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Palaeogeography, Palaeoclimatology, Palaeoecology

journal homepage: www.elsevier.com/locate/palaeo

Taphonomic modes on uvial deposits of the Monte Hermoso Formation

(early Pliocene), Buenos Aires province, Argentina
Rodrigo L. Tomassini a,, Claudia I. Montalvo b
a
b

CONICET, Departamento de Geologa Universidad Nacional del Sur, San Juan 670, 8000 Baha Blanca, Buenos Aires, Argentina
Facultad de Ciencias Exactas y Naturales, Universidad Nacional de La Pampa, Avenida Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina

a r t i c l e

i n f o

Article history:
Received 21 May 2012
Received in revised form 23 October 2012
Accepted 27 October 2012
Available online 6 November 2012
Keywords:
Fluvial deposits
Taphonomic modes
Vertebrates
Monte Hermoso Formation
Pliocene
Argentina

a b s t r a c t
The sedimentary deposits of the Monte Hermoso Formation, outcropping at its type locality (Farola Monte
Hermoso, Buenos Aires province, Argentina), accumulated through uvial processes acting on highly sinuous
rivers. The Monte Hermoso Formation comprises a rich vertebrate fauna corresponding to the Montehermosan
Stage/Age (early Pliocene), which includes numerous and diverse vertebrate remains. Comparative taphonomic
analysis indicated that the faunal assemblages coming from two different fossiliferous levels, FL 1 and FL 2, recognized in oodplain deposits and channel deposits respectively, are composed of remains affected by diverse
taphonomic processes and therefore representing distinct taphonomic histories. The variations recorded between both fossiliferous levels, with respect to their taphonomic attributes and sedimentary context, allowed
the recognition of two distinct taphonomic modes: oodplain in the FL 1 and channel-lag in the FL 2. These
taphonomic modes represent different conditions within the range of taphonomic histories for the Monte
Hermoso Formation assemblage formed by attritional accumulation of remains. At the moment, continental systems linked to uvial environments have been poorly analyzed for the South American Neogene; this study contributes new information on taphonomic and paleoenvironmental characteristics of this type of deposit.
2012 Elsevier B.V. All rights reserved.

1. Introduction
Several taphonomic studies have evaluated mammal assemblages
from the Neogene in Argentina, providing knowledge of the
organisms present, their relation with the environment and the processes affecting their remains at different stages following their
death (e.g. Tauber, 1997a,b; Montalvo, 2002b; Montalvo et al., 2008;
Pomi and Scanferla, 2008; Pomi, 2009; Tomassini and Montalvo,
2010b; Tomassini et al., 2010a,b; Tomassini, 2012). Many of the Neogene fossil deposits in central and south of the Pampean Region (Argentina), including Farola Monte Hermoso, are interpreted as uvial
deposits (e.g. Deschamps, 2005; Zrate, 2005; Folguera and Zrate,
2009). In some case, these deposits show similar features to those observed in plain river systems present in the area today. However, so
far, the abundant fossils from these deposits were analyzed mainly
from a systematic and biochronostratigraphic viewpoint (Cione and
Tonni, 2005).
Fluvial environments present a number of characteristic features
related to channel size, sediment load and variations in water discharge, promoting the development of different sub-environments
that constitute potential scenarios for vertebrate fossil preservation
(Behrensmeyer and Hook, 1992). This is interesting from a taphonomic
point of view, given that the abundance and taxonomic diversity of
Corresponding author. Tel.: +54 291 4595101x3063.
E-mail address: rodrigo.tomassini@yahoo.com.ar (R.L. Tomassini).
0031-0182/$ see front matter 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.palaeo.2012.10.035

fossil remains and the processes affecting them vary according to the
type of sub-environment, thus generating different taphonomic histories
for each (e.g. Badgley, 1986a,b; Behrensmeyer, 1988; Behrensmeyer and
Hook, 1992; Smith, 1993; Pereda-Suberbiola et al., 2000; Wilson, 2008;
Csiki et al., 2010).
The Monte Hermoso Formation (early Pliocene), at its type locality
of Farola Monte Hermoso, in southeastern Buenos Aires province
(Pampean Region, Argentina), has been interpreted as sedimentary
rocks deposited through uvial dynamics, similar to those of the
Miall (1985) model muddy ne-grained rivers (Zavala, 1993;
Zavala and Navarro, 1993). This is a well known paleontological locality from the Argentinean Neogene, visited and studied by several
authors (e.g. Darwin, 1846; Ameghino, 1887; Bonaparte, 1960), who
highlighted the abundance and diversity of its vertebrate remains.
However, there are few mentions about the features of preservation
of the remains (e.g. Tomassini and Montalvo, 2010b; Tomassini et
al., 2010b). In a recent study of the faunal assemblage collected in
this formation, which includes an extensive taphonomic analysis
(Tomassini, 2012), signicant differences were recognized in the attributes of fossil vertebrate remains from different fossiliferous levels.
The main goal of this paper is to describe, analyze and compare the
characteristics of remains recovered from two fossiliferous levels (FL),
the FL 1 (recovered from oodplain deposits) and FL 2 (recovered
from channel deposits), of this stratigraphic unit. The focus was placed
on these two levels because they were the only ones in which a representative number of specimens were found to allow such an evaluation.

