Journal of South American Earth Sciences 101 (2020) 102588

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Journal of South American Earth Sciences

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New outcrop with vertebrate remains from Solimões Formation T

(Eocene–Pliocene), Southern Solimões Basin, Acre State, Northern Brazil
Mauro Bruno da Silva Lacerdaa,b,∗, Lucy Gomes de Souzac, Leonardo S. Lobod,
Carlos Ernesto G.R. Schaefere, Pedro Seyferth R. Romanoa
a
Departamento de Biologia Animal, Universidade Federal de Viçosa, Viçosa, Minas Gerais, Brazil
b
Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil
c
Museu da Amazônia, Manaus, Amazonas, Brazil
d
Departamento de Geologia e Paleontologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil
e
Departamento de Solos, Universidade Federal de Viçosa, Viçosa, Minas Gerais, Brazil

ARTICLE INFO ABSTRACT

Keywords: Here we describe fossil vertebrate remains from a new Solimões Formation (Eocene-Pliocene) locality, near the
Cenozoic highway BR 364 at Sena Madureira municipality (Acre, Brazil). The findings comprise more than forty-four
Crocodyliformes isolated fragments from which twenty-three specimens could be properly identified anatomically and system-
Notoungulata atically. The materials include at least three different Caimaninae genera (Caiman, Melanosuchus, and
Proto-Amazon
Mourasuchus), two distinct side-necked turtles (Chelus and Podocnemididae) and one mammal (Toxodontidae).
Taphonomy
Testudines
The studied fossils could be systematized into two taphonomic classes based on their preservational status,
indicating a sinuous fluvial meandering paleoenvironment. This new locality seems to be relevant regarding not
only its fossil vertebrate diversity but also because of its geographic position: on a highway side, which allows
access at any season of the year, as contrary to most common riverine outcrops from Solimões Formation, which
are restricted to collections during the dry seasons.

1. Introduction
The State of Acre, northern Brazil, presents rocks that preserve some
of the latest South American faunas before the turnover caused by the
Great American Biotic Interchange event (Negri et al., 2010). The
Acrean fossil vertebrates have been early mentioned in literature by
Agassiz (1868) and, since then, the age of this fauna and its relation-
ships with other fossil assemblages from South America have been
discussed (e.g., Cozzuol, 2006; Latrubesse et al., 1997, 2007; 2010;
Hoorn et al., 2010a, b; Riff et al., 2010; Cidade et al., 2019a). Despite
this, some doubtfulness regarding the stratigraphic correlation between
different localities persists, mostly because of the continuous and dense
vegetation coverage, which restrict the access to outcrops, that are
generally limited to natural (e.g., rivers) or artificially (e.g., highways)
devastated areas (Negri et al., 2010). As a consequence of this strati-
graphic unknowledge, the uncertainties regarding the Acrean portion of
Solimões Basin, corresponding to the west portion of the state
(Menegazzo et al., 2016; Sá et al., 2020), lead to a debate, that still in
progress, about its age (e.g., Cozzuol, 2006; Latrubesse et al., 2007;
Bissaro-Júnior et al., 2019), paleoenvironment (e.g., Räsänen et al.,
1995; Hoorn et al., 2010a; Latrubesse et al., 2010; Gross et al., 2011;
Nogueira et al., 2013), and the sedimentary depositional history and
dynamic of these beds (Horbe et al., 2019). The influence of Andean
orogeny and the genesis of current Amazon hydrographic basin influ-
enced the landscapes, ecosystems, and also the faunal diversification in
the Amazonian region in the Early Cenozoic, being the current diversity
patterns rooted in the diversity of the Neogene biota (Hoorn et al.,
2010a,b; Bicudo et al., 2019). The direct influence of the Andes up-
lifting to the Amazonian paleoenvironment is evidenciated based on the
analysis of sedimentological data by Horbe et al. (2019), see also Hoorn
et al. (2017), which concluded the Andean provenance of the sedi-
mentary cover in Solimões Formation.
The deposition of proto-Andean and Andean sediments over the
Acre and Solimões Basins resulted in the genesis of the rocks known as
Solimões Formation, which is a well-known Cenozoic lithostratigraphic
unit due to its biodiverse fossiliferous content comprising microfossils,
plants, invertebrate, and vertebrate (e.g., Cozzuol, 2006; Latrubesse
et al., 2010; Gross et al., 2013; Nogueira et al., 2013; Souza-Filho and
Guilherme, 2015; Haag and Henriques, 2016; Kloster et al., 2017; Sá
and Carvalho, 2017; Sá et al., 2020). This formation extends from

∗
Corresponding author. Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil.
E-mail address: mbslacerda@ufmg.br (M.B.d.S. Lacerda).

