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New fossils shed light on the Late Cretaceous terrestrial community in the
Caribbean and the First American Biotic Interchange

Article  in  Cretaceous Research · October 2021

DOI: 10.1016/j.cretres.2021.105067

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 Cretaceous Research 130 (2022) 105067

Contents lists available at ScienceDirect

Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

New fossils shed light on the Late Cretaceous terrestrial community in

the Caribbean and the First American Biotic Interchange

zaro W. Vin
La
pez a, *, Ignacio A. Cerda b, Julian Correa-Narvaez a, Laura Codorniú c,
~ ola-Lo
Carlos R. Borges-Selle
n d, Alberto F. Arano-Ruiz d, Yasmani Ceballos-Izquierdo e
a
Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800, USA
b
n en Paleobiología y Geología, Museo Carlos Ameghino, Universidad Nacional de Río Negro, Belgrano 1700, Paraje Pichi
CONICET-Instituto de Investigacio
Ruca (predio Marabunta), Cipolletti, 8300, Río Negro, Argentina
c
CONICET, Departamento de Geología, Facultad de Ciencias Físico, Matema
ticas y Naturales, Universidad Nacional de San Luis, San Luis, Argentina, Av.
Ej

ercito de Los Andes 950
d
Sociedad Cubana de Geología, Cienfuegos, Cuba
e
Instituto de Geofísica y Astronomía, Cuba

a r t i c l e i n f o a b s t r a c t

Article history: The Caribbean islands are one of the most important hotspots of endemism and biodiversity globally, and
Received 7 February 2021 the scenario of unique examples of biological radiations. Although our knowledge of the current and
Received in revised form recently extinct diversity in the area is strong, the origin and evolution of most groups in the region
26 September 2021
remain obscure because of the absence of fossils from deep time periods. The existence of temporal
Accepted in revised form 14 October 2021
Available online 25 October 2021
islands on the Caribbean plate can be traced back to the late Mesozoic, but little evidence of the paleo-
communities that once inhabited the archipelago and their relationship with the older lineages in the
region has been discovered. Simultaneously, the relationship of the early Caribbean archipelago with the
Keywords:
Caribbean
Late Cretaceouseearly Paleogene biotic interchange between the Americas has remained unsolved. Here
Biogeography we describe the first evidence of a Late Cretaceous terrestrial community in the region based on several
Late Cretaceous remains recovered at three upper Campanianelower Maastrichtian localities in Central Cuba. The fossil
Pterosaur assemblage includes four specimens referable to a midsize pterosaur on the base of morphological and
Cuba paleohistological characters, as well as seeds and casts of leafy shoots of plants of the families Cupres-
saceae and Lauraceae. Fossils fruits of a new taxon closely related to Chlorocardium are of particular
interest because they correspond to the first direct evidence of the role played by the Caribbean seaway
and islands in the First American Biotic Interchange.
Published by Elsevier Ltd.

1. Introduction islands on the Caribbean became permanently exposed (Iturralde-

The origin of the present and past biota of the West Indies has
been a question surrounded by vigorous debate for over a century
without a clear consensus. The extensive bias of the fossil record on
the islands, where remains of the first species established on the
twilight of the Mesozoic are virtually absent, has impaired any
attempt to assess the diversity of the early arc of islands. The origin
of the Caribbean Plate can be traced back to the Late Jurassic when
its oceanic crust began to form east of the North and South America
plates (Pindell and Kennan, 2009). By the late Eocene, some of the
* Corresponding author.
~ ola-Lo
E-mail address: lwvl94@gmail.com (L.W. Vin
pez).
Vinent and MacPhee, 1999; Iturralde-Vinent, 2006), but the exis-
tence of older temporal islands is well documented based on
geological and, to a lesser degree, paleontological evidence (Rosen,
1975; Iturralde-Vinent and MacPhee, 1999; Iturralde-Vinent, 2006;
Pindell and Kennan, 2009). During the Cretaceous, a volcanic
island-arc was situated on the Caribbean plate's eastern leading
margin in the gap between North and South America (Pindell and
Kennan, 2009) (Fig. 1). Nearly 70 Mya, the extinction of the vulca-
nism in part of the arc system resulted in the extensive uplift of
some islands and their areal increment (Iturralde-Vinent, 2006;
Garcia-Casco et al., 2008). Coincidentally, this is when the first
pieces of evidence of a direct biotic exchange between North and
South America are found in the fossil record after the drift between
Laurasia and Gondwana. Extensive evidence of this event is known

