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New records of the enigmatic Neotropical fossil fish Acregoliath rancii

(Teleostei incertae sedis) from the middle Miocene Honda Group of Colombia

Article  in  AMEGHINIANA · December 2019

DOI: 10.5710/AMGH.17.09.2019.3266

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 NEW RECORDS OF THE ENIGMATIC
NEOTROPICAL FOSSIL FISH ACREGO-
LIATH RANCII (TELEOSTEI INCERTAE
SEDIS) FROM THE MIDDLE MIOCENE
HONDA GROUP OF COLOMBIA

GUSTAVO A. BALLEN1
JORGE W. MORENO-BERNAL2

1
Museu de Zoologia da Universidade de São Paulo, CEP 04263-000, São Paulo, SP, Brazil.
2
Grupo de Investigación en Geociencias GEO4, Universidad del Norte, Km 5 Vía Puerto Colombia, Barranquilla, Atlántico, Colombia.

Submitted: June 10th, 2018 - Accepted: September 17th, 2019 - Published online: October 7th, 2019

To cite this article: Gustavo A. Ballen, and Jorge W. Moreno-Bernal (2019). New records of the enigmatic
neotropical fossil fish Acregoliath rancii (Teleostei incertae sedis) from the middle Miocene Honda Group of
Colombia. Ameghiniana 56: 431–440.
To link to this article: http://dx.doi.org/10.5710/AMGH.17.09.2019.3266

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Scales of the enigmatic neotropical Gastropods, bivalves, and sepulids Experimental dissolution of diatoms
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in the Miocene of South America. diversity related to salinity conditions. to changes in water chemistry.
 AMEGHINIANA - 2019 - Volume 56 (6): 431–440 ARTICLES

ISSN 0002-7014

NEW RECORDS OF THE ENIGMATIC NEOTROPICAL FOSSIL FISH

ACREGOLIATH RANCII (TELEOSTEI INCERTAE SEDIS) FROM THE
MIDDLE MIOCENE HONDA GROUP OF COLOMBIA
GUSTAVO A. BALLEN1, AND JORGE W. MORENO-BERNAL2

1
Museu de Zoologia da Universidade de São Paulo, CEP 04263-000, São Paulo, SP, Brazil. gaballench@gmail.com
2
Grupo de Investigación en Geociencias GEO4, Universidad del Norte, Km 5 Vía Puerto Colombia, Barranquilla, Atlántico, Colombia. jwmoreno@uninorte.edu.co

Abstract. Fossil scales of the enigmatic teleost Acregoliath rancii Richter, collected in 1945 and 1946, are described for the first time from fine-
grained sediments of the La Victoria Formation of the Honda Group in central Colombia. The specimens include one isolated but nearly com-
plete scale from the anterior region of the body, and one isolated scale fragment. Although fragmentary, these specimens provide relevant
anatomic and biogeographical information. Both specimens agree in most details with the type material of A. rancii; however, the almost com-
plete scale differs in focus position and outline, thus suggesting topological variation. This aspect of lepidosis in A. rancii was previously unknown
and could be relevant for future reassessment of the interrelationships of this taxon. The presence of A. rancii in the middle to late Miocene
fossil assemblages from La Venta, Fitzcarrald, and Acre suggests a relationship between these areas during the middle Miocene. The rele-
vance for paleodrainage evolution in northern South America is discussed in the context of these findings.
Key words. Lepidosis. Freshwater. Fitzcarrald. Acre. Biogeography.