https://doi.org/10.1016/j.jsames.2020.102588
Received 8 January 2020; Received in revised form 18 March 2020; Accepted 28 March 2020
Available online 10 April 2020
0895-9811/ © 2020 Elsevier Ltd. All rights reserved.
M.B.d.S. Lacerda, et al.
Manaus municipality (Amazonas State, Brazil) to the South limits of
Acre State, Brazil (Latrubesse et al., 2010). Despite the divergences
about dating and paleoenvironment interpretations (e.g., Latrubesse
et al., 2010 contra Räsänen et al., 1995), the paleontological im-
portance of Solimões Formation is unquestionable, providing several
specimens of a variety of Tetrapoda clades, such as anurans (Muniz
et al., 2016), lizards and snakes (Hsiou and Albino, 2009; Hsiou et al.,
2009, 2010), turtles (Oliveira and Romano, 2007; Riff et al., 2010),
crocodilians (Cidade et al., 2019a,b), birds (Alvarenga and Guilherme,
2003; Negri et al., 2010), and several mammalian orders (Cozzuol,
2006; Negri et al., 2010, Ribeiro et al., 2013). This plethora of fossil
organisms provide relevant and essential information on the evolution
of neotropical fauna and flora from the Amazon region (Hoorn et al.,
2010a,b; Bissaro-Júnior et al., 2019).
The present work intend to increase our knowledge regarding the
vertebrate diversity of the Solimões Formation through the description
of some fossils found on a new outcrop situated within Sena Madureira
municipality, State of Acre. This work provide brief comments about
the systematics and taphonomy of those specimens.
1.1. Geological background
Situated within the Acre, Solimões, and Madre de Dios basins, the
Solimões Formation is exposed in northwestern of Brazil, east of Peru,
and north of Bolivia (Duarte, 2011). Its rocks are characterized by
several facies of conglomerates, sandstones, and mudstones with fining
upward vertical profiles, showing great lateral heterogeneities (Caputo
et al., 1971; Brasil, 1976), also including claystone, with calcareous
concretions, calcite, and gypsum veins (Latrubesse et al., 2010). This
horizontal heterogeneity of the rocks made previous authors recognize
the Solimões Formation as a series of different formations (see Brasil,
1976 for a historical perspective). Dating and thickness of the Solimões
Formation adopted here follows those proposed by Cunha (2007),
which corresponds to the temporal range Eocene–Pliocene age, with an
estimated thickness of 2200 m. The Solimões Formation presents sev-
eral reverse faults related to the Andean Orogeny (Brasil, 1976) and
possible diachronism during the Late Miocene sedimentary deposition
(Bissaro-Júnior et al., 2019). The paleoenvironment was attributed to
sandbars, channels, and floodplains formed in fluvial or fluviolacustrine
systems (Caputo et al., 1971, Räsänen et al., 1995, Gross et al., 2011,
also see Latrubesse et al., 2010 for a historical perspective of pa-
leoenvironment interpretation). Souza et al. (2016), refined these flu-
vial interpretation, at least for the uppermost portion of the Solimões
Formation, as a high sinuosity fluvial meandering paleoenvironment,
interpretation that is followed here.
2. Material and methods
2.1. New fossiliferous locality
This new locality is situated, in State of Acre, within Sena Madureira
municipality in the exit to Manuel Urbano municipality, on highway BR
364. The fossils were collected in the region of the uppermost part of
the Solimões Formation. The geographical coordinates of this new
outcrop are: 9°02′43,20″ S 68°46′18,12″ W (Fig. 1).
The site have been exposed due to scarping of the BR 364 highway.
It represents a strata of expansive clays, greenish gray, rich in phos-
phates and sulphates. It is possible to see whitish concreted areas on
this level, where several bone remains abound, many of which have
been rolled or transported. The specimens were collected by free
walking along the site, at a height of 10 m above road level.
2.2. Studied materials
Carlos E. G. R. Schaefer collected at least forty-four isolated fossils
in July of 2010. From those fossils, twenty-three were described,
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Journal of South American Earth Sciences 101 (2020) 102588
whereas the remaining twenty-one are too fragmentary to perform a
precise systematic and anatomical identification and were only eval-
uated for taphonomical considerations (see session 3.2). Those mate-
rials are housed in the paleontological collection of the Laboratório de
Paleontologia e Osteologia Comparada (LAPOC), of the Museu de
Zoologia João Moojen (MZUFV), Departamento de Biologia Animal
(DBA), Universidade Federal de Viçosa (UFV), Minas Gerais State,
Brazil. The materials are under the acronym MZUFV-P and each spe-
cimen received its own register number. The studied materials, that are
described here were compared to extant and fossil specimens from
several species, those samples were accessed from osteological collec-
tions and specialized literature (see SM to details on the compared
specimens and information about the institutions where the materials
are deposited).
2.3. Anatomical nomenclature
The Crocodyliformes cranial nomenclature was based on Mook
(1921a) and Iordansky (1973) and the postcranial nomenclature on
Mook (1921b), Carvalho (1951), Hoffstetter and Gasc (1969), Frey
(1988), Vieira et al. (2016) and modifications on vertebral nomen-
clature proposed by Souza (2018). The Testudines nomenclature was
based on Gaffney (1990), Cadena et al. (2008), and Sereno and ElShafie
(2013). The Mammalia nomenclature follows Paula-Couto (1979).
2.4. Measurement of specimens
The measurements of all bones and structures were performed with
a manual Mitutoyo® digital caliper. The angles of morphological
structures (e.g., processes, zygapophysis) were measured using photo-
graphs of the specimens using the software ImageJ 1.8.0 (Rasband,
1997). All measurements were taken by the same person (MBSL) and
those were used in morphological descriptions and to assist in speci-
mens’ identifications and comparisons.
3. Results
3.1. Morphological descriptions and systematic
CROCODYLIFORMES Hay (1930).
CROCODYLIFORMES Indet.
Referred materials: MZUFV-P 0005; MZUFV-P 0010; MZUFV-P
0011; MZUFV-P 0015; MZUFV-P 0037; MZUFV-P 0038; MZUFV-P
0039; MZUFV-P 0040; MZUFV-P 0044; MZUFV-P 0045; MZUFV-P
0046; MZUFV-P 0047; MZUFV-P 0048; MZUFV-P 0049.
Morphological description (osteoderms): Seven specimens
(MZUFV-P 0005, MZUFV-P 0040, MZUFV-P 0045, MZUFV-P 0046,
MZUFV-P 0047, MZUFV-P 0048 and MZUFV-P 0049) are isolated os-
teoderms. Based on the ornamentation (rounded and shallow pits at
dorsal surface and parallel oriented scar lines forming grids in ventral
surface) and general shape (sub-rectangular), those osteoderms can be
attributed to the Crocodyliformes.
MZUFV-P 0005 (Fig. 2A) is completely preserved. It has a rectan-
gular shape with 36 mm of maximum length and 32 mm of maximum
width. The dorsal surface of the dermal plaque has a small elevation
(not forming a true crest) in the medial region, with small pits at its
dorsal surface. Joint sutures with other osteoderms are visible on left
and right sides. The rectangular shape of MZUFV-P 0005 with rounded
corners, the sutures morphology and its position are compatible with an
osteoderm from the main dorsal roll.
MZUFV-P 0040 (Fig. 2B) has a triangular shape in lateral view, the
apice is rounded and posterodorsally inclined. The medial surface is
more plane than the lateral, which is concave laterally. The base of this
osteoderm is not completely preserved and the most ventral part of it is
30 mm of maximum width and 26 mm of maximum height. It belongs
M.B.d.S. Lacerda, et al.
Fig. 1. Map of South America, highlighting the Acre State, north of Brazil. The dot symbol indicates the new fossiliferous outcrop (9°02′43,20″S 68°46′18,12″W).
to the caudal series of osteoderms, representing an isolated crest of one
of the first caudal series (dentiform caudal verticiles sensu Carvalho,
1951).
The remaining osteoderms (MZUFV-P 0045, MZUFV-P 0046,
MZUFV-P 0047, MZUFV-P 0048 and MZUFV-P 0049) are markedly
fragmented. The grooves on the dorsal side seen in MZUFV-P 0045 and
MZUFV-P 0046 allow us to identify it as lateral or dorsal trunk osteo-
derms. The specimens MZUFV-P 0047, MZUFV-P 0048, and MZUFV-P
0049 do not have these ornamentations grooves, but, as seen in all
osteoderms collected, they exhibit small scars in ventral surface, which
indicate the joint between the dermal plaque and the derm of the
specimen.
Morphological description (ribs): MZUFV-P 0010 represents a
medial region of a fragmentary isolated rib (Fig. 2D). The rib has
73 mm of maximum length and 23 mm of maximum width in its most
expanded region. The ventral margin is slightly curved, being convex.
The morphology of its anterior and posterior margins, forming crests,
suggests that it represents a dorsal rib fragment.
Another rib fragment is MZUFV-P 0044 (Fig. 2C), this specimen
refers to part of the proximal region. It has 51 mm of maximum length
and 45 mm of maximum width in its most proximal region and 38 mm
of maximum width in the most distal region. MZUFV-P 0044 has a
relatively anteroposterior flattened, being slightly concave in the dorsal
region with a maximum thickness not exceeding 8 mm. The tubercular
articulation is dorsoventrally elliptical, whereas capitular articulation is
elliptical anteroposteriorlly. The tubercular articulation is larger than
the capitular. The capitular and tubercular processes are almost
equivalent in size (capitulum maximum length 14 mm; tuberculum
maximum length 10 mm, when measured from the base of the notch
capitulum-tuberculum), and the notch has a well-defined angular
aperture. Due to the length similarities between the capitulum and
tuberculum processes of MZUFV-P 0044 we identify it tentatively as the
first left dorsal rib.
Morphological description (vertebra): MZUFV-P 0011 is a well-
preserved caudal vertebra, missing most part of the neural spine
(Fig. 2E1-E3). It has 56 mm of maximum length, from cotyl to condyle
and 38 mm of maximum height up to the preserved part of the base of
3
Journal of South American Earth Sciences 101 (2020) 102588
neural spine. The specimen presents the right and left basis of the fused
caudal rib (Fig. 2E3), the base of the spinous process (Fig. 2E1) and the
right prezygapophysis (Fig. 2E1). The vertebral body is flattened la-
teromedially and this flattening is most evident in its medial portion,
below the fused ribs (Fig. 2E2). The vertebral condyle is well-procelic
(Fig. 2E1-2E2). Below the condyle, it is possible to observe a scar, in-
terpreted as the contact between the ventroposterior region of the
vertebral body and the chevron/hemal arch (Fig. 2E2). The pre-
zygapophysis is dorsally elongated, narrowing in the anteroposterior
direction and willing horizontally in relation to the vertebral center in
lateral side. The specimen MZUFV-P 0011 also presents a surface of
articulation with the pre- and postzygapophysis slightly medially la-
teralized, thus the articulation with the posterior vertebra is more in-
ternalized (Fig. 2E3). The length of the caudal rib is 26 mm, which is
about 46% of the total length of the vertebra (Fig. 2E2), allows the
identification of MZUFV-P 0011 as a caudal vertebra from the anterior
portion of the tail. In addition, its general morphology and ven-
troposteriorlly scars of the centrum, indicating the chevron articulation,
corroborate to this conclusion.
MZUFV-P 0015 (Fig. 2G) is a fragment of neural spine, with 45 mm
of maximum length and 40 mm of maximum width. The fossil consists
on the dorsal region of a neural spine and is lateromedially flattened
and relatively thin, with about 5 mm of maximum thickness. The apical
region of the neural spine has a lateral extension forming an elliptical
structure in dorsal view. The anteroposterior length of MZUFV-P 0015
and the lateral expansions on its apex allows identifying it as probably a
posterior dorsal or anterior lumbar vertebra of a large specimen.
Morphological description (crania): MZUFV-P 0037 (Fig. 2H1-
H2) is a medial fragment of the right maxillary, and it has 76 mm of
maximum length and its maximum width is 30 mm. The palatal surface
is smooth and nearly straight with the presence of a foramen in most
posterior region (Fig. 2H2). This foramen is similar to the foramina of
the palatal region observed in some Caimaninae specimens such as
Caiman latirostris, and Melanosuchus niger. The dorsal region of MZUFV-
P 0037 is irregular, with a dorsally projecting structure that forms the
inner region of the maxilla (Fig. 2H1), but the most dorsal region of the
bone is not preserved. The anterior and posterior edges of the specimen
M.B.d.S. Lacerda, et al. Journal of South American Earth Sciences 101 (2020) 102588