https://doi.org/10.1016/j.cretres.2021.105067
0195-6671/Published by Elsevier Ltd.
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al.
from the fossil record in the continent and has been supported by
phylogenetic studies on the modern biota of the region. The biotic
interchange involved the migration of hadrosaurs, condylarth and
didelphimorph mammals, land snails, plants, and other groups
from North to South America, while titanosaurs colonized the
boreal continent from the south (Bonaparte, 1984; Rage, 1986;
Gayet et al., 1992; Chanderbali et al., 2001; Santiago-Valentin and
Olmstead 2004; Pascual, 2006; Uit De Weerd, 2008; Valieri et al.,
2010; Goin et al., 2016). The dispersal between the two isolated
landmasses supposedly took place through the evanescent islands
of the Caribbean arc in the Campanian e Paleocene, coinciding with
the Late Cretaceous transgression (Chanderbali et al., 2001;
Iturralde-Vinent, 2006; Ortiz-Jaureguizar & Pascual, 2011; Goin
et al., 2016). Here we refer to this event as the First American Bi-
otic Interchange (FABI).
Although the evidence of the interchange between the Americas
is unequivocal, the role of the Caribbean is not completely clear
given the lack of fossils from terrestrial environments in the island-
arc. Most Upper Cretaceous fossil-bearing beds in the region are
marine (Rojas-Consuegra et al., 1995; Mutter et al., 2005;
Underwood and Mitchell, 2020; Vega et al., 2020) lacking any ev-
idence of land-dwelling species, except for some undescribed
plants from the upper Maastrichtian of Jamaica and Cuba (Mitchell
and Blissett, 2001; Villegas-Martin and Rojas-Consuegra, 2011).
Meanwhile, according to molecular studies (Noonan et al., 2013;
Springer et al., 2018), the origin of the oldest lineages currently
Fig. 1. Paleogeographic reconstruction of the Caribbean in the latest Cretaceous modified from Iturralde-Vinent (2006). Note the future Antilles were localized at the eastern leading
margin of the Caribbean Plate in the gap between North and South America. The arrows indicate the direction of migrations of different lineages during the FABI, and the star the
general location of the localities studied.
2
Cretaceous Research 130 (2022) 105067
inhabiting the Antilles can be traced back to the Late Cretaceous
(but see Iturralde-Vinent and MacPhee, 1999).
Here, we bring attention upon a collection of fossil vertebrates
and plants recovered from three upper Campanianelower Maas-
trichtian outcrops in Central Cuba, representing the first remains of
pterosaurs and plants of the families Lauraceae and Cupressaceae
described from the Upper Cretaceous in the Antilles. The discovery
of these fossils provides a much-needed insight into the biota that
once inhabited those ephemeral islands and their relationship with
the FABI (Iturralde-Vinent, 2006). In particular, fossil fruits of a new
taxon closely related to the extant genus Chlorocardium strongly
supports the hypothesis that the Caribbean seaway and islands
formed along the latitude of Central America were involved in the
FABI.
2. Materials and methods
The specimens here described are housed at the Museo
Municipal de Rodas (MMR) in Rodas, Cienfuegos, Central Cuba.
Thin sections of two specimens of pterosaur (MMR A-1 and
MMR A-2) were prepared at the Paleohistological Laboratory of the
Museo Provincial Carlos Ameghino (Cipolletti, Río Negro Province,
Argentina). The cross-section slices were prepared using standard
methods in paleohistology (Chinsamy-Turan and Raath, 1992) and
studied using a petrographic polarizing microscope (BestScope and
Nikon E200 pol). The nomenclature and definitions of structures
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al.
s et al., 1991;
used in this study are standardized in the field (Ricqle
Chinsamy-Turan, 2005).
Also, fossils fruit were CT scanned to help visualize the
morphology. The CT scan was performed at the University of Florida
College of Engineering Nanoscale Research Facility (NRF) with a GE
Phoenix V|tome|xm240 CT Scanner, using a Tungsten reflection
target and 0.5 mm copper filter, at 200 kV and 240 mA, with 1500
images of a single specimen for a voxel size of 0.858154 mm.
3. Geological setting
The specimens here studied were recovered from three outcrops
of the upper Campanianelower Maastrichtian Monos Formation in
the localities Presa Damuji, Aguaditas, and Puente Potrerillo, in
Cienfuegos Province, Central Cuba (Fig. 2). The Monos Formation
was deposited during part of a transgressive cycle characterized by
a sequence of polymictic gravels that transition into sandstones
originated by the erosion of the Antillean Cretaceous Volcanic Arc.
Simultaneously, siltstone, claystone, and calcareous beds can also
be found intercalated in some sections. These clastic sediments
were deposited in a post-volcanic basin during the upper
Campanianelower Maastrichtian (Iturralde-Vinent, 2014). The
input of land-derived sediments decreases upward in the formation
