Geobios 48 (2015) 309–320

Available online at

ScienceDirect
www.sciencedirect.com

Original article

Historical relationship of the Caribbean and Amazonian Miocene

ichthyofaunas: A hypothesis reviewed under a biogeographical
approach§
Amparo Echeverry *, Valéria Gallo
Laboratório de Sistemática e Biogeografia, Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro (UERJ),
Pavilhão Haroldo Lisboa da Cunha, sala 228, Rua São Francisco Xavier 524, Maracanã, 20550-013 RJ, Brazil

A R T I C L E I N F O A B S T R A C T

Article history: South-American foreland basins are related to the compressional tectonics that built the Andes starting
Received 29 November 2014 in the Mesozoic; then the Neogene uplift of the Eastern Cordillera directly affected the sedimentary
Accepted 9 June 2015 history of Amazonia, changing the course of the proto-Amazon/Orinoco Rivers. Marine influence in South
Available online 26 June 2015
America has a long history with episodic transgressions and regressions from the Campanian to the Late
Miocene, with the Caribbean and/or the Pacific connections considered as the most likely sources.
Keywords: Previous studies on Neogene fish assemblages regarding historical relationships of the Caribbean and
Miocene
Amazonian regions based their conclusions on non-biogeographical methods. Here, we analyze the
Ichthyofauna
Palaeobiogeography
distribution of 241 Miocene fish taxa from 32 litostratigraphic units in northern South America and the
Caribbean Caribbean using a biogeographical method. We identify eleven distributional patterns and nine
Amazonian biogeographical nodes. Our results show a close relationship between the Caribbean and Amazonian
South America Miocene ichthyofaunas. Furthermore, they indicate that this relationship has to be separated into two
Pacific distinct components: western Amazonian/Caribbean, and eastern Amazonian/Caribbean. We hypothe-
size that patterns and nodes that involve the Pacific margin of northern South America or southern
Central America reflect pre-Andean (older) geographical configurations pointing to the eastern Pacific as
a relevant component in the history of diversification of Amazonian fish biota prior to and during the
Neogene.
ß 2015 Elsevier Masson SAS. All rights reserved.

1. Introduction
Neotropical fish diversity has an ancient history with some
higher-level endemic clades extending back into the Cretaceous.
Fossils show that some modern, generic-level clades had
differentiated by the Paleogene, and that the fish fauna was
essentially modern by the Late Miocene (Lundberg et al., 1998).
Although the majority of Neotropical fishes have a long history of
evolution dating back to the final rupture of the South American
and African continents (e.g., Siluriformes and Characiformes;
Lundberg et al., 1998, 2010), it is thought that some lineages
colonized South America much later, as a product of marine
incursions in the Amazon during the Oligocene and Miocene
(Hoorn, 1993; Webb, 1995; Lovejoy et al., 1998, 2006; Lundberg
§
Corresponding editor: Gilles Escarguel.
* Corresponding author.
E-mail address: echeverryceae@gmail.com (A. Echeverry).
et al., 1998; Monsch, 1998; Boeger and Kritsky, 2003; Wesselingh
and Macsotay, 2006; Cooke et al., 2012).
Marine influence in South America has a long history with
episodic transgressions and regressions from the Campanian up to
the Late Miocene (Lundberg et al., 1998; Brito et al., 2007). The
Western Amazon was affected by marine incursions at least twice
during the Neogene (Hoorn, 1993, 1994a,b, 1996). The extent and
timing of Neogene marine incursions in Amazonia is currently
controversial. Some authors have suggested that marine incursions
were long lasting and widespread (Räsänen et al., 1995, 1998;
Webb, 1995; Gingras et al., 2002). Others have suggested that the
predominant depositional context of Miocene Amazonia was
freshwater, interspersed with brief and geographically limited
bouts of marine influence (Hoorn, 1996; Hoorn et al., 2010; Vonhof
et al., 1998, 2003; Wesselingh et al., 2002; Linhares et al., 2011).
Gross et al. (2013) proposed a fluvial depositional system for
Western Amazonia during the Late Miocene. Latrubesse et al.
(2007) refuted the existence of an intracontinental seaway through
southwestern Amazonia during the Late Miocene (11–10 Ma),

http://dx.doi.org/10.1016/j.geobios.2015.06.001
0016-6995/ß 2015 Elsevier Masson SAS. All rights reserved.
310 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320
connecting the Caribbean Sea with the Paraná Basin (Räsänen et al.,
1995). Instead, they held that the depositional environment of the
entire Late Miocene was continental, and that the deposits in
Peruvian Amazonia attributed to a marine Miocene tidal environ-
ment are fluvial sediments (Latrubesse et al., 2010).
The Caribbean is usually considered the most likely gateway of
the Miocene marine influences and incursions into the Llanos Basin
(Bayona et al., 2008; Gómez et al., 2009) and upper Amazon (Hoorn
et al., 1995; Wesselingh et al., 2002; Vermeij and Wesselingh,
2002), but Nuttall (1990) also considered three other possible
sources: an eastern connection along the course of the current
Amazon river, a southern connection via the Paraná river basin,
and a western connection to the Pacific. According to the summary
of Neogene palaeogeography presented by Nuttall (1990), and
partially from his own palaeontological evidence, the Caribbean
and/or the Pacific connections was thought the most likely, the last
one thought to have lain in the Peruvian-Ecuadorian border area.
Non-Caribbean marine incursions during the Miocene were
considered unlikely by Hoorn et al. (1995) and Lundberg et al.
(1998) for a variety of reasons, but Lovejoy et al. (2006) noted the
presence of marine sister taxa in the alternative marine incursion
areas.
The sedimentary basins of the Amazonia are fundamental
elements that dictate the present landscape of the region. The
geological evolution of the basins established the topography and
the geomorphology of the rivers, lakes and marine systems that
filled the depressions over the last 500 million years. The extensive
continental lowlands of Amazonia interacted dramatically with
the uplift of the Andes during the last 23.5 million years (Neogene
times), creating a special and unique region in terms of drainage,
fauna and flora (Ribeiro Wanderley-Filho et al., 2010).
The history of the sub-andean foreland basins in particular is of
major relevance in understanding how past geographical config-
urations shaped the fish biotas in western Amazonia. Foreland
basins are related to the compressional tectonics that built the
Andes, which in turn were initiated by the collision of the
Caribbean Oceanic Plateau with the Northern Andean margin
(75–65 Ma) and break-up of the Farallon Plate during the
Oligocene (27–25 Ma) (Martin-Gombojav and Winkler, 2008).
Although tectonic pulses in the Andes are documented from
Mesozoic time onwards, it is mainly the Late Paleogene and
Neogene uplift of the Eastern Cordillera and Cordillera de Merida,
the Cordillera Real, and Nazca Ridge subduction that directly
affected the sedimentary history of Amazonia (Hoorn et al., 2010
and references therein). Foreland basins extend from Argentina to
Venezuela along the eastern side of the Andes.
The history of the Amazonian drainage basin shows a
geochronological progression that proceeds from west to east.
During the Late Cretaceous, western Amazonia was a backarc basin
in a predominantly marine setting, with a discontinuous proto-
Andean island arc located to the west (Wesselingh et al., 2010).
From the Late Cretaceous to the Late Miocene, the history of the
basin was closely linked to the evolution of the Andes. Foreland
basins formed east of the Andes, where the south-north trending
fluvial system was located during Middle Eocene-Early Oligocene
(43–34 Ma). During the Oligocene (34–24 Ma), the major drainage
divided between west and east flowing Amazonian drainages
system shifted from Monte Alegre in eastern Amazonia to the Purus
Arch in central Amazonia. By the Early-Middle Miocene
(24–11 Ma), Amazonia was mostly part of the south-north trending
river system that became a complex of lakes, swamps, tidal
channels and marginal marine embayments that was directed
towards the Caribbean. Finally during the Late Miocene, (11–7 Ma),
the transcontinental drainage of the Amazon River was established
and the first Andean sediment influx was registered in the Amazon
Foz basin and the Ceara Rise (Figueiredo et al., 2009).
Comparative studies on Neotropical Miocene marine ichthyo-
faunas remain scarce and descriptive (Longbottom, 1979; Gillette,
1984). Some studies on Neogene fish assemblages and the
palaeobiogeography of the Caribbean and Amazonian regions
based their conclusions regarding historical relationships on
Principal Component Analyses (Aguilera et al., 2011; Aguilera
and Páes, 2012). Most of the studies involving the Miocene fossil
record provided taxonomic descriptions of new fossil taxa (fish or
other groups), sometimes with brief discussions about taxonomic
and geographical affinities (Deynat and Brito, 1994; Laurito, 1996;
Távora and Garrafielo Fernandes, 1999; Aguilera and Rodrigues de
Aguilera, 2001, 2004a,b; Brito and Deynat, 2004; Laurito, 2004;
Fernandes dos Reis, 2005; Aguilera et al., 2008; Laurito and Valerio,
2008; Laurito et al., 2008; Ferreira Costa et al., 2009; Aguilera et al.,
2013a,b; Pimiento et al., 2013). However, none of the previous
studies have used a biogeographical method to analyze the
distribution of the fossil taxa involved.
Historical biogeography analyzes patterns of species and
supraspecific taxa at large spatial and temporal scales; it deals
mainly with processes that occur over long periods of time
(Morrone, 2009). These processes are responsible for the historical
pathways of changes in biogeographical distributions. The main
factors involve changes in the Earth’s surface: creation or
destruction of seas and seaways on cratonic areas, plate move-
ments and realignments, creation of mountain ranges, and
oceanographic and climatic changes (Cecca, 2002).
Biogeographical studies using only fossil taxa are rare (Fielitz,
1996; Lieberman, 2003; Cecca et al., 2005; Gallo et al., 2007).
Studies adopting a biostratigraphic approach only allude to the
palaeobiogeography of taxa over a short interval of time and often
overlook historical relationships among taxa and areas (Silva and
Gallo, 2007). On the other hand, integration of fossil and living taxa
in the same analysis has been successfully undertaken (Heads,
1989, 2006, 2012; Candela and Morrone, 2003).
Interpretations of historical biogeographical patterns stress the
steps of biological evolution in its spatial dimension. This allows
the study of phyletic relationships relative to the evolution of their
geographic distributions, as in cladistic biogeography, for example,
but also means that geological work can exploit biogeographical
data for palaeogeographic reconstructions (Cecca, 2002). Through
a biogeographical approach, we aim to test the hypothesis of a
historical relationship between the Caribbean and Amazonian
Miocene ichthyofaunas.
2. Material and methods
2.1. Units of analysis
Fifty-seven sedimentary basins were used as units of analysis
(Fig. 1). They contain the 32 Miocene formations from which
distributional fossil records were taken. Sedimentary basins
constitute historical entities that have developed under particular
tectonic regimes that account for their origin, evolution and
depositional history through geological time. Fluvio-lacustrine
hydrographic systems have been determined by changing
mountain slopes and foreland basin geometry. Table 1 lists the
sedimentary basins in northern South America and the Caribbean
by type, country, and Miocene formation(s) retrieved in each basin.
2.2. Taxa
The palaeo-ichthyofauna studied includes 241 taxa of fishes:
62 species of Chondrichthyes (31 genera and 16 families), and
59 species of Osteichthyes (82 genera and 48 families). In several
cases, specimens were identified only to the family or subfamily level
(e.g., Potamotrygonidae, Gobiidae, Loricariidae, Pseudopimelodidae,
 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320 311