are flat, and no suture border is preserved. Even if fragmented, the
presence of the foramen and the morphology of the ventral region of the
MZUFV-P 0037 allow us to conclude that it is a maxillary fragment
assigned to Crocodyliformes.
MZUFV-P 0039 (Fig. 2I1-I2) is probably a right ectopterygoid with
46 mm of maximum length and 22 mm of maximum width. The ventral
portion, in the region of contact with pterygoid is broken, enabling to
visualize the internal structure of the bone (Fig 2I-2). The rugose sur-
faces preserved seen in lateral portion are probably the suturement
region with maxillary and jugal (Fig 2I-1).
Morphological description (appendicular skeleton): MZUFV-P
0038 (Fig. 2F) is a fragment epiphysis, probably from first right meta-
tarsus. It has a maximum length of 31 mm and an elliptical shape in
cross-section. On the lateral side, the specimen has two smooth digs in
the epiphysis region. The articulation of distal region with the first
phalange is planar and triangular when seen in distal view.
Remarks: Price et al. (1977) and Souza et al. (2016) pointed out the
presence of at least two Crocodyliformes clades in Solimões Formation:
Sebecidae and Crocodylia. Given the absence of enough diagnostic
features distinguishing the above-mentioned fragmentary material,
conservatively we identify those specimens as indeterminate Crocody-
liformes.
CROCODYLIA Gmelin, 1789 (sensu Benton and Clark, 1988)
ALLIGATOROIDEA Gray (1844) (sensu Brochu, 2003)
ALLIGATORIDAE Gray (1844) (sensu Norell et al., 1994)
CAIMANINAE Brochu (2003).
CAIMANINAE indet.
Referred material: MZUFV-P 0014.
Morphological description: MZUFV-P 0014 is a fragmented neural
arch (Fig. 3A1-A3). It has 57 mm of maximum length and 28 mm of

Fig. 2. Crocodyliformes: MZUFV-P 0005 (A) dorsal view; MZUFV-P 0040 (B) right lateral view; MZUFV-P 0044 (C) right lateral view; MZUFV-P 0010 (D) ventral
view; MZUFV-P 0011 (E1) right lateral view; (E2) ventral view; (E3) anterior view; MZUFV-P 0038 (F) dorsal view; MZUFV-P 0015 (G), lateral view; MZUFV-P 0037
(H1) lateral view, (H2) palatal view; MZUFV-P 0039 (I1) medial view; (I2) caudal view. Scale bars = 20 mm. Anatomical abbreviations: af-anterior facet; cap-
capitulum; cr-caudal rib; tub-tuberculum; prz-anteriorzygapophyses; for-foramen.