as the upper Campanian transgression advanced. Foraminifera
from the marine sections of the formation supports the age of the
sediments; they include Globotruncana arca, Globotruncanita cf.
G. stuarti, Sulcoperculina cf. S. dickersoni, S. globosa, Praeglobo-
truncana havanensis, Pseudotextularia cornuta, Rugoglobigerina
rugosa, and Vaughanina barkeri (Gil-Gonzales, 2013).
Remains of pachydiscid and well-preserved baculitid ammon-
ites (Baculites paradoxus, B. sp.), gastropods, rudists, other bivalves,
echinoids, starfish, decapod crustaceans (Vegaranina rivasi, unde-
scribed raninidae), elasmobranch vertebra, plants (trunks, seeds,
and leaves), pterosaurs, among others have been recovered from
sediments of the Monos Formation (Arano et al., 2018; Ceballos-
Izquierdo et al., 2021). Long bones of pterosaur were filled with
polymictic sandstone, suggesting fragmentation and transportation
before the final burial. The sedimentology of the formation and the
association of abundant fossil plants, raninid crustaceans, and
pterosaurs support that part of the sequence was deposited in the
distal part of fluvial systems under the influence of marine and
terrestrial environments. Furthermore, considering that fruits and
leaves tend to spread close to their source (Dutra, 2007), the
abundance and preservation of these fossils reinforce the local
origin of part of the assemblage.
3.1. Presa Damuji
More than 60 m of upper Campanianelower Maastrichtian
sediments of the Monos Formation are exposed in the east bank of
the Damuji river (N 22
160 51.11400 , W 80
320 52.508), south of the
Presa Damuji's spillway in the municipality of Rodas, Cienfuegos
Province, Central Cuba (Fig. 2). The outcrop can reach up to 4 m in
thickness in some sections and is covered discontinuously by a
deposit of alluvial and geothermal Quaternary sediments.
The Upper Cretaceous section at Presa Damuji is dominated by
dark beige to brown, friable, and very fine polymictic sandstone
rich in clay, which grades into siltstone in some sections. Fossils
found in the sandstone include a fragment of the diaphysis of
pterosaurs, raninid crustaceans, bivalves (Inoceramus), gastropods,
echinoids, and abundant remains of plants consisting in fragments
of charcoal, pith casts, leafy shoots of the family Cupressaceae, and
large seeds of new taxa closely related with Chlorocardium. Within
the polymictic sandstone, there are medium-sized lenses of organic
rick black siltstone and mudstone with abundant fossils of small
3
Cretaceous Research 130 (2022) 105067
gastropods, bivalves, fragments of plants, and less commonly
decapod crustaceans including shrimps. On the south edge of the
outcrop, there is a bank of oysters and Inoceramus insitu.
The combination of very fine sandstone derived from the
erosion of the extinct volcanic island with the faunal association
suggests they were deposited in the distal section of a fluvial sys-
tem with high sediment input under the influence of continental
and marine environments. Also, small channels cut intermittently
thru this bed and were filled with sediments rich in organic matter
under low energy conditions.
3.2. Aguaditas
Most outcrops at this locality are found in a creek that cuts
through different facies of the Monos Formation at Aguaditas farm
(N 22
180 31.31300 , W 80
300 26.62500 ), 5 km southeast of the town
Rodas in Cienfuegos Province, Central Cuba (Fig. 2). The base sec-
tion is characterized by the presence of polymictic sandstone rich in
clay and cemented by carbonates. The sandstone is rich in bioclast
derived from bivalves, gastropods, and echinoids, which are tightly
packed, well stratified, and in some cases, show a preferential
orientation resulted from the current direction. Other fossils found
in the sandstone include a nearly complete starfish, rudists, and
shark vertebrae. Higher in the section, fossiliferous beds of friable
fine sandstone and siltstone are intercalated with deposits of
mudstone and claystone while the top of the section contains
calcareous and biodetritic sandstone, interbedded with very fine
polymictic sandstone, siltstone, and claystone layers. Fossils of
pterosaurs, plants, and a new raninid crustacean (also known from
the Damuji outcrop), have been recovered from the polymictic
sandstone in the upper section at this location.
3.3. Puente Potrerillo
The deposit is localized 120 m west of the bridge that crosses the
Caunao river (N 22
170 12.93400 , W 80
90 19.62800 ), north of Potrerillo,
in the province of Cienfuegos in Central Cuba. The outcrop has a
thickness of approximately 1.5 m and extends over 20 m on the
south bank of the river (Fig. 2). It is dominated by dark beige and
brown mudstone and siltstones poorly stratified with some lenses
of very fine polymictic sandstone derived from the erosion of rocks
from the extinct volcanic arc. Fossils of plants, including seeds and
fragments of charcoal, were found in association with solitary