Resumen. NUEVOS REGISTROS DEL ENIGMÁTICO PEZ FÓSIL ACREGOLIATH RANCII (TELEOSTEI INCERTAE SEDIS) DEL GRUPO HONDA, MIOCENO
MEDIO DE COLOMBIA. Se describen por primera vez escamas fósiles del enigmático teleósteo Acregoliath rancii Richter colectadas en 1945 y
1946 en sedimentos de grano fino de la Formación La Victoria, Grupo Honda, en Colombia central. Los especímenes corresponden a una es-
cama aislada de la porción anterior del cuerpo y a un fragmento aislado de escama. Aunque fragmentarios, estos especímenes proporcionan
valiosa información anatómica y biogeográfica. Ambos especímenes concuerdan en la mayoría de detalles con el material tipo de A. rancii,
sin embargo, la escama más completa difiere en la posición del foco y la forma general, sugiriendo variación topológica. Este aspecto de la le-
pidosis en A. rancii era previamente desconocido y podría ser relevante para futuras reinterpretaciones sobre las interrelaciones del taxón. La
presencia de A. rancii en ensambles del Mioceno medio a tardío de La Venta, Fitzcarrald y Acre sugiere una relación entre dichas áreas durante
el Mioceno medio. Se discute la relevancia de dicho hallazgo en la evolución de los paleodrenajes en el norte de América del Sur.
Palabras clave. Lepidosis. Agua dulce. Fitzcarrald. Acre. Biogeografía.

WITH more than 300 species per million km2, South America
harbors the richest extant freshwater fish fauna, followed
by Tropical Asia with almost half of that density (Albert and
Reis, 2011). This enormous diversity evolved through time
on a relatively stable continental plate that has remained
isolated from other land masses since the breakup of Gond-
wana during the early Cretaceous (Cavin, 2008). The fossil
record of freshwater fishes is, however, mostly restricted
to the Cenozoic (Lundberg et al., 2010) with scattered oc-
currences of Cretaceous age (Alveş et al., 2016; Alveş et al.,
2019). The fossil record of the Cenozoic suggests that the
South American freshwater fish fauna is highly conserva-
tive in composition, with virtually the same genera and fami-
lies represented since the Paleocene to the present
(Lundberg et al., 1986, 2010; Gayet et al., 2001). In some
cases, fossils recovered from Neogene deposits have been
referred to extant species (Lundberg et al., 1986; Lundberg
and Chernoff, 1992; Lundberg, 1997).
Among South American freshwater fish fossils, the most
enigmatic example is Acregoliath rancii, an incertae sedis in
the Teleostei (Richter, 1989), represented only by isolated
scales collected in Amazonian localities of Miocene age
(Lundberg et al., 2010; Tejada-Lara et al., 2015). Given the
fragmentary nature of its remains (partial to complete
scales) very little is known about its morphology and inter-
relationships. Studies on scale morphology have suggested
similarities with Dipnoans, Osteoglossomorphs, Rhipidis-
tians, Cladistians, and Loricarioid siluriforms, based on his-