Fig. 1. Sedimentary basins as units of analysis. 1. Middle Magdalena; 2. Upper Magdalena; 3. Cesar-Racheria; 4. Lower Magdalena; 5. Sinú-San Jacinto; 6. Chocó; 7. Tumaco;
8. Guajira; 9. Eastern Cordillera; 10. Llanos; 11. Caguán-Putumayo; 12. Vaupés-Amazonas; 13. Chucunaque; 14. Panamanian Canal; 15. Panamanian Gulf; 16. Bocas del Toro;
17. Southern Limon; 18. Northern Limon; 19. Tempisque; 20. Parrita; 21. Térraba; 22. Mosquitia; 23. Jamaica; 24. Northern Cuba; 25. Western Cuba; 26. Central Cuba;
27. Eastern Cuba; 28. Enriquillo; 29. Azua; 30. Puerto Rico; 31. Grenada; 32. Trinidad & Tobago; 33. Barbados; 34. Cariaco; 35. Oriental; 36. Barinas-Apure; 37. Maracaibo-
Falcón; 38. Borbón; 39. Progreso; 40. Manabi; 41. Oriente; 42. Marañon; 43. Talara; 44. Huallaga; 45. Ene; 46. Ucayali; 47. Pisco; 48. Madre de Dios; 49. Acre; 50. Solimões;
51. Amazonas; 52. Tapajós; 53. Bragança-Viseu-Ilha Nova; 54. Pará-Maranhão; 55. Marajo; 56. Amazon Foz; 57. Guyana-Suriname. Asterisks (*) indicates the presence of
Miocene formations.

Anostomidae, Serrasalmidae, and Tetragonopterinae). Distributional
records of Miocene fossil fishes were taken from published studies
(Appendix B).
2.3. Analysis
Parsimony analysis of endemicity (PAE) was used to identify
distributional patterns or generalized tracks (Craw et al., 1999;
Morrone, 2009, 2014). PAE was proposed by Rosen and Smith
(1988) in a paleontological context. These authors analyzed the
distributions of taxa from sampling localities in different geologi-
cal horizons. PAE became a useful tool in historical biogeography,
because it allows the analysis of historical information (spatial
distribution of taxa) when phylogenetic analyses are lacking. In its
panbiogeographical implementation, PAE highlights the spatial
information by using individual tracks (instead of single sample
localities) as the basis for the construction of the data matrix from
which generalized tracks are identified (Echeverry and Morrone,
2010).
Individual tracks are line graphs drawn on a map that connect
the different localities where a given taxon is observed according
to their geographical proximity. In the present study, individual
tracks for each taxon were constructed using TRAZOS (Rojas-Parra,
2007) as an extension of ARCVIEW GIS 3.2. Generalized tracks
result from the spatial congruence of two or more individual tracks
and represent ancestral biotic components that became fragmen-
ted by geological or tectonic events. In order to identify generalized
tracks (distributional patterns), a matrix r  c was constructed,
where the rows (r) represent sedimentary basins and the columns
(c) represent individual tracks (taxa). Each matrix entry is ‘‘1’’
when an individual track is present in, or crosses, a sedimentary
basin, and ‘‘0’’ if it is absent or does not cross it. A hypothetical area
coded ‘‘0’’ for all the columns is included to root the resulting
cladogram(s). The matrix of 241 individual tracks by 57 sedimen-
tary basins (Appendix B) was analyzed using NONA (Goloboff,
1998) through WINCLADA 1.00.08 (Nixon, 1999), with the
heuristic search option (tree bisection and reconnection, 100 repli-
cations). Clades supported by at least two individual tracks were
mapped as a generalized track (distributional pattern). Last, nodes
were identified manually at sites where different generalized
tracks intersected. Biogeographic nodes represent tectonic and
biotic convergence zones, areas of ancient geography around
which evolution has taken place (Heads, 2004).
Through the identification of distributional patterns and
biogeographic nodes, we expect to test the hypothesis of a
historical relationship between the Caribbean and Amazonian
Miocene ichthyofaunas. If the distributional patterns obtained are
supported by taxa found in both areas (connecting the Caribbean
and Amazonian Miocene ichthyofaunas), it would mean that our
results are inconsistent with our null hypothesis that ‘‘there is not a
historical relationship between the Caribbean and Amazonian
Miocene ichthyofaunas’’, in which case we consider our null
hypothesis to be falsified and the alternative hypothesis con-
firmed.
It is worth mentioning that all the deep-time distributions
(individual and generalized tracks) were plotted on the modern
Earth’s map. The rationale behind this choice is stated clearly by
Rosen (1988: 46): ‘‘Since there is always uncertainty about the
312 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320

Table 1
Sedimentary basins by type, country, references and Miocene formations.

Basin type Basin name Country Reference Miocene formation(s)

Forearc Borbón (38) Ecuador Deniaud et al., 1999 Borbón-Onzole Fm.

Hasui, 2012 Jama Fm.
Manabi (40) Ecuador Deniaud et al., 1999 Loyola/Guapán/Mangán
Hasui, 2012 Fm. (Cuenca Fauna)
Progreso (39) Ecuador Higley, 2004 __________
Hasui, 2012
Talara (43) Peru Klein et al., 2011 Miramar Fm.
Hasui, 2012
Pisco (47) Peru Klein et al., 2011 Pisco Fm.
Hasui, 2012 Chilcatay Fm.
Térraba (21) Costa Rica Astorga et al., 1991 Curré Fm.
Escalante & Astorga, 1992
Parrita (20) Costa Rica Astorga et al., 1991 Punta Judas Fm.
Escalante & Astorga, 1992
Tempisque (19) Costa Rica Astorga et al., 1991 Punta Carballo Fm.
Escalante & Astorga, 1992
Tumaco (7) Colombia Barrero et al., 2007 __________
Chocó (6) Colombia Barrero et al., 2007 __________

Foreland Madre de Dios (48) Brazil Zalán, 2004 Solimões/Pebas Fm.

Peru Klein et al., 2011 (Rio Acre Fauna)
Bolivia Hasui, 2012
Acre (49) Brazil da Cruz Cunha, 2007 __________
Hasui, 2012
Marañón (42) Peru Klein et al., 2011 Solimões/Pebas Fm.
Hasui, 2012
Ucayali (46) Peru Klein et al., 2011 Solimões/Pebas Fm.
Hasui, 2012 (Utuquina Fauna)
Oriente (41) Ecuador Baby et al., 1999 __________
Caguán-Putumayo (11) Colombia Barrero et al., 2007 __________
Cesar-Rancheria (3) Colombia Barrero et al., 2007 __________
Llanos (10) Colombia Barrero et al., 2007 __________
Barinas-Apure (36) Venezuela Hasui, 2012 Cantaure Fm.
Maracaibo-Falcón (37) Venezuela Hasui, 2012 Castillo Fm.
Urumaco Fm.
Oriental (35) Venezuela Hasui, 2012 __________
Panamanian Gulf (15) Panama Montes et al., 2011 __________

Passive margin with inversión Western Cuba (25)a Cuba AAPG Datapages, 2014 Güines Fm.
Lagunitas Fm.
Central Cuba (26)a Cuba AAPG Datapages, 2014 __________
Eastern Cuba (27)a Cuba AAPG Datapages, 2014 __________
Mosquitia (22) Honduras/Nicaragua AAPG Datapages, 2014 __________
Jamaica (23) Jamaica AAPG Datapages, 2014 Montpelier Fm.
Enriquillo (28) Hispaniola AAPG Datapages, 2014 __________
Azua (29) Hispaniola AAPG Datapages, 2014 Gurabo Fm.
Cercado Fm.
Puerto Rico (30) Puerto Rico AAPG Datapages, 2014 Aymamon Fm.

Intracratonic Solimões (50) Brazil Ribeiro Wanderley- __________

Filho et al., 2007
Tapajós (52) Brazil Zalán, 2004 __________
Amazonas (51) Brazil da Cruz Cunha et al., 2007 __________
(?) Vaupés-Amazonas (12) Colombia Barrero et al., 2007 __________

Backarc Bocas del Toro (16) Panama Escalante & Astorga, 1992 __________
Southern Limon (17) Costa Rica Astorga et al., 1991 Uscari Fm.
Escalante & Astorga, 1992 Banano Fm.
Northern Limon (18) Costa Rica Astorga et al., 1991 __________
Escalante & Astorga, 1992

Retro-arc foreland Panama Canal (14) Panama AAPG Datapages, 2014 Gatún Fm.
(ocean-ocean)
Culebra Fm.
Barbados (33) Barbados Bisex Hill Fm.
AAPG Datapages, 2014 Oceanique Fm.

(ocean-continent) Northern Cuba (24)a Cuba AAPG Datapages, 2014 Husillo/Cojimar Fm.

Aborted rift Bragança-Viseu- Brazil Zalán, 2007 Pirabas Fm.

Ilha Nova (53)
Marajó (55) Brazil Zalán & Matsuda, 2007 __________

Poly-historic, rift to broken Upper Magdalena (2) Colombia Barrero et al., 2007 La Victoria/Villavieja Fm.
foreland
(La Venta Fauna)
Middle Magdalena (1) Colombia Barrero et al., 2007 __________
 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320 313

Table 1 (Continued )

Basin type Basin name Country Reference Miocene formation(s)

Intermontane Huallaga (44) Peru Klein et al., 2011 __________

Ene (45) Peru Klein et al., 2011 __________

Transform margin Amazon Foz (56) Brazil Picanço de Figueiredo et al., 2007 Pirabas Fm.
Pará-Maranhão (54) Brazil Fernandes Soares et al., 2007 __________

Trans Lower Magdalena (4) Colombia Barrero et al., 2007 __________

tensional
Guajira (8) Colombia Barrero et al., 2007 Castilletes Fm.

Retro-arc post-extensional sag Cariaco (34) Venezuela AAPG Datapages, 2014 Cubagua Fm.
(ocean-continent)
Trinidad & Tobago (32) Trinidad & Tobago AAPG Datapages, 2014 Tamana Fm.
Springvale Fm.

Inverted graben/fold belt Eastern Cordillera (9) Colombia Barrero et al., 2007 __________

Accretionary prism Sinú-San Jacinto (5) Colombia Barrero et al., 2007 __________

Inverted intra-arc Chucunaque (13) Panama Montes et al., 2007 __________

Retro-arc extensional Grenada (31) Grenada AAPG Datapages, 2014 __________

(ocean-continent)

Passive margin Guyana-Suriname (57) Guyana/Suriname AAPG Datapages, 2014 __________

a
These units do not represent sedimentary basins but general geological divisions.

accuracy of a palaeogeographic reconstruction, there is a beguiling
objective merit in using the modern configuration of the Earth as a
biogeographical frame of reference, even for fossil distributions. In
fact, the logical conclusion of this argument is initially to disregard
the geographical frame of reference altogether and handle
distributional data simply in terms of disconnected, contempora-
neous sample points. It is a common practice however to plot past
distributions on palaeogeographic reconstructions and then look
for empirical evidence of geological cause and distributional effect.
From an applied point of view, this raises the difficulty that unless
this is done in the particular context of a test or comparison, it
prejudges (or obscures) any palaeogeographical conclusions that
might be drawn from the distributions’’.
Patterns identified by panbiogeographical analyses represent
pre-existing ancestral biotic components that became fragmented
by geological or tectonic events, independently of what time plane
(past or present) distributional records are taken from. It follows
that the use of a panbiogeographical approach along with the use
of palaeomaps would be redundant and lead to circular reasoning.
In the words of Cecca (2002: 9): ‘‘It must be stressed that if the use
of biotic distributions is aimed at palaeogeographical reconstruc-
tion, the plotting of fossil distributions on palaeogeographical
maps may entail circular reasoning reflecting the hypothetical
character of palaeogeography’’.
3. Results
The cladistic analysis produced 190 equally most parsimonious
cladograms with 552 steps, a consistency index of 0.30 and a
retention index of 0.58. The strict consensus cladogram had
613 steps, a consistency index of 0.27 and a retention index of
0.52. Seven generalized tracks (3 of them nested) were identified
by the parsimony analysis (Figs. 2 and 3(a, b): T1, T2, T3 and T4).
Additionally, another six generalized tracks were identified
manually on the map (Fig. 3(a, b): T5, T6, T8, T9, T10 and T11).
In the case of these tracks, the parsimony analysis showed
perfectly congruent individual tracks as homoplasies (instead of
synapomorphies). Their spatial distributional congruence was
confirmed on the maps and, based on this evidence, we retrieved
those patterns that otherwise would have been overlooked.
Fig. 3(c) summarizes the nine biogeographical nodes identified
and the generalized tracks involved. Table 2 lists the supportive
taxa (individual tracks) of each generalized track, as well as their
habitat (inferred from geology/fossil fauna and extant relatives)
and fossil age range.
4. Discussion
4.1. Distributional patterns, biogeographic nodes, and
palaeogeography
Wesselingh et al. (2002) and Hoorn et al. (2010) document the
different phases of a long-standing (24–7 Ma) mega-wetland in the
upper Amazon (often referred to as ‘‘Lake Pebas’’ or the ‘‘Pebas
System’’). This consisted of fluvial systems that originated on the
Amazonian Craton and were directed towards the sub-Andean
zone and the Caribbean via the Llanos basin. This connection
existed until the Late Miocene. The central part of the mega-
wetland was situated in Western Amazonia. Typical exposures of
the sediments of upper Pebas and Solimões formations (Middle to
early Late Miocene) are found along the Amazon River between
Nauta (Peru) and Leticia (Colombia), along the Peruvian Napo
River, and the Javari River in Brazil. Other outcrops are known from
the Putumayo and Caquetá rivers in Colombia. Isolated outcrops
are reported from as far south as Shepahua (Ucayali, Peru) and as
far north as the Serranı́a de la Macarena (Colombia). During the
Middle Miocene, the wetland system reached its maximum
development (an area of over one million km2) and formed a
complex of lakes, embayments, swamps, rivers and fluvio-tidal
environments (as the Pebas Basin described by Nuttall, 1990).
Along its periphery the system was bordered by the alluvial plains
of Andean rivers in the west, and lowland rivers on the craton to
the east, whereas in the northeast, the wetland extended onto the
cratonic rocks (Wesselingh et al., 2006). Most of the distributional
patterns and nodes identified in this paper seem to conform to the
general palaeogeographic scenario of the mega-wetland docu-
mented by Hoorn et al. (2010). We discuss them along with other
broader patterns found in the neighborhood areas.
Nodes 7 and 5 (N7, N5; Fig. 3(c)) are located in areas where the
Pebas System was located during its phase of major expansion
(Middle Miocene). The four generalized tracks that intersect in N7
(T6, T4, T5 and T9) are briefly described. Track 6 (Table 2) is a
pattern supported by an extinct freshwater species of Scianidae
(yXenotolithus sasakii), a recently described extinct species of
Ariidae (yCantarius nolfi) (Aguilera et al., 2013a), and other taxa
reported by Nelson (2006) as: marine but rarely occurring in
314 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320
Fig. 2. Strict consensus cladogram. Marks on nodes indicate clades supported by
two or more individual tracks and represent generalized tracks (T1, T2, T3, T4). T2a,
T3a and T4a are smaller generalized tracks nested within T2, T3 and T4,
respectively.
freshwater and ascending rivers (Pristidae); marine, brackish and
freshwater (Dasyatidae and Sciaenidae), and mainly marine but
often in fresh or brackish water (Ariidae). The genus Pachypops
(Sciaenidae) enters estuaries in South America. Pachypops fourcroi
is one of the taxa that support T6. The material identified by
Monsch (1998: 43) as Arius sp. and Ariidae indet. corresponds to
what Aguilera et al. (2013a) described as yC. nolfi. Based on this, the
fossil record of this species expanded to the Solimões/Pebas
formations at the boundaries of Brazil, Peru and Colombia
(Solimões, Marañón and Amazonas basins, respectively). Accord-
ingly, their habitats inferred from geology and fossil fauna are
freshwater (Solimões/Pebas formation) and marine (Cantaure,
Castillo and Castilletes formations) (Lundberg et al., 2010; Aguilera
et al., 2013a). Track 4 describes a pattern in which all the taxa
involved (including the freshwater stingrays Potamotrygonidae)
are freshwater groups. Most of the genera involved in pattern T5
have a fossil record that extends as far back as the Cretaceous in the
American continent (Table 2). Track T9 is a pattern supported by
freshwater taxa. Modern Leporinus species are presently wide-
spread from the Magdalena River through the Orinoco, Guianas
and Amazon to the Rio de La Plata (Lundberg et al., 2010).
The intersection of patterns T10 and T4 forms node 5 (N5). The
area where this node is located corresponds to the upper
Magdalena basin. Taxa supporting T10 are all either freshwater,
marine/freshwater (Carcharhinus aff. falciformis) or marine/brack-
ish taxa (Sphyrna cf. lewini). The pattern includes some of the fossil
taxa of the Middle Miocene Honda Group (La Venta fauna from
Colombia). This fossil fauna includes fluvial and lacustrine deposits
proximal to the Central Cordillera in the west, and connected at
that time in the east with the Pebas Formation deposits. According
to Lundberg et al. (1998), this unit includes several fish species that
at present occur in the Magdalena, Orinoco and Amazonas
hydrographic basins, suggestive of an earlier connection between
these areas. The orogeny of the Eastern Cordillera of Colombia (12–
11 Ma; Hoorn et al., 1995; Guerrero, 1997) isolated the Magdalena
drainage basin. The extant Arapaima gigas occurs throughout the
Amazon lowlands and in Guyana, but Arapaima (part of track T4) is
now absent from the Magdalena region. Track T10 highlights the
overall resemblance of the vertebrate faunas of La Venta (Upper
Magdalena Basin) and Urumaco (Maracaibo-Falcón Basin in
Venezuela).
Nodes 3 and 4 (N3, N4; Fig. 3(c)), while spatially related to the
Pebas system, can be associated with two events later in time: the
orogeny of the Merida Andes that isolated the Maracaibo Basin
(10–8 Ma; Duque-Caro, 1990; Diaz de Gamero, 1996; Colletta et al.,
1997; Lundberg, 1998) in the case of N3, and the shift towards the
East Venezuela Basin of the lowland corridors between the Llanos
Basin and the Caribbean in the case of N4 (Wesselingh et al., 2006).
Mora et al. (2010) consider the Vaupés Swell (Arch) to be a key
element preventing the northern Amazon tributaries from flowing
towards the Caribbean (as previously documented by Hoorn et al.,
1995), and playing a role in directing the Amazon drainage basin
towards the Atlantic. Activity on the Vaupés Swell is Late Miocene
in age, although the area was emergent since the Late Cenomanian.
The Swell is a western, low-elevation structural high where the
Guiana Shield is exposed or overlapped by a thinner sedimentary
cover. The Macarena, Chiribiquete and Araracuara low-elevation
ranges represent the surface expression of the Vaupés Swell, where
the Guiana Shield basement rocks rise up to 1000 m a.s.l.
Node N3 represents the intersection of tracks T6, T10 and T2.
The pattern described by T2 resembles the mollusk-based area
erected by Petuch (1988) as the ‘‘Puntagavilanian Subprovince’’,
which corresponds roughly to Woodring’s (1974) ‘‘Colombian-
Venezuelan-Trinidad Subprovince’’. Based on Pliocene gastropod
data, Landau et al. (2008) recognized the same area along the
northern coasts of South America as having highly endemic taxa. In
addition to the areas described by the preceding authors, our track
includes the Central American isthmian region to the west (Costa
Rica and Panama), and it does not include Trinidad Island. Based on
a multivariate analysis of fossil fish genera, Aguilera et al. (2011)
described a similar pattern that they called ‘‘The Venezuelan
subprovince’’, also described by Robertson and Cramer (2014) as a
southern province (northern South America).
The Pirabas Formation in northeastern Brazil is involved in two
different patterns. One extends to northeastern Venezuela via the
western Atlantic (track T3); the other is a transcontinental track
extending towards the Pacific and to Panama (track T8). Track T3
and partially track T8 are nested in a larger pattern that connects
northeastern Brazil (Pirabas) with the Caribbean (Greater Antilles)
 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320 315