corals, echinoids, annelids (Rotularia sp., Pentaditrupa sp.), sca-
phopods (Dentalium sp.), bivalves (Inoceramus, ostreids, pinnids),
and osteichthyian fishes. Juveniles of Rotularia were found
cemented to fragments of echinoids while adults were free living.
Species in the genus Dentalium, Inoceramus, and Rotularia are
commonly considered bioindicators of soft-bottom, and the last
genus is found in environments where medium to high energy
predominate (Borja et al., 2000; Uppsala, 1995).
4. Systematic paleontology
Reptilia Laurenti, 1786
Archosauria, Cope, 1869
Pterosauria Kaup, 1834
Gen. and sp. Indet.
Figs. 3, 4
Referred material. Three tubular fragments of diaphysis of limb bones
MMR A-1, MMR A-2, MMR A-3 collected at Aguaditas and a fourth
diaphysis fragment MMR PD-4 recovered from Presa Damuji. Both
localities are from the Monos Formation (upper Campanianelower
Maastrichtian).
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al.
Fig. 2. Geologic map of central Cuba with the location of the three Upper Cretaceous outcrops of the Monos Formation from where specimens were recovered. Scale bar, 10 km.
Description. The specimens are fragments of diaphysis of limb bones
preserved in three dimensions, and only MMR A-4 is partially
collapsed (Fig. 3). The cortical bone is not completely preserved in
specimens MMR A-1 and MMR A-2, probably because of rework
and weathering. Surface coloration varies from amber (MMR A-1,
MMR A-2) to light brown (MMR A-3) and black (MMR PD-4). The
four specimens are characterized by having a large narrow cavity
and extremely thin cortical bone, which accounts only for 27% of
the transverse cross-section area. The cross-section areas possess
an oval to D shape with the following dimensions 22.4  14.4 mm
(MMR A-1), 19  10.5 mm (MMR A-2), 32  20 mm (MMR A-3) mm,
and 27  18 mm (MMR PD-4). No internal trabecular structure has
been noticed, but the visibility of the narrow cavity is limited by the
sediment filling it.
Paleohistology. The transverse sections of the two specimens
selected for paleohistological analysis (MMR: A-1, MMR A-2) are
composed of a wide central medullary cavity surrounded by a
narrow, compact cortex. Cancellous bone is absent from the med-
ullary cavity, and the cortical bone thickness varies between 0.72
and 1.41 mm. The compacta in MMR A-1 is formed almost entirely
by poorly vascularized primary bone tissue (Fig. 2A). Vascular
spaces are organized as primary osteons and longitudinally ori-
ented (Fig. 4B). Oblique canals are also present, but uncommon. At
least seven lines of arrested growth (LAGs) are recorded in the
compact tissue. Secondary remodeling is evidenced by the pres-
ence of a few Haversian osteons and resorption cavities scattered in
the inner half cortex (Fig. 4C). The primary bone matrix exhibits a
stratified pattern resulted from the spatial arrangement of the
intrinsic fibers (Fig. 4D).
4
Cretaceous Research 130 (2022) 105067
The intrinsic fibers of the primary bone matrix of MMR A-2 also
exhibit two main orientations, parallel and concentric to the main
axis of the shaft (Fig. 4E, F). The arrangement is, however, not as
ordered as in MMR A-1. In this regard, the intrinsic fibers are not
organized in distinct layers. Instead, patched with different fibrillar
orientation are mostly irregularly distributed in the compacta.
Vascular spaces are more abundant than those recorded in specimen
MMR A-1. These spaces are simple, and they are mostly longitudi-
nally arranged, occasionally showing anastomosis with oblique ca-
nals (Fig. 4E). Three possible lines of arrested growth are recorded in
the compacta, but they cannot be traced along the cortex.
The most noticeable histological feature of the sampled individuals is
related to the intrinsic fibers' spatial arrangement, which exhibits at
least two different orientations. This pattern, which is more pro-
nounced in the specimen MMR A-1, strongly resembles the plywood
organization described for other pterosaurs (De Ricqle s et al., 2000).
The plywood arrangement of the intrinsic fibers from the primary
bone matrix has been considered a diagnostic feature of Pterosauria
(De Ricqle s et al., 2000). The narrow band of birefringent bone
described in the specimen MMR A-1 corresponds to the circumfer-
ential lamellae described for some pterosaur long bones (Steel,
2008). The absence of an External Fundamental System in both
sampled bones suggests that the individual was not somatically
mature at the time of death (Chinsamy-Turan, 2005). At least three
possible LAGs in MMR A-2 and seven in MMR A-001 suggest a
minimum age of three and seven years old for each individual.
Given the extensive reduction of the cortical thickness by an
increased development of the medullary cavity (De Ricqle s et al.,
2000; Steel, 2008), a large part of the growth record was lost,
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al. Cretaceous Research 130 (2022) 105067

