Journal of South American Earth Sciences 76 (2017) 257e263

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Journal of South American Earth Sciences

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Origin of bonebeds in Quaternary tank deposits

Hermnio Ismael de Arajo-Jnior a, *, Kleberson de Oliveira Porpino b,
Llian Paglarelli Bergqvist c
a
Departamento de Estratigraa e Paleontologia, Universidade do Estado do Rio de Janeiro, 20550-013, Rio de Janeiro, RJ, Brazil
b
Departamento de Ci^ gicas, Universidade do Estado do Rio Grande do Norte, 59610-090, Mossoro
encias Biolo , RN, Brazil
c
Departamento de Geologia, Instituto de Geoci^
encias, Universidade Federal do Rio de Janeiro, 21941-916, Rio de Janeiro, RJ, Brazil

a r t i c l e i n f o a b s t r a c t

Article history: Tank deposits are an exceptional type of fossiliferous deposit and bear a remarkably fossil record of the
Received 22 December 2016 Pleistocene megafauna of South America, particularly of Brazil. The taphonomy of vertebrate remains
Received in revised form preserved in this type of environmental context was clearly driven by climate, similarly to most of the
24 March 2017
Quaternary continental fossil record. The formation of the vertebrates fossil record in tank deposits was
Accepted 24 March 2017
Available online 27 March 2017
inuenced by the climate seasonality typical of arid climate. The taphonomic history of most tank de-
posits is a consequence of this seasonality and, as a result, the paleoecological data preserved in their
fossil assemblages is reliable with respect to paleobiological and paleoenvironmental settings of the
Keywords:
Bonebeds
Quaternary ecosystems of the Brazilian Intertropical Region (BIR). Other tank deposits experienced an
Tank deposits unusual taphonomic history that, besides climate, was affected by recurrent events of reworking pro-
Quaternary duced by the depositional agents dominant in the surrounding alluvial plains. The conclusions obtained
Megafauna here concerning the main taphonomic settings and formative processes that characterize fossil verte-
Taphonomy brate assemblages of tank deposits will help further studies aimed to recover information on the
Paleoecology paleoecology of Quaternary fauna collected in such deposits by allowing a better understanding of their
time and spatial resolutions and other potential biases.
2017 Elsevier Ltd. All rights reserved.

1. Introduction et al., 2013a, 2015).

Tank deposits e one of the most singular type of fossiliferous
deposit in South America e are small sedimentary bodies of Qua-
ternary age that ll natural depressions (called natural tanks) on
basement rocks in northeastern Brazil (Arajo-Jnior et al., 2013a).
These sedimentary bodies are usually stratied and fossiliferous,
preserving remains of diverse representatives of the Quaternary
megafauna, such as huge ground sloths, glyptodonts, mastodons,
toxodonts, saber-toothed cats and macraucheniids (Paula-Couto,
1980; Mabesoone et al., 1990; Bergqvist et al., 1997; Cartelle,
1999). In some cases, megafaunal remains are associated with
few fossils of small- to mid-sized mammals, crocodiles, lizards,
birds and anurans (Arajo-Jnior and Moura, 2014). The genesis of
these deposits has been interpreted as deposited in contexts of
debris ows and ash oods (Mabesoone et al., 1990; Arajo-Jnior
* Corresponding author.
E-mail addresses: herminio.ismael@yahoo.com.br (H.I. Arajo-Jnior),
kleporpino@yahoo.com.br (K.O. Porpino), bergqvist@geologia.ufrj.br
(L.P. Bergqvist).
Tank deposits provide some of the best evidences for under-
standing paleoenvironments and paleoecology of the Brazilian
Intertropical Region (BIR) during the Quaternary. However, these
deposits have preserved different paleoecological snapshots from
ancient vertebrate communities and their associated ecosystems
(Arajo-Jnior et al., 2011a), and these differences only began to be
scrutinized in the last decade by taphonomy-based studies (Santos
et al., 2002; Alves et al., 2007; Silva, 2008; Arajo-Jnior et al.,
2013a, 2013b, 2015).
Currently, tank deposits are the most taphonomically studied
type of vertebrate-bearing deposit in Brazil, but some studies have
revealed a wide range of variation in the preservational patterns in
their fossil concentrations (Arajo-Jnior et al., 2013a, 2013b, 2015;
Arajo-Jnior and Moura, 2014). Therefore, a comparative tapho-
nomic analysis among tank deposits would be crucial to better
delineate the main taphonomic processes and pathways that led to
these perceived differences in the preservation of vertebrate re-
mains in such deposits.
Here we present the results of a regional-scale taphonomic
analysis seeking to describe and explain the variation in