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AMEGHINIANA - 2019 - Volume 56 (6): 431–440
tological similarities between Acregoliath rancii and both ex-
tant and extinct representatives of each of these groups
(Richter, 1989). Most of these similarities, however, are su-
perficial, ambiguous, or of no comparative value, making
Acregoliath rancii a very distinctive taxon due to the mor-
phology of its scales, but of uncertain relationships. Macro-
scopically, scales of Acregoliath can be distinguished by its
large size, thickness, and presence of a posterior field cov-
ered by bright tissue comprising 0.5 to 0.33 of the scale area
and bearing thick subparallel ridges (“ribs” in the original
description of Richter, 1989). This morphology is similar to
that seen in the scales of extant Amazonian osteoglosso-
morphs, and to some degree, extant dipnoans. However,
ornamentation ridges in the scales of those taxa form a
reticulated, rather than subparallel pattern, and lack the
characteristic layer of bright tissue on the posterior field.
These characters were already discussed by Richter (1989)
when assessing the possible relationships of Acregoliath rancii.
Fossils from the La Victoria and Villavieja formations of
the Honda Group in the Upper Magdalena Valley of Colom-
bia comprise one of the most complete continental verte-
brate assemblages in the Neogene of tropical South
America (Kay et al., 1997). The La Venta fauna, as this as-
semblage is known, includes a wide diversity of terrestrial
and freshwater vertebrate taxa, and it has been interpreted
as one of the best-preserved examples of Amazonian envi-
ronments in the fossil record (Lundberg et al., 1998, 2010).
On the other hand, La Venta is located outside the modern
Amazon basin, in an area without drainage or landscape
continuity with Amazonian drainages. Together with the in-
formation on past Amazonian environments, La Venta fauna
and its geologic context have provided evidence that allows
inferring past drainage connections between areas now
separated by the Andes. This is particularly true for the case
of fossil fishes, for which all taxa preserved in the La Venta
fauna have been referred to groups now restricted to
drainages east of the Andes, particularly in the Amazon and
Orinoco basins (Lundberg et al., 1986, 2010; Lundberg and
Chernoff, 1992; Lundberg, 1997, 2005). The high diversity
and geographic position of La Venta fauna have triggered
a considerable field of study that has systematically re-
covered relationships between areas east and west of the
Andes, thus conforming to the geological drainage model
suggested by this assemblage (Albert et al., 2006, 2011;
432
Rodríguez-Olarte et al., 2011).
The aim of this work is to expand our current knowledge
on the La Venta freshwater fossil assemblage, with the ad-
dition of previously unreported remains of Acregoliath rancii,
from the La Victoria Formation of the Honda Group.
MATERIALS AND METHODS
The fossil specimens herein studied are housed at the
University of California Museum of Paleontology and are
part of earlier collections made by Royo y Gómez, Stirton,
and others (Royo y Gomez, 1946; Stirton, 1953a). Duke lo-
calities (e.g., Duke locality 26) are named according to the
code numbers in Kay et al. (1997). A recent specimen of
Arapaima gigas (MZUSP 21476) was examined for compari-
son and characterization of scale shape associated to spa-
tial location. Lateral-line scale location is termed LL along
with the scale number (e.g., LL34 is the 34th scale along the
lateral line). Stratigraphic nomenclature follows Guerrero
(1997). The referral of these specimens to Acregoliath rancii
is based on observations of the external morphology, while
histological examination was not performed due to the
scarcity of specimens. Therefore, the taxonomic assignment
is limited to only one of the evidence lines used in the origi-
nal description of the species by Richter (1989). Institutional
abbreviations are University of California Museum of Paleon-
tology, California, USA (UCMP) and Museu de Zoologia da
Universidade de São Paulo, São Paulo, Brazil (MZUSP).
GEOLOGICAL CONTEXT, PALEOENVIRONMENT, AND AGE
Specimens described here were collected in UCMP lo-
cality V4531, near the hill known as Cerro Gordo in La Venta
area, municipality of Villavieja, Huila Department, Colombia
(Fig. 1). Sedimentary rocks of the Honda Group crop out
in La Venta area, forming the badlands known as “Tatacoa
Desert” (Royo y Gomez, 1946; Stirton, 1953a). The Honda
group is largely comprised by volcanic litharenites, mud-
stones and conglomerates. This unit unconformably
contacts Mesozoic and Paleogene units of the Upper Mag-
dalena valley. In the La Venta area, the Honda Group over-
lies the Jurassic andesites of the Saldaña Formation, which
comprise the Cerro Gordo and Chacarón hills (Guerrero,
1997).
Two stratigraphic units can be distinguished in the
Honda Group (Fig. 2). The La Victoria Formation is composed
 BALLEN AND MORENO-BERNAL: ACREGOLIATH SCALES FROM COLOMBIA

Figure 1. 1, Map showing the geographical positions of Acregoliath rancii occurrences in equatorial South America: 1, middle Miocene La Venta
Fauna, Honda Group, Huila Department, Colombia; 2, Acre Fauna, late Miocene Solimoes Formation, Acre State, Brazil; 3, middle Miocene Fitz-
carrald Fauna, Ipururo Formation, Madre de Dios Department, Peru; 2, Geographical position of the La Venta area in the Huila Department of
Colombia. 3, Simplified geologic map (modified from Guerrero, 1997; Fuquen et al., 2003), showing the position of UCMP locality V4531.