Fig. 3. Undetermined pterosaur bones recovered from Aguadita (A, B, D) and Presa Damuji (C) along their main axes and transverse section. (A) MMR A-2. (B) MMR A-1. (C) MMR
PD-4. (D) MMR A-3. Dashed lines indicate the sites from thin sections where obtained. Scale bars, AeD: 10 cm.

indicating the age is underestimated. Considering that the distri-
bution of histological characters among Pterosauria has not been
evaluated in detail and the specimens are too fragmentary, it is not
possible to assign them to a less inclusive clade.
Remarks. The analysis of bone microstructure gives unparalleled
insights into the developmental patterns hitherto unknown in
pterosaurs. For example, studies in Argentinian pterosaur shows
that upon hatching, Pterodaustro juveniles grew rapidly for about
two years until they reached approximately 53% of their mature
body size, at which point they attained sexual maturity. Thereafter,
growth continued for at least another 3e4 years at comparatively
slower rates until larger adult body sizes were attained. The bone
histology analysis of the ontogenetic series of the cretaceous
pterosaur Pterodaustro supports the earlier hypothesis that small
Jurassic pterodactyloid took several years to reach adult body size
(Bennett, 1996; Chinsamy et al., 2008, 2009). Considering previous
studies in other pterosaurs, and the observation of two diagnostic
microstructural characteristics (such as plywood organization and
circumferential lamellae) support that these fragments of ptero-
saurs from the Upper Cretaceous of Cuba belonged to long bones of
a subadult individual of Pterosauria. Given that the last pterosaurs
were mainly large animals, with wingspans typically around 4e5 m
(possibly reaching up to10 m or more), the new material could
represent at least a medium-sized pterosaur.
PLANTAE Haeckel, 1866
Fig. 5C-F
Referred materials. Seven fragments of trunks and branches (MMR
PD-18, MMR A-15, MMR A-16, MMR A-17, MMR A-18, MMR A-19,
MMR A-20) recovered from Aguaditas and Presa Damuji, the
specimens are housed at the MMR collection. However, there are
over a hundred specimens of different dimensions at the Aguaditas’
outcrop.
Description. Most of the samples were invaded by burrowing bi-
valves and their internal structure was lost, limiting their identifi-
cation. The wood tracheids' architecture and the lack of preserved
vessel elements in one of the specimens (MMR A-15) suggest it
belongs to gymnosperm incertae sedis.
PINALES Gorozh, 1904
? CUPRESSACEAE Gary, 1821
Gen. and sp. indet.
Fig. 5A-B
Referred materials. Four leafy-shoot casts (MMR PD-8, MMR PD-9,
MMR PD-10, MMR PD-13) collected at Presa Damuji. The speci-
mens are housed at the MMR collection.
Description. Four fragments of leafy shoot (MMR PD-8, MMR PD-9,
MMR PD-10, MMR PD-13) that bear spiral, to near opposite, scale-
like leaves (Fig. 3A, B). Two of the specimens (MMR PD-8, MMR PD-
10) are branched. The leaves are linear and about 2.5e6.0 cm long,
while the shoot fragments do not exceed 7.5 cm. Stomata and other
epidermal characters are not preserved on the specimens, but the
general arrangement in which the leaves occur is associated with
members of the family Cupressaceae (Schulz et al., 2005). Within
Cupressaceae, the fossils resemble closely the leaves and shoots of
the extant genera Widdringtonia, Glyptostrobus, Sequoia, and
Athrotaxis (Schulz et al., 2005). However, other families like Podo-
carpaceae and Araucariaceae occasionally present similar charac-
ters (Judd et al., 1999; Andruchow-Colombo et al., 2019).
LAURALES Juss. ex Bercht & Presl (1820)
LAURACEAE Juss., 1789, nom. Cons.