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Although other fossiliferous levels of this formation show particular

taphonomic characteristics, it was considered that the number of specimens recovered from each of them was insufcient for inclusion in this
analysis. As sedimentary and taphonomic processes are intertwined
(Behrensmeyer, 1991), even though the sedimentary characterization
is independent from the type of preservation, our approach was to
consider both the stratigraphic context together with the taphonomic
analysis.
The taphonomic evaluations realized in uvial deposits from South
American Neogene are very scarce. Differences between the two
fossiliferous levels in their taphonomic attributes, the stratigraphic
context and their lithologic characteristics, point to two taphonomic
modes (sensu Behrensmeyer, 1988; Behrensmeyer and Hook, 1992)
for the Monte Hermoso Formation. Different taphonomic histories
are interpreted linked to the presence of assemblages formed from
the attritional accumulation of remains in plain uvial environments.
2. Geographic and stratigraphic setting
Farola Monte Hermoso is located on the coast of southern Buenos
Aires province, Argentina (S 385801, W 614143), 53 km east of
Baha Blanca city (Fig. 1). From a morphostructural point of view
this area is located in the Positivo Bonaerense, unit encompassing
Tandilia and Ventania mountain ranges and also the Claromec
basin (Yrigoyen, 1975; Ramos, 1984; Folguera and Zrate, 2009).
Coastal deposits form a three km marine cliff, 15 m maximum height,
in which fossil-bearing Neogene sediments of the Monte Hermoso
Formation are exposed. The complete sedimentary sequence includes
three stratigraphic units (Zavala, 1993): the Monte Hermoso Formation (LowerMiddle Pliocene), Puerto Belgrano Formation (Upper
Pleistocene) and Punta Tejada Formation (Upper PleistoceneMiddle
Holocene).
The Monte Hermoso Formation (Zavala, 1993) crops out at the
abrasion platform and the lower part of the cliff, along its entire extension, with a maximum thickness of up to 6 m. This formation is

283

very important due to the abundance and diversity of vertebrate fossils and because it is the type locality of biostratigraphic units of the
South American mammal bearing Pliocene, the Montehermosan and
Lower Chapadmalalan stages/ages (Cione and Tonni, 1995, 2005).
Several authors (e.g. Wichmann, 1916; Kantor, 1922; Vignati, 1925;
Kraglievich, 1946) recognized in this formation the existence of two
levels, differentiated by color and separated by a discontinuity. They
proposed that these levels are temporarily different. With regard to
this, Cione and Tonni (1995, 1996, 2001, 2005) published revisions of
the biostratigraphy and biochronology of Monte Hermoso Formation
at Farola Monte Hermoso. These authors recognized two chronologically successive biozones, the Biozone of Trigodon gaudryi, biostratigraphic
basis of the Montehermosan Stage/Age (late Mioceneearly Pliocene),
and the Biozone of Neocavia depressidens, biostratigraphic basis of the
Lower Chapadmalalan Stage/Age (early Pliocene). However, other authors (e.g. Frenguelli, 1928; Bonaparte, 1960; Deschamps et al., 2012;
Olivares et al., 2012; Tomassini, 2012) emphasized that the stratigraphic and paleontological differences are minimal, so it would not be possible to dene with certainty two levels with different ages.
Zavala and Navarro (1993) interpreted that the Monte Hermoso
Formation was deposited through a uvial dynamic of high sinuosity
muddy ne-grained rivers. These authors identied architectural elements of channel, lateral accretion deposits and overbank deposits.
From a taphonomic point of view, the most interesting sedimentary
deposits of this formation (oodplain and channel deposits) include
the FL 1 and the FL 2. Both fossiliferous levels have abundant and diverse vertebrate remains.
The FL 1 is included in deposits mainly composed by muddy siltstones, with colors that range from reddish brown to yellowish
brown. Reddish brown sandy siltstones and silty sandstones are
present too. These deposits have a massive structure in the middle
upper sector, while in the lower one they show a very ne lamination.
Thickness ranges from a few centimeters to a maximum of 4 m
(Fig. 2). In some sections there are common burrows produced by rodents (Actenomys priscus) and probably by other micromammal taxa

Fig. 1. Location map showing Farola Monte Hermoso, Buenos Aires province, Argentina.

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3. Materials and methods

Fossil remains were collected with an exhaustive control of their
stratigraphic provenance. They are deposited in the Museo Municipal
de Ciencias Naturales Carlos Darwin collection (Punta Alta, Buenos
Aires), under the acronym MD-FM. The present study only considered
remains collected at the FL 1 and the FL 2.
The following taphonomic features were taken into account:

Fig. 2. Stratigraphic section of the Monte Hermoso Formation in Farola Monte Hermoso
locality.
Modied from Zavala and Navarro (1993).

(Tomassini et al., 2008), fossil footprints produced by Xenarthra

mammals (Aramayo and Manera de Bianco, 1996) and coprolites
assigned to carnivore mammals (Tomassini and Montalvo, 2010a).
In addition, desiccation cracks, rhizoliths, and traces attributed to
Taenidium barretti (Bradshaw, 1981) were identied. Tomassini
(2012) indicated that this level represents oodplain deposits originated from the settling from suspension materials provided by the
channel during ooding events. The presence of fossil footprints,
coprolites and burrows indicates a favorable environment for the development of terrestrial vertebrates, in accordance with what usually
happens in this type of sub-environment (Behrensmeyer and Hook,
1992).
The FL 2 is included in deposits ranging in thickness from 5 to
60 cm, presenting dark brown to greenish hues and located few
meters apart from each other. These are clast-supported breccias,
composed by embedded silty and sandy intraclasts over 15 cm in diameter, some of them rounded off, and a silt-sandy matrix. In some
sections, these deposits have a massive structure; while others have
a planar cross-bedding (Fig. 2). Generally, the deposits rest on erosion
surface over previously described oodplain deposits. Based upon the
features described, Tomassini (2012) pointed out that these levels
represent channel deposits.

