuela. The sequence of
Cubagua Formation (late Miocene to early Pliocene), Barrigo´n
Formation (Pliocene), Cumana´ Formation (Pliocene), Castillo de
Araya Formation (Pleistocene) exhibits high mollusk diversity
(see: Bermudez, 1966; Macsotay, 1965). Also, the presence of
large specimens (at least 30 cm length) of Lyropecten and Os-
traea, and taxa that typically inhabit deep water or superficial cold
waters, are indicative of higher phytoplankton productivity.
In particular the mesopelagic and bathypelagic Cubagua For-
mation ichthyofauna from the Araya Peninsula horizon, contrast
markedly with the neritic assemblages from the insular area (Cub-
agua and Margarita islands). These differences in faunal compo-
sition are probably also the result of the influence of submarine
topography (the Araya Peninsula horizon close to the Cariaco
trench, and Cubagua and Margarita islands surrounded by shallow
continental platform).
Based on this preliminary evidence, coastal upwelling was ap-
parently persistent in the geological past and probably varied in
intensity.
Compared with other Caribbean Neogene fish assemblages that
are reported in the literature, such as Cantaure Formation in Ven-
ezuela (Nolf and Aguilera, 1998: 61 teleostean taxa; herein: 18
elasmobranch taxa), Gatun Formation in Panama (Gillette, 1984:
24 teleostean and 17 elasmobranch), Bowden Formation in Ja-
maica (Stringer, 1998: 68 teleostean; Kruckow and Thies, 1990:
one elasmobranch), Nariva, Brasso, Manzanilla, and Tarama for-
mations in Trinidad (Nolf, 1976: 66 teleostean; Kruckow and
Thies, 1990: 35 elasmobranch), and Baitoa, Cercado, Gurabo and
Mao Formations (Nolf and Stringer, 1992: 65 teleostean taxa; no
record of elasmobranch is available), the Cubagua Formationwith
140 species, and approximately 714 elasmobranch teeth and
20,105 teleostean otoliths, is the more diverse and abundant Ca-
ribbean Neogene fish assemblages.
NOTE OF SPECIES OCCURENCES
Additional comments are made below regarding the distribution
of epipelagic, mesopelagic, and benthopelagic elasmobranchs and
teleosts in the upwelling assemblages. It was observed that some
fossils of living species are not present in the Recent Caribbean
ichthyofauna. Figures 6 and 7 show thirty one species that are
distinctive and support the upwelling interpretation.
The sharpnose sevengill shark (Fig. 6.1, 6.2), Heptranchias
perlo (Bonnaterre) is the first fossil record for the Caribbean late
740 JOURNAL OF PALEONTOLOGY, V. 75, NO. 3, 2001
Miocene to early Pliocene. An earlier Caribbean species, Hep-
tranchias tenuidens (Leriche, 1938) was recorded from the late
Oligocene of Venezuela. The first recorded catch of living Hep-
tranchias, H. perlo, was made in 1998 along the Venezuelan coast
from a depth of 166 fathoms (Crevigo´n and Alcala´, 1999).
The broadnose sevengill shark (Fig. 6.3, 6.4), Notorynchus sp.
is the first fossil record for the Caribbean late Miocene to early
Pliocene, although the recent monotypic Notorynchus cepedianus
(Peron) is known in mostly temperate seas (Compagno, 1984).
Deep water squaliform sharks (Fig. 6.5–6.12), Dalatias sp.,
Centrophorus sp., Deania sp., Etmopterus sp., Isistius aff. trian-
gulus (Probst) and Squalus sp. (no figure), represent additional
fossil records for the Caribbean late Miocene to early Pliocene.
The Miocene species Centrophorus sp., Squalus stehleni (Leri-
che), Etmopterus acutidens Casier, and Isistius triangulus (Probst)
are the only previous Caribbean records cited by Kruckow and
Thies (1990).
The present record of monotypic squalid shark Trigonognathus
aff. kabeyai Mochisuki and Ohe (Fig. 6.12) is the first in the
Caribbean late Miocene to early Pliocene. Cappetta and Sylvain
(H. Cappetta, personal commun., 1999) reported an early record
of Trigonognathus in the Paleogene of southern France. The liv-
ing species is known from only two specimens collected from the
coastal waters of Japan at depths of 330 m and 360 m, respec-
tively (Mochisuki and Ohe, 1990).
The longnose sawshark (Fig. 6.13, 6.14) Pristiophorus sp. is
the first fossil record for the Caribbean late Miocene to early
Pliocene. This genus is represented in the fossil record from Eu-
rope, Japan, New Zealand, U.S.A., and South America (Pacific
of Peru) (Cappetta, 1987). The living Atlantic species Pristiopho-
rus schroederi Springer and Bullis, is known only from the Ba-
hamas region (Compagno, 1984).