mainly by lithic arenites rich in volcanic fragments, together
with reddish brown and greenish grey mudstones. Thick
sandstone beds allow defining divisions within the unit. The
fossils included in this study were collected in strata be-
tween the Chunchullo Sandstone beds and the Tatacoa
Sandstone beds (Fig. 2). The uppermost bed of the La Vic-
toria Formation is a thick, clast-supported conglomerate,
which allows correlating the Honda Group in several locali-
ties of the Upper Magdalena Valley (Guerrero, 1997). The
top of this Cerbatana Conglomerate marks the contact be-
tween La Victoria Formation and the overlaying Villavieja
Formation.
The Villavieja Formation comprises mostly red mud-
stones alternating with reddish yellow sandstones. Gray
greenish mudstones and grey volcanic litharenites occur in
minor proportion. The Villavieja Formation is divided in
two members, on the basis of lithological differences. The
Baraya Member is predominantly composed by grey and red
mudstones, with intercalated beds of volcanic arenites. In
the Cerro Colorado Member, there are thick horizons of red
mudstones and fine sandstones, together with volcanic
litharenites and chert litharenites (Guerrero, 1997). The top
of the Villavieja Formation is overlain by the predominantly
conglomeratic Neiva Formation.

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AMEGHINIANA - 2019 - Volume 56 (6): 431–440

A middle Miocene (Serravalian to early Tortonian: 13.8–
10.1 Ma) age has been assigned to the Honda Group, on the
basis of magnetostratigraphy and 40
Ar/ 39 Ar analyses
(Flynn et al., 1997). The age of the specimens included in
this study is further constrained by radiometric dates of
13.767±0.052 Ma and 13.342±0.408 Ma obtained from
both the Chunchullo Sandstone beds and levels closer to the
Tatacoa Sandstone beds (Flynn et al., 1997; Guerrero, 1997).
Most deposits of the Honda Group represent fluvial de-
positional environments, dominated by meandering rivers.
The Cerbatana Conglomerate at the top of the La Victoria
Formation was likely deposited by braided rivers, whereas
anastomosing stream deposits are found towards the top
of the Villavieja Formation (Guerrero, 1997). Freshwater fish

Figure 2. Stratigraphic column of the Honda Group (modified from Guerrero, 1997), showing the stratigraphic position of the Acregoliath rancii
specimens, together with previously published fish occurrences (Lundberg, 1997; 2005; Lundberg and Chernoff, 1992).

434
 BALLEN AND MORENO-BERNAL: ACREGOLIATH SCALES FROM COLOMBIA

fossils have been recovered from different levels of the La
Victoria and Villavieja formations (Fig. 2). Three of the taxa
reported by Lundberg (1997), occur in Duke Localities be-
tween the Chunchullo and Tatacoa Sandstone beds (Hy-
drolycus sp., Duke Locality 38; Lepidosiren paradoxa, Duke
Localities 39, 62, 81, 84, 88, 90, 102; and cf. Leporinus, Duke
Locality 26). Acregoliath rancii (UCMP Locality V5431) adds
to the occurrences known from the stratigraphic interval
between these two sandstone beds. Among fossil fresh-
water fishes described for the Honda Group, only A. rancii
and cf. Leporinus are so far restricted to the La Victoria For-
mation (Fig. 2). Other vertebrate fossils from UCMP locality
V4531 include the holotypes of the Turtle Podocnemis
pritchardi and the Litoptern Villarroelia totoyoi, as well as re-
ferred specimens of Podocnemis sp., Chelus colombianus, and
Miocochilius anomopodus (Cifelli and Guerrero, 1997; Stirton,
1953b; Wood, 1976, 1997).
SYSTEMATIC PALEONTOLOGY
Infraclass TELEOSTEI Müller, 1845
Family ACREGOLIATHIDAE Richter, 1989
Genus Acregoliath Richter, 1989
Type species. Acregoliath rancii Richter, 1989, Teleostei incertae sedis,
middle to late Miocene. Type by original designation.