1. Anatomic and taxonomic representation. Frequency and diversity of

taxa were estimated with the following indexes. The number of recovered remains (NR) represents the sum of undetermined fragments and specimens identied anatomically and taxonomically
(Blasco Sancho, 1995). The number of identied specimens per
taxon (NISP) was calculated with the remains (bone, tooth or fragments of either) identied anatomically and taxonomically
(Badgley, 1986b). A third index is the minimal number of individuals (MNI) (Badgley, 1986b), calculated from the most abundant
skeletal part. Most remains from both fossiliferous levels belong
to mammals, and teeth are the most abundant elements. Therefore, the most frequent tooth was chosen to obtain the MNI for
each taxon. Dasypodids and glyptodonts (Xenarthra, Cingulata),
as well as tortoises (Reptilia, Chelonii), are mainly represented
by few isolated osteoderms. On the other hand, Osteichthyes are
mainly represented by vertebrae. The MNI value, in these cases,
was considered one for each taxa. In the case of other vertebrates,
MNI was calculated after the most frequent skeletal element. The
vertebrates were classied into two categories, according to the
estimated body mass, calculated after Vizcano and Faria (1999)
and Montalvo (2004). Category 1 includes microvertebrates with an
estimated body mass b5 kg and category 2 includes macromammals
with an estimated body mass >5 kg. These categories allow evaluating whether there is a bias linked to the size of the remains.
The index isolated teeth/vertebrae (Behrensmeyer, 1975;
Behrensmeyer and Dechant-Boaz, 1980) was calculated considering only mammalian remains. This index takes into account differences in structural density between these two
elements and provides a criterion for determining if a group of
bones was selectively biased before burial. It therefore can be considered like an indicator of the degree of hydrodynamic sorting. An
index of 1 would indicate the absence of sorting, while values close
to 1 represent low development of sorting, and values higher than
1 indicate a greater degree of sorting. On the other hand, the possible
dispersion and transport have been focused on evaluating the effect
of rearrangement of remains by uvial transport. It was evaluated
using skeletal elements categories, according to their susceptibility
to transport, proposed by Voorhies (1969), Dodson (1973) and
Korth (1979). Thus, an assemblage composed mostly of elements included in any of these groups represents evidence of selection processes triggered by the original thanatocoenoses (Behrensmeyer,
1975).
2. Geometric shape. This attribute was evaluated from relationships
between length (D1), width (D2) and maximum thickness (D3)
of fossil remains. These measures evaluated in each specimen
allowed their assignment to one of the three geometric categories:
planar (D1 ~ D2, D2 D3), cubic (D1 ~ D2, D2 ~ D3) and elongated
(D1 D2, D2 ~ D3). These shapes were determined and plotted
into Flinn's (1978) diagrams. This attribute is used to assess
whether the assemblage was biased before burial.
3. Weathering degree. Characteristic atmospheric weathering taking
place before burial was taken into account. The following categories
proposed by Andrews (1990) were considered for micromammals;
0: unweathered remains; 1: remains showing surface slight splitting,
parallel to the brous structure; 2: remains showing a greater
number of splitting, but little exfoliation; and 3: remains showing
deep splitting and loss of segments between splits. In the case of

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4.

5.

6.

7.

8.

9.

macromammals, the categories proposed by Alcal (1994) were considered; 1: intact remains; 2: remains showing surface loss of bone
material; and 3: remains showing deep loss of bone material.
Abrasion degree. This attribute was determined following the categories proposed by Alcal (1994), for both micro- and macromammals;
namely: 1: intact remains; 2: rounded remains; and 3: polished
remains.
Degree and type of breakage. The degree of breakage was evaluated
following categories proposed by Alcal (1994); 0: complete
bones; 1: incomplete bones. The types of fracture were only evaluated for long bones, and established on the basis of the angle
formed by the main dimension axis and the border of fracture;
longitudinal (029), oblique (3059) and transverse (6090)
fractures were considered.
Degree of bioerosion. Two kinds of bioerosive trace fossils were recognized. One of them is due to root growth on the bone element
sediment interface (category 1), and assigned to the Corrosichnia
ethological category (Mikul, 1999; Montalvo, 2002a). The second kind of trace fossil is due to soil fungal and/or bacterial activity
(category 2) (Montalvo, 2004). Some materials presented both
types of bioerosive trace fossils (category 3).
Skeletal articulation. This feature was evaluated according to the
stages of articulation of skeletal elements proposed by Behrensmeyer
(1991); articulated, disarticulated but associated, and isolated and
dispersed.
Spatial density. This attribute was evaluated from number of remains by surface unit (Behrensmeyer, 1991). In FL 1 the density
was not determinate due to the wide spatial distribution of the remains and the distance between them. In FL 2 three areas of different size were delimited (A = 1 m 2; B = 3.6 m 2; and C = 2.8 m 2).
The areas were a few meters apart from each other and correlated
to each other from the stratigraphic point of view.
Orientation and dip/trend. Orientation data were plotted in a 360
rose diagram, at 10 intervals; and dip/trend data were plotted in
a stereographic projection.