http://dx.doi.org/10.1016/j.jsames.2017.03.012
0895-9811/ 2017 Elsevier Ltd. All rights reserved.
258 H.I. Arajo-Jnior et al. / Journal of South American Earth Sciences 76 (2017) 257e263

taphonomic and paleoecological signatures among tank deposits, NISP 118; LG NISP 230; JC NISP 556; JI NISP 1405). They are
shedding light on the origin of the bonebeds preserved in this housed at Museu Nacional, Rio de Janeiro City, Brazil, and Museu de
environmental context and its implications for the paleoecology of -histo
Pre  ria de Itapipoca, Itapipoca City, Brazil. Their faunal lists are
Quaternary of BIR. presented in Appendix A. All fossil specimens (NISP 2640)
recovered from those tank deposits were evaluated in order to
recognize and interpret taphonomic features, providing a consis-
2. Material and methods
tent background for comparative analyses and to infer the paleo-
environmental conditions related to their origin.
Fossil assemblages of tank deposits from ve paleontological
Fossil assemblages were evaluated following classical vertebrate
sites located at different sections of BIR (Fig. 1) were analyzed
taphonomy methods (Behrensmeyer, 1978, 1991; Shipman, 1981;
seeking for common patterns and distinctive features: (i) Jirau
Frison and Todd, 1986; Lyman, 1994; Rogers et al., 2007). Tapho-
Paleontological Site (JI; 3
2102300 S 39
420 2000 W); (ii) Joa ~o Cativo
nomic attributes evaluated were: (i) physical integrity (complete,
Paleontological Site (JC; 3
300 1700 S 39
400 2400 W); (iii) Campo Alegre
partial or fragment); (ii) degree of disarticulation (articulated or
Paleontological Site (CA; 7
15018.5500 S 36
440 26.3500 W); (iv) Curi-
disarticulated); (iii) desiccation marks (weathering stages of
mata ~s Paleontological Site (CM; 7
070 3600 S 36
0704800 W); and (v)
Behrensmeyer, 1978; from 0 to 5); (iv) degree of abrasion (ac-
Lage Grande Paleontological Site (LG; 8
250 2700 S 36
430 2000 W).
cording to Shipman, 1981; no abrasion, moderate abrasion, heavy
Although a large amount of tank deposits (approximately 36; see
abrasion); (v) tooth marks (presence/absence); (vi) trample marks
Fig. 1 of Arajo-Jnior, 2016) have been excavated in Brazil during
(presence/absence); (vii) anthropogenic signs (presence/absence);
the last 80 years, a few ones experienced a controlled excavation,
and (viii) bioclastic sorting according to the three groups of Fluvial
including those analyzed in this study.
Transport Index (FTI groups of Frison and Todd, 1986).
The material analyzed in this work consists of all identiable
Furthermore, correspondence and cluster analyses (Q- and R-
specimens recovered in previous excavations (CA NISP 331; CM