4
M.B.d.S. Lacerda, et al.
maximum width. Both postzygapophyses are preserved, with the left
one missing part of its dorsal extremity (Fig. 4A1). The right post-
zygapophysis exhibits a rectangular shape and is inclined poster-
olaterally in dorsal view. Only the right prezygapophysis is preserved
and has its articulation surface arranged in a horizontal position with a
subcircular shape. The basis of the right diapophyseal process is also
preserved and located over the lateral surface of the prezygapophyseal
process (Fig. 3A1, 3A3). The shape of the vertebral canal is preserved
and has 12 mm in width (Fig. 3A2). The peduncle and part of the la-
mina of the neural arch is also preserved on the right side of the spe-
cimen. In ventral view, the neurocentral junction of the neural arch
peduncle has rugose texture (Fig. 3A2) characteristic of the contact
between neural arch and vertebral body. The presence of a more hor-
izontalized diapophyseal process with two well-developed ridges con-
necting with prezygapophysis and postzygapophysis allows identifying
it as an anterior dorsal vertebra, probably being the second dorsal
vertebra (sensu Souza, 2018).
Fig. 3. Caimaninae: MZUFV-P 0014 (A1) dorsal view; (A2) ventral view; (A3) left lateral view. Scale bar = 20 mm. Anatomical abbreviations: dpr-diapophyseal
process; poz-posteriorzygapophyses; prz-anteriorzygapophyses; nc-neural channel.
Comparisons: Despite the fragmentary nature of MZUFV-P 0014, it
preserves a key feature for taxonomic identification, the diapophyseal
process (Fig. 3A1 and 3A3). In this specimen (MZUFV-P 0014), the
diapophyseal process is located on the lateral surface of the pre-
zygapophyseal process (Fig. 3A3). The anterior margin of the diapo-
physeal process present a ridge, connecting it to the dorsal portion of
the prezygapophyseal process. In the posterior margin of the diapo-
physeal process, a posterodorsal ridge connects it with the anterior
margin of the postzygapophyseal process. This set of features is similar
to those observed in extant Caimaninae species Caiman crocodilus,
Caiman latirostris, Caiman yacare, Melanosuchus niger, Paleosuchus pal-
pebrosus, and Paleosuchus trigonatus. On the other hand, the remaining
extant species of non-Caimaninae observed, such as Alligator mis-
sissippiensis (Daudin, 1802), Alligator sinensis (Fauvel, 1879), Crocodylus
niloticus Laurenti, 1768, Crocodylus porosus (Schneider, 1801), Gavialis
gangeticus, and Tomistoma schlegelii (Müller, 1838), differ by having the
diapophyseal process more posteriorly located, with the anterior ridge
that connects it with prezygapophyseal process more horizontalized
and also, by having the posterior ridge shorter and almost vertical.
Remarks: The presence of an anterior displaced diapophyseal
process in the second dorsal vertebra is a feature exclusively shared
with extant Caimaninae species.
Mourasuchus Price, 1964
Type-species: Mourasuchus amazonensis Price (1964)
Mourasuchus sp.
Referred material: MZUFV-P 0004; MZUFV-P 0009.
Morphological description: MZUFV-P 0004 is a well-preserved
and nearly complete isolated osteoderm (Fig. 4A1-A2). It has a sub-
circular base with 42 mm of maximum length and 26 mm of maximum
width. The lateroposterior border is broken and partially lost. It does
5
Journal of South American Earth Sciences 101 (2020) 102588
not show any joints or sutures to other osteoderms. The dorsal surface
(Fig. 4A1) presents irregular pits and grooves ornamentation, being
concentrated to the anterior portion of the osteoderm, but also present
in the lateral surface of the crest. The median crest is well-developed
dorsally, with 32 mm of maximum length and 21 mm of maximum
high, with an angle of 148° in relation to the base of the osteoderm
(Fig. 4A2). The crest is compressed lateromedially, projecting poster-
odorsally and presenting a rounded apex. The ventral surface of the
specimen has parallel line scars, formed due to the attachment of the
osteoderm with the body specimen.
MZUFV-P 0009 (Fig. 4B1-B4) is an almost complete posterior dorsal
vertebra. This vertebral body is longer than it is tall (Fig. 4B2), having
56 mm of maximum length (from cotyl to condyle) and 78 mm of
maximum height (measured to mid dorsal line from the pre-
zygapophyses uppermost edge to the ventral edge of the cotyl)
(Fig. 4B1). The vertebral body has a medial constriction which reaches
the smallest thickness (29 mm) in the midventral part (Fig. 4B4). The
anterior part of the cotyl is slightly convex and wider than tall
(Fig. 4B2). Only the basis of the neural spine, which covers the entire
dorsal surface of the neural arch is preserved, reaching 50 mm in
maximum length (measured from the base) (Fig. 4B3). The pre-
zygapophyses of MZUFV-P0009 have a rectangular shape in dorsal view
(Fig. 4B) with a slight medial arching and the dorsal edges straightened.
The articular surface of the prezygapophyses has an angle of 52°
(anterior view) in relation to the sagittal plane (Fig. 4B1). The post-
zygapophyses also have a rectangular shape, with rounded edges
(Fig. 4B3-4B4), the articulation surface of the postzygapophyses, pro-
trudes anteriorly to the vertebral body and have a greater area of ar-
ticulation in relation to prezygapophyses. Both transverse processes are
preserved, but the left one is distorced due to taphonomic bias. Based
on the right transverse process it is possible to infer that these structures
are planar and flattened dorso-ventrally, not exceeding 10 mm of
thickness (Fig. 4B3-4B4). There is no evidence that the neural arch and
the vertebral body are completely fused, however, the region of medial
contact between vertebral body and neural arch is not entirely pre-
served due to taphonomic bias, presenting some excavations of the
bone surface (Fig. 4B2).
Comparisons: The morphology of the osteoderm MZUFV-P 0004 is
compatible with osteoderms referred to Mourasuchus arendsi Langston
(1965), as pointed out by Langston (2008), and Mourasuchus sp. as
inferred by Souza et al. (2016). It has the following Mourasuchus di-
agnostic features: small and subcircular base; well-developed medial
crest with anterior margin inclined; and rounded apical extremity.
MZUFV-P 0004 probably is an osteoderm of the lateral rolls of the
trunk, given its general morphology and the previous proposal made by
Langston (2008) and reiterated by Souza et al. (2016).
MZUFV-P 0009 is larger than all extant Crocodylia species ver-
tebrae. It is compatible in size with, at least, three extinct genera: the
Caimaninae Mourasuchus and Purussaurus and the Gavialoidea
M.B.d.S. Lacerda, et al. Journal of South American Earth Sciences 101 (2020) 102588

Fig. 4. Mourasuchus sp.: MZUFV-P 0004 (A1) dorsal view; (A2) lateral view; MZUFV-P 0009 (B1) anterior view; (B2) lateral view; (B3) dorsal view; (B4) ventral view.
Scale bars = 20 mm. Anatomical abbreviations: tp-transverse process; poz-posteriorzygapophyses; prz-anteriorzygapophyses.

Fig. 5. – Melanosuchus cf. M. niger: MZUFV-P 0016 (A1) proximal view; (A2) lateral view; (A3) medial view; (A4) posterior view. Scale bar = 20 mm.

Gryposuchus. Since there are no post cranium axial skeletal elements
described for Gryposuchus so far, it is impossible to compare MZUFV-P
0009 to this genus. Purussaurus dorsal vertebra have condyle and cotyl
with similar size (Gervais, 1876), differing from the observed in
MZUFV-P 0009 (Fig. 4B2) and Mourasuchus. Also, in Purussaurus, the
ventral portion of vertebral body is arched with the greatest
constriction in the medial region of the vertebral body, differing from
MZUFV-P 0009 (Fig. 4B2) and Mourasuchus, which is straight. MZUFV-
P 0009 has a lateromedial compression on the vertebral body, seen in
ventral view (Fig. 4B4), and similar to that described by Langston
(2008) for Mourasuchus arendsi. Moreover, MZUFV-P 0009 and Mour-
asuchus arendsi also share corresponding angulation of prezygapophyses

6
M.B.d.S. Lacerda, et al.
in relation to the sagittal plan. According to Langston (2008), the first
lumbar vertebra of Mourasuchus arendsi has an angulation of 48° and
the fifth, 50°. The prezygapophyses of MZUFV-P 0009 have an angu-
lation of 52°. It is not possible to identify which posterior dorsal ver-
tebra MZUFV-P 0009 is, but it is secure to identify it as Mourasuchus
based on the above-mentioned features as well as the differences
pointed out to Purussaurus and all extant Crocodylia species.
Melanosuchus Spix, 1825
Type-species: Melanosuchus niger (Spix, 1825)
Melanosuchus cf. M. niger
Referred material: MZUFV-P 0016.
Morphological description: MZUFV-P 0016 (Fig. 5A1-A4) is a
fragmentary left tibia. Only the proximal epiphysis is preserved, with
37 mm of maximum length (Fig. 5A2-A4) and the maximum width at its
most proximal portion, is 33 mm. The posterior process of the tibia is
broken (Fig. 5A4), but in proximal view (Fig. 5A1) the inner margin of
it is visible, and in posterior view (Fig. 5A1), the origin of the ventral
portion slope is also visible (Fig. 5A1). The posterior process has a
subtle leaning, which we interpret as evidence that it does not range
extensively. MZUFV-P 0016 general shape, in proximal view is elliptical
with a straight anterior border (Fig. 5A1). On the lateral side, the
proximal tibiofibular joint facet presents a shallow concavity between
the lateral process and the posterior process (Fig. 5A2). The medial
process projects slightly occupying most of the medial margin of the
tibia (seen in proximal view). Between the medial and the posterior
process, lies a shallow concavity (Fig. 5A3), the intercondylar sulcus
(sensu Vieira et al., 2016) of the tibia, that is located between the base
of the posterior, lateral and medial processes (Fig. 5A1) in a medio-
laterally portion of the epiphysis, in proximal articulation.
Comparisons: MZUFV-P 0016 is identified as Melanosuchus cf. M.
niger by the following features: a well-developed posterior process that
is laterally and medially delimited by two concavities (seen in proximal
view); these concavities isolate the posterior process from the lateral
and medial process, respectively; both lateral and medial process points
respectively to lateral and medial directions; also the anterior margin is
slightly arched. Those features seen in Melanosuchus niger and MZUFV-P
0016 differs from Caiman species by having a medial process facing
medioanteriorly seen in proximal view. Also, in Caiman there is an
additional process, facing medioposteriorly, between the medial and
posterior process that is not seen in MZUFV-P 0016. In Paleosuchus the
lateral process is pronounced and limited to the anterior most portion of
the lateral surface, whereas in Gavialis the anterior margin is straight
but with two small concavities, the posterior margin is well-developed
and its lateral and medial processes are not developed, both differ from
the morphology seen in Melanosuchus. Other extinct Caimaninae of the
Solimões Formation, such as Acresuchus (Souza-Filho et al., 2019),
Caiman brevirostris Souza-Filho (1987) (Fortier et al., 2014) and Mour-
asuchus do not have any preserved tibia, and therefore, the comparisons
with these taxa is not possible. Thus, although poorly preserved,
MZUFV-P 0016 preserves key features that allow us to identified as a
specimen of Melanosuchus cf. M. niger.
Caiman Spix, 1825
Type-species: Caiman crocodilus (Linnaeus, 1758)
Caiman aff. C. crocodilus
Referred material: MZUFV-P 0012.
Morphological description: MZUFV-P 0012 is the 1st sacral ver-
tebra (Fig. 6B1-B4). It has 88 mm of maximum length measured from
the vertebral body to the rib/ilium contact, and 47 mm of maximum
width in region of vertebral body. Only the left side of the vertebra is
preserved, including a fusioned sacral rib, prezygapophysis and a small
dorsal portion of the vertebral centrum (Fig. 6B1-B4). Part of neural
canal is preserved and has a diameter of 9 mm. The sacral rib and the
7
Journal of South American Earth Sciences 101 (2020) 102588
transverse process are completely fused, without any visible suture.
However, the suture between the vertebral body and neural arch re-
mains open, with 15 mm of maximum length, situated above the ver-
tebral centrum. MZUFV-P 0012 presents the sacral rib with dorsal
margin horizontalized, seen in anterior view (Fig. 6A4). In dorsal view,
there is a concavity in the lateral margin of the rib forming the ar-
ticulation with the left ilium (Fig. 6A1). The prezygapophysis is well-
developed, with 28 mm of maximum length and 13 mm of maximum
width, and its articular surface is elliptical in shape (Fig. 6A3). In dorsal
view (Fig. 6A1), the lateral portion of the articular surface of the pre-
zygapophysis is more posteriorlly than medial, becoming gradually
more inclined along the anteroposterior axis. In anterior view
(Fig. 6A4), the articulation surface of the prezygapophysis is relative
inclined to the sagittal plane of the vertebra, with approximately 62°.
The tuberculum of sacral rib is extended anteriorly, as seen in anterior
view. This tuberculum fuses to the lateral border of the vertebral cen-
trum, and in MZUFV-P 0012 is a well-developed structure (Fig. 6A2).
The tuberculum position in MZUFV-P 0012, in relation to rib, presents
an angulation of 131° and extends anteriorly to the anterior limits of the
prezygapophysis (Fig. 6A1-A3).
Comparisons: The rib of MZUFV-P 0012 presents its dorsal surface
horizontalized in relation to the vertebra-ilium contact, this condition is
opposite to that observed in Gavialis gangeticus, which has the inclined
rib with its distal region more ventrally positioned. In Paleosuchus
palpebrosus, the sacral rib also has the dorsal surface slightly inclined
towards the ilium, but this inclination is less evident than in Gavialis
gangeticus. In Melanosuchus niger, the dorsal surface of the rib is arched
and forms a convex structure. This feature differs from the observed in
MZUFV-P 0012. Caiman species have the dorsal surface of the sacral rib
relatively flatter and more horizontal, when compared to Gavialis,
Paleosuchus, and Melanosuchus. Among Caiman species, C. crocodilus
presents the straightest rib in its dorsal surface (seen in anterior view),
being this condition similar to those observed in MZUFV-P 0012.
The left prezygapophysis of MZUFV-P 0012 is well-developed, being
a relatively large structure. This condition differs from Gavialis gang-
eticus and Paleosuchus, because both have relatively small pre-
zygapophyses with a smaller articulation surface. Melanosuchus and
species of Caiman have the most developed prezygapophyses, being
observed in Caiman crocodilus the greater prezygapophyses between
extant Caimaninae. In anterior view, MZUFV-P 0012, Caiman crocodilus
and Caiman latirostris, have the prezygapophyses occupying a great part
of the neural arch, being its base broadly enlarged. The inclination of
this structure in MZUFV-P 0012, based on its dorsal surface of articu-
lation with the last lumbar vertebra and relative to the sagittal plane of
the vertebra is 62°. The dorsal end of the prezygapophysis is laterally
inclined in MZUFV-P 0012. The opposite can be observed in Gavialis
gangeticus which have an angulation of 43° and Caiman yacare, 44°. In
these two species, the edge of the prezygapophyses are situated in a
more dorsal position when compared with MZUFV-P 0012. Caiman
crocodilus and Melanosuchus niger show the most inclined laterally
prezygapophyses, with 55° and 56° of inclinations, respectively,
whereas Mourasuchus arendsi has an angulation of the prezygapophyses
with 58°, according to Langston (2008).
A feature observed in MZUFV-P 0012 that differs from any living
Crocodylia is the position of the contact between the tuberculum and
the vertebral body. This structure is positioned in a more anterior re-
gion and forms an angulation of 131° in relation to the anterior border
of the rib (Fig. 6A1). In general, extant Crocodylia have the tuberculum
that does not exceed the anterior line of the prezygapophysis. However,
this feature is unknown in Purussaurus and Acresuchus.
Remarks: The dorsal surface of the first sacral ribs of MZUFV-P
0012 is horizontalized, which seems to be a feature exclusively shared
with Caiman species. Therefore, MZUFV-P 0012 is securely assigned to
Caimaninae. Due to the overall similarities and the above-mentioned
shared features of MZUFV-P 0012 and Caiman crocodilus, we tentatively
indentify it as Caiman aff. C. crocodilus.
M.B.d.S. Lacerda, et al.
Fig. 6. – Caiman aff. C. crocodilus: MZUFV-P 0012 (A1) dorsal view; (A2) lateral view; (A3) medial view; (A4) anterior view. Scale bar = 20 mm. Anatomical
abbreviations: tub-tuberculum; prz-anteriorzygapophysis.
TESTUDINES Linnaeus, 1758.
PLEURODIRA Cope (1865).
PLEURODIRA indet.
Referred material: MZUFV-P 0042.
Morphological description: MZUFV-P 0042 is a fragmentary right
xiphiplastron (Fig. 7A1-7A2). It has 39 mm of maximum length and
42 mm of maximum width. There is no suture demarcation in MZUFV-P
0042 with the hypoplastron. In visceral view, the specimen MZUFV-P
0042 presents the pubic scar nearly preserved (Fig. 7A1), with a
semicircular shape, and slightly elevated on its most anterior portion.
The ventral surface of the bone shows the femoral-anal sulcus that is
positioned above the pubic scar (Fig. 7A2).
Comparisons: The femoral-anal sulcus in MZUFV-P 0042 is similar
to those observed in chelids such as Chelus fimbriata (Schneider, 1783)
and Phrynops geoffroanus (Schweigger, 1812) in both position and
shape, but differs from the podocnemidid Podocnemis expansa
(Schweigger, 1812) that have the femoral-anal sulcus more anteriorly
positioned. The position of femoral-anal sulcus could be interpreted as
diagnostic to Chelidae, but the fragmentary condition of MZUFV-P
0042 does not allow us to properly compare it. However, the pubic scar
observed in MZUFV-P 0042 allows us to identify it as a side-necked
turtle specimen.
PODOCNEMIDIDAE Cope (1868).
PODOCNEMIDIDAE indet.
Referred material: MZUFV-P 0013.
Morphological description: MZUFV-P 0013 is probably the 7th or
8th cervical vertebra (Fig. 7B1-7B2). It has 18 mm of maximum length,
13 mm of maximum height, and 11 mm of width. Only the neural arch
of the vertebra is preserved. The right side of neural arch presents the
8
Journal of South American Earth Sciences 101 (2020) 102588
fragmentary base of the transverse process. The neural arch also pre-
sents both postzygapophyses and the base of neural spine, which is
quite fragmented. Only the posterior portion of neural spine is pre-
served and is dorsally elongated (Fig. 7B1). The postzygapophyses have
an elliptical shape in articulation surface, being taller (9 mm of height,
and 6 mm of length), and clearly visible in dorsal view (Fig. 7B2). The
angulation of the postzygapophyses are 34° in relation to the sagittal
plane, therefore, the articular surface is laterally positioned (Fig. 7B1).
Part of the neural channel shape is preserved and presents a maximum
diameter of 5.5 mm (Fig. 7B1).
Comparisons: Even though not completely preserved, the neural
spine of MZUFV-P 0013 is probably well developed, as seen in Chelidae
and Podocnemididae turtles. In Chelidae, the postzygapophyses show
smaller angulation aperture, with the articulation surface more ven-
trally positioned. Also, Chelidae species have smaller joint surfaces, if
compared with MZUFV-P 0013 and Podocnemididae species. On the
other hand, Podocnemididae species also have articular surface of
postzygapophyses more developed than Chelidae's, as well as greater
angular aperture, with the surface of articulation more laterally posi-
tioned. The aforementioned Podocnemididae features are observed in
MZUFV-P 0013, allowing us to assign it to this family. Since the post-
zygapophyses of 7th and 8th cervical vertebra of Podocnemis and
Stupendemys do not differ from the general morphology of remaining
podocnemidids and other South American genera, and Caninemys
(Gaffney et al., 1998; Meylan et al., 2009) and Bairdemys (Gaffney and
Wood, 2002) do not have preserved cervical vertebra, it is impossible to
identify which genus and species MZUFV-P 0013 could belong.
CHELIDAE Gray (1825).
Chelus Duméril, 1806
Type-species: Chelus fimbriata (Schneider, 1783)
Chelus sp.
M.B.d.S. Lacerda, et al. Journal of South American Earth Sciences 101 (2020) 102588

Referred material: MZUFV-P 0006.
Morphological description: MZUFV-P 0006 is a partially pre-
served left 8th costal bone (Fig. 7C1-C3). It has 69 mm of maximum
length and 41 mm of maximum width. Only the medial and lateral
portion is preserved. The suture surface with the left 10th and 11th
peripherals are visible and a groove can be seen in visceral medial re-
gion of the specimen. This groove has been interpreted as a taphonomic
damage on the rib contact with 10th peripheral bone (Fig. 7C1). The
anterior portion of MZUFV-P 0006 preserves the posterior edge of the
dorsal ridge (Fig. 7C3), being notably thicker (22 mm of thickness) than
the lateral portion (5 mm of thickness) of it (Fig. 7C2). The posterior
region is also more arched, notably on the visceral surface, a feature
consistent with 8th costal morphology (of Chelus). The ventral surface of
the specimen preserves the medioposteriorlly portion of the iliac scar,
with 26 mm of maximum width (Fig. 7C2).
Comparisons: The iliac scar allows the identification of MZUFV-P
0006 as a Pleurodira. Moreover, the presence of a partially preserved
dorsal ridge enables to assert it to Chelus, since this structure is only
known in this genus (Wood, 1976; Cadena et al., 2008; Ferreira et al.,
2016). The shape of the preserved portion of the dorsal ridge seems to
be continuous, as in Chelus colombianus, and differing from the Chelus
fimbriata and Chelus lewisi. However, given the fact that only the most
posterior portion of this structure is preserved, we prefer to keep a more
conservative identification due to the lack of other specific diagnostic
feature.

Fig. 7. Testudinata: MZUFV-P 0042 (A1) visceral view; (A2) ventral view; (B1) MZUFV-P 0013, posterior view; (B2) dorsal view; (C1) MZUFV-P 0006, dorsal view;
(C2) ventral view; (C3) anterior view. Scale bars = 20 mm. Anatomical abbreviations: pbs-pubic scar; a-f sc-anal-femoral scar; nsp-neural spine; poz-poster-
iorzygapophyses; phc-peripheral contact; ils-iliac scar; dri-dorsal ridge.

9
M.B.d.S. Lacerda, et al.
Fig. 8. – Toxodontidae: MZUFV-P 0008; (A1) proximal view; (A2) distal view; (A3) lateral view. Scale bars = 20 mm.
MAMMALIA Linnaeus, 1758.
NOTOUNGULATA Roth (1903).
TOXODONTIA Owen (1853).
TOXODONTIDAE Owen (1845).
TOXODONTIDAE indet.
Referred material: MZUFV-P 0008.
Morphological description: MZUFV-P 0008 is a complete right
scaphoid (Fig. 8A1-A3). It has 39 mm of length, 69 mm of height, and
45 mm and 32 mm of width in anterior and posterior regions, respec-
tively. The proximal view (Fig. 8A1) presents a plane articular facet that
becomes concave in the posterior region. This articular facet also pro-
jects a tuberosity in the anterolateral region. Laterally, there is an
anterior facet plane onto the tuberosity which is followed posteriorly by
a small, circular, and slight convex facet. A third facet, displaced dis-
tally, is the biggest of the three, and deeply concave at this posterior
half. In the distal view (Fig. 8A2) the biggest facet of the scaphoid is
separated by a smooth crest. The posterior facet of the distal region
presents a deeply concave region that becomes convex in the posterior
region. The medial region presents a slight concave facet displaced
posteriorly (Fig. 8A3).
10
Journal of South American Earth Sciences 101 (2020) 102588
Comparisons: The Notoungulata order is the unique group which
has taxa that reach this dimension of carpal bones in outcrops of
Solimões Formation. Apart from that, the overall morphology of
MZUFV-P 0008 is compatible with the description of Toxodon made by
Roth (1898) and is also similar to the specimen MCL 23330/19 of
Toxodon platensis. The MZUFV-P 0008 is slightly smaller than the
Toxodon described by Roth (1898) and the specimen of Toxodon pla-
tensis housed in MCL/PUC. The MZUFV-P 0008 has the distal facets less
concave than in Toxodon platensis MCL 23330/19, and the facet placed
laterally onto the tuberosity is more delimited than Toxodon platensis
MCL 23330/19. Cozzuol (2006) and Latrubesse et al. (2007) list the
occurrence of eleven genera of Notoungulata, specifically the family
Toxodontidae, for outcrops from the Solimões Formation. Most of these
listed taxa are described and have key features only in dental and
cranial remains, such as Abothrodon (Paula-Couto, 1944) and Trigodo-
nops (Kaglievich, 1930), but do not have, to our knowledge, any formal
description of carpal and tarsal bones to the Notoungulata from Acre
State outcrops. Thus, the general morphology described above allow us
to identify MZUFV-P 0008 as a Toxodontidae.
M.B.d.S. Lacerda, et al.
3.2. Taphonomy of collected specimens
Based on the studied material, some taphonomical comments can be
provided to explain the differential material preservation, and to in-
terpret their correlation with possible effects generated by the inferred
depositional paleoenvironment of the newly described locality. The
fossils described here can be systematized into two distinct taphono-
mical classes:
Class 1: The materials are usually small and fragmented, but do not
present rounding edges. The break is usually acute and the external
surface is well-preserved without wear signals. This category is inter-
preted as low energy transport of dismembered carcasses and/or iso-
lated bones, explaining the absence of transport marks. This class ma-
terials are probably deposited in overbank fines or floodplains being
subjected to trampling, explaining the acute breaks. Regarding the well-
preserved surface, two hypotheses are equally plausible: i) the materials
were not exposed to the atmosphere, being covered by thin layers of
sand or mud; or ii) the materials were exposed to the atmosphere,
however, the high humidity and the vegetal cover (preventing direct
sun influence) protected the decomposing bones from desiccation. The
specimens showing these features are: MZUFV-P 0004, MZUFV-P 0005,
MZUFV-P 0006, MZUFV-P 0007, MZUFV-P 0008, MZUFV-P 0010,
MZUFV-P 0011, MZUFV-P 0012, MZUFV-P 00013, MZUFV-P 0014,
MZUFV-P 0015, MZUFV-P 0016, MZUFV-P 0037, MZUFV-P 0038,
MZUFV-P 0039, MZUFV-P 00042, MZUFV-P 0043, MZUFV-P 00045,
MZUFV-P 0046, MZUFV-P 0047, MZUFV-P 0048, MZUFV-P 0049,
MZUFV-P 0050, MZUFV-P 0051, MZUFV-P 0052, MZUFV-P 0053,
MZUFV-P 0054, MZUFV-P 0055, MZUFV-P 0056, MZUFV-P 0057,
MZUFV-P 0058, MZUFV-P 0059, MZUFV-P 0060, and MZUFV-P 0061.
Class 2: The materials present deformations in their external sur-
face, being usually broken in acute angles. Here we interpret that those
fossils suffered similar pre-diagenetic processes as described by class 1.
However, during the early fossildiagenesis those materials were more
superficially deposited enabling the calcite recrystallization and dis-
placement of the superficial bone tissue (see Holz and Schultz, 1998 for
further information on taphonomy). This interpretation is based on the
presence of expanded clays into the bones. The specimens showing
these characteristics are: MZUFV-P 0009, MZUFV-P 0044, MZUFV-P
0062, MZUFV-P 0063, MZUFV-P 0064, MZUFV-P 0065, MZUFV-P
0066, MZUFV-P 0067, MZUFV-P 0068, and MZUFV-P 0069.
4. Discussion
4.1. New fossiliferous locality
The Solimões Formation has a great diversity of fossil species and a
large number of outcrops, located mainly in Acre and Southern
Amazonas State (Campos et al., 1976; Price et al., 1977; Cozzuol, 2006;
Ribeiro et al., 2013). However, several outcroppable localities in the
region refer to sites that are situated on the river slopes and along their
course (e.g., Bergqvist et al., 1998; Alvarenga and Guilherme, 2003;
Cozzuol, 2006; Cozzuol et al., 2006; Hsiou et al., 2009; Fortier et al.,
2014; Souza et al., 2016, 2018). The development of the Amazonia
rainforest over the rocks and soils derived from Solimões Formation
make the outcrops more easily recognizable in regions with more ero-
sion such as river margins and roads/highways cuts (e.g., Bocquentin
and Melo, 2006; Cozzuol, 2006; Hsiou et al., 2010). Consequently, very
few works in the northern region of Brazil present outcrops that are not
situated around watercourses (e.g., Cozzuol, 2006). This becomes more
evident when we look at the several localities that are described in the
pioneering work developed by Campos et al. (1976) and Price et al.
(1977), covering an extensive area of the Solimões Formation. How-
ever, the new fossiliferous locality described here represents one of the
few exceptions for the Solimões Formation, being located in a highway
section with a region of trifling vegetation cover, that makes this out-
crop more accessible, with less physical boundaries limiting access.
11
Journal of South American Earth Sciences 101 (2020) 102588
4.2. Paleobiodiversity
The diversity of taxonomic groups from the outcrops of the Solimões
Formation is unquestionable, and the fossil record of this formation is
the one of the best records of the Cenozoic biota providing extensive
knowledge about the evolution of proto-Amazonian fauna and flora
(Cozzuol, 2006; Hoorn et al., 2010a,b; Ribeiro et al., 2013; Sá and
Carvalho, 2017; Cidade et al., 2019a; Sá et al., 2020). Some localities in
the Solimões Formation have more representativity of this biological
diversity, such as the Talismã and Patos localities (see Bergqvist et al.,
1998; Cozzuol, 2006; Hsiou and Albino, 2009; Negri et al., 2010; Souza-
Filho and Guilherme, 2015; Souza et al., 2016). Both assemblages have
been well sampled and provide much of the biodiversity that allows us
to understand the development of the environments of the proto-
Amazonian. The new fossiliferous locality described here (see session
2.1), although not presenting records of several animal groups (so far),
still presents great diversity of vertebrates, comprising the record of
crocodiles, turtles, and mammals in a single locality (see session 4.1),
which is not common in other Solimões Formation outcrops, but was
represented at this new site in a single collection effort.
Crocodylia is greatly representated in Solimões Formation, being
well known, abundant, and diverselly recorded (Souza-Filho, 1987;
Fortier et al., 2014; Souza et al., 2016, 2018; Cidade et al., 2019a,b,c;
Souza-Filho et al., 2017, 2019). Some of the specimens described here,
such as the genus Mourasuchus (Fig. 4) and Caiman (Fig. 6) are rela-
tively well recorded in the Solimões Formation (e.g., Mourasuchus –
Price, 1964; Souza et al., 2016; Cidade et al., 2019b,c; Caiman – Souza-
Filho, 1987; Cozzuol, 2006; Fortier et al., 2014). However, the Mela-
nosuchus (Fig. 5) described here represents the first postcranial record
of its genus for the Early Cenozoic of Solimões Formation (described in
a previous note by Lacerda et al., 2017), since the other known record
of this genus is represented by cranial remains (Souza-Filho et al.,
2017). We reinforce that Melanosuchus fossil record extend, at least, to
the Paleogene/Neogene of Solimões Formation, and these records are
still restricted to the outrcrops of the northern Brazil (Lacerda et al.,
2017; Souza-Filho et al., 2017; Cidade et al., 2019a). Those Melano-
suchus specimens found in Solimões Formation are in agreement with
the fossil occurrence of this genus in other formations in South America,
such as Urumaco Formation, from Upper Miocene of Venezuela
(Medina, 1976; Foth et al., 2017; Cidade et al., 2019a).
The side-necked turtle findings (Fig. 7) extend geographically the
diversity of both, Chelidae and Podocnemididae, record in the Solimões
Formation, which are already widely registered (e.g., Campos et al.,
1976; Gaffney et al., 1998, 2002; Carvalho et al., 2002; Bocquentin
et al., 2001; Bocquentin and Melo, 2006; Meylan et al., 2009; Ferreira
et al., 2016). In addition, it reinforces the notion that Chelus represents
one of the most abundant representants in this formation, with several
occurrences (e.g., Campos et al., 1976; Bocquentin et al., 2001; Ferreira
et al., 2016).
Finally, the occurrence of a new specimen of Toxodontidae (Fig. 8),
although is not new, considering the diversity and abundance of this
clade in the Solimões Formation (Cozzuol, 2006; Latrubesse et al.,
2007; Negri et al., 2010), represents another piece in the paleobio-
geography history of this group, expanding its occurrence to the
northern region of Acre State, and to south limits of Solimões basin,
confirming the abundance of the group in rocks proceeding from the
Solimões Formation.
As discussed above, the records presented by this new locality
confirm the fossil potential of this outcrop adding more scientific in-
formation related to the proto-Amazonian biota, and also adding an-
other piece in the historical scope of the studies carried out in the
Solimões Formation, which have been developed for over a century.
Thus, new controlled excavations can further sample the proto-
Amazonian biota in this specific locality, hitherto unknown biological
information may emerge.
M.B.d.S. Lacerda, et al.
4.3. Taphonomy
The fossils described in this paper have distinct preservation pat-
terns, being systematized into two distinct classes according to their
preservation (see session 3.2). Regarding the taphonomical scenario of
the remaining localities of the Solimões Formation, some of those lo-
calities have their own taphonomical explanations, such as the pre-
sented by Souza et al. (2016) for outcrops along the Acre River. How-
ever, taphonomic studies and interpretations in outcrops of the
Solimões Formation are scarce and remain restricted to the southern
part of the Acre State (see Souza et al., 2016 and references therein).
Furthermore, complete taphonomical studies, similar to those carried
out for Bauru Group (e.g., Azevedo et al., 2013; Bandeira et al., 2018)
including several biological information such as plants, invertebrates,
and vertebrates, besides lithological studies and characterization of the
sediments in different outcrops from Solimões Formation still in need to
better understand the origin and preservation patterns of these Paleo-
gene/Neogene fossils. The taphonomic hypotheses for the two classes
described in this work might suggest a high sinuosity fluvial mean-
dering paleoenvironment, and this scenario is consistent with ob-
servations at the south limits of Acre State (Souza et al., 2016). How-
ever, this sedimentary deposition scenario cannot be generalized to
other outcrops from the Solimões Formation considering its extensive
horizontal area and possible different processes that may have occurred
throughout the depositional history of the formation, being our ta-
phonomic conclusions restricted to the area under study of this new
outcrop in the southern limits of Solimões basin.
5. Conclusion
The fossil findings from the new locality, presented in this work,
address some important informations for the paleobiodiversity and
paleogeographical distribution of some mammals and reptiles groups
from the Paleogene/Neogene of Solimões Formation. Furthermore, we
reinforce four important considerations: 1) the new locality is on a
highway side, in a slope area, increasing our knowledge about the
Solimões Formation in the interfluvial regions; 2) the outcrop seems to
be a rich fossil tetrapod assemblage, presenting fossils of side-necked
turtles (Chelidae and Podocnemididae), notoungulates (Toxodontidae),
and crocodiles (Caimaninae), including the first postcranial fossil ma-
terial attributed to the genus Melanosuchus from the Solimões
Formation; 3) even though some materials have a fragmentary nature,
it was possible to carry out a broad systematic study, allowing the ac-
cess to key morphology features of some taxonomic groups and thereby
presenting an accurate description of the morphology; and, 4) based on
the preservational pattern of the studied material, a local taphonomical
systematization is proposed in the form of two distinct classes of pre-
servation related to distinct fossilization processes that seem to have
been preserved in a fluvial meandering paleoenvironment. We highlight
the importance of the findings in this new outcrop from interfluvial
regions (e.g., highways and farms), which make possible the execution
of field works in the Acre region all year long, not being restricted to the
dry seasons as the more common riverine outcrops usually explored in
Solimões Formation, both in Acre and Amazonas States.
Author statement
We declare that this manuscript is original, the results have not been
published before and is not currently being considered for publication
elsewhere. We provide the confirmation that the manuscript has been
read and approved by all the listed authors to be published in Journal of
South American Earth Sciences.
We further confirm that the order of authorship listed in the
manuscript has been approved by all of us and the contribution of each
author is detailed hereafter in the order of authorship: Mauro B. S.
Lacerda: Conceived and designed the research, performed the
12
Journal of South American Earth Sciences 101 (2020) 102588
identifications and morphological descriptions, prepared figures and/or
tables and wrote the manuscript, reviewed drafts of the paper, ap-
proved the final draft. Lucy G. Souza: Conceived and designed the
research, performed the identifications and morphological descriptions,
wrote the manuscript, reviewed drafts of the paper, approved the final
draft. Leonardo S. Lobo: Performed the identifications and morpho-
logical descriptions and wrote the manuscript, reviewed drafts of the
paper, approved the final draft. Carlos E. G. R. Schaefer: Collected the
samples, performed the geological descriptions, reviewed drafts of the
paper, approved the final draft. Pedro S. R. Romano: Wrote the
manuscript, reviewed drafts of the paper, approved the final draft and
provided the supervision.
Declaration of competing interest
The authors declare no conflict of interest.
Acknowledgments
The authors wish to thank the curators who gave us access to the
collections: Rodrigo Machado (DNPM); Matthew A. Brown and J. Chris
Sagebiel (TMM); Luciana Barbosa de Carvalho and Manoela Woitovicz
Cardoso (MN/UFRJ); Kevin de Queiroz, Kenneth A. Tighe and Addison
Wynn (USNM); David C. Blackburn and Coleman M. Sheehy III (UF);
David A. Kizirian and Lauren Vonnahme (AMNH); and Castor Cartelle
(PUC-MG). We also thank to Kamila L. N. Bandeira (MN/UFRJ) for
taphonomical comments and discussion about all specimens; Natália
Benevenuto (UFV) and Thiago Mariani (MN/UFRJ) for comments about
turtle specimens; Giovanne Cidade (UFSCar) for comments about
Mourasuchus; and also to Camila B. Pinto (UFMG) for revision of the
manuscript draft and help with grammar. The author MBSL thanks for
fundings provided by the Fundação de Amparo à Pesquisa do Estado de
Minas Gerais (FAPEMIG) and Coordenação de Aperfeiçoamento de
Pessoal de Nível Superior (CAPES); LGS thanks for the PhD scholarship
granted by CAPES and to support granted to Paleo and Geohistory ex-
hibition at Museu da Amazônia [PRONAC 183808].
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.jsames.2020.102588.
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