Fig. 3. Generalized tracks and biogeographical nodes identified. a: T1, T2, T4, T6, T10; b: T3, T5, T8, T9, T11; c: summary of generalized tracks and nodes.
316 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320

Table 2
Generalized tracks (T1–T11), supportive taxa (individual tracks), habitat and fossil age range.

Generalized tracks/supportive taxa Geology/fossil faunaa Extant relatives Fossil age range (Ma)b

T1
Carcharodon carcharias Marine/Brackish 23.03–0.012
yGaleocerdo aduncus Marine/Brackish 33.9–4.9
Negaprion aff. brevirostris Marine/Brackish 23.03–0.012
Negaprion Marine/Brackish 40.4–0.012
Aetobatus Marine/Brackish 55.8–0.012

T2
Galeocerdo cuvier Marine/Freshwater Marine/Fresh/Brackish 5.332–0.012
yAetobatus arcuatus Marine/Freshwater Marine 20.43–7.246
Equetus Marine/Freshwater Marine 23.03–3.6
Cynoscion Marine/Freshwater Marine/Fresh/Brackish 11.608–0.012
Bairdiella Marine/Fresh/Brackish 11.608–0.012
Paraconger Marine 48.6–3.6
Anchoa Marine/Fresh/Brackish 11.608–3.6
Cetengraulis Marine/Fresh/Brackish 11.608–7.246
Engraulis Marine/Brackish 37.2–2.588

T3
Sphyraena Marine Marine/Brackish 55.8–0.012
Sphyrna Marine/Brackish 66.043–0.012
Diodon Marine 55.8–0.012
Arius Marine/Freshwater Marine/Freshwater 66.043–0.012

T4
Potamotrygonidae Freshwater Freshwater 13.65–7.246
Arapaima Freshwater Freshwater
cf. Hoplosternum Freshwater Freshwater 9.0–7.246
Colossoma macropomum Freshwater Freshwater
Serrasalmus/Pygocentrus/ Freshwater Freshwater
Pristobrycon Freshwater Freshwater 70.6–48.6
Lepidosiren paradoxa Freshwater Freshwater
Loricariidae Freshwater Freshwater
Anostomidae Freshwater Freshwater

T5
yIsurus hastalis Marine 58.7–0.781
Isurus cf. oxyrinchus Marine 20.43–0.781
Carcharias taurus Marine 28.4–0.012
Sphyrna cf. media Marine 28.4–3.6
Heterodontus Marine 242.0–3.6
Pristiophorus Marine 85.8–1.806
Plagioscion Freshwater Freshwater 23.03–15.97

T6
cf. Pristis Freshwater Marine/Freshwater 161.2–0.3
cf. Dasyatis Freshwater Marine/Freshwater 136.4–0.012
Pachypops fourcroi Freshwater Freshwater
yXenotolithus sasakii Freshwater ?
yCantarius nolfi Marine/Freshwater Marine/Freshwater

T7
yCarcharodon megalodon Marine/Freshwater Marine 28.4–0.781
yHemipristis serra Marine/Freshwater Marine 55.8–0.3
Myliobatis Marine/Freshwater Marine 84.9–0.012

T8
yCarcharhinus priscus Marine/Freshwater 20.43–2.588
Carcharhinus Marine/Freshwater Marine/Freshwater 55.8–0.012
yScoliodon taxandriae

T9
cf. Leporinus Freshwater Freshwater
cf.Tetragonopterinae Freshwater Freshwater
Hoplias Freshwater Freshwater 66.043–61.7

T10
Carcharhinus aff. falciformis Freshwater/Marine Freshwater/Marine 23.03–0.0
Sphyrna cf. lewini Marine/Brackish 15.97–3.6
Freshwater
Brachyplatystoma cf. vaillantii Freshwater Freshwater 15.97–11.608
Serrasalmidae Freshwater Freshwater