5
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Fig. 4. Bone microstructure of specimens MMR A-1 (AeD) and MMR A-2 (E, F). (A) General view of the cortical bone tissue. (B) Detailed view of the inner half portion of the cortex.
(C) Detailed view of the perimedullary cortex showing a single secondary osteon and some resorption cavities. (D) The primary bone matrix is formed by thin layers of

6
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al.
Gen. and sp. indet.
Fig. 6
Referred materials. Eight large fruit molds and one cast (MMR PD-6,
MMR PD-7, MMR PD-11, MMR PD-12, MMR PD-15, MMR PD-16,
MMR PD-17, MMR PD-18) collected at Presa Damuji and a fruit
mold (MMR PP-1) from Puente Potrerillo.
Description. The single berry or drupe fruits are relatively large and
preserved as molds (Fig. 4). The specimens are 4e6 cm long by
5e7 cm wide by 2e5.5 cm deep, have a globose, ovoid-elliptical
shape, wider than taller, and often compressed on one of their
sides; the fruits have a prominent fleshy, moderately shallow,
cupule holding them. At the base, where the fruit connects to the
cupule, there is a large circular scar, possibly the chalaza e where it
detached from the cupule. The apical region, opposite the scar, has a
prominent “dimple-like” depression, for either the hilum e the
foramen where the funicle entered the seed e or the adjacent
micropyle e the pollen tube opening. A fleshy outer layer (possible
pericarp) was present but not well recovered, seen only from traces
of the mold. The fruits are homogenous in shape and traits, mostly
affected by size and deformation from fall or diagenesis. The
presence of a cupule, along with the scar on the fruit and the
depression-foreman opposite (usually considered a hilum in Laur-
aceae), are prominent characteristics commonly associated with
Lauraceae fruits (Reid and Chandler, 1933; Little et al., 2009). The
fossil also shares characteristics (chalaza, hilum, globose) similar to
the Arecaceae, a family of monocots that has fossils present by the
Upper Cretaceous of North and South America (Harley 2006;
Manchester et al., 2010). The primary difference lies in the cupule,
where Arecaceae have persistent bracts at the base of the fruit; the
bracts of Arecaceae are small & thin, arranged in an imbricate
pattern The cupule.observed here more closely resembles that of
the Lauraceae - large fleshy/woody structures. The combination of
characters indicates that this fossil may belong to a new taxon.
Considering the large size of the fruit and shallow cupule, this taxon
more closely resembles the modern genus Chlorocardium (Rohwer
1991); though the cupule in the fossil appears far more enlarged.
5. Discussion
5.1. Pterosaur distribution in the Americas during the Late
Cretaceous
Late Cretaceous pterosaurs from the Cenomanian to the Maas-
trichtian, maintained a global distribution, but their diversity was
somewhat lower. The reduced diversity is partially a reflection of
the putative extinction of several clades by the end of the Cen-
omanian. It seems highly probable that a decrease in pterosaurian
diversity during the Late Cretaceous is a real phenomenon (Buttler
et al., 2013). As it turned out, Late Cretaceous pterosaurian faunas
are almost exclusively composed of members of the Pter-
anodontidae, Nyctosauridae, Azhdarchidae (Upchurch et al., 2015),
and the enigmatic Piksi barbarulna (Agnolin and Varricchio, 2012;
Longrich et al., 2018). At present, confirmed pteranodontids are
only known from the Upper Cretaceous of North America, although
one species also occurs in South America. The Nyctosauridae were
similarly restricted; it was a small clade of pterosaurs apparently
endemic to North America during the Coniacian and Santonian
(Frey et al., 2006), but later found in the Maastrichtian of South
America (Price, 1953). This distribution pattern for nyctosaurids
circumferentially oriented fibers, which alternate with thicker layers of longitudinally oriented fibers. General view (E) and detail (F) of the cortical bone tissue. Note the variable
arrangement of the intrinsic fibers evidenced by the optical properties of the matrix and the shape and arrangement of the osteocyte lacunae. Arrowheads indicate the position of
possible lines of arrested growth. (AeC) Plane-polarized light. A: cross-polarized light with lambda compensator; BeC: normal transmitted light. cf: circumferentially oriented
intrinsic fibers; lf: longitudinally oriented intrinsic fibers; mc: medullary cavity; ol: osteocyte lacunae; po: primary osteons; rc: resorption cavity; so: secondary osteons. Scale bars,
A: 0.7 mm; BeC: 0.1 mm; D and F: 0.2 mm; E: 0.5 mm.
7
Cretaceous Research 130 (2022) 105067
might reflect no more than the patchiness of the fossil record.
However, it is interesting to note that their apparent dispersal from
North America to South America during the Maastrichtian is
consistent with the faunal exchange hypothesis proposed
(Bonaparte and Kielan-Jaworowska, 1987; Gayet, 2001; Sullivan
and Lucas, 2000). On the other hand, although azhdarchids had a
low diversity they reached a global distribution during the latest
Cretaceous, occurring in Africa, Central, and East Asia, Europe, and
North America, and South America (Codorniú and Gianechini,
2016; David et al., 2018; Novas et al., 2012).
Although pterosaurs' records from the Upper Cretaceous in
South America are fragmentary, the number of finds has consid-
erably increased in recent years. Non-azhdarchid pterosaurs from
the latest Cretaceous are represented by Campanian pter-
anodontids; the pterodactyloid Nyctosaurus lamegoi (Price, 1953)
from the Maastrichtian of Brazil; and the tapejaride Caiuajara
dobruskii (Manzig et al., 2014) from the Campanian of Brazil, this
last one recovered from one of the few pterosaur bone beds
known so far. Notably, in Argentina, several Upper Cretaceous
pterosaur fossils from Patagonia are represented by fragmentary
bones assigned to Pterodactyloidea and ?Azhdarchoidea
(Codorniú and Gianechini, 2016). At the same time, two azh-
darchid pterodactyloid pterosaurs have been reported for Neu-
quen Basin in Argentina. This includes Aerotitan sudamericanus
(Novas et al., 2012) from the upper Campanianelower Maas-
trichtian Allen Formation, and a complete right humerus referred
to Azhdarchoidea from the upper Coniacianelower Santonian
Plottier Formation (David et al., 2018). These recent findings
further support the cosmopolitan distribution of azhdarchid
pterosaurs.
5.2. Biogeographical and paleogeographic implications
Understanding the diversity of the flora and fauna that inhabi-
ted the Caribbean archipelago during the Cretaceous not only in-
forms about the origin of the oldest members of the Antillean
terrestrial biota (Francisco-Ortega et al., 2007; Noonan et al., 2013;
Springer et al., 2018), but also sheds light into the provenance of
multiple lineages that flourished in the Americas for millions of
years, and in some cases still do. Remains of pterosaurs and plants
of the families Cupressaceae and Lauraceae described above doc-
uments the first shreds of evidence of the community that once
occupied the region in the upper Campanianelower Maastrichtian.
Fossils of Jurassic plants and two species of rhamphorhynchid
pterosaurs, Nesodactylus hesperius and Cacibupteryx caribensis, are
documented from Western Cuba. However, these species inhabited
the coast of Laurasia and are not related to the biota that occupied
the Caribbean later in the Cretaceous (Areces-Mallea, 1989, 1990;
Gasparini et al., 2004). Other shreds of evidence of dry land in the
Caribbean during the Mesozoic comes from several genera of plants
(including Gleichenites, Zamites, Phoenicopsis, Yuccites, Podozamites,
Frenelopsis, and Brachyphyllum) reported from the Lower Creta-
ceous Los Ranchos Formation in the Dominican Republic (Smiley,
2002). Whether some of these taxa survived until the Late Creta-
ceous in other parts of the archipelago is unknown. However,
sediments of Los Ranchos Formation are overlaid by Albian lime-
stone from the Rio Hatillo Formation (Iturralde-Vinent and
MacPhee, 1999), like other Cretaceous and Paleocene terrestrial
and fluvial deposits across the Antilles, indicating that those islands
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al. Cretaceous Research 130 (2022) 105067