The angleshark (Fig. 6.15, 6.16) Squatina aff. dumerili Le Seur,
is an uncommon fossil occurrence for the Caribbean (Kurckow
and Thies, 1990). The living counterpart can be found in water
depths reaching 1,390 m (Compagno, 1984).
The horn shark (Fig. 6.17) Heterodontus sp. is the first fossil
record for the Caribbean late Miocene to early Pliocene. Earlier
fossil occurrences in the Caribbean of Heterodontus (H. pineti
Case, H. cf. woodwardi Casier and H. janefirdae Case) were re-
ported by Kruckow and Thies (1990) from Eocene and early Mio-
cene deposits of the U.S.A. The living species is known only from
the eastern and western Pacific, and western Indian Ocean (Com-
pagno, 1984).
The small-tooth sand tiger shark (Fig. 6.18, 6.19) Odontaspis
ferox (Risso) is the first fossil record for the Caribbean late Mio-
cene to early Pliocene. Landini (1977) reported an early record
of Odontaspis ferox in the early Pliocene of Tuscani, Italy. Al-
though this species is also known from the northeastern Atlantic;
eastern, central and western Pacific, and the Western Indian Ocean
(Compagno, 1984).
The crocodile shark (Fig. 6.20, 6.21) Pseudocarcharias ka-
moharai Matsubara, is represented in the Mediterranean fossil re-
cord (Cigala-Fulgoi, 1992). This present fossil record is the first
for the Caribbean late Miocene to early Pliocene. The living spe-
cies is also known in the eastern and western south Atlantic; east-
ern, central and western Pacific, and in the western Indian Ocean
(Compagno, 1984; Lessa et al., 1991). Alcala´ (1993) reports a
single Caribbean record of a live specimen of Pseudocarcharias
kamoharai caught off the Venezuelan coast. Crevigo´n and Alcala´
(1999) comment that additional specimens are occasionally ob-
tained by sport-fisherman.
The bigeye thresher (Fig. 6.22, 6.23) Alopias superciliosus
Lowe, represents an additional fossil record for the Caribbean
Neogene. The depth distribution of the living counterpart ranges
from the surface to at least 500 m of depth (Compagno, 1984).
The mako (Fig. 6.24, 6.25) Isurus sp. is a genus common in
all Recent Oceans (Compagno, 1984). Miocene to middle Plio-
cene species are widely distributed in Europe, Africa, Australia,
North and South America (Cappetta, 1987; Kruckow and Thies,
1990). Leriche (1938) reported an earlier Venezuelan record of
Isurus cf. desori from the late Oligocene San Lorenzo Formation.
The dusky smooth hound (Fig. 6.26) Mustelus sp. is a new
fossil record for the Caribbean Pliocene. According to Compagno
(1984), it is found in tropical waters ranging from shallow inshore
and intertidal zones most commonly down to 200 m of depth, but
occasionally down to 579 m.
The skates are represented by a deep tropical batoid, Raja sp.
(Fig. 6.27, 6.28). The fossil record shows them to be uncommon
in the entire Caribbean area (Kruckow and Thies, 1990).
There is a high diversity and abundance of lantern fish (Fig.
7), such as, Diaphus dumerili (Bleeker), D. splendidus (Brauer),
approximately four additional undetermined Diaphus species,
Electrona rissoi (Cocco), Hygophum macrochir (Gunther), Lam-
panyctus cupriarius (Taaning), L. aff. latesulcatus Nolf and Steur-
baut, and Symbolophorus sp., represent additional fossil records
for the Caribbean Neogene.
Special attention is given to a new extinct species of lantern
fish, Lampadena jacksoni n. sp., because of its abundance and
distribution in the Cubagua Formation (Araya Peninsula horizon,
Cerro Barrigo´n, faunule 4), which perhaps might be used as a
fossil indicator of Miocene-Pliocene upwelling events in the Ca-
ribbean Sea.
Associated with the myctophids (lantern fishes) is the presence
of marine benthopelagic gadiforms (Fig. 7.23–7.26) such as, the
hollowsnout grenadier, Coelorhinchus aff. coelorhinchus (Risso),
the hake, Merluccius sp., the luminous hake, Steindachneria ar-
gentea Goode and Bean, and the codlets, Bregmaceros aff. can-
tori Miliken and Houde are indicative of deep-water in the tropics
(Cohen et al., 1990).