Fig. 1. Location map of the paleontological sites of Brazilian Intertropical Region evaluated in this study.
 H.I. Arajo-Jnior et al. / Journal of South American Earth Sciences 76 (2017) 257e263 259

modes) of the ve tank deposits were carried out in order to
identify both typical and atypical categories of taphonomic features
and taphonomic contexts of the deposits. We produced a data
matrix for the categories of taphonomic attributes listed above,
which were scored using percentage values for each paleontolog-
ical site (Appendix B). The resulting data matrix was submitted to
cluster analyses (Q-mode and R-mode) using the weighted pair
group method with simple arithmetic averages (UPGMA) and to a
correspondence analysis. In all analyses, we used the Euclidean
distance.
In the Q-mode cluster analysis, the paleontological sites were
clustered based on the similarity of their categories of taphonomic
signatures and this clustering was used to identify sites with similar
taphonomic contexts in the BIR. In the R-mode analysis, the cate-
gories of taphonomic signatures were clustered based on the sim-
ilarity of the percentages with which they occur among the
paleontological sites. In both R- and Q-mode analyses, an ANOSIM
permutation test (one-way) was employed to evaluate the signi-
cance of the differences between the groups obtained. This test
provides an R statistics that varies from 1 to 1, where R values
equal to 1 are obtained only when all samples within the groups
are more similar to each other than to any sample of other groups
(Clarke and Warwick, 2001). The signicance value (p) of the
clusterings was also obtained from this test. All multivariate ana-
lyses were performed using the software Paleontological Statistics
version 2.17 (PAST; Hammer et al., 2001).
The correspondence analysis species the Chi-squared distances
(x2) between the paleontological sites of BIR, using the Chi-squared
distances (x2) between the main taphonomic categories in the
assemblages as their background. According to Hammer (2013, pg.
97), [] If your data are well behaved, taxa [in our case, paleon-
tological sites] typical for an association should plot in the vicinity of
that association []. In this study, the paleontological site(s)
plotted out of the vicinity of the main clustering of sites, were
considered to represent taphonomic outliers, indicating a possible
exotic (time- or spatial-averaged) assemblage in comparison with
the remaining taphocoenoses, which represent the typical assem-
blages (see Hammer et al., 2001 and Hammer and Harper, 2006 for
additional information on the application of correspondence
analysis to paleontological data).
3. Comparative taphonomy
Our taphonomic analysis reveals a wide array of taphonomic
processes that have interacted during the genesis of tank assem-
blages, such as disarticulation, weathering, transport, abrasion,
sorting and biogenic phenomena (Fig. 2A); the percentages of
taphonomic signatures for each analyzed tank are presented in
Appendix B. The correspondence analysis allowed the identica-
tion of three main taphonomic settings for tank deposits (tapho-
nomic settings A, B and C; Fig. 3).
The most common taphonomic signatures in tank deposits are
disarticulated and fragmented specimens, specimens with stage 1
of weathering and with moderate abrasion, representing the
taphonomic setting A and Group A (Fig. 3B and A, respectively).
Disarticulation and fragmentation occur in a context of subaerial
exposure prior to burial (Hill and Behrensmeyer, 1984;
Behrensmeyer, 1991) and, therefore, suggest that bones were

Fig. 2. Comparative taphonomic analysis of tank deposits; A. Taphogram showing of percentages of specimens according to taphonomic categories for each tank deposit; B. MN
3638-V, proximal end of right femur of Leopardus sp. (Mammalia, Carnivora, Felidae) from Curimat~as Paleontological Site e note the high stage of fragmentation; C. MN 2819-V,
right hemimandible of Ozotocerus bezoarticus (Mammalia, Artiodactyla, Cervidae) from Joa~o Cativo Paleontological Site e note the aking of the bone surface which is placed in the
weathering stage 1; D. MN 3612-V, osteoderm of dorsal carapace of Panochthus sp. (Mammalia, Cingulata, Glyptodontidae) from Curimata ~s Paleontological Site e note the moderate
abrasion; E. MN 2853-V, diaphysis of right femur of Palaeolama major (Mammalia, Artiodactyla, Camelidae) from Joa ~o Cativo Paleontological Site e in close-up, note the parallel and
v-shaped grooves on the surface; CA e Campo Alegre, CM e Curimata ~s, LG e Lage Grande, JI e Jirau, JC e Jo~ao Cativo; scale bar 2 cm.
260 H.I. Arajo-Jnior et al. / Journal of South American Earth Sciences 76 (2017) 257e263