Figure 3. 1–3. Specimens of Acregoliath rancii from the La Victoria Fm; 1, complete scale UCMP 38082; 2, reconstruction of overlapping scale
outlines on the preserved ornament margins of UCMP along with the position of the focus (*); shaded contour lines indicate portions of the
scale that are covered underneath adjacent scales; 3, partial scale UCMP 38835 preserving the area between the anterior and posterior
fields. Scale bars= 10 mm.

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AMEGHINIANA - 2019 - Volume 56 (6): 431–440
Acregoliath rancii Richter, 1989
Figure 3.1–3
Remains examined. UCMP 38082, 1, nearly complete scale,
coll. J. Royo y Gómez, 1945; UCMP 38835, 1, fragment of
scale, coll. J. Royo y Gómez, 1945.
Locality and Horizon. Both specimens were collected in
UCMP locality V4531, near Cerro Gordo hill, ca. 13.5 Km
northeast from the town of Villavieja, Huila Department,
Upper Magdalena Valley, Colombia (Fig. 1). La Victoria
Formation, unit between the Chunchullo and Tatacoa
sandstone beds (Fig. 2), middle Miocene, 13.76 to 13.34 Ma
(Flynn et al., 1997; Guerrero, 1997).
Description of external morphology. Isolated cycloid scales
one nearly complete (UCMP 38082) and fragmentary
(UCMP 38835). Scale outline roughly tetragonal with ante-
rior margin concave and posterior margin convex (Fig. 3.1).
Circuli asymmetrical, wider and more developed on poste-
rior portion. Focus located roughly about 0.2 of scale length
along anteroposterior axis, barely visible, still receiving all
radii (Fig. 3.2). Posterior field comprising nearly 0.33 of
scale. Birefringent tissue (Richter, 1989) defining posterior
field, forming longitudinal ridges of varied degrees of con-
tinuity; some ridges interleaved by aligned tubercles (Fig.
3.3). Three scales superposed on each scale as suggested
by the marks on posterior field (Fig. 3.2).
DISCUSSION
Scale morphology
Very little is known about the lepidosis pattern in Acre-
goliath rancii, and UCMP 38082 sheds some light on the
pattern or scale variation along the body. As illustrated by
Richter (1989), in the holotype of Acregoliath rancii the
shape of the unornamented portion of the scale indicates
the presence of three overlapping scales. This pattern of
overlap coincides with that of UCMP 38082 (Fig. 3.2). Other
aspects of the holotype morphology differ from that of the
scales herein described. Richter (1989) described Acrego-
liath rancii based on scales with a symmetrical focus and
widely oval outline, comparable to midbody scales of other
teleosts.
On the other hand, UCMP 38082 is higher than wide,
with an anteriorly displaced focus, and considerably nar-
rower circuli on the region anterior to the focus. Even
436
though these differences may suggest that UCMP 38082
represents a different species within the genus, we interpret
this particular morphology as the result of topological origin.
Scales from the anterior portion of the body are commonly
narrower and asymmetrical when compared to those on the
sides of the body at mid-length level in teleosts, and this
can be seen in Arapaima gigas as a functional analog to Acre-
goliath rancii (Fig. 4). The portion of the scale anterior to the
focus, much narrower than the regions posterior to that
landmark, indicates that there was no room for accommo-
dating a wider scale, due to its proximity to the pectoral
girdle or cranium (compare Figs. 3.1 and 4.4).
Connections among middle and late Miocene faunas in
Amazonia
The genus Acregoliath was originally restricted to the
Lula Locality of the middle to late Miocene Solimões For-
mation, west of the Sena Madureira municipality in Acre
(Brazil). Lula is part of a set of localities along the border be-
tween the states of Acre and Amazonas, globally referred
to as the Acre (local) fauna (Kay and Cozzuol, 2006; Aguilera
et al., 2008; Souza-Filho et al., 2019). A Late Miocene age
has been attributed to the outcrops representing the upper
part of the Solimões Formation, based on palynology and
vertebrate biostratigraphy (Latrubesse et al., 2010). In par-
ticular, mammal fossil assemblages from some localities
have strong affinities with Huayquerian (9–6.5 Ma) South
American Land Mammal Age (SALMA) faunas from Argen-
tina (Cozzuol, 2006). However, correlations between lo-
calities in the Acre basin are difficult due to the limited
lateral continuity of outcrops, as well as to the extensive re-
verse faulting associated to the Andean orogeny (Cunha,
2007; Souza-Filho and Guilherme, 2015).
Based on U-Pb dating of detrital zircons, Bissaro-Júnior
et al. (2019) constrained the maximum depositional age of
two fossil vertebrate localities of the Solimões Formation:
Niterói at 8.5±0.5 Ma and Talismã at 10.89±0.13 Ma. Ac-
cording to Souza-Filho and Guilherme (2015), the Lula Lo-
cality is located between the localities Talismã and Niterói
(although closer to Talismã). Therefore, vertebrate fossils
from the Lula locality are likely to be Late Miocene (Tor-
tonian: 11.63 to 7.24 Ma) in age.
Acregoliath has also been reported, together with other
fishes of Amazonian distribution, as part of the vertebrate
 BALLEN AND MORENO-BERNAL: ACREGOLIATH SCALES FROM COLOMBIA