SYSTEMATIC PALEONTOLOGY
Family M
YCTOPHIDAE
Gill, 1893
Genus L
AMPADENA
Goode and Bean in Gill, 1893
L
AMPADENA JACKSONI
new species
Figure 7.15–7.22
Diagnosis.Otolith uniform oval to oblong-ovate shape, with
a markedly straight dorsal margin, distinctly notched in the pos-
tero-dorsal angle and well developed and sharply rostrum and
antirostrum.
Description.Otolith oval to oblong-ovate. Moderately thin
and slightly convex. Dorsal margin markedly straight; sculptures
irregular, and distinctly notched in the postero-dorsal angle. Ven-
tral margin rounded and serrate, with nine to ten denticles. Pos-
terior margin rounded, irregular, and notched dorsally. Elongated
slightly and flares ostium, anteriorly with large, low, and oblong
colliculum. Short cauda, with oblong colliculum. Ostio-caudal
differentiation slightly ventrally constricted. Ostium/cauda rela-
tion 1:0.5. Pseudocolliculum, as long as posterior colliculum. No
collum. Crista superior low ridges from ostium to mid-cauda,
poorly developed posteriorly, and absent at caudal tip. Crista in-
ferior poorly developed along entire sulcus, and absent under cau-
da. Oval and shallow dorsal depression. No ventral depression,
and shallow groove near margin. Large, broad, and angled ros-
trum, with sharply rounded tip. Small, narrows, and sharply
rounded antirostrum. Excisura moderately wide, notch shallow,
and angle acute.
Etymology.Dedicated to Dr. J. Jackson, Smithsonian Tropical
Research Institute, Panama, for his valuable contribution in the
Panama Paleontology Project.
Types.The holotype, UNEFM-PF-035, and six paratypes,
741AGUILERA AND AGUILERA—A CARIBBEAN NEOGENE FISH ASSEMBLAGE
UNEFM-PF-032 to UNEFM-PF-034, and UNEFM-PF-036 to
UNEFM-PF-038, are deposited in the Universidad Francisco de
Miranda, Coro, Edo. Falco´n, Venezuela.
Occurrence.The material was collected from the late Mio-
cene to early Pliocene Cubagua Formation, Cerro Verde Member,
north of Cerro Barrigo´n, Araya Peninsula, northeasternVenezuela
at locality PPP3055 (Fig. 2).
Discussion.This otolith can be distinguished from the otoliths
of recent species Lampadena chavesi Collett, L. luminosa (Gar-
man), L. notialis Nafpaktitis and Paxton, and L. speculigera Goo-
de and Bean [Smale et al., 1995 (pl. 21, fig. E1–G2; pl. 22, fig.
A1–A3)], and from the fossil species Lampadena dea Fraser-
Brunner, L. speculigeroides Brzobohaty and Nolf, and L. gracile
(Schubert) [Brzobohaty and Nolf, 1996 (pl. 4, fig. 1–10, 12–16)]
by the uniform oval to oblong-ovate shape otolith, with a mark-
edly straight dorsal margin, distinctly notched in the postero-dor-
sal angle, and well developed and sharply rostrum and antiros-
trum. Lampadena jacksoni n. sp. occurs on the lower part of the
Cubagua Formation (Cerro Verde Member) exposed at Araya
Peninsula (Appendix 1, faunule 4) associated with high diversity
upwelling assemblages. Its unique presence and abundance in a
specific horizon suggests that it may be a useful fossil indicator
of Caribbean upwelling events during the late Miocene to early
Pliocene.
ACKNOWLEDGMENTS
This study would not have been possible without the encour-
agement and support of A. Coates and J. Jackson. We are also
grateful to H. Fortunato, V. Padron, J. Reyes, and R. Sanchez for
their help with the fieldwork. J. Reyes also assisted with the lab-
oratory work. H. Cappetta and D. Nolf contributed greatly in the
original identification and/or verification of elasmobranch teeth
and teleostean otoliths, respectively. L. Collins provided age de-
termination. W. Campos and J. Zeballos assisted us with the Scan-
ning Electronic Microscope. M. Diaz assisted with light photog-
raphy. M. Mendoza, D. Miller, and J. Moody assisted with lin-
guistic corrections. T. Hazen assisted with editorial corrections.
We also thank M. L. de Gamero and J. Lundberg for helpful
comments. Discussions with J. Jackson helped to organize our
manuscript. This work was supported by the National Geographic
Society, the Walcott and Scholarly Studies Funds of the Smith-
sonian Institution, the Smithsonian Tropical Research Institute,
and the Francisco de Miranda University, Venezuela.
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CCEPTED
15 D
ECEMBER
2000
APPENDIX I
Elasmobranchs and teleosteans from the Cubagua Formation discussed
in the text. Frequencies are available on the Journal of Paleontology’s
Supplemental Database (http://www.journalofpaleontology.org).