Only mammal remains were considered for the evaluation of the

degree of weathering, abrasion, breakage and bioerosion, given that
they were the only vertebrates in which the number of specimens
found was representative in both levels considered.
To evaluate the attributes of these two assemblages, we determined whether the remains were accumulated, resedimented or
reelaborated. According to Efremov (1940), accumulation is a unique
and unrepeatable process involving the passage from the biosphere to
the lithosphere. Two other taphonomic processes may affect the remains after being accumulated: resedimentation and reelaboration.
The rst means the displacement of remains prior to the burial.
Reelaboration (or taphonomic reworking) means the exhumation
and displacement of remains. Both processes may be iterative and
generate different degrees of alteration (Fernndez-Lpez, 1991,
2000). We also analyzed whether the remains were autochthonous,
para-autochthonous or allochthonous (Fernndez-Lpez, 1990, 2000).

4. Taphonomic analysis
The FL 1 registered a NR = 195; 90.77% of these were identiable
specimens. A total of 28 taxa were identied, 22 of which were mammals, two avian, two reptiles, one amphibian and one bony sh. The
NISP = 177, was mainly represented by mammals (91.53%), and
the NMI was 45 (Table 1). The FL 2 registered a NR = 473; 49.26%
of these were identiable specimens. A total of 15 taxa were included, 11 of which were mammals, one avian, two amphibians
and one bony sh. The NISP = 233, was mainly represented mainly
by mammals (66.52%) and bony sh (30.47%), and the NMI was 25
(Table 1).

285

Table 1
List of the vertebrates represented in both levels and NISP and MNI values for each
taxon.
FL 1

FL 2

NISP

MNI

NISP

MNI

Clase Osteichthyes
Osteichthyes indet.

71

Clase Amphibia
Ceratophrys sp.
Rhinella cf. R. pisanoi

5
1

2
1

Clase Reptilia
Tupinambis sp.
Chelonoidis cf. Ch. australis

3
2

1
1

Clase Aves
Dendrocygna sp.
Chunga incerta

1
1

1
1

Clase Mammalia
Rodentia indet.
Necromys bonapartei
Auliscomys formosus
Pithanotomys sp.
Neophanomys sp.
Actenomys priscus
Caviidae indet.
Palaeocavia sp.
Orthomyctera sp.
Lagostomus (Lagostomopsis) sp.
Paramyocastor diligens
Phugatherium cataclisticum
Eoauchenia aff. E. primitiva
Macraucheniidae indet.
Toxodontidae indet.
Toxodon chapalmalensis
Pseudotypotherium sp.
Paedotherium sp.
Paedotherium bonaerense
Argyrolagidae indet.
Proscelidodon cf. P. patrius
Lestodon sp.
Macrochorobates chapadmalensis
Doellotatus inornatus
Plaina cf. P. intermedia
Glyptodontidae indet.
Plohophorus guratus
Eleutherocercus antiquus
Mammalia indet.
Total

18
1
2
2
1
14
1
1
2
2
1
1
1
2
6
1
11
38
8

6
1
1
2

7
4
2
26
177

1
1
1
1
7

1
2
2
1
1
1
1
1
1
2

1
1
1
1

2
2

45

36

13
8
1

3
2
2

3
31

4
1
1

1
48
233

1
1
1

1
3

1
1

25

Microvertebrate remains (category 1) predominated in both levels

(71.22% in FL 1 and 93.56% in FL 2), including the following mammalian families: Cricetidae, Octodontidae, Chinchillidae, Caviidae and
Echimyidae (Rodentia), Didelphidae (Didelphimorphia), Argyrolagidae
(Polydolopimorphia), Hegetotheriidae (Notoungulata) and Dasypodidae
(Cingulata). The other taxa included in this category were Osteichthyes,
Amphibia (Bufonidae, Ceratophryidae), Reptilia (Teiidae, Testudinidae)
and Aves (Cariamidae, Anatidae).
Macromammal remains (category 2), including representatives
of the families Hydrochoeridae (Rodentia), Mesotheriidae and
Toxodontidae (Notoungulata), Proterotheriidae and Macraucheniidae
(Litopterna), Mylodontidae (Tardigrada) and Glyptodontidae and
Pampatheriidae (Cingulata), were abundant in the FL 1 but very
scarce in the FL 2.
Different skeletal elements assigned to mammals were represented in both assemblages (Fig. 3). In the FL 1, jaws and isolated teeth
being the most frequent, while skull elements, vertebrae, metapodials
and phalanges, were less represented. On the other hand, humerii,
tibiae and femora predominated among long bones. Isolated teeth
were the most frequent in the FL 2, while jaws, metapodials and
phalanges were also well represented. Humerii and femora were

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Fig. 3. Skeletal element distribution from both fossiliferous levels.

noteworthy among long bones. The remaining skeletal elements were

present in low frequencies in both levels. Thus, the different groups
suggested by Voorhies (1969), Dodson (1973) and Korth (1979), for
the categorization of mammalian skeletal elements according to
their susceptibility to hydrodynamic transport, were present at both
levels. On the other hand, the values obtained for the teeth/vertebrae
index were 1.33 in the FL 1 and 3.75 in the FL 2.
Very few skeletal elements (NR = 15) were assigned to nonmammalian vertebrates in the FL 1, while they were more abundant
in the FL 2 (NR = 78) and were mainly represented by Osteichthyes vertebrae (87.18%).
Cubic shapes predominated in both levels, although this was more
evident in the FL 2 (Fig. 4). The FL 1 had a greater diversity in shapes,
with a higher proportion of elongated and planar shapes.
In the FL 1, 25% of the micromammal skeletal elements (Fig. 5A)
present only slight evidences (categories 1 and 2) of weathering
(Fig. 6A,B). Among macromammals, 33.33% showed signs of slight
weathering (category 2), while 9.26% showed signs of extreme
weathering (category 3) (Fig. 7A,B). The most affected skeletal elements were vertebrae, ribs and isolated osteoderms. In the FL 2,
only 11.43% of micromammal skeletal elements were affected, all
with slight evidences (category 1). The 13.33% of macromammal skeletal elements were affected with slight signs (category 2). The most
affected skeletal elements were vertebrae and ribs.
In the FL 1, only 5.56% of micromammal (Fig. 6C,D) and 7.41% of
macromammal (Fig. 7C,D) skeletal elements presented modications
due to abrasion (Fig. 5B), all of them rounded (category 2). In the FL 2,
33.57% of micromammal remains showed slight modications (category
2), and 10.72% showed an extreme degree of abrasion (category 3); while
26.67% of macromammal remains showed slight modications, in all
cases corresponding to category 2.
At both levels, the skeletal elements least affected by breakage
were teeth and metapodials. In the FL 1, 26.85% and 48.15% of
micromammal and macromammal remains respectively were preserved whole (Fig. 5C). Among the incomplete macromammal remains predominate the ones that have a single fracture, while most
of the micromammal remains evidenced several fractures. In both
cases (Figs. 6EG; 7EG), longitudinal fractures were the most abundant (68.42% and 72.72% respectively), followed by oblique (26.32%

and 18.18% respectively) and transverse (5.26% and 9.10% respectively).

Few skeletal elements which have longitudinal fractures showed slight
signs of rounding on the fracture surface.
At the FL 2, only 8.57% and 13.33% of the micro- and macromammal
remains respectively were preserved whole (Fig 5C). Among the incomplete micro- and macromammal remains predominate the ones that
have several fractures. In both cases, transverse fractures were the
most common type (71.42% and 66.67% respectively), while longitudinal (14.29% and 22.22% respectively) and oblique fractures (14.29%
and 11.11% respectively) were much less frequent. More than 50% of

Fig. 4. Flinn diagrams showing geometric shape distribution of remains from both fossiliferous levels. D1 = maximum length; D2 = maximum width; D3 = maximum
thickness.

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287

Fig. 5. Pie diagrams showing different taphonomic variables evaluated in both fossiliferous levels. A. Weathering. B. Abrasion. C. Breakage. D. Bioerosion.

remains with transverse fractures showed signs of rounding and polish

on the fracture surface.
Bioerosive traces were recognized at both levels (Fig. 5D). Particularly, trace fossils assigned to the Corrosichnia ethological category
(category 1) were frequent in remains from FL 1. The trace fossils
due to soil fungal and/or bacterial activity (category 2) were equally
frequent at both levels (Fig. 8A,B). Some remains, especially those
from the FL 1, jointly presented both types of traces (category 3).
The 82.73% of specimens found at the FL 1 were isolated and dispersed, 12.23% were articulated, and 5.04% were disarticulated but associated. The articulated specimens include carapaces, limbs and
vertebrae from micro- and macromammals (Fig. 9). All specimens
found at FL 2 were isolated and dispersed.

The density of specimens could not be determined for the FL 1,

due to their wide spatial distribution and the distance from one another. On the other hand, the density of specimens for the FL 2 varied
between 38.93 and 52.86 remains/m 2.
Polymodal rose diagrams showing orientation were obtained for
both levels (Fig. 10). In the FL 1, there was no preferential orientation.
Data for FL 2 were mostly concentrated between values of 180270.
With reference to dip (Fig. 11), horizontal or semi-horizontal (15)
remains dominated in the FL 1. Few specimens were inclined >10,
whereas the maximum measured angle was 19. Conversely, inclination of remains was highly variable in the FL 2, with several specimens lying near the horizontal (between 1 and 10), as well as
other with high dip values, in general within the 6090 range.

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Fig. 6. Taphonomic attributes in skeletal elements of micromammals. A. Actenomys priscus, jaw with slight weathering (category 1). B. Actenomys priscus, jaw with slight weathering
(category 2). C. Mammalia indet., metapodial with slight abrasion (category 2). D. Mammalia indet., diaphysis with extreme abrasion (category 3). E. Mammalia indet., femur with
longitudinal fracture. F. Mammalia indet., femur with oblique fracture. G. Mammalia indet., metapodial with transverse fracture. Scale bar = 0.5 cm.

5. Discussion
5.1. Fossiliferous level 1 (FL 1)
The assemblage recovered from FL 1 showed, according to NISP, a
high degree of anatomical and taxonomic resolution. It included a rich
vertebrate fauna that consisted mainly of micro- and macromammals
and to a lesser extent, birds, sh, amphibians and reptiles. Moreover,
the highest MNI at this level, in relation to the estimate of channel deposits (FL 2), coincides with that expressed by Badgley (1986a,b) regarding this index, who said that in the oodplain the preserved
remains have a high probability of being part of the original assemblage, so that the data reect a good approximation of the real conditions. As indicated, the different groups suggested by Voorhies
(1969), Dodson (1973) and Korth (1979) were present at this level.
Jaws and isolated teeth predominate among those elements that are
more resistant to transport, while metapodials, phalanges and vertebrae are among those that are more easily transported by water.
However, the largest number of skeletal elements corresponding to

groups II and III indicates some selection for those remains with less
susceptibility to be mobilized (Behrensmeyer, 1975). The index was
1.33 pointing to an assemblage with a very low sorting degree. In
turn, this might allow us to infer that the water ows originated during the ooding events were mainly low-energy surface runoff, with
scarce ability to move and sort the remains. The high diversity of
shapes and sizes of remains supports the interpretation that sorting
processes had a low importance at this level (Alcal, 1994), in agreement with oodplain sub-environment.
The occurrence of different weathering categories, both microand macromammal skeletal elements, indicates that the remains
have been gradually incorporated to the substrate. However, the
dominance of specimens with intact surfaces might suggest that, in
these cases, the time of exposure was not very long (Behrensmeyer,
1978). Considering that the percentage of affected micromammal remains was low and that, in turn, all exhibited very low intensity, it is
estimated that small dimensions thereof would have triggered rapid
burial (Behrensmeyer, 1978, 1982; Andrews, 1990), which is essential for preservation because they have little resistance to destructive

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289

Fig. 7. Taphonomic attributes in skeletal elements of macromammals. A. Pseudotypotherium sp., phalange with slight weathering (category 2). B. Mammalia indet., metapodial with extreme weathering (category 3). C. Plohophorus guratus, osteoderm with slight abrasion (category 2). D. Mammalia indet., rib with slight abrasion (category 2). E. Pseudotypotherium sp.,
with longitudinal fracture. F. Pseudotypotherium sp., humerus with oblique fracture. G. Macraucheniidae indet., tibia with transverse fracture. Scale bar=0.5 cm.

processes (Behrensmeyer, 1991). By contrast, the most affected remains, both in percentage and intensity, belonged to macromammals,
for which it could be interpreted that due to their larger size, the subaerial exposure period must have been longer.
Only a few micro- and macromammal remains have slight evidence of abrasion. Therefore, it could be estimated that the residence
time of remains under abrasive conditions generated by sedimentary
particles in motion, might have been very short (Behrensmeyer,
1991).
Whole skeletal elements are abundant at this level, mainly teeth
and metapodials, which are skeletal elements with high structural

densities (Lyman, 1994). However, the predominance of incomplete

skeletal elements indicates that in many cases the time of exposure
to different destructive taphonomic processes must have been long.
The marked dominance of longitudinal fractures associated with
weathering, suggests that the breakage occurred at a pre-burial
stage (biostratinomic), when they were still exposed at the surface
(Behrensmeyer, 1978; Behrensmeyer et al., 1989; Marshall, 1989).
Oblique fractures might have happened at this same stage; however
there is no concrete evidence suggesting a relationship with some
process in particular. The few transverse fractures might have originated when the remains were already mineralized (fossil diagenetic

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Fig. 8. Bioerosion in skeletal elements. A. Mammalia indet., rib with bioerosion trace (category 1). Arrow indicates the affected area. B. Detail of the affected area. C. Mammalia
indet., humerus with bioerosion trace (category 2). Circle indicates the affected area. D. Detail of the affected area. Scale bar = 0.5 cm.

stage), probably due to the lithostatic load generated by overlying

sediments (Alcal and Martn Escorza, 1988, 1998).
The recorded bioerosion trace fossils indicate that the remains
were affected by soil processes after their burial. The presence of
trace fossils assigned to Corrosichnia suggests a temporary shallow
burial in a substrate supporting vegetation (Montalvo, 2004), in
agreement with oodplain sub-environment.

Fig. 9. Articulated specimens from FL 1. A. Paedotherium sp., posterior limb. B.

Mammalia indet., vertebrae and pelvic girdle. Scale bar = 3 cm.

Fig. 10. Rose diagrams showing orientations of elongated skeletal elements from both
fossiliferous levels.

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291

5.2. Fossiliferous level 2 (FL 2)

Fig. 11. Stereographic diagrams showing dip/trend distribution of remains from both
fossiliferous levels.

The joint occurrence of specimens with diverse articulation states

might point to the development of an assemblage formed from the
gradual incorporation of bone elements to the substrate. Exposure
time and the rate of burial depend on the frequency and intensity of
ooding events and the distance between the channel and the area
where the remains were deposited. The prevalence of isolated and
dispersed remains might indicate that, in most cases, the exposure
time was long enough for different processes (e.g. transport by
water ows, trampling, activity of predators and/or scavengers) to
separate and disperse skeletal elements. However, the occurrence of
several articulated specimens and other disarticulated but associated
ones might indicate rapid burial events and the presence of connective tissue at the time of burial (Hill, 1979; Hill and Behrensmeyer,
1984). This kind of specimen might represent evidence for accumulation (Fernndez-Lpez, 2000), without important later transport.
The wide spatial distribution of remains at this level might be related
to the absence of physical or biological processes favoring the concentration of remains. The lack of preferential orientations indicates shallow and low-energy water currents, with a weak inuence on the
disposition of remains (Alcal et al., 1999; Fernndez-Lpez, 2000).
The few dip oriented remains recorded, all of them with very low
angle, reveal that the original horizontal position was not changed. Considering that most inclined materials were in spatial proximity to burrows produced by the rodent A. priscus, it could be interpreted that
the burrowing activity might have affected already-buried remains.
While inclination produced by trampling is common in remains recovered in oodplain deposits (Behrensmeyer et al., 1989; Fiorillo, 1989),
no diagnostic characters (e.g. lines and grooves) indicative of this process were recorded at FL 1.

For the assemblage recovered in the FL 2, NISP values reveal a low

degree of anatomical and taxonomic resolution. Most of the specimens recovered correspond to micromammals and Osteichthyes. On
the other hand, the low value of MNI obtained is in agreement with
Badgley (1986a,b), who noted that the remains present in the channel deposits have, generally, very diverse temporal and spatial origins
and are subjected to continually transport and reworking. Thus, it is
very unlikely that the same individual remains end up in the same
fossil concentration.
Like in the FL 1, the different groups for mammalian skeletal elements proposed by Voorhies (1969), Dodson (1973) and Korth
(1979) were represented in this level. Though the transport resistant elements such as isolated teeth and jaws dominated, other elements easily transported by water currents, such as phalanges and metapodials,
were also found. The predominance of skeletal elements included
in groups II and III indicate a selection towards remains with low susceptibility to be mobilized (Behrensmeyer, 1975). The isolated teeth/
vertebrae index was 3.75. This value is higher than 1, indicating an assemblage with a high degree of hydrodynamic sorting. The latter
might indicate that the currents developed in the uvial channel had
a marked inuence over the remains, affecting them differentially
according to their characteristics. The marked dominance of small,
cubic materials supports the interpretation that sorting processes
were of great importance at this level.
No evidence for sub-aerial exposure was found in channel deposits;
it is thus inferred that those remains showing signs of weathering were
incorporated from other areas. One explanation for that might imply
transport of remains before burial, from the oodplain towards the
channel's interior, and therefore these remains might represent evidences of resedimentation (sensu Fernndez-Lpez, 1991, 2000). Another option might be the exhumation of fossil remains from older
oodplain levels due to the lateral migration of the channel and the
erosion of the margins, representing, in that case, evidence for
reelaboration (sensu Fernndez-Lpez, 1991, 2000).
The high percent of elements with abrasion evidence, extreme in
several micromammal remains, is consistent with the high energy assumed for channel deposits. It is estimated that once the materials
were incorporated to the channel, many of them were exposed for
prolonged periods of time to the abrasive action of hydraulic currents
and the transported sedimentary particles (Behrensmeyer, 1991)
and, therefore were affected with more intensity.
The abundance of incomplete remains, both of micro- and
macromammals, in several cases with an extreme degree of breakage,
might be related to a prolonged time of exposure to diverse destructive
taphonomic processes; this is consistent with the inferred energy to
such sub-environment. With regard to that, the few complete remains
were represented mainly by skeletal elements with high structural densities (Lyman, 1994). The marked dominance of transverse fractures
suggests that most remains were affected after the burial stage (Alcal
and Martn Escorza, 1988, 1998). Like in the FL 1, one possibility is the
lithostatic load generated by overlying sediments. However, considering that many of these remains had signs of abrasion on the fracture surface, another possibility involves exhumation from older oodplain
levels and subsequent incorporation into the cannel. Some longitudinal
and oblique fractures were also identied, which might have been
originated by diverse biostratinomic processes (e.g. weathering and
trampling), probably while the remains were exposed in the oodplain,
previously to their transport into the channel.
The remains that exhibited bioerosion trace fossils linked to soil
processes might have been incorporated to the channel deposits
through exhumation from oodplain deposits and therefore would
represent another evidence of reelaboration.
The absence of articulated and/or associated specimens might
be related to the high energy of hydraulic ows developed in the

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channel, and its capacity to disarticulate, transport and disperse the

remains.
The high density of remains suggests the identication of FL 2 as
bonebeds (Behrensmeyer, 1991; Rogers et al., 2007). This concentration, derived from channel deposits, might have been originated by
diverse processes, among them: unevenness or depressions in the substrate, obstruction by other materials acting as barriers, and/or rapid
ow deceleration causing a lower hydraulic competency and the deposition of materials. However, no concrete evidence linking these high
concentrations to a particular process was recognized.
The occurrence of preferential orientation, with most of the data
falling into the 180270 quadrant, is partially coincidental with
paleocurrent data recorded by Zavala and Navarro (1993) from crossbedding analysis. These authors determined a main WSW paleorunoff
and approximately 120 of dispersion. In line with the paleogeography
of the region, the main drain would be from the Positivo Bonaerense
to the Colorado basin (Yrigoyen, 1975; Zavala and Navarro, 1993). The
inclination of the remains, which reached 90 in many cases, might be
linked to the transport generated by water currents and events of
rapid deceleration/deposition taking place within the uvial channel
(Voorhies, 1969; Alcal et al., 1999, 2002).
5.3. General interpretations
The assemblages found at both fossiliferous levels, FL 1 and FL 2,
are mainly made up of resedimented remains. Moreover, some accumulated materials were recovered from the FL 1, while in the FL 2
several remains that had been reelaborated from more ancient oodplain deposits, were identied. Most of the elements in the FL 1 are
autochthonous or paraautochthonous, and only a few (remains from
Osteichthyes) might be considered allochthonous. On the other
hand, paraautochthonous and allochthonous elements were recognized for the FL 2.
Both assemblages can be dened as mixed (sensu Fernndez-Lpez,
2000), given that they are made up of elements corresponding to biological entities coming from different environments. Terrestrial and
aquatic taxa occur together in both levels. Moreover, the assemblages
can be considered as condensed (sensu Fernndez-Lpez, 2000),
given that they originated from the gradual incorporation of remains
corresponding to temporally-successive biological entities. However,
the taxonomic evidence registered does not reveal important faunal
changes that indicate signicant timing differences between the two
fossiliferous levels.
Behrensmeyer (1988) dened taphonomic mode as a recurring
pattern of preservation of organic remains in a particular sedimentary

context, accompanied by characteristic taphonomic features. Later,

Behrensmeyer and Hook (1992) indicated that taphonomic mode is
a set of fossil occurrences that result from similar physical, chemical,
and biological processes. The differences found between the fossiliferous levels (FL 1 and FL 2) recovered from the Monte Hermoso Formation allowed the identication of two distinct taphonomic modes,
oodplain and channel-lag taphonomic modes (Table 2); both related
to the different processes that took place before and after burial in the
sub-environments in which they were recognized. These taphonomic
modes are partially similar to those described by Behrensmeyer
(1988) and Behrensmeyer and Hook (1992).
6. Conclusions
The assemblages recovered from two fossiliferous levels, FL 1and
FL 2, of the Monte Hermoso Formation, are composed of remains
that were affected by diverse biostratinomic and fossil-diagenetic
processes, related to the oodplain and channel deposits, and therefore present distinct taphonomic histories. The variations recorded
between this fossiliferous levels, allowed us to recognize two taphonomic modes.
The oodplain taphonomic mode was recognized in the FL 1. The
taphonomic characteristics of this mode partially correspond to
those of model I (rapidly accumulated and buried elements), and
model II (resedimented elements, with considerable exposure before
burial), proposed by Johnson (1960) for putative formation mechanisms of fossil assemblages. The channel-lag taphonomic mode was
recognized in the FL 2. This taphonomic mode partially corresponds
to model III (resedimented and reelaborated elements, with considerable lateral transport) proposed by Johnson (1960).
Other fossiliferous levels of the Monte Hermoso Formation present particular sedimentary and taphonomic characteristics. Even
when those levels probably comprise different taphonomic modes,
we opted for not dening them in this study because more information is needed with regard to their faunistic content, sedimentary
context and the taphonomic history of their remains.
The taphonomic modes described here represent different conditions within the range of probable taphonomic histories for assemblages generated from the attritional accumulation of remains in a
uvial environment. Moreover, the taxonomic evidence can assign
both fossiliferous levels to the early Pliocene.
Considering that the taphonomic analyses related with uvial deposits are very scarce for the South American Neogene, the data obtained
in this comparative study and the distinction between two taphonomic
modes provide new information related to the paleoenvironmental

Table 2
Characteristics of the two taphonomic modes recognized in the Monte Hermoso Formation.
Monte Hermoso Formation
Floodplain taphonomic mode

Channel-lag taphonomic mode

Sedimentological context
Lithology
Anatomic and taxonomic representation

Floodplain deposits
Muddy siltstones, sandy siltstones and silty sandstones
Clear predominance of identiable skeletal elements.
Predominance of micromammal remains

Index isolated teeth/vertebrae

Geometric shape
Weathering degree
Abrasion degree
Degree and type of breakage

1.33
Great diversity of shapes, but cubic forms were predominated
High proportion of remains with different weathering degree
High proportion of remains without evidences of abrasion
High proportion of whole skeletal elements.
Longitudinal fractures predominate
High proportion of remains with evidences of bioerosion
High proportion of isolated and dispersed skeletal elements
Could not be assessed
No preferential orientations were recorded
High proportion of remains with horizontal position
or very low dip (b19)

Channel deposits
Clast-supported breccias
Similar proportions of identiable skeletal elements and
unidentiable fragments. Predominance of micromammal
and bony sh remains
3.75
Clear predominance of cubic forms
High proportion of remains without evidences of weathering
High proportion of remains with different abrasion degree
High proportion of incomplete skeletal elements. Transverse
fractures predominate
Low proportion of remains with evidences of bioerosion
All specimens isolated and dispersed
Density variable between 38.93 and 52.86 remains/m2
Some remains oriented by paleocurrents
Great diversity of dip values, between 0 and 90

Bioerosion degree
Skeletal articulation
Spatial density
Orientation
Dip

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characteristics of this continental ecosystem. On the other hand, these

new data provide a reference for the comparison and evaluation of
other local and regional deposits presenting similar problems and faunistic associations, particularly those related to rivers typical of plain
environments.
Acknowledgments
We thank R. Caputo, director of Museo Carlos Darwin for allowing
us access to the materials studied. T. Manera and C. Oliva helped during
eldwork. J. Pan and E. Braun helped with the English version. Valuable
suggestions and comments from C. Deschamps, R. Melchor, A.M.
Umazano, G. Visconti and two anonymous reviewers on an early version, and detailed editorial work, helped to improve this manuscript.
This paper was partially supported by FCEyN UNLPam 209 grant.
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