Figure 4. 1–8. Scales of Arapaima gigas from different portions of the body in the specimen MZUSP 21476; 1, first scale of the mid-dorsal
series just posterior to the supraoccipital; 2, one scale above LL3; 3, two scales above LL18; 4, two scales below LL3; 5, first ventral scale
before insertion of the pectoral-fin in ventral view; 6, first scale posterior to pectoral-fin insertion; 7, first scale below LL4; 8, fourth scale
below LL34. Scale bars= 1 mm.

fossil assemblage form the Ipururo Formation of Peru, known
as the Fitzcarrald Arch local fauna (Tejada-Lara et al., 2015).
Based on paleomagnetic data and mammal occurrences
shared with the La Venta fauna, a late middle Miocene
(Laventan SALMA: 13.8–11.8 Ma) age has been attributed
to the Ipururo Formation by Antoine et al. (2007). The pres-
ent record of Acregoliath reinforces the faunistic similarity
between the assemblages of the Ipururo Formation and the
Honda Group.
The La Venta fauna of the Honda Group is one of the
richest Neogene fossil freshwater fish assemblage with
23 taxa (Lundberg, 1997, 2005; Lundberg et al., 2010), in-
cluding the occurrences of Acregoliath rancii herein reported.
The composition of this assemblage is an important tool to
reconstruct ancient environments (Lundberg, 1997), and
most important, as evidence for persistent hydrographic
connections in equatorial South America during the Miocene
(Lundberg et al., 2010). The fish assemblage from the Honda
Group records a phase where the Eastern Cordillera had not
completely separated drainage systems east and west of
the Andes. Despite the uncertain systematic position of
Acregoliath rancii, its presence in the La Venta, Fitzcarrald,
and Acre faunas (Fig. 1.1) provides additional support for
drainage connections between the Magdalena and Amazon
basins in the middle to late Miocene.