T11
Deania Marine 55.8–48.6
Centropomus Marine/Fresh/Brackish 33.9–0.3
yParascombrops 33.9–28.4
Lonchopisthus Marine 11.608–7.246
Apogon Marine 55.8–3.6
Epigonus Marine
 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320
Table 2 (Continued )
Generalized tracks/supportive taxa Geology/fossil faunaa
Lactarius
Eucinostomus
Haemulon
Pomadasys
Mugil
Polyipnus
Saurida
Ariosoma selenops
Hildebrandia flava
Bregmaceros
Lepophidium
yOtophidium robinsi
Porichthys
Opisthonema
a
From Lundberg et al. (2010) and www.fishbase.org (version 02/2014).
b
From http://fossilworks.org.
and the Pacific, ending in Peru (Pisco) in a strictly coastal path.
Given the latest reconstructions of the Central American seaway’s
closure (Montes et al., 2012; Boschman et al., 2014; Montes et al.,
2015), the amphi-American broad pattern that connects eastern
Amazonia, Caribbean and Pacific may be older than the one
established between western Amazonia and Caribbean (through
the Pebas System) during the Miocene.
In a study of Miocene mollusks of tropical America, Petuch
(1988) proposed to divide the Miocene Gatunian province into an
Atlantic and a Pacific component. According to him, the northern
limit of the Gatunian Province was located north of Baja California
(Pacific) and the southern limit was positioned in southern Brazil
(Atlantic). Gillette (1984) concluded that the Miocene ichthyofau-
na from Panama shows similarities with other Miocene faunas
from Ecuador, the Caribbean, and North Carolina. He pointed out
that these occurrences collectively represent the vertebrate fauna
of the Tertiary Caribbean Province.
In this spatiotemporal context, track T3 (from the Pirabas
Formation to northeastern Venezuela) seems to reflect a distinct
and probably younger pattern than the broader one relating
Pirabas with the Greater Antilles and the Pacific. The first one (T3)
agrees with the Tropical Western Central Atlantic Subprovince
erected by Aguilera and Páes (2012) based on the Neogene faunal
assemblage from the Pirabas Formation. This is reported by these
authors as having specific and unique features that distinguish it
from the rest of the Proto-Caribbean subprovince. On the other
hand, the broader track that relates Pirabas with the Greater
Antilles agrees with the relationship reported by de Távora and
Garrafielo Fernandes (1999).
We hypothesize that patterns and nodes that include or involve
the Pacific margin of northern South America or southern Central
America reflect pre-Andean (older) geographical configurations
(tracks T1, T5, T8 and T9; nodes N2, N6, N9). T5 and T9 involve
forearc basins (Pisco and Manabi, respectively) of pre-Andean age
at the Pacific margin and intersect in node N7 with younger, post-
Andean patterns that conform to the geography of the foreland
basins caused by Andean uplift (T4 and T6). The intersection of the
four tracks at node N7 occurs in the area where the Pebas mega-
wetland is supposed to have existed during the Middle Miocene.
Node N2 coincides with a node identified by Echeverry and
Morrone (2013) in southern Central America that involves a
chocoan-amazonian track.
The biogeographic patterns described by T5, T6, T10, and T3 are
supported by fossil taxa shared by Caribbean and Amazonian
Miocene formations. Caribbean and Amazonian Miocene ichthyo-
faunas were spatially related. The biogeographic nodes obtained
(N7, N5, N3, and N4) represent tectonic and biotic convergence
zones and can be related with the area where the Pebas
mega-wetland existed. Our results thus corroborate the hypothesis
317
Extant relatives Fossil age range (Ma)b
Marine/Brackish 48.6–5.332
Marine/Fresh/Brackish 5.332–3.6
Marine 23.03–3.6
Marine/Fresh/Brackish 48.6–7.246
Marine/Fresh/Brackish 23.03–0.012
Marine
Marine 48.6–7.246
Marine 37.2–7.246
Marine
Marine 48.6–2.588
Marine 48.6–3.6
11.608–3.6
Marine 11.608–0.012
Marine
of a historical relationship between the Miocene ichthyofaunas of
the Caribbean and Amazonian regions. Furthermore, they indicate
that this relationship has to be separated into two distinct
components: a western Amazonian/Caribbean (T5, T6, T10 and N7,
N5, N3, N4) and an eastern Amazonian/Caribbean (T3) one.
4.2. Marine/freshwater sister lineages
The history of marine incursions in Amazonia is not
straightforward. Besides the impressive amount of evidence that
shows the Caribbean as the main source of marine incursions,
pathways from the Pacific have been proposed to have occurred
not only in the Miocene (Nuttall, 1990; Steinmann et al., 1999),
but also much earlier. Wesselingh et al. (2010) mention a short-
lived but widespread marine incursion that covered most of the
foreland basin during the Early Paleocene (60–61 Ma). Even
though these authors do not point to the Pacific as the most likely
source of this marine incursion, Lundberg et al. (1998) considered
that before significant uplift of the western margin of South
America, drainage from the continental shields was likely
predominantly westward into the Pacific Ocean during Aptian/
Albian times (112–105 Ma).
Lovejoy (1996) and McEachran et al. (1996) presented stingray
phylogenies based on morphological data, and Lovejoy et al.
(1998), Marques (2000), and Dunn et al. (2003) proposed
molecular-based trees. These studies identified a pair of marine
stingrays, Himantura schmardae from the western Atlantic (WA)
and H. pacifica from the eastern Pacific (EP) as the sister taxa of
freshwater potamotrygonids (Potamotrygonidae is part of our
track T4). Lovejoy et al. (2006) noted that the biogeographic
pattern in which a freshwater South American group has a marine
sister lineage with representatives in both the Atlantic and Pacific
is repeated in other taxa: several clades of helminth parasites on
stingrays (Brooks et al., 1981; Lovejoy, 1997; Hoberg et al., 1998);
the freshwater Potamorrhaphis/Belonion lineage (Belonidae), relat-
ed to a clade that includes WA/EP species (Lovejoy et al., 2006); the
freshwater species Jurengraulis juruensis, most closely related to
the two marine species (Atlantic and Pacific) of Cetengraulis (in T2)
(Nelson, 1984); and the freshwater genus Plagioscion (in T5) and its
western Atlantic/eastern Pacific marine sister genus Paralonchurus
(Boeger and Kritsky, 2003; Boeger et al., 2015).
Based on geographic distribution, age estimates and phyloge-
nies for these taxa, Lovejoy et al. (2006) concluded that ‘‘Miocene
marine incursions prompted the evolutionary transition from
marine to freshwater habitats’’. Cooke et al. (2012), based on
phylogenetic results, concluded that marine incursions between
Late Oligocene and Early Miocene were responsible for the
adaptation to freshwaters in Plagioscion species (Scianidae). More
recently, Boeger et al. (2015) based on phylogenetic analysis
318 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320
rejected the hypothesis of gradual adaptation to freshwater in
favor of the hypothesis of ecological fitting in Scianidae. Based on a
molecular phylogeny of anchovies (Engraulidae), Bloom and
Lovejoy (2012) concluded that New World freshwater anchovies
are the product of a single (instead multiple) marine to freshwater
transition, supporting a pattern of biome conservatism.
Nevertheless, inferences about adaptive evolution (in this case
from marine to freshwater habitats or Marine-Derived Lineages)
should not be directly derived from a phylogeny; nor should a basal
clade be interpreted as ancestral to its sister as it is interpreted by
Cooke et al. (2012). In modern dispersal theory, the sequence of
nodes in a phylogeny is read as a sequence of dispersal events, with
taxa invading a new region, differentiating there, and then
invading another region (Bloom and Lovejoy, 2012). The model
has been criticized by vicariance biogeographers because allopatry
among clades might not be due to dispersal, but instead to a
sequence of in situ differentiation events in an ancestor that was
already widespread. The phylogeny would reflect a sequence of
breaks among the areas involved. A basal clade does not
necessarily indicate a geographic center of origin or an ancestral
habitat (Heads, 2012).
An alternative explanation to the biogeographic pattern in
which a freshwater South American group has a marine sister
lineage with representatives in both the Atlantic and Pacific is
ecological stranding (a mechanism that results in changes in the
habitat of a biota over time without requiring movement or
dispersal of members of the biota; Craw et al., 1999). Isolation, or
stranding through events of Earth history (e.g., changes in sea
level or altitude) can result in a former euryhaline species being
restricted to freshwater. Such isolation does not require or imply
invasion of that habitat, although such isolated taxa are often
described as having ‘‘invaded’’ freshwater. There is no evidence
for extraordinary movement or dispersal: some taxa persist in one
habitat, and others in another habitat (Parenti and Ebach, 2009).
Most of the patterns identified in this analysis are supported by
a combination of freshwater and marine taxa or belong to groups
that have extant relatives living in one of those environments or
both (Table 2). Although these patterns were identified from the
Miocene fossil record, they may reflect older, ancestral patterns
that became fragmented by major changes in northern South
American geography. During Miocene time, the foreland Andean
basins and the aquatic biota of the mega-wetland of western
Amazonia underwent geographical fragmentation caused by
orogenetic events: the main uplift of the eastern Cordillera that
caused the isolation of the Magdalena Basin, the orogeny of the