Fig. 3. Multivariate analyses using percentages of specimens according to taphonomic categories for each tank deposit; A. UPGMA cluster analysis with measures of Euclidean
distances in Q-mode; B. Correspondence analysis with measures of Euclidean distances; C. UPGMA cluster analysis with measures of Euclidean distances in R-mode.

deposited inside natural tanks after a period of carcasses decay. This
is supported by the occurrence of weathered specimens with slight
modications, as any weathering occurs only in circumstances of
subaerial exposure (Behrensmeyer, 1978).
Moderate abrasion can be generated by hydraulic transport (i)
during a moderate to long distance or (ii) during a moderate time
interval but over a short distance (around and/or inside the tanks).
Analysis of bioclast sorting e here employing analysis of Fluvial
Transport Index (Frison and Todd, 1986) e is the most appropriate
mode of evaluating the transportation distance for Quaternary
large-sized mammals (see discussion in Arajo-Jnior et al., 2012),
because sorting occurs only in context of moderate- to long-dis-
tance transport (Aslan and Behrensmeyer, 1996). Considering tank
assemblages, the rst hypothesis is plausible for those with large
amounts of bones placed into FTI >75 Group (Campo Alegre,
Curimata ~s and Lage Grande), that probably experienced some
uvial inuence. Inversely, the second hypothesis explains the
fossil assemblage where the three FTI groups are represented (Joa ~o
Cativo); probably, the thanatocoenoses from which this deposit
derived were transported by a non-sorting agent, such as gravita-
tional ows or ash oods. Thus, two modes of hydraulic transport
may have occurred during the genesis of tank-deposit bonebeds.
Only Jirau assemblage is uneven regarding the pattern of FTI groups
if compared to the other fossil assemblages. The rarity of FTI <75
elements can be related to a spreading of the most transportable
elements away from the tank, leaving only the lesser transportable
ones (FTI 50e75 and FTI < 50 elements) inside it, regardless of
the transport agent. Considering that this particular tank deposit
was likely formed under a debris-ows regime (Arajo-Jnior et al.,
2013a), we suggest that the debris ows that transported the
 H.I. Arajo-Jnior et al. / Journal of South American Earth Sciences 76 (2017) 257e263
specimens to the tank had such a high energy that the slighter el-
ements continued to be transported and scattered after passing by
the tank depression (i.e. they were not deposited inside the tank
due to the high energy of the transport agent).
Taphonomic settings B and C include the rarest taphonomic
features of tank deposits: biogenic traces (trample, teeth and
anthropogenic marks); specimens in weathering stages 3e5, and
non-abraded and heavy-abraded specimens (Fig. 3B). These cate-
gories are distributed into two subgroups (B1 and B2) within Group
B (Fig. 3A). Teeth marks occur in few specimens from two assem-
blages and were likely caused by a midsized canid scavenger
(Arajo-Jnior et al., 2011b). In the correspondence and cluster
analyses (Fig. 3), both teeth and trample marks were plotted near
the highest weathering stages, suggesting that scavenging and
trampling occurred only in circumstances of prolonged subaerial
exposure. Therefore, it appears that, in tank bonebeds, the condi-
tions allowing the origin of such marks took place only at low rates
of burial, probably during periods when rainfalls were very scarce.
Anthropogenic signs are rare in vertebrate fossils from the
Quaternary of Brazil (Vilhena-Vialou and Vialou, 2008; Dantas et al.,
2012; Be lo and Oliveira, 2013; Mothe  et al., 2014). In this study,
anthropogenic marks were observed only in a single specimen of
the camelid Palaeolama major (Fig. 2E). Like other biogenic signs,
anthropogenic modications of bones (by either scavenging activ-
ity by hunters-gatherers or hunting and processing the prey) in the
thanatocoenoses is only possible in conditions of delayed burial. A
detailed description and interpretation of this anthropogenic
alteration is in progress elsewhere by some authors of this paper.