8
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al. Cretaceous Research 130 (2022) 105067

Fig. 6. CT scans of large Chlorocardium-like fruits of the family Lauraceae from Presa Aguaditas (AeE), and photographic image of cupule of MMR PD-7. (A, F). Specimen showing the
typical shape and proportions; MMR PD-6, MMR PD-16. (B) Detail of apical region showing hilum; MMR PD-6. (C) Detail of basal region showing chalaza; MMR PD-6. (D) Second
specimen showing shape variation and more prominent hilum; MMR PD-15. (E) Specimen showing fruit with cupule and chalaza; MMR PD-7. (F) Cupule of MMR PD-7. C, chalaza;
Cu, cupule; h, hilum. 3D of the cupule of MMR PD-7 can be found at https://sketchfab.com/3d-models/f2ed5e3bda9e4456a533d6d7edcfd648. Scale bars, AeF: 5 cm.

and their tenants had an ephemeral existence (Iturralde-Vinent and
MacPhee, 1999; Iturralde-Vinent, 2006).
Paleogeographic reconstructions of the Caribbean plate during
the Late Cretaceous situates the island arch on the eastern leading
edge of the plane, bridging the gap between North and South
America (Iturralde-Vinent, 2006; Garcia-Casco et al., 2008). By the
upper Campanianelower Maastrichtian, the uplift following the
extinction of the vulcanism in part of the arc exposed large por-
tions of the archipelago, laying the ground conditions for the FABI.
Molecular and paleontological studies agree that this biotic
interchange was bidirectional but with a larger number of taxa
dispersing from North American into South America (Goin et al.,
2016). This event, preceding the better documented Great Amer-
ican Biotic Interchange at least by 50 million years (Gayet, 2001),
involved hadrosaurs, titanosaurs, pterosaurs, condylarth and
didelphimorph mammals, urocoptid snails, several groups of
plants, among others (Bonaparte, 1984; Chanderbali et al., 2001;
Uit De Weerd, 2008; Valieri et al., 2010). Current evidence suggests
that FABI spanned from the Campanian to the Early Paleocene and
was likely asynchronous and pulsed. The reduced number of lin-
eages involved in the inter-American journey may be the result of
our poor understanding of the diversity in the tropics during the
Late Cretaceous but also because these ephemeral Caribbean
islands acted as a filter. No evidence of a continuous land
connection between the archipelago and the continents are
known, which further support the idea that the islands were no
more than a series of steppingstones (Iturralde-Vinent and
MacPhee, 1999; Iturralde-Vinent, 2006; Goin et al., 2016). At the
same time, a land bridge would have allowed the flow of entire
communities between the Americas rather than a few selected
species. Also, it would have resulted in the disruption of water
circulation patterns between the Eastern Pacific and the Atlantic,
causing local extinctions, along with depositional and climatic
changes at a larger scale.
The discovery of not fully mature midsize pterosaur on the
Caribbean archipelago is not surprising given the capacity of this
group to disperse into new territories and their common asso-
ciation with insular deposits in the CampanianeMaastrichtian
(Dalla-Vecchia and Gau, 2015). The insular condition may have
favor gigantism among Maastrichtian pterosaurs (Dalla-Vecchia
and Gau, 2015), but smaller species are also known from
similar environments. The Caribbean islands sheltered ptero-
saurs (and potentially other groups) that disperse from the
mainland over water, providing them with resources and limited
competition. The origin of these island-dwelling pterosaurs and
their alleged relationship with the FABI is uncertain and requires
the discovery of further fossils. Nevertheless, the dispersion of
nyctosaurid in the Maastrichtian from the Mexican Gulf area into
Brazil may have been thru the Caribbean into the eastern shore
of South America.

Fig. 5. Fossil plants from the Upper Cretaceous of Central Cuba. (A) Leafy shoot of indeterminate genus and species of Cupressaceae showing scale-leaves; organization on the shoot;
MMR PD-9. (B) Leafy shoot of Cupressaceae with bifurcating shoots; MMR PD-8. (C) Fragment of wood with several perforations caused by Teredolites; MMR A-17. (D) Cross-section
of indeterminate fossil good invaded by Theredolites; MMR A-16. (E) Gymnosperm-like wood sample; MMR A-15. (F) Transverse section of gymnosperm-like wood showing the lack
of preserved; MMR A-15. Scale bars, A and E: 3 cm; BeD: 5 cm; F: 1 cm.