Merida Andes responsible for the isolation of the Maracaibo Basin,
and the rise of the Vaupés Arch that separated the Orinoco and
Amazon rivers into their modern drainage basins. Although there is
still some controversy about the salinity regimes of the Pebas
System, Hovikoski et al. (2005, 2007) and Hoorn et al. (2006)
believe that while the repeated brackish water phases were
temporally less significant than the freshwater phases, tidal
processes were important and indicate periods of prolonged
connection between the sea and the depositional system.
Based on our biogeographic results (distributional patterns and
nodes) and considering the changes in northern South American
geography during the Miocene, we postulate that geological
events were the major forces causing geographical changes and
fragmentation/isolation of habitats of the Miocene ichthyofauna
over time (ecological stranding). We conclude that the Eastern
Pacific is a relevant component in the history of diversification of
Amazonian fish lineages prior to and during the Neogene (tracks
T1, T5, T8 and T9; nodes N2, N6, N9). It also puts into a broader
spatiotemporal perspective the ‘‘marine-derived lineages’’
hypothesis. The biogeographic pattern in which a freshwater
South American lineage has a marine sister lineage with
representatives in both the Atlantic and Pacific may indicate that
the ancestor of the entire group could have occupied both
freshwater and marine habitats, and undergone sequential
differentiation linked to the geographical breaks in the area. In a
vicariance approach, such freshwater lineages are interpreted as
‘‘marine-related lineages’’ rather than ‘‘marine-derived lineages’’.
Furthermore, Miocene marine incursions do not explain the origin
and diversification of all South American marine-related lineages.
Some lineages might have diversified more recently, such as
Odontesthes (Beheregaray et al., 2002) and freshwater Tetraodon-
tidae (Amaral, 2014), or be older than Miocene and products of
earlier marine incursions or freshwater invasions (Brito and
Deynat, 2004; Lovejoy et al., 2006; Adnet et al., 2014).
We believe that a better understanding of the historical
relationships of the Tropical American Neogene ichthyofaunas
can be achieved by adopting broad spatiotemporal scales. This
allows the consideration of life and Earth evolving together and
helps reconciliation of apparently contradictory results as in the
case of the Pirabas Formation’s relationships.
Acknowledgments
We thank Orangel Aguilera, Michael Heads and one anonymous
reviewer for providing valuable comments on the manuscript. This
research has been supported by grants from the Fundação Carlos
Chagas de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ),
Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico
(CNPq), and UERJ. A. Echeverry thanks CNPq for financial support
through a Post-Doctoral grant. V. Gallo acknowledges research
fellowship grants from the CNPq (Brazilian Federal Government)
and from the PROCIÊNCIA (Rio de Janeiro State Government).
David M. McHenry made useful suggestions to improve the English
of the manuscript.
Appendix A. Supplementary data
Supplementary data associated with this article can be found,
in the online version, at http://dx.doi.org/10.1016/j.geobios.2015.
06.001.
References
AAPG Datapages, 2014. Fugro Tellus Sedimentary Basins of the World Map. ,
Available at: http://www.datapages.com/AssociatedWebsites/GISOpenFiles/
FugroTellusSedimentaryBasinsoftheWorldMap.aspx.
Adnet, S., Salas Gismondi, R., Antoine, P.-O., 2014. Comparisons of dental morphol-
ogy in river stingrays (Chondrichthyes: Potamotrygonidae) with new fossils
from the middle Eocene of Peruvian Amazonia rekindle debate on their evo-
lution. Naturwissenschaften 101, 33–45.
Aguilera, O.A., Rodrigues de Aguilera, D., 2001. An exceptional coastal upwelling fish
assemblage in the Caribbean Neogene. Journal of Paleontology 75, 732–742.
Aguilera, O.A., Rodrigues de Aguilera, D., 2004a. Giant-toothed white sharks and
wide-toothed Mako (Lamnidae) from the Venezuela Neogene: their role in the
Caribbean, shallow-water fish assemblage. Caribbean Journal of Science 40,
368–382.
Aguilera, O.A., Rodrigues de Aguilera, D., 2004b. Goliath grouper (Pisces: Serrani-
dae) from the Upper Miocene Urumaco Formation, Venezuela. Journal of
Paleontology 78, 1202–1206.
Aguilera, O.A., Páes, E., 2012. The Pirabas Formation (Early Miocene from Brazil) and
the Tropical Western Central Atlantic subprovince. Boletim do Museu Paraense
Emilio Goeldi-Ciências Naturais 7, 29–45.
Aguilera, O.A., Bocquentin, J., Lundberg, J.G., Maciente, A., 2008. A new cajaro catfish
(Siluriformes: Pimelodidae: Phractocephalus) from the Late Miocene of south-
western Amazonia and its relationship to yPhractocephalus nassi of the Urumaco
Formation. Paläontologische Zeitschrift 82, 231–245.
Aguilera, O.A., Ramos, M.I., Páes, E., Costa, S., Sánchez-Villagra, M., 2011. The
Neogene tropical American fish assemblage and the paleobiogeography of
the Caribbean region. Swiss Journal of Palaeontology 130, 1–24.
Aguilera, O.A., Moraes-Santos, H., Costa, S., Ohe, F., Jaramillo, C., Nogueira, A., 2013a.
Ariid sea catfishes from the coeval Pirabas (Northeastern Brazil), Cantaure,
Castillo (Northwestern Venezuela), and Castilletes (North Colombia) forma-
tions (early Miocene), with description of three new species. Swiss Journal of
Palaeontology 132, 45–68.
 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320
Aguilera, O.A., Lundberg, J., Birindelli, J., Sabaj Pérez, M., Jaramillo, C., Sánchez-
Villagra, M.R., 2013b. Palaeontological evidence for the last temporal occur-
rence of the ancient western Amazonian River outflow into the Caribbean. PLOS
One 8, e76202.
Amaral, C.R.L., 2014. Filogenia e Biogeografia dos Tetraodontidae, com ênfase no
gênero Colomesus (Gill, 1885). Tese de Doutorado, Programa de Pós-graduação
em Biociências, Universidade do Estado de Rio de Janeiro (UERJ), unpubl.
Astorga, A., Fernández, J.A., Barboza, G., Campos, L., Obando, J., Aguilar, A., Obando,
L.G., 1991. Cuencas Sedimentarias de Costa Rica: evolución geodinámica y
potencial de hidrocarburos. Revista Geológica de América Central 13, 25–59.
Baby, P., Rivadeneira, M., Christophoul, F., Barragan, R., 1999. Style and timing of
deformation in the Oriente Basin of Ecuador. In: Fourth ISAG, October 4–6,
Goettingen, Germany.
Barrero, D., Pardo, A., Vargas, C.A., Martı́nez, J.F., 2007. Colombian Sedimentary
Basins: Nomenclature, Boundaries and Petroleum Geology, a New Proposal.
Agencia Nacional de Hidrocarburos (ANH), Bogotá, Colombia.
Bayona, G., Valencia, A., Mora, A., Rueda, M., Ortiz, J., Montenegro, O., 2008.
Estratigrafı́a y procedencia de las rocas del Mioceno en la parte distal de la
cuenca antepaı́s de los Llanos de Colombia. Geologı́a Colombiana 33, 23–46.
Beheregaray, L.B., Sunnucks, P., Briscoe, D.A., 2002. A rapid fish radiation associated
with the last sea-level changes in southern Brazil: the silverside Odontesthes
perugiae complex. Proceedings of the Royal Society London B 269, 65–73.
Bloom, D.D., Lovejoy, N.R., 2012. Molecular phylogenetics reveals a pattern of bioma
conservatism in New World anchovies (family Engraulidae). Journal of Evolu-
tionary Biology 25, 701–715.
Boeger, W.A., Kritsky, D.C., 2003. Parasites, fossils and geologic history: historical
biogeography of the South American freshwater croakers, Plagioscion spp.
(Teleostei, Sciaenidae). Zoologica Scripta 32, 3–11.
Boeger, W.A., Marteleto, F.M., Zagonel, L., 2015. Tracking the history of an invasion:
the freshwater croakers (Teleostei: Sciaenidae) in South America. Zoologica
Scripta 44, 250–262.
Boschman, L.M., van Hinsbergen, D.J.J., Torsvik, T.H., Spakman, W., Pindell, J.L., 2014.
Kinematic reconstruction of the Caribbean region since the Early Jurassic. Earth-
Science Reviews 138, 102–136.
Brito, P.M., Deynat, P.P., 2004. Freshwater stingrays from the Miocene of South
America with comments on the rise of potamotrygonids (Batoidea, Mylioba-
tiformes). In: Arratia, G., Wilson, M.V.H., Cloutier, R. (Eds.), Recent Advances in
the Origin and Early Radiation of Vertebrates. Verlag, München, Germany, pp.
575–582.
Brito, P.M., Meunier, F.J., Leal, M.E.C., 2007. Origine et diversification de l’ichtyo-
faune néotropicale : une revue. Cybium 31, 149–163.
Brooks, D.R., Thorson, T.B., Mayes, M.A., 1981. Fresh-water stingrays (Potamotry-
gonidae) and their helminth parasites: testing hypotheses of evolution and
coevolution. In: Funk, V.A., Brooks, D.R. (Eds.), Advances in Cladistics. New York
Botanical Garden, New York, pp. 147–175.
Candela, A., Morrone, J.J., 2003. Biogeografı́a de puercoespines neotropicales
(Rodentia, Hystricognathi): integrando datos fósiles y actuales a través de un
enfoque panbiogeográfico. Ameghiniana 40, 361–378.
Cecca, F., 2002. Palaeobiogeography of Marine Fossil Invertebrates: Concepts and
Methods. Taylor and Francis, London and New York.
Cecca, F., Vrielynck, B., Lavoyer, T., Gaget, H., 2005. Changes in the ammonite
taxonomical diversity gradient during the late Jurassic-early Cretceous. Journal
of Biogeography 32, 535–547.
Cooke, G.M., Chao, N.L., Beheregaray, L.B., 2012. Marine incursions, cryptic species
and ecological diversification in Amazonia: the biogeographic history of the
croaker genus Plagioscion (Sciaenidae). Journal of Biogeography 39, 724–738.
Colletta, B., Roure, F., DeToni, B., Loureiro, D., Passalacqua, H., Gou, Y., 1997. Tectonic
inheritance, Crystal architecture, and contrasting structural styles in the Vene-
zuela Andes. Tectonics 16, 777–794.
Craw, R.C., Grehan, J.R., Heads, M., 1999. Panbiogeography: tracking the history of
life. Oxford University Press, New York.
da Cruz Cunha, P.R., 2007. Bacia do Acre, in: Boletim de Geociências da Petrobras.
Cartas estratigráficas 15, 207–213.
da Cruz Cunha, P.R., Gonçalves de Melo, J.H., Braga da Silva, O., 2007. Bacia do
Amazonas, in: Boletim de Geociências da Petrobras. Cartas estratigráficas 15,
227–251.
Deniaud, Y., Baby, P., Basile, C., Ordoñez, M., Mascle, G., Montenegro, G., 1999.
Neogene evolution of the main Ecuadorian fore-arc sedimentary basins and
sediment mass-balance inferences. In: Fourth ISAG, October 4–6, Goettingen,
Germany.
Deynat, P.P., Brito, P.M., 1994. Révision des tubercules cutanés de raies (Chon-
drichthyes, Batoidea) du bassin du Paraná, Tertiaire d’Amerique du Sud. Annales
de Paléontologie 80, 237–251.
Diaz de Gamero, M.L., 1996. The changing course of the Orinoco River during the
Neogene: a review. Palaeogeography Palaeoclimatology Palaeoecology 123,
385–402.
Dunn, K.A., McEachran, J.D., Honeycutt, R.L., 2003. Molecular phylogenetics of
myliobatiform fishes (Chondrichthyes: Myliobatiformes), with comments on
the effects of missing data on parsimony and likelihood. Molecular Phyloge-
netics and Evolution 27, 259–270.
Duque-Caro, H., 1990. Major Neogene events in panamaic South America. In:
Tsuchi, R. (Ed.), Pacific Neogene Events, their Timing, Nature and Interrelation-
ships. University Press, Tokyo, pp. 101–114.
Echeverry, A., Morrone, J.J., 2010. Parsimony analysis of endemicity as a panbio-
geographical tool: An analysis of Caribbean plant taxa. Biological Journal of the
Linnean Society 101, 961–976.
319
Echeverry, A., Morrone, J.J., 2013. Generalized tracks, area cladograms and tectonics
in the Caribbean. Journal of Biogeography 40, 1619–1637.
Escalante, G., Astorga, A., 1992. Geologı́a del este de Costa Rica y el Norte de Panamá.
Revista Geológica de América Central, vol Especial. Terremoto de Limón 1–14.
Fernandes dos Reis, M.A., 2005. Chondrichthyan fauna from the Pirabas Formation,
Miocene of northern Brazil, with comments on paleobiogeography. Anuário do
Instituto de Geociências-UFRJ 28, 31–58.
Fernandes Soares, E., Zalán, P.V., Picanço de Figueiredo, J.J., Trosdtorf Junior, I., 2007.
Bacia do Pará-Maranhão. Boletim de Geociências da Petrobras. Cartas estrati-
gráficas 15, 321–329.
Ferreira Costa, S.A.R., Richter, M., Mann de Toledo, P., Moraes-Santos, H.M., 2009.
Shark teeth from Pirabas Formation (Lower Miocene), northeastern Amazonia,
Brazil. Boletim do Museu Paraense Emilio Goeldi-Ciências Naturais 4, 221–230.
Fielitz, C., 1996. A Late Cretaceous (Turonian) ichthyofauna from Lac des Bois,
Nothwest Territories, Canada, with paleobiogeographic comparisons with Tur-
onian ichthyofaunas of the Western Interior Seaway. Canadian Journal of Earth
Sciences 33, 1375–1389.
Figueiredo, J., Hoorn, C., van der Ven, P., Soares, E., 2009. Late Miocene onset of the
Amazon River and the Amazon deep-sea fan: Evidence from the Foz do
Amazonas Basin. Geology 37, 619–622.
Gallo, V., Cavalcanti, M.J., Silva, H.M.A., 2007. Track analysis of the marine palaeofauna
from the Turonian (Late Cretaceous). Journal of Biogeography 34, 1167–1172.
Gillette, D.D., 1984. A marine ichthyofauna from the Miocene of Panama, and the
Tertiary Caribbean faunal province. Journal of Vertebrate Paleontology 4, 172–186.
Gingras, M.K., Räsänen, M.E., Pemberton, G., Romero, L.P., 2002. Ichnology and
sedimentology reveal depositional characteristics of bay-margin parase-
quences in the Miocene Amazonian foreland basin. Journal of Sedimentary
Research 72, 871–883.
Goloboff, P., 1998. NONA ver. 2.0., Available at : http://www.cladistics.com/about_
nona.htm.
Gómez, A.A., Jaramillo, C.A., Parra, M., Mora, A., 2009. Huesser horizon: a lake and a
marine incursión in Northwestern South America during the early Miocene.
Palaios 24, 136–147.
Gross, M., Ramos, M.I., Caporaletti, M., Piller, W.E., 2013. Ostracods (Crustacea) and
their palaeoenvironmental implication for the Solimoes Formation (Late Mio-
cene; Western Amazonia/Brazil). Journal of South American Earth Sciences 42,
216–241.
Guerrero, J., 1997. Stratigraphy, sedimentary environments and the Miocene uplift
of the Colombian Andes. In: Kay, R.F., Madden, R.H., Cifelli, R.L., Flynn, J.J. (Eds.),
Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta,
Colombia. Smithsonian Institution Press, Washington DC, pp. 15–43.
Hasui, Y., 2012. In: Hasui, Y., Dal Ré Carneiro, C., Marques de Almeida, F.F., Bartorelli,
A. (Eds.), Tectônica de placas. Geologia do Brasil, Petrobras, São Paulo, Brasil,
pp. 66–97.
Heads, M., 1989. Integrating earth and life sciences in New Zealand natural history:
the parallel arcs model. New Zealand Journal of Zoology 16, 549–585.
Heads, M., 2004. What is a node? Journal of Biogeography 31, 1883–1891.
Heads, M., 2006. Panbiogeography of Nothofagus (Nothofagaceae): analysis of the
main species massings. Journal of Biogeography 33, 1066–1075.
Heads, M., 2012. Molecular panbiogeography of the tropics. University of California
Press, Berleley, CA.
Higley, D.K., 2004. The Progreso Basin Province of Northwestern Peru and South-
western Ecuador: Neogene and Cretaceous-Paleogene Total Petroleum Systems.
U.S. Geological Survey Bulletin 2206-B, Available at: http://pubs.usgs.gov/bul/
b2206-b/.
Hoberg, E.P., Brooks, D.R., Molina-Ureña, H., Erbe, E., 1998. Echinocephalus janzeni
n.sp. nematoda: (Gnathostomatidae) in Himantura pacifica (Chondrichthyes:
Myliobatiformes) from the Pacific coast of Costa Rica and Mexico, with histori-
cal biogeographic analysis of the genus. Journal of Parasitology 84, 571–581.
Hoorn, C., 1993. Marine incursions and the influence of Andean tectonics on the
Miocene depositional history of northwestern Amazonia: results of a palynos-
tratigraphic study. Palaeogeography Palaeoclimatology Palaeoecology 105,
267–309.
Hoorn, C., 1994a. Fluvial palaeoenvironments in the intracratonic Amazonas Basin
(Early Miocene-Early Middle Miocene, Colombia). Palaeogeography Palaeocli-
matology Palaeoecology 109, 1–54.
Hoorn, C., 1994b. An environmental reconstruction of the palaeo-Amazon river
system (Middle-Late Miocene, NW Amazonia). Palaeogeography Palaeoclima-
tology Palaeoecology 112, 187–238.
Hoorn, C., Guerrero, J., Sarmiento, G.A., Lorente, M.A., 1995. Andean tectonics as a
cause for changing drainage patterns in Miocene northern South America.
Geology 23, 237–240.
Hoorn, C., 1996. Miocene deposits in the Amazonian foreland basin. Science 273,
122–123.
Hoorn, C., Aalto, R., Kaandorp, R.J.G., Lovejoy, N.R., 2006. Miocene semidiurnal tidal
rhythmites in Madre de Dios, Peru: Comment. GSA, Forum. Geology
doi:10.1130/G22115.1.
Hoorn, C., Wesselingh, F.P., Hovikoski, J., Guerrero, J., 2010. The development of the
Amazonian mega-wetland (Miocene; Brazil, Colombia, Peru, Bolivia). In: Hoorn,
C., Wesselingh, F. (Eds.), Amazonia: landscape and species evolution. A look into
the past. Wiley-Blackwell, UK, pp. 123–142.
Hovikoski, J., Räsänen, M.E., Gingras, M., Roddaz, M., Brusset, S., Hermoza, W., 2005.
Miocene semidiurnal tidal rhythmites in Madre de Dios, Peru. Geology 33, 177–180.
Hovikoski, J., Gingras, M., Räsänen, M.E., Rebata, L., Guerrero, J., Ranzi, A., 2007. The
nature of Miocene Amazonian epicontinental embayment: High-frequency
shifts of the low-gradient coastline. GSA Bulletin 119, 1506–1520.
320 A. Echeverry, V. Gallo / Geobios 48 (2015) 309–320
Klein, G.D., Zúñiga Rivero, F.J., Hay-Roe, H., Alvarez-Calderón, E., 2011. A reappraisal
of the Mesozoic/Cenozoic tectonics and sedimentary basins of Peru. Search and
Discovery Article #10332. BPZ Resources, Inc. AAPG.
Landau, B., Vermeij, G., Marques da Silva, C., 2008. Southern Caribbean Neogene
palaeobiogeography revisited. New data from the Pliocene of Cubagua, Vene-
zuela. Palaeogeography Palaeoclimatology Palaeoecology 257, 445–461.
Latrubesse, E.M., da Silva, S.A.F., Cozzuol, M., Absy, M.L., 2007. Late Miocene
continental sedimentation in southwestern Amazonia and its regional signifi-
cance: Biotic and geological evidence. Journal of South American Earth Sciences
23, 61–80.
Latrubesse, E.M., Cozzuol, M., da Silva-Caminha, S.A.F., Rigsby, C.A., Absy, M.L.,
Jaramillo, C., 2010. The Late Miocene paleogeography of the Amazon Basin and
the evolution of the Amazon River system. Earth-Science Reviews 99, 99–124.
Laurito, C.A., 1996. El género Isistius (Squalidae) en el alto Guayacán. Formación
Uscari (Mioceno Superior-Plioceno Inferior), Provincia de Limón, Costa Rica.
Revista Geológica de América Central 19/20, 87–92.
Laurito, C.A., 2004. Ictiofauna de la Formación Punta Judas, Mioceno Medio, Pro-
vincia de Puntarenas, Costa Rica. Brenesia 62, 57–74.
Laurito, C.A., Valerio, A.L., 2008. Ictiofauna de la localidade de San Gerardo de
Limoncito, Formación Curré, Mioceno Superior, Cantón de Coto Brus, Provincia
de Puntarenas, Costa Rica. Revista Geológica de América Central 39, 65–85.
Laurito, C.A., Valerio, A.L., Ovares, E., Hernández, A.C., Pizarro, D., 2008. Peces fósiles
de la localidad Lomas de Siquirres, cauce del Rı́o Reventazón, Formación Rı́o
Banano, Mioceno Superior, Provincia de Limón, Costa Rica. Revista Geológica de
América Central 38, 11–25.
Lieberman, B.S., 2003. Paleobiogegraphy: the relevance of fossil to biogeography.
Annual Review of Ecology and Systematics 34, 51–69.
Linhares, A.P., Feijó Ramos, M.I., Gross, M., Piller, W.E., 2011. Evidence for marine
influx during the Miocene in southwestern Amazonia, Brazil. Geologı́a Colombi-
ana. Edición Especial 36, 91–104.
Longbottom, A.E., 1979. Miocene sharks’ teeth from Ecuador. Bulletin of the British
Museum (Natural History) 32, 57–70.
Lovejoy, N.R., 1996. Systematics of myliobatoid elasmobranchs: with emphasis on
the phylogeny and historical biogeography of Neotropical freshwater stingrays
(Potamotrygonidae: Rajiformes). Zoological Journal of the Linnean Society 117,
207–257.
Lovejoy, N.R., 1997. Stingrays, parasites, and neotropical biogeography: a closer
look at Brooks et al.’s hypotheses concerning the origins of neotropical fresh-
water rays (Potamotrygonidae). Systematic Biology 46, 218–230.
Lovejoy, N.R., Bermingham, E., Martin, A.P., 1998. Marine incursion into South
America. Nature 396, 421–422.
Lovejoy, N.R., Albert, J.S., Crampton, W.G.R., 2006. Miocene marine incursions and
marine/freshwater transitions: evidence from Neotropical fishes. Journal of
South American Earth Sciences 21, 5–13.
Lundberg, J.G., Marshall, L.G., Guerrero, J., Horton, B., Malabarba, M.C., Wesselingh,
F., 1998. The stage for Neotropical fish diversification: a history of tropical South
American rivers. In: Malabarba, L.R., Reis, R.E., Vari, R.P., Lucena, Z.M.S., Lucena,
C.A.S. (Eds.), Phylogeny and classification of Neotropical fishes. EDIPUCRS,
Porto Alegre, Brazil, pp. 13–48.
Lundberg, J.G., Sabaj Pérez, M.H., Dahdul, W.M., Aguilera, O.A., 2010. The Amazonian
Neogene fish fauna. In: Hoorn, C., Wesselingh, F. (Eds.), Amazonia: landscape
and species evolution. A look into the past. Wiley-Blackwell, UK, pp. 281–301.
Marques, F.P.L., 2000. Evolution of neotropical freshwater stingrays and their
parasites: taking into account space and time. PhD Thesis, University of Toronto,
unpubl.
Martin-Gombojav, N., Winkler, W., 2008. Multiple recycling of Proterozoic crust in
the Andean Amazon foreland of Ecuador: implications for orogenetic develop-
ment of the Northern Andes. Terra Nova 20, 22–31.
McEachran, J.D., Dunn, K.A., Miyake, T., 1996. Interrelationships of the batoid fishes
(Chondrichthyes: Batoidea). In: Stiassny, M.L.J., Parenti, L.R., Johnson, G.D.
(Eds.), Interrelationships of Fishes. Academic Press, San Diego, pp. 63–84.
Monsch, K.A., 1998. Miocene fish faunas from the northwestern Amazonia basin
(Colombia, Peru, Brazil) with evidence of marine incursions. Palaeogeography
Palaeoclimatology Palaeoecology 143, 31–50.
Montes, C., Cardona, A., Farris, D., Strong, N., Bayona, G., Jaramillo, C., 2011. The
Panamá Geology Project. Cenozoic Evolution of the Panamá Isthmus. Smithso-
nian Tropical Research Institute, Available at: http://www.stri.org.
Montes, C., Bayona, G., Cardona, A., Buchs, D.M., Silva, C.A., Morón, S., Hoyos, N.,
Ramı́rez, D.A., Jaramillo, C.A., Valencia, V., 2012. Arc-continent collision and
orocline formation: closing of the Central American seaway. Journal of Geo-
physical Research 117, B04105.
Montes, C., Cardona, A., Jaramillo, C., Pardo, A., Silva, J.C., Valencia, V., Ayala, C.,
Pérez-Angel, L.C., Rodriguez-Parra, L.A., Ramirez, V., Niño, H., 2015. Middle
Miocene closure of the Central American seaway. Science 348, 226–229.
Mora, A., Baby, P., Roddaz, M., Parra, M., Brusset, S., Hermoza, W., Espurt, N., 2010.
Tectonic history of the Andes and sub-Andean zones: implications for the
development of the Amazon drainage basin. In: Hoorn, C., Wesselingh, F.
(Eds.), Amazonia: landscape and species evolution. A look into the past. Wiley
Blackwell, UK, pp. 38–60.
Morrone, J.J., 2009. Evolutionary Biogeography: an integrative approach with case
studies. Columbia University Press, New York.
Morrone, J.J., 2014. Parsimony analysis of endemicity (PAE) revisited. Journal of
Biogeography 41, 842–854.
Nelson, G., 1984. Identity of the anchovy Hildebranchthyes setiger with notes on
relationships and biogeography of the genera Engraulis and Cetengraulis.
Copeia 1984, 422–427.
Nelson, J.S., 2006. Fishes of the World, 4th ed. John Wilet and Sons, New Jersey.
Nixon, K.C., 1999. About Winclada. Ithaca, N.Y. , Available at:http://www.cladistics.
com/about winc.
Nuttall, C.P., 1990. A review of the Tertiary non-marine molluscan faunas of the
Pebasian and other inland basins of north-western South America. Bulletin of
the British Museum (Natural History) Geology Series 45, 165–371.
Parenti, L.R., Ebach, M.C., 2009. Comparative Biogeography. University of California
Press, Los Angeles.
Petuch, E.J., 1988. Neogene history of tropical American mollusks, Biogeography
and evolutionary patterns of tropical western Atlantic Mollusca. Coastal Edu-
cation and Reserch Foundation, Charlottesville, Virginia.
Picanço de Figueiredo, J.J., Zalán, P.V., Fernandes Soares, E., 2007. Bacia da Foz do
Amazonas. Boletim de Geociências da Petrobras. Cartas estratigráficas 15, 299–309.
Pimiento, C., Gonzalez-Barba, G., Hendy, A.J.W., Jaramillo, C., MacFadden, B.J.,
Montes, C., Suarez, S.C., Shippritt, M., 2013. Early Miocene chondrichthyans
from the Culebra Formation, Panama: a window into marine vertebrate faunas
before closure the Central American seaway. Journal of South American Earth
Sciences 42, 159–170.
Räsänen, M.E., Linna, A.M., Santos, J.C.R., Negri, F.R., 1995. Late Miocene tidal
deposits in the Amazonian foreland basin. Science 269, 386–390.
Räsänen, M.E., Linna, A.M., Irion, G., Hernani, L.R., Huaman, R.V., Wesselingh, F.,
1998. Geologia y geoformas de la zona de Iquitos. Annales Universitatis Tur-
kensis Ser. A II 114, 59–137.
Ribeiro Wanderley-Filho, J., Fernandes Eiras, J., Tenório Vaz, P., 2007. Bacia do
Solimões. Boletim de Geociências da Petrobras. Cartas estratigráficas 15, 217–223.
Ribeiro Wanderley-Filho, J., Fernandes Eiras, J., da Cruz Cunha, P.R., van der Vem,
P.H., 2010. The Paleozoic Solimões and Amazonas basins and the Acre foreland
basin of Brazil. In: Hoorn, C., Wesselingh, F. (Eds.), Amazonia: landscape and
species evolution. A look into the past. Wiley-Blackwell, UK, pp. 29–37.
Robertson, D.R., Cramer, K.L., 2014. Defining and dividing the Greater Caribbean:
insights from the biogeography of shorefishes. PLoS One 9, e102918.
Rojas-Parra, C.A., 2007. Una herramienta automatizada para realizar análisis pan-
biogeográficos. Biogeografı́a 1, 31–33.
Rosen, B.R., 1988. Biogeographic patterns: a perceptual overview. In: Myers, A.A.,
Giller, P.S. (Eds.), Analytical biogeography: an integrated approach to the study
of animal and plantdistributions. Chapman and Hall, London, pp. 23–55.
Rosen, B.R., Smith, A.B., 1988. Tectonics from fossils? Analysis of reef-coral and sea-
urchin distributions from late Cretaceous to Recent, using a new method. In:
Audley-Charles, M.G., Hallam, A. (Eds.), Gondwana and Tethys, 37, Geological
Society Special Publication, pp. 275–306.
Silva, H.M.A., Gallo, V., 2007. Parsimony analysis of endemicity of enchodontoid
fishes from the Cenomanian. Carnets de Géologie Letter 2007/01, 1–8.
Steinmann, M., Hungerbühler, D., Seward, D., Winkler, W., 1999. Neogene tectonic
evolution and exhumation of the southern Andes: a combined stratigraphy and
fission-track approach. Tectonophysics 307, 255–276.
Távora, V., Garrafielo Fernandes, J.M., 1999. Estudio de los foraminı́feros de la
Formación Pirabas (Mioceno Inferior), Estado de Pará, Brasil y su correlación con
las faunas del Caribe. Revista Geológica de América Central 22, 63–74.
Vermeij, J., Wesselingh, F.P., 2002. Neogastropod mollusks from the Miocene of
western Amazonia, with comments on marine to freshwater transitions in
mollusks. Journal of Paleontology 76, 265–270.
Vonhof, H.B., Wesselingh, F.P., Ganssen, G.M., 1998. Reconstruction of the Miocene
western Amazonian aquatic system using molluscan isotopic signatures.
Palaeogeography Palaeoclimatology Palaeoecology 141, 85–93.
Vonhof, H.B., Wesselingh, F.P., Kaandorp, R.J.G., Davies, G.R., van Hinte, J.E., Guerrero, J.,
Räsänen, M., Romero-Pittman, L., Ranzi, A., 2003. Paleogeography of Miocene
western Amazonia: isotopic composition of molluscan shells constrains the influ-
ence of marine incursions. Geological Society of America Bulletin 115, 983–993.
Webb, S.D., 1995. Biological implications of the middle Miocene Amazon seaway.
Science 269, 361–362.
Wesselingh, F.P., Macsotay, O., 2006. Pachydon hettneri (Anderson, 1982) as indica-
tor for Caribbean-Amazonian lowland connections during the early-middle
Miocene. Journal of South American Earth Sciences 21, 49–53.
Wesselingh, F.P., Räsänen, M., Irion, G., Vonhof, H.B., Kaandorp, R., Renema, W.,
Pitmann, L.R., Gingras, M., 2002. Lake Pebas: a palaeoecological reconstruction
of a Miocene, long-lived lake complex in western Amazonia. Cainozoic Research
1, 35–81.
Wesselingh, F.P., Guerrero, J., Räsänen, M., Romero Pitmann, L., Vonhof, H., 2006.
Landscape evolution and depositional processes in the Miocene Amazonian
Pebas lake/wetland system: evidence from exploratory boreholes in northeast-
ern Peru. Scripta Geologica 133, 323–361.
Wesselingh, F.P., Hoorn, C., Kroonenberg, S.B., Antonelli, A., Lundberg, J.G., Vonhof,
H.B., Hooghiemstra, H., 2010. On the origin of Amazonian landscapes and
biodiversity: a synthesis. In: Hoorn, C., Wesselingh, F. (Eds.), Amazonia:
landscape and species evolution. A look into the past. Wiley-Blackwell, UK,
pp. 421–431.
Woodring, W.P., 1974. The Miocene Caribbean Faunal Province and its Subpro-
vinces. Verhandlungen der naturforschenden Gesellschaft in Basel 84, 209–213.
Zalán, P.V., 2004. Evolução fanerozóica das bacias sedimentares brasileiras. In:
Mantesso Neto, V., Bartorelli, A., Carneiro, C.D.R., de Brito-Neves, B.B. (Eds.),
Geologia do Continente Sul-Americano: Evolução da obra de Fernando Flávio
Marques de Almeida. São Paulo, Brasil, pp. 595–612.
Zalán, P.V., 2007. Bacias de Bragança-Viseu, São Luı́s e Ilha Nova, in: Boletim de
Geociências da Petrobras. Cartas estratigráficas 15, 341–345.
Zalán, P.V., Matsuda, N.S., 2007. Bacia do Marajó. Boletim de Geociências da
Petrobras. Cartas estratigráficas 15, 311–319.