Based on the clustering of CA, CM and LG into the group Sites A
(Fig. 3C) and their plotting into the taphonomic setting A (Fig. 3B) e
which comprises the commonest taphonomic features of tank de-
posits e we consider these assemblages as typical for tank deposits
and their taphonomic signatures are the most diagnostic. On the
other hand, JC and JI (Sites B in Fig. 3C) represent particular and
unusual tank assemblages, produced when environments around
the tanks probably underwent variations in their rates of sedi-
mentation (decreasing), reworking (increasing) or biogenic activity
(increasing) (see Arajo-Jnior et al., 2013a for a full interpretation
of the taphonomic history of the JI assemblage).
Finally, despite the differences related to the physical integrity
and the presence/absence of biogenic traces in the specimens, the
array of taphonomic processes seems to have been very similar
among the tank deposits evaluated (Fig. 2A), varying only in their
intensity. Therefore, despite possible minor differences in the age of
the deposits, the taphonomic processes identied were recurrent
throughout the Pleistocene-Holocene boundary in BIR. However,
the cyclicity of such processes probably varied, as suggested by
evidences pointing to variations in rates of subaerial exposure of
the bioclasts in some tanks (see weathering stages and abrasion
analyses in Fig. 2A).
4. Taphonomic-paleoenvironmental scenarios for the origin
of tanks bonebeds
We hypothesize the following taphonomic-paleoenvironmental
scenario to explain the features of the typical fossil assemblages of
tank deposits, represented by Sites A: megamammals, small-
sized mammals, reptiles and avians died in dry periods around
the tanks, which represented the last water reservoirs during these
periods; in parallel, anurans died within the tank depressions
(Arajo-Jnior and Moura, 2014). After death, the carcasses un-
derwent necrolysis followed by disarticulation. Later, bones un-
derwent slight weathering (weathering stage 1eBehrensmeyer,
1978) in the dry periods, for a few years, resulting in a few super-
cial longitudinal cracks on the bone surface. At the same time,
261
bones experienced constant mobilization (e.g. by sporadic rain-
waters), reaching the stage of moderate abrasion (see Shipman,
1981). During rainfall seasons, the alluvial plains experienced
gravitational ows and oods, which deposited peripheral ele-
ments (sensu Arajo-Jnior, 2016) e the disarticulated, weathered
and abraded bones e inside the tanks, mixing them: (i) with newly
inputted remains into the thanatocoenoses and (ii) with in situ
preserved remains (sensu Arajo-Jnior, 2016), like anurans, pro-
ducing a mild spatial-mixing.
The fact that specimens with weathering stage 1 are over-
represented in all tank assemblages suggests that the time span of
subaerial exposure for most elements was around a few years
(according to Behrensmeyer and Miller, 2012), indicating seasonal
burial of bones inside the tanks. As in present days, this deposi-
tional seasonality can have occurred due to an alternation between
dry and rainfall periods in an arid or semiarid climate, in which
oods and gravitational ows are typical of rainfall events (Nichols,
2009). In fact, an arid/semiarid climate in a wide area of BIR is
supported by isotopic studies of mammalian bones and teeth pre-
served in other Quaternary deposits in northeastern Brazil (Dantas
et al., 2013) and in speleothems of caves in Rio Grande do Norte and
Bahia states (Wang et al., 2004; Cruz et al., 2009).
A taphonomic scenario characterized by low rates of sedimen-
tation and high rates of reworking was suggested for both JI
(Arajo-Jnior et al. (2013a)) and JC (Arajo-Jnior et al., 2014),
based on fossildiagenetic analyses, and on evaluation of types of
breakage. This scenario would explain the unusual features asso-
ciated to Sites B: initially, vertebrate bones were subaerially
exposed for a prolonged time span around the tanks prior to burial,
reaching the stages of weathering 3 to 5. Meanwhile, the bones
were disarticulated and modied by other physical and biogenic