9
 ~ ola-Lo
L.W. Vin
pez, I.A. Cerda, J. Correa-Narvaez et al.
In contrast, fossil plants present a more reliable perspective on
the dispersion of species between the continents. The family
Cupressaceae can be traced back to the Jurassic of Patagonia,
Argentina (Escapa et al., 2008). By the Cretaceous, it was already
worldwide established with records from Asia (Shi et al., 2014),
Europe (Rothwell et al., 2011), North America (Atkinson et al.,
2014), and South America (Escapa et al., 2008). Currently, Cupres-
saceae still has a worldwide distribution and can be found in a wide
variety of habitats, including the West Indies (Judd et al., 1999).
Similarly, Lauraceae had a high diversity and wide distribution
range by the Late Cretaceous (Poole et al., 2000; Frumin et al., 2004)
comprising Gondwana (Poole et al., 2000), North America (Little
et al., 2009), and Asia (Frumin et al., 2004). In the present, the
family is predominantly found in tropical-subtropical latitudes of
southeast Asia and northern South America (Judd et al., 1999;
Chanderbali et al., 2001). The origin and evolution of some clades of
Lauraceae are believed to be tied with important geological events
like the break-up of Gondwana and the Caribbean's subaerial
exposition (Chanderbali et al., 2001). Specifically, the divergence of
the sister clades Perseeae-Laureae and Chlorocardium-Mezila is set
to have occurred in the middleeLate Cretaceous, when their an-
cestors migrated between the Americas through the precursor of
the Greater Antilles. The discovery of fossils of a new taxon closely
related to Chlorocardium from the Upper Cretaceous of Cuba is
consistent with this phylogenetic hypothesis and documents the
first direct evidence of the role the Caribbean played in the
dispersion of the group. If this is also true for other groups of ani-
mals and plants, as has been hypothesized in the past, we expect to
find their fossils on the islands. The interchange of species during
FABI had a strong repercussion in the biota of the continents,
inducing biodiversity changes that lasted well into the Mesozoic.
The arrival of condylarths and didelphimorphs to South America
resulted in the extinction of endemic non-tribosphenic and pre-
tribosphenic mammals, followed by the radiation of the formers
across the continent in what is known as the First Great Turnover
(Pascual, 2006; Ortiz-Jaureguizar and Pascual, 2011; Goin et al.,
2016). Snails of the family Cerionidae and several lineages of
plants, including some in the family Lauraceae, are among the
groups that successfully radiated and survived until the present
(Chanderbali et al., 2001; Uit De Weerd, 2008).
On the other hand, the instability of the Caribbean archipelago
until the Eocene, combined with the impact of the meteorite 66
million years ago relatively close to the islands, had dire conse-
quences for the biota on the region (Iturralde-Vinent and MacPhee,
1999). However, phylogenetic studies suggest that the ancestors of
some extant lineages in the West Indies may have arrived at the
archipelago in the Late Cretaceous and survived until the present
(Santiago-Valentin and Olmstead, 2004; Francisco-Ortega et al.,
2007; Noonan et al., 2013; Springer et al., 2018). A more compre-
hensive exploration of Upper Cretaceous and Paleogene deposits in
Cuba and other regions in the Circum-Caribbean may shed light on
some of these biogeographical disputes.
6. Conclusions
New fossils of vertebrates and plants from the Upper Cretaceous
of Cuba support the existence of subaerial islands during the late
Campanianeearly Maastrichtian in the gap between South and
North America. The specimens were recovered from three localities
with exposed deposits of the Monos Formation in Central Cuba and
were deposited in the distal part of fluvial systems under the in-
fluence of marine and terrestrial environments. The morphology of
fossil leaves collected at Presa Damuji indicates that they may
belong to the family Cupressaceae whereas large seeds collected at
Presa Damuji and Puente Potrerillo are identified as an undescribed
10
Cretaceous Research 130 (2022) 105067
taxon of Lauraceae, closely related with the extant genus Chlor-
ochardium from South America. On the other hand, the external
morphology and microanatomy of the vertebrate remains indicate
these belong to a midsize pterosaur, corresponding to the first
member of this group to be found in Cretaceous deposits in the
Antilles.
The existence of an archipelago in the eastern margin of the
Caribbean plate at the end of the Mesozoic coincides with the
estimated ages for the origin of several lineages of vertebrates and
plants in the Caribbean, and the biotic interchange between the
Americas. More importantly, fossils of the taxa closely related to
Chlorocardium are relevant because they represent the first direct
evidence of the role played by the Caribbean seaway and islands in
the FABI. Recent discoveries in the West Indies are changing our
current understanding of the biota that once occupied the region
millions of years ago and how it evolved throughout time. This
finding invites to the systematic exploration and study of deeps
time deposits across the West Indies.
Acknowledgments

Antonio Pin~ ero Castello

n and Jesús
Our sincere thanks to Jose
Servilio Quintero V
azquez for their assistance in the field. We are
also in debt to Dr. Zulma Gasparini for helping us to establish this
international collaboration and would like to extend our gratitude
also to Manuel Iturralde-Vinent, Richard Carr, Reinaldo Rojas, and
Johanset Orihuela for their comments and suggestions on the early
stages of the manuscript. The final version of the manuscript was
improved thanks to the revision of the editor Eduardo Koutsoukos
and an anonymous reviewer.
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