Journal of South American Earth Sciences 100 (2020) 102551

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Journal of South American Earth Sciences

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Goliath catfish Brachyplatystoma Bleeker, 1862 (Siluriformes: Pimelodidae) T

from the Miocene of Argentina
Federico L. Agnolina,b,c, Sergio Bogana,∗
a
Fundación de Historia Natural “Félix de Azara”, Departamento de Ciencias Naturales y Antropología, Universidad Maimónides, Hidalgo 775 piso 7 (1405BDB), Buenos
Aires, Argentina
b
Laboratorio de Anatomía Comparada y Evolución de los Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Av. Ángel Gallardo 470 (1405),
Buenos Aires, Argentina
c
CONICET, Argentina

ARTICLE INFO ABSTRACT

Keywords: Brachyplatystoma is a genus of freshwater catfishes endemic of South America that includes one the largest
Brachyplatystoma siluriform species of the neotropics. The fossil record of the genus is restricted to the Miocene of Colombia and
Amazonas Venezuela and the extant species are distributed along Orinoco and Amazonas basins. The present works aims to
Paraná describe a new species of Brachyplatystoma from late Miocene deposits at Entre Ríos Province, Argentina. The
Argentina
present report constitutes the first finding of Brachyplatystoma in the Paleoparaná Basin, which is in agreement
Late Miocene
with previous hypothesis indicating hydrographic connections of this river basin with the Amazon basin until at
least the early late Miocene times.

1. Introduction Brachyplatystoma and Platynematichthys Bleeker, 1858.

The Order Siluriformes is a diverse clade of ostariophysan tele-
osteans with more than 3.900 species that are widespread around the
world (e.g., Arratia et al., 2003; Nelson, 2006; van der Laan et al.,
2020). The fossil record of catfishes in the New World is patchy and
biased, the oldest records coming from the Late Cretaceous of Perú,
Bolivia, Brazil, and Argentina (Arratia and Cione, 1996; Gayet and
Meunier, 2003; Alves et al., 2016). The Family Pimelodidae is one of
the most diverse catfish clade from the current Neotropical ichthyo-
fauna (Reis et al., 2003). The first undoubted fossil record of pimelodids
dates back to the late Oligocene-early Miocene of Taubaté, Brazil, with
two species of the genus Steindachneridion Eigenmann and Eigenmann,
1919 (Lundberg, 1998).
Within extant Pimelodidae, the subfamily Sorubiminae was coined
by Schultz (1944) with the aim to nest diverse catfishes, including the
genera Zungaro Bleeker, 1858, Hemisorubim Bleeker, 1862, Sorubim
Cuvier, 1829, Hypophthalmus Cuvier, 1829, and the groups Pseudopla-
tystoma-Sorubimichthys and Platysilurus-Platystomatichthys, as well as the
Tribe Brachyplatystomatini. This grouping was partially confirmed by
recent molecular analyses that included brachyplatistomines within
Sorubiminae (Lundberg and Akama, 2005; Lundberg et al., 2011).
Lundberg and Akama (2005) presented a phylogenetic analysis of the
Brachyplatystomatini Tribe, in which they include the extant genera
The genus Brachyplatystoma Bleeker, 1862 is the largest-sized of
freshwater catfishes inhabiting South America, and includes the species
commonly known as Piraíba (B. filamentosum Lichtenstein, 1819) which
reaches up to about 3.6 m in length and 200 kg of weight (Lundberg
and Littmann, 2003). Like other large pimelodid catfishes, some species
of Brachyplatystoma (B. vaillantii, B. filamenosum, and B. rousseauxii)
make large migrations along freshwater routes, with the aim to spawn
in the western Amazon basin, as far upstream as the Andes (Barthem
et al., 2017).
At present, the genus Brachyplatystoma is endemic to northern and
central South America and includes seven extant species (i.e., B. jur-
uense Boulenger, 1898, B. platynemum Boulenger, 1898, B. tigrinum
Britski, 1981, B. vaillantii Valenciennes, 1835–1847, B. capapretum
Lundberg and Akama, 2005, B. filamentosum Lichtenstein, 1819, B.
rousseauxii Castelnau, 1855) and one fossil form (i.e., †B. promagdalena
Lundberg, 2005; also see Nass, 1991). These taxa are grouped in the
subgenera Brachyplatystoma and Malacobagrus [the latter including B.
rousseauxii, B. filamentosum and B. capapretum (Lundberg and Akama,
2005; Lundberg et al., 2011; Sullivan et al., 2013)].
The fossil record of the genus Brachyplatystoma dates back to the
middle Miocene of La Venta Group in central Colombia. Lundberg
(1998, 2005) reported the extinct species B. promagadalena and remains
identified as B. cf. vaillantii. More recently, records of B. cf. vaillantii

∗
Corresponding author.
E-mail addresses: fedeagnolin@yahoo.com.ar (F.L. Agnolin), sergiobogan@yahoo.com.ar (S. Bogan).

https://doi.org/10.1016/j.jsames.2020.102551
Received 7 January 2020; Received in revised form 19 February 2020; Accepted 28 February 2020
Available online 06 March 2020
0895-9811/ © 2020 Published by Elsevier Ltd.
F.L. Agnolin and S. Bogan Journal of South American Earth Sciences 100 (2020) 102551

were reported for Miocene outcrops of northern Colombia and Vene-

zuela (Aguilera et al., 2013). However, most fossil remains determined
as B. cf. vaillantii consist of isolated and somewhat poorly preserved
bones, whereas B. promagadalena is based on a single, incomplete We-
berian apparatus (Lundberg, 2005).
In this contribution we describe remains referable to the genus
Brachyplatystoma coming from Late Miocene beds of Entre Ríos pro-
vince, Argentina. The specimens represent an important increment on
Brachyplatystoma diversity, since constitutes the first record for the
genus outside Orinoco and Amazon basins.

2. Material and methods

The examined fossil specimens are housed in the Vertebrate

Paleontological Collection of the Museo Argentino de Ciencias
Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (MACN Pv).
Under the same number (MACN Pv 16052) there are included three
Weberian apparatus belonging to different individuals.
Dry skeletons of living species belong to the following comparative
collections: ANSP, Academy of Natural Sciences of Philadelphia, USA.
DUF, Duke University, Vertebrate Collection, Durham, North Carolina,
USA. USNM, National Museum of Natural History, Smithsonian
Institution, Department of Vertebrate Zoology, Washington D.C. USA.
MZUSP, Museu de Zoologia da Universidade de São Paulo, São Paulo,
Brazil. Platynematichthys notatus, ANSP 178258; Brachyplatystoma fila-
mentosum, ANSP 187105, DUF 1052; Brachyplatystoma capapretum,
MZUSP 53262; Brachyplatystoma vaillantii, DUF 1109, DUF 1034 (avail-
able at http://catfishbone.ansp.org/Pimelodidae/Brachyplatystoma/
vaillantii/dry.skeleton.html); Brachyplatystoma tigrinum, ANSP 179236,
USNM 280746; Brachyplatystoma juruense, ANSP 178514;
Brachyplatystoma platynemum, ANSP 187321; and Brachyplatystoma
rousseauxi, DUF 1057.
Osteological nomenclature follows Lundberg and Akama (2005) and
osteological traits of the Weberian apparatus follow Lundberg (2005). Fig. 1. Map showing distribution of living species of genus Brachyplatystoma
and fossil localities. 1, Brachyplatystoma cf. B. vaillantii; Alta Guajira Peninsula,
northern Colombia; Upper Castilletes Formation, Early Pliocene (Aguilera et al.,
3. Geological settings
2013). 2, Brachyplatystoma cf. B. vaillantii; North Corralito, Urumaco trough,
Northwestern Venezuela; Urumaco Formation, Late Miocene (Aguilera et al.,
The material here studied was collected at the end of the 19th and 2013). 3, Brachyplatystoma promagdalena; La Venta, Colombia; La Venta For-
beginning of 20th centuries in the Paraná riverside cliffs near Paraná mation, Late Miocene (Lundberg, 2005). 4, Brachyplatystoma elbakyani; Paraná
city, Entre Ríos Province, Argentina (Fig. 1). According to Azpelicueta city, Entre Ríos province, Argentina; Paraná Formation, Late Miocene.
and Cione (2016) fossils from this site actually come from the Toma
Vieja locality and other nearby sites.
Type species: Platystoma vaillanti Valenciennes, 1835–1847. Type
The fossil-bearing bed of the new Brachyplatystoma species is the
by original designation.
“Conglomerado osífero” which is a well-known fossiliferous bed since
Brachyplatystoma elbakyani nov. sp.
the 19th century (Bravard, 1858; Ameghino, 1889; Bogan et al., 2012).
Holotype. MACN Pv 16091, fronto-ethmoid/vomerine region of
Cione et al. (2000, 2009) analyzed in detail the stratigraphical position
skull (Figs. 2 and 3).
of the freshwater fish remains from those old collections and concluded
Referred material. MACN Pv 15989, incomplete ethmoid/vo-
that they found at the base of the late Miocene Ituzaingó Formation
merine region of skull (Figs. 3 and 5).
(Aceñolaza, 2000; Brunetto et al., 2013). Fossil mammals indicate that
Type locality. Paraná riverside cliffs near Paraná city (31°43′ S,
the Ituzaingó Formation ranges from about 9 to about 6 Ma, probably
60°31′ W), Entre Ríos Province, Argentina.
representing a Tortonian age (late Miocene; Cione et al., 2000;
Diagnosis. Giant catfish diagnosable on the basis of the following
Brandoni, 2013).
combination of characters (exclusive features marked by an asterisk*):
Unfortunately, as noted by Cione et al. (2009) most fossils lack
dorsal surface of frontal bones ornamented with ridges and tubercles;
adequate provenance. Labels in collections usually only state that the
single vomerine tooth plate; transversely narrow single dental patch on
material comes from the base of the cliffs near Paraná. In this sense,
the tooth plate with a continuous posterior margin; median posterior
recent fieldwork has confirmed that almost all of the Miocene terrestrial
process of vomerine tooth patch pointed and with dentate surface*;
and freshwater aquatic vertebrates come from the “Conglomerado
anterior margin of vomerine tooth plate gently convex; and vomerine
osífero” at the base of the Ituzaingó Formation (Cione et al., 2009).
tooth patch with well-developed posterolateral lobes.
Etymology. The species is dedicated to Alexandra Elbakyan, for her
4. Systematic paleontology
efforts in order to make available the scientific literature for researchers
around the world.
SILURIFORMES Hay, 1929.
Description. The skull is elongate, being longer than transversely
PIMELODIDAE sensu Lundberg and Littmann (2003).
wide (Fig. 2; maximum preserved length: 131 mm; maximum preserved
BRACHYPLATYSTOMATINI sensu Lundberg and Akama (2005).
width through the cornua: 73 mm; minimum transverse width of me-
Brachyplatystoma Bleeker 1862.
sethmoid: 57 mm; maximum length of mesethmoid 78 mm; length
Brachyplatystoma Bleeker, 1862 (in Bleeker, 1862–63): 10.

2
F.L. Agnolin and S. Bogan
Fig. 2. Brachyplatystoma elbakyani nov. sp. Holotype specimen (MACN Pv
16091) in A, dorsal, B, ventral views. Abbreviations: af, anterior fontanelle; cor,
cornua; fr, frontal; le, lateral ethmoid; mb, mesethmoid bulge; mes, mesetho-
moid, ob, orbithosphenoid bridge; or, orbithosphenoid; par, parasphenoid; v,
vomerine tooth plate. Scale bar: 1 cm [planned for single column].
between the anterior margin of mesethmoid and the anterior margin of
cranial fontanelle: 61 mm; skull height at the level of the mesethmoid
bulge: 12 mm; maximum preserved height of skull 23 mm; maximum
length of the vomerine tooth patch 28 mm; maximum transverse width
of vomerine tooth patch 47 mm; minimum transverse width of para-
sphenoid: 11.5 mm). In dorsal view, the skull roof is almost flat with
exception of the mesethmoid, which is convex and dorsally raised. The
snout is notably spatulate. The skull roofing bones articulate with
shallow to deeply interdigitating suture.
The mesethmoid is a robust and wide bone that shows a gently
convex anterior margin. It exhibits a nearly smooth and shallow cres-
cent-shaped transverse groove that delimits a dorsal bulge of the bone.
The bone surface ornamentation at the anterior portion of the skull is
markedly striated, especially at level of the mesethmoid bulge. The
mesethmoid bears robust but short cornua. There is no median notch.
The lateral ethmoid is notably anteroposteriorly elongate and inserts
between the frontal and the mesethmoid. Its lateral margin is not
complete, and thus, the orbital notch morphology is not certain.
The anterior cranial fontanelle is completely preserved (MACN Pv
16091). This fontanelle is narrowly open and is located within a
broader depression. It begins at the posterior rear of the mesethmoid
and is posteriorly delimited by the epiphyseal bar.
The frontal ornamentation is made up by subparallel ridges that
originate at the posterior end of the bone and slightly diverge ante-
riorly. The ridges bear tubercles along their length.
In ventral view the preserved portion of skull is homogeneously flat.
The premaxillae were not preserved and only their articular surfaces are
present in the anterior portion of the mesethmoid.
The vomer shows a single median tooth patch that is strongly an-
kylosed to the posterior portion of the mesethmoid. This patch is
3
Journal of South American Earth Sciences 100 (2020) 102551
Fig. 3. Brachyplatystoma elbakyani nov. sp. A, line drawing of modern B. vail-
lantii with superimposed shaded approximation of fossil specimen (MACN Pv
16091) in dorsal view; B–C, incomplete ethmoid/vomerine region of skull of
referred specimen MACN Pv 15989 in B, dorsal, and C, ventral views.
Abbreviations: ex, extrascapular; fr, frontal bone; le, lateral ethmoid; mes,
mesethmoid; pte, pterotic; soc, supraoccipital; sph, sphenotic. Scale bar: 1 cm
[planned for two columns].
anteroposteriorly long and transversely compressed. The anterior
margin of the patch is gently convex. The posterior margin of vomer
forms a median pointed process that exhibits a dentate surface, which is
also present, albeit smaller, in Brachyplatystoma capapretum (see
Lundberg and Akama, 2005, Fig. 7). This process is relatively short and
is laterally delimited by narrow but deep notches. The diameters of the
tooth bases indicate that the teeth were subequal in size and shape.
The parasphenoid is ventrally flat and shows anteriorly divergent
margins, resulting in a fan-shaped contour. The orbitosphenoid broadly
contacts the frontals dorsally and the parasphenoid ventrally, its side-
walls are relatively deep and conforms a longitudinal subhorizontal
shelf.
In addition to the holotype specimen a second incomplete individual
(MACN Pv 15989) is available (Figs. 3 and 5; maximum preserved
length: 78 mm; maximum preserved width through the cornua: 97 mm;
minimum transverse width of mesethmoid: 69 mm; skull height at the
level of the mesethmoid bulge: 12 mm; maximum length of the vo-
merine tooth patch 26 mm; maximum transverse width of vomerine
tooth patch 55 mm). The material is very similar to the holotype, but
can be distinguished because the vomerine tooth patch shows the
posterolateral projections strongly posteriorly extended.
Brachyplatystoma sp.
Referred material. MACN Pv 16052, three incomplete Weberian
apparatus with neural arches, spines, and most of the transverse pro-
cesses missing (Fig. 4).
Description. The Weberian apparatus (MACN Pv 16052; Fig. 4) are
incompletely preserved (Fig. 4). The base of the transverse process and
the anterolateral process are seen in lateral view.
F.L. Agnolin and S. Bogan
Fig. 4. Brachyplatystoma sp. Webberian complexes (MACN Pv 16052). A,D, anterior; B,E, left lateral; and C,F, ventral views. Abbreviations: aacf, aortic canal anterior
foramen; c1-cc, First centrum-compound centrum joint; cc-c5, compound centrum-fifth centrum joint; c5-c6, fifth centrum-sixth centrum joint; mve, midventral
expansion; na, neural arch; nc, neural canal; tr, transverse ridge; pacf aortic canal posterior foramen. Scale bar: 2 cm [planned for two columns].
The bone texture of the complex is densely laminar, with exception
of the first vertebra that shows a strongly spongy bone texture. The first,
compound, fifth and sixth vertebrae are enlarged and tightly jointed
together. The first centrum articulation surface for the basioccipital is
heart-shaped, slightly concave and surrounded by a thick rim, as
usually occurs in Brachyplatystoma (see Aguilera et al., 2013). As in
other catfishes, the first vertebra comprises a centrum only without
neural arches or transverse processes. The anterior articular surface of
the first vertebra shows the focus of growth rings located above the
center of the centrum at about one third of the depth below the dorsal
rim (Fig. 4 A,D).
In lateral view, only the base of the anterior and posterior limbs of
the fourth transverse process is observed. The anterior limb base is
anterolaterally oriented, whereas the posterior one is subhorizontally
extended. The sixth transverse process is much expanded and forms an
enlarged fossa that covers the posterior end of the gas bladder.
In dorsal view the bases of neural arches flank the neural canal
across the compound, fifth and sixth vertebrae. Ventrally the centrum
of the compound vertebra contains the large anterior foramen for
passage of the dorsal aorta into the aortic canal that is embedded within
the compound, fifth and sixth centra.
5. Discussion
The clade Sorubiminae include several genera, as well as the Tribe
Brachyplatystomatini (Lundberg and Akama, 2005). Despite the ab-
sence of clear osteological synapomorphies in the skull roof of the genus
4
Journal of South American Earth Sciences 100 (2020) 102551
Brachyplatystoma, the morphology of skull bones of this taxon is dis-
tinctive, and strikingly different from other sorubimines, specially re-
garding the mesethmoid shape (see details in Lundberg and Akama,
2005). In this way, specimen MACN Pv 16091 is referred to Brachy-
platystoma on the basis of the following combination of characters:
elongate and dorsoventrally flat skull with notably spatulate snout,
presence of a profusely striated mesethmoid bulge, and a single large
vomerine tooth patch. In other related genera, such as Platynema-
tichthys, the mesethmoid bulge is nearly smooth and is not prominent,
and vomerine tooth patch is much smaller and crescent-shaped, very
different from that reported for Brachyplatystoma (see Lundberg and
Akama, 2005, Fig. 6).
The presence of a single vomerine tooth patch is a feature that B.
elbakyani nov. shares with most species of Brachyplatystoma with the
exception of B. trgrinum, which shows a vomerine dentition that is
dispossed in two different patches (Lundberg and Akama, 2005). On the
other side, B. elbakyani nov. shows a wide tooth patch with a con-
tinuous posterior margin, that is different from B. juruense which shows
a deep and edentulous median notch. This notch is also present in B.
rousseauxii and B. platynemun that further differ from B. elbakyani nov.
in having a strongly convex anterior edge of the tooth patch, that results
in a subtriangular contour. Further, B. elbakyani nov. shows a robust
and transversely narrow tooth patch, that is different from the ante-
roposteriorly short and transversely wider condition exhibited by B.
capapretum and B. filamentosum.
Brachyplatystoma elbakyani nov. shows the presence of well-devel-
oped posterolateral lobes on the vomerine tooth patch (Fig. 5). This
F.L. Agnolin and S. Bogan
Fig. 5. Comparison between the vomerine region of different Brachyplatstoma species en ventral view. A, Brachyplatystoma elbakyani nov. sp. (holotype; MACN Pv
16091); B, Brachyplatystoma elbakyani nov. sp. (MACN Pv 15989); C, B. platynemum (ANSP 187321); D, B. rousseauxi (DUF 1057); E, B. juruense (ANSP 178514); F, B.
tigrinum (ANSP 179236). Scale bar: 1 cm. C–F, photographs courtesy of Kyle Lunckenbill. [planned for two columns].
condition shared with most species of the genus with exception of B.
vaillantii and B. tigrinum (Lundberg and Akama, 2005) (Fig. 5).
Furthermore, B. elbakyani nov. differs from other species in having
the posterior margin of vomer that shows a median pointed process that
exhibits a dentate surface. This projection is present in other species,
but lacks any sign of teeth.
Furthermore, the skull of B. elbakyani nov. differs from most
Brachyplatystoma species (i.e., B. juruense, B. platynemum, B. capapretum,
B. filamentosum, and B. rousseauxii) with exception of B. vaillantii and B.
tigrinum in having a rugose ornamentation on skull roof bones
(Lundberg and Akama, 2005, Fig. 5).
In addition, the weberian complexes here referred as
Brachyplatystoma sp. share with B. promagdalena, B. rousseauxii, B. fi-
lamentosum, and B. capapretum the apomorphic presence of spongy
texture of the first vertebral centrum (Lundberg, 2005). These elements
differ from the extinct B. promagdalena in lacking the fifth centrum with
a coarsely sculptured prominent midventral expansion (Lundberg,
2005). Further, fossil specimens here described differ from B. pro-
magdalena in some subtle features, including the anterolateral process
that does not extends as far anteriorly and in that the first centrum
seems to be anteroposteriorly longer (see Lundberg, 2005; Aguilera
et al., 2013).
Because the genus Brachyplatystoma includes six species that are
almost sympatric in distribution, and due to the dissociated nature of
the Weberian apparatuses here reported, it is not possible to refer these
elements to the specific level.
5
Journal of South American Earth Sciences 100 (2020) 102551
5.1. Palaeobiogeographical implications
Different data sources indicate that during the Paleocene
(65–56 Ma) or probably by the Late Cretaceous (Iriondo and Paira,
2007; Aguilera et al., 2013) freshwater fishes populated the proto
Amazon-Orinoco river valley that drained the Sub-Andean foreland
(Lundberg et al., 1998; Albert et al., 2018). Further, seasonal connec-
tions between the modern La Plata and Amazon Basins occurred at least
at two different places (Iriondo and Paira, 2007; Brea and Zucol, 2011).
This resulted that the Orinoco-Amazonas-Paraná-La Plata basin ex-
tended over 4000 km, from modern northern Argentina to Venezuela
and Colombia (Mora et al., 2010), resulting in a common ichthyofaunal
realm (Lundberg et al., 1998).
During the Neogene, tectonics, and climatic changes resulted on
important modifications in geographical connectivity between these
mega-wetlands, ranging from more connected in the late Miocene to
more fragmented by Pliocene to Recent times (Albert et al., 2018). The
above mentioned drainage connections allowed fishes to disperse be-
tween northern South America and Parana basins (Albert et al., 2011,
2018; Dias et al., 2014; Evans et al., 2017).
The genus Brachyplatystoma is nowadays distributed along the
lowlands of Orinoco and Amazonas basins and major rivers of the
Guianas (Lundberg and Akama, 2005).
The occurrence of Paraná fossils belonging to Brachyplatystoma, as
well as, other aquatic taxa such as Phractocephalus, are in agreement
with ancient wider-ranging Amazona and Orinoco faunas than today
F.L. Agnolin and S. Bogan
(Lundberg, 1997, 1998; Azpelicueta and Cione, 2016; Bogan and
Agnolín, 2019), and constitute the southernmost records for those gi-
gantic catfishes. In this sense, the co-occurrence of Phractocephalus and
Brachyplatystoma is currently restricted to the Amazonas and Orinoco
basins (Lundberg and Akama, 2005).
The presence of large-sized specimens Phractocephalus and
Brachyplatystoma is typical of large river channels (Lundberg, 2005;
Bogan and Agnolín, 2019). The co-occurrence of both taxa, suggests
that an important part of the fossil record at the Paraná site belonged to
main water courses of the paleobasin.
These, together with recent findings from Paraná (see Azpelicueta
and Cione, 2016) are playing an important role for understanding the
evolution and geographical distribution of Neotropical freshwater cat-
fishes during the Neogene.
CRediT authorship contribution statement
Federico L. Agnolin: Conceptualization, Data curation, Writing -
original draft, Investigation, Writing - review & editing. Sergio Bogan:
Conceptualization, Data curation, Writing - original draft, Investigation,
Writing - review & editing.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgements
Special thanks to M. Ezcurra, L. Chornogubsky and A. Martinelli
(MACN) who allowed access to the collections under his care. To A.
Giacchino (Fundación de Historia Natural “Félix de Azara”-Universidad
Maimónides) for his support during the confection of the present con-
tribution. We acknowledge to John Lundberg, Mark Sabaj Pérez and
especially Kyle Lunckenbill (Drexel University) for sending unpublished
information and photographs of diverse osteological specimens of
Platystomatichthys and Brachyplatystoma species. Special thanks to the
editor F.J. Veiga, and J. Lundberg, and two anonymous reviewers for
their enlightening comments on the Manuscript.
References
Aceñolaza, F.G., 2000. La Formación Paraná (Mioceno medio): estratigrafía, distribución
regional y unidades equivalentes. El Neógeno de Argentina: Ser. Correlación Geol. 14,
9–28.
Aguilera, O., Lundberg, J., Birindelli, J., Perez, M.S., Jaramillo, C., Sánchez–Villagra,
M.R., 2013. Palaeontological evidence for the last temporal occurrence of the ancient
western Amazonian River outflow into the Caribbean. PloS One 8 (9), e76202.
Albert, J.S., Petry, P., Reis, R.E., 2011. Major biogeographic and phylogenetic patterns.
In: Albert, J.S., Reis, R.E. (Eds.), Historical Biogeography of Neotropical Freshwater
Fishes. University of California Press, Berkeley, pp. 21–59.
Albert, J.S., Val, P., Hoorn, C., 2018. The changing course of the Amazon River in the
Neogene: center stage for Neotropical diversification. Neotrop. Ichthyol. 16,
e180033.
Alves, Y.M., Bergqvist, L.P., Brito, P.M., 2016. New occurrences of microvertebrate fossil
accumulations in Bauru group, late cretaceous of western Sao Paulo state, Brazil. J. S.
Am. Earth Sci. 69, 80–90.
Ameghino, F., 1889. Contribución al conocimiento de los mamíferos fósiles de la
República Argentina. Actas de la Academia Nacional de Ciencias de Córdoba 6,
1–1027.
Arratia, G., Kapoor, B., Chardon, M., Diogo, R., 2003. Catfishes, vols. 1 and 2 Science
Publishers, Enfield, NH.
Arratia, G., Cione, A.L., 1996. The record of fossil fishes of southern South America.
Müncher Geowissenschaftliche Abhandlungen. Reihe A. Geologie und Paläontologie
30, 9–72.
Azpelicueta, M.M., Cione, A.L., 2016. A southern species of the tropical catfish genus
Phractocephalus (Teleostei: Siluriformes) in the Miocene of South America. J. S. Am.
Earth Sci. 67, 221–230.
Barthem, R.B., Goulding, M., Leite, R.G., Cañas, C., Forsberg, B., Venticinque, E., Petry,
P., Ribeiro, M.L.B., Chuctaya, J., Mercado, A., 2017. Goliath catfish spawning in the
far western Amazon confirmed by the distribution of mature adults, drifting larvae
6
Journal of South American Earth Sciences 100 (2020) 102551
and migrating juveniles. Sci. Rep. 7, 41784.
Bleeker, P., 1858. De visschen van den Indischen Archipel. Beschreven en toegelicht.
Siluri. Verhandelingen der Koninklijke Natuurkundige Vereeniging in Nederlandsch
Indië 4, 1–370.
Bleeker, P., 1862. Descriptions de quelques espèces nouvelles de Silures de Suriname.
Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen
(Afdeeling Natuurkunde) 14, 371–389.
Bogan, S., Agnolín, F.L., 2019. Phractocephaline catfishes from the late Miocene of
Argentina, with the description of a new taxon. J. Vertebr. Paleontol. https://doi.org/
10.1080/02724634.2019.1676254.
Bogan, S., Sidlauskas, B., Vari, R.P., Agnolin, F.L., 2012. Arrhinolemur scalabrinii
Ameghino, 1898, of the late Miocene: a taxonomic journey from the Mammalia to the
Anostomidae (Ostariophysi: characiformes). Neotrop. Ichthyol. 10 (3), 555–560.
Boulenger, G.A., 1898. On a collection of fishes from the Rio Jurua, Brazil. Trans. Zool.
Soc. Lond. 14, 421–428.
Brandoni, D., 2013. Los mamíferos continentales del “Mesopotamiense” (Mioceno tardío)
de Entre Ríos, Argentina. Diversidad, edad y paleobiogeografía. In: In: Brandoni, D.,
Noriega, J.I. (Eds.), El Neógeno de la Mesopotamia Argentina, Asociación
Paleontológica Argentina, Publicación Especial, vol. 14. pp. 179–191 Buenos Aires.
Bravard, A., 1858. Monografıa de los terrenos marinos terciarios del Paraná. Diario oficial
del Gobierno. El Nacional Argentino, Buenos Aires.
Brea, M., Zucol, A.F., 2011. The Paraná–Paraguay basin: geology and paleoenvironments.
In: Albert, J.S., Reis, R.E. (Eds.), Historical Biogeography of Neotropical Freshwater
Fishes. University of California Press, Berkeley, pp. 69–87.
Britski, H.A., 1981. Sobre um novo gênero e espécie de Sorubiminae da Amazônia (Pisces,
Siluriformes). Papeis Avulsos de Zoologia 34, 109–114.
Brunetto, E., Noriega, J.I., Brandoni, D., 2013. Sedimentología, estratigrafía y edad de la
Formación Ituzaingó en la provincia de Entre Ríos, Argentina. In: Brandoni, D.,
Noriega, J.I. (Eds.), El Neógeno de la Mesopotamia Argentina, Asociación
Paleontológica Argentina, Publicación Especial 14, pp. 13–27 Buenos Aires.
Castelnau, F.L., 1855. Poissons. In: Bertrand, P. (Ed.), Animaux Nouveaux or Rares
Recueillis Pendant l'Expédition dans les Parties Centrales de l'Amérique du Sud, de
Rio de Janeiro à Lima, et de Lima au Para; Exécutée par Ordre du Gouvernement
Français Pendant les Années 1843 a 1847, pp. 1–112 Bertrand, Paris.
Cione, A.L., Azpelicueta, M.M., Bond, M., Carlini, A.A., Casciotta, J.R., Cozzuol, M.A., de
la Fuente, M., Gasparini, Z., Goin, F.J., Noriega, J., Scillato–Yané, G.J., Soibelzon, L.,
Tonni, E.P., Verzi, D., Vucetich, M.G., 2000. Miocene vertebrates from Entre Ríos
province, eastern Argentina. El Neógeno de Argentina: Ser. Correlación Geol. 14,
191–237.
Cione, A.L., Dahdul, W., Lundberg, J., Machado–Allison, A., 2009. Megapiranha para-
nensis, a new genus and species of serrasalmidae (characidae, Teleostei) from the
upper Miocene of Argentina. J. Vertebr. Paleontol. 29, 350–358.
Cuvier, G., 1829. Le Règne Animal Distribué d'après son Organisation pour Servir de Base
à l'Histoire Naturelle des Animaux et d'Introduction à l'Anatomie Comparée. In:
Nouvelle Éd., rev. et augm., Tome 2–Les Reptiles et les Poissons: Deterville, Paris.
Dias, M.S., Oberdorff, T., Hugueny, B., Leprieur, F., Jézéquel, C., Cornu, J.F., Brosse, S.,
Grenouillet, G., Tedesco, P.A., 2014. Global imprint of historical connectivity on
freshwater fish biodiversity. Ecol. Lett. 17 (9), 1130–1140.
Evans, K.M., Crampton, W.G.R., Albert, J.S., 2017. Taxonomic revision of the deep
channel electric fish genus Sternarchella (Teleostei: Gymnotiformes: Apteronotidae)
with descriptions of two new species. Neotrop. Ichthyol. 15 (2), e160168.
Gayet, M., Meunier, F.J., 2003. Palaeontology and palaeobiogeography of catfishes. In:
Arratia, G., Kapoor, B.G., Chardon, M., Diogo, R. (Eds.), Catfishes. Science Publishers,
Inc., Enfield, NH, USA, pp. 491–522.
Hay, O.P., 1929. Second bibliography and catalogue of the fossil Vertebrata of North
America. Publ. Carnegie Instit. Wash. 390, 1–2003.
Iriondo, M.H., Paira, A.R., 2007. Physical geography of the basin. In: Iriondo, M.H., Paggi,
J.C., Parma, M.J. (Eds.), The Middle Paraná River: Limnology of a Subtropical
Wetland. Springer–Verlag, Berlin Heidelberg, pp. 7–31.
Lichtenstein, M.H.C., 1819. Ueber einige neue Arten von Fishen aus der Gattung Silurus.
Zoologisches Magazin 1, 57–63.
Lundberg, J.G., 1997. Fishes of the Miocene La Venta Fauna: additional taxa and their
paleobiotic implications. 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, pp. 67–91.
Lundberg, J.G., 1998. The temporal context for the diversification of Neotropical fishes.
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, Porto Alegre, Brazil. EDIPUCRS,
pp. 49–68.
Lundberg, J.G., 2005. Brachyplatystoma promagdalena, new species, a fossil goliath catfish
(Siluriformes: Pimelodidae) from the Miocene of Colombia. S. Am.: Neotrop.
Ichthyol. 3 (4), 597–605.
Lundberg, J.G., Akama, A., 2005. Brachyplatystoma capapretum: a new species of goliath
catfish from the Amazon basin, with a reclassification of allied catfishes
(Siluriformes: Pimelodidae). Copeia 2005 (3), 492–516.
Lundberg, J.G., Littmann, M.W., 2003. Pimelodidae (Long–whiskered catfishes). In: Reis,
R.E., Kullander, S.O., Ferraris, C.J. (Eds.), Check List of the Freshwater Fishes of
South and Central America, pp. 432–446 Edipucrs.
Lundberg, J.G., Marshall, L.G., Guerrero, J., Horton, B., Malabarba, M.C.S.L., 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, Porto Alegre, Brazil.
EDIPUCRS, pp. 13–48.
Lundberg, J.G., Sullivan, J.P., Hardman, M., 2011. Phylogenetics of the South American
catfish family Pimelodidae (Teleostei: Siluriformes) using nuclear and mitochondrial
gene sequences. Proc. Acad. Nat. Sci. Phila. 161, 153–190.
F.L. Agnolin and S. Bogan
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 develop-
ment of the Amazon drainage basin. In: Hoorn, C.M., Wesselingh, F.P. (Eds.),
Amazonia, Landscape, and Species Evolution: A Look into the Past. John Wiley &
Sons, New York, pp. 38–60.
Nass, P., 1991. Anatomia comparada del bagre cunagaro Brachyplatystoma juruense
(Boulenger, 1898), incluyendo un analisis filogenético de la familia Pimelodidae.
Unpublished Ph. D. dissertation. Universidad central de Venezuela, Caracas, pp. 262.
Nelson, J.S., 2006. Fishes of the World, fourth ed. John Wiley, New York, pp. 624.
Reis, R.E., Kullander, S.O., Ferraris, C.J., 2003. Check List of the Freshwater Fishes of
South and Central America. Edipucrs.
Schultz, L.P., 1944. The catfishes of Venezuela, with descriptions of thirty-eight new
forms. Proc. U. S. Natl. Mus. 94, 173–338.
Journal of South American Earth Sciences 100 (2020) 102551
Sullivan, J.P., Muriel–Cunha, J., Lundberg, J.G., 2013. Phylogenetic relationships and
molecular dating of the major groups of catfishes of the Neotropical superfamily
Pimelodoidea (Teleostei, Siluriformes). Proc. Acad. Nat. Sci. Phila. 162, 89–110.
Valenciennes, A., 1835–1847. Poissons; Catalogue des principales espèces de Poissons,
rapportées de l'Amérique méridionale. In: d'Orbigny, A. (Ed.), Voyage dans
L’Amérique Méridionale (le Brésil, la République Orientale de l'Uruguay, la
République Argentine, la Patagonie, la République du Chili, la République de Bolivia,
la République du Pérou), Exécuté Pendant les Années 1826, 1827, 1828, 1829, 1830,
1832 et 1833. Bertrand et Levrault, Paris.
van der Laan, R., Fricke, R., Eschmeyer, W.N., 2020. Eschmeyer's Catalog of Fishes.
scientists/catalog–of–fishes–classification/Electronic version. http://www.
calacademy.org/, Accessed date: 17 February 2020.