factors, such as trampling, scavenging and, eventually, human ac-
tivity. These conditions were disrupted during eventual rainfall
increase periods, which established conditions of high energy that
transported new carcasses and reworked skeletal elements previ-
ously buried outside the tanks, to inside the tanks. It is also possible
that, during these episodes, some of the specimens already in the
taphocoenosis had also been reworked. Such events mixed
different taphonomic settings into a single fossil assemblage,
resulting in time-averaged and internally complex assemblages.
This time-averaging is evidenced by the co-occurrence e in the
assemblages of JC e JI sites e of several taphonomic signatures in
markedly distinct states, such as: expressive variation in stages of
fragmentation; very-weathered skeletal specimens together with
non-weathered ones; and non-abraded, moderate-abraded and
heavy-abraded specimens.
Considering that most taphonomic features observed in the tank
deposits analyzed here occur in dryer conditions, the taphonomic-
paleoenvironmental scenarios proposed herein may be associated
with the dry phases e between ~20 ky and ~12 ky e identied in
the northeastern Brazil by Cruz et al. (2009) through isotopic an-
alyses of speleothems. Indeed, most of the dated tank assemblages
include fossils from this time interval (Dantas et al., 2013, 2014;
Ribeiro et al., 2014). In this time span, the periods with relatively
more sedimentary supply led to a relatively short subaerial expo-
sure of the thanatocoenoses, as inferred for bonebeds of Sites A.
Conversely, the cases in which the thanatocoenoses underwent a
much longer subaerial exposure (bonebeds of Sites B) may be
related to periods with relatively less sedimentary supply.
5. The quality of the paleoecological data in tank assemblages
The different scenarios proposed for the studied assemblages
imply differences in the quality of the fossil record in tank deposits.
Therefore, the paleoecological data preserved in tank bonebeds can
262 H.I. Arajo-Jnior et al. / Journal of South American Earth Sciences 76 (2017) 257e263
vary considerably.
Tank deposits formed only in the taphonomic setting A e such
as Sites A (Fig. 3C) e were formed in a context of low rates of
time-averaging and spatial-mixing. Thus, these tank deposits pre-
sent high time and spatial resolutions and, therefore, preserve
more reliable paleoecological data. For these fossil assemblages, the
paleobiological and paleoecological information recovered, such as
taxonomic composition and relative abundance, can be considered
reliable relative to the past biocoenosis from which the fossil as-
semblages derived.
On the other hand, tank deposits resulting from the mixing of
the three taphonomic settings e such as Sites B (Fig. 3C) e
incorporate taphonomic biases introduced during the recurrent
events of reworking and temporal-mixing between vertebrate re-
mains inside the tanks. Therefore, these tank deposits probably
provide more biased paleoecological information from the ancient
ecosystems. Thus, paleontologists must be careful when inter-
preting past conditions from remains preserved in such type of
fossil assemblage.
Acknowledgments
The authors are grateful to Museu Nacional/UFRJ (especially to
D.D.R. Henriques) and Museu de Pre -histo
ria de Itapipoca (espe-
cially to C.L. Ximenes and A.S.T. Santos) for allowing access to the
analyzed material; to C.L. Mello, D.D.R. Henriques, I.S. Carvalho,
L.C.M.O. Ponciano and P.F.F. Dal'Bo for useful comments and sug-
gestions that helped to improve an early version of the manuscript
and to reviewers for their thoughtful suggestions and criticisms;
HIAJr thanks to Conselho Nacional de Desenvolvimento Cientco e
Tecnolo gico (CNPq; process # 140497/2012-9) and Funda~ ao Carlos
Chagas Filho de Amparo  a Pesquisa do Estado do Rio de Janeiro
(FAPERJ; process # E-26/100.221/2014) for the nancial support to
this work.
Appendix A. Supplementary data
Supplementary data related to this article can be found at http://
dx.doi.org/10.1016/j.jsames.2017.03.012.
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