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AMEGHINIANA - 2019 - Volume 56 (6): 431–440
Paleoecology
Fish fossils from the Honda Group comprise an assem-
blage typical of Neotropical lowland fluvial systems, in-
cluding taxa occupying a wide diversity of niches in
heterogeneous aquatic environments (Lundberg, 1997;
Lundberg et al., 2010). Most of the La Victoria Formation
was deposited by meandering rivers leaving channel sand-
stones and muddy overbank deposits. In particular, the unit
between the Chunchullo and Tatacoa sandstone beds is
predominantly fine-grained unit, sandstone channels within
this interval have little lateral extent and thickness (Gue-
rrero, 1997). The most common taxon in the unit between
the Chunchullo and Tatacoa sandstone beds is the lungfish
Lepidosiren paradoxa. The lungfish is restricted to still and
stagnant waters of flooded forests, swamps and lagoons in
the Amazon and Paraná basins as well as in the Guyanas
(Almeida-Val et al., 2011). The presence of this taxon in this
part of the La Victoria Formation is consistent with the low-
energy conditions indicated by the fine-grained sediments
comprising the unit between the Chunchullo and Tatacoa
sandstone beds. Other taxa from the same stratigraphic in-
terval are less indicative of a particular environment, either
due to taxonomic uncertainty (e.g., cf. Leporinus), whereas
others, such as Hydrolycus sp., are known to dwell in a variety
of freshwater habitats (Toledo-Piza et al., 1999).
Body size and scale morphology of the largest living
Amazonian fish, the osteoglossiform Arapaima, make this
species a likely ecological analog for Acregoliath. Arapaima is
present in both river channels and the still waters of
swamps, lagoons, and flooded forests. However, Arapaima
shows preference for lentic environments and its presence
in lotic systems seems to be a consequence of the periodi-
cal level decrease in the Amazon (Castello, 2008). If Acrego-
liath was strongly dependent on large, lentic environments,
its extinction may be linked to hydrographic changes asso-
ciated with Andean uplift and subsequent drainage isola-
tion. These changes include the disappearance of the
mega-wetland Pebas and Acre systems, together with the
onset of the modern river systems east and west of the
Andes (Wesselingh et al., 2009). A relationship between hy-
drographic change and Neogene extinctions in equatorial
South America has also been proposed for crocodylians, a
group for which high diversity levels have been linked to the
438
presence of mega-wetland systems (Scheyer and Moreno-
Bernal, 2010; Aureliano et al., 2015; Moreno-Bernal et al.,
2016). Acregoliath rancii is restricted to the late middle to
late Miocene of equatorial South America, a geographic and
stratigraphic distribution of this taxon that coincides with
that of high-diversity crocodilian faunas.
Despite the very fragmentary nature of the fossil re-
mains from which Acregoliath rancii is currently known, its
stratigraphic and geographic context reveals its relevance
to environmental changes in Amazonia during geological
time. We expect that a better understanding of its relation-
ships among Teleostei may help to refine these inferences,
allowing a better understanding of the ecological role of this
enigmatic freshwater fish in South America.
ACKNOWLEDGMENTS
GAB was funded through a doctoral scholarship (process 2014/
11558-5) a BEPE internship (process 2016/02253-1), and a the-
matic project (process 2016/19075-9) by the Fundação de Amparo
à Pesquisa do Estado de São Paulo (FAPESP), and the Böhlke en-
dowment of the Academy of Natural Sciences of Drexel University.
JWMB was funded by the Doris O. and Samuel P. Welles Research
Fund (University of California Museum of Paleontology). We thank
P.A. Holroyd (University of California Museum of Paleontology) for
access to the La Venta collections at UCMP. GAB thanks J. Lundberg,
M. de Pinna, and M. Pastana for discussions on the morphological
affinities of Acregoliath and other Neotropical Neogene fossil fishes,
and S. Reinales for discussion and support. JWMB thanks M. Molina
and M. Moreno for their continuous support. The authors thank the
diligence and insightful reviews by A. Lopez Arbarello, N. Toledo, and
two anonymous reviewers.
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