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A new Early Cretaceous brachiosaurid (Dinosauria,
Neosauropoda) from northwestern Gondwana (Villa de
Leiva, Colombia)
José L. Carballidoa, Diego Pola, Mary L. Parra Rugeb, Santiago Padilla Bernalb, María E.
Páramo-Fonsecac & Fernando Etayo-Sernad
a Consejo Nacional de Investigaciones Científicas y Técnicas-Museo Paleontológico Egidio
Feruglio, Fontana 140, 29100 Trelew, Argentina,
b Centro Investigaciones Paleontológicas, Kilometer 5.2 en vía a Santa Sofía, Villa de Leyva,
Colombia,
c Departamento de Geociencias, Universidad Nacional de Colombia, Carrera 30 No. 45-03,
Bogotá D.C., Colombia,
d Servicio Geológico Nacional–Museo Geológico José Royo y Gómez, Dg 5334-53, Bogotá DC,
Colombia,
Published online: 27 Aug 2015.
To cite this article: José L. Carballido, Diego Pol, Mary L. Parra Ruge, Santiago Padilla Bernal, María E. Páramo-Fonseca &
Fernando Etayo-Serna (2015): A new Early Cretaceous brachiosaurid (Dinosauria, Neosauropoda) from northwestern Gondwana
(Villa de Leiva, Colombia), Journal of Vertebrate Paleontology
To link to this article: http://dx.doi.org/10.1080/02724634.2015.980505
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ARTICLE
A NEW EARLY CRETACEOUS BRACHIOSAURID (DINOSAURIA, NEOSAUROPODA) FROM
NORTHWESTERN GONDWANA (VILLA DE LEIVA, COLOMBIA)
JOS
E L. CARBALLIDO,*
,1
DIEGO POL,
1
MARY L. PARRA RUGE,
2
SANTIAGO PADILLA BERNAL,
2
MAR
IA E. P
ARAMO-FONSECA,
3
and FERNANDO ETAYO-SERNA
4
1
Consejo Nacional de Investigaciones Cient
ıficas y T
ecnicas-Museo Paleontol
ogico Egidio Feruglio, Fontana 140, 29100 Trelew,
Argentina, jcarballido@mef.org.ar; dpol@mef.org.ar;
2
Centro Investigaciones Paleontol
ogicas, Kilometer 5.2 en v
ıa a Santa Sof
ıa, Villa de Leyva, Colombia, mlparra@centropaleo.com;
spadilla@rochembiocare.com;
3
Departamento de Geociencias, Universidad Nacional de Colombia, Carrera 30 No. 45-03, Bogot
a D.C., Colombia,
meparamof@unal.edu.co;
4
Servicio Geol
ogico Nacional–Museo Geol
ogico Jos
e Royo y G
omez, Dg 5334-53, Bogot
a DC, Colombia, fetayos@gmail.com
ABSTRACT—Brachiosaurid sauropods achieved a broad distribution during the Late Jurassic, which has been considered to
provide evidence of their origins during the Middle Jurassic, prior to the breakup of Pangea. In contrast to their broad
geographic distribution during the Late Jurassic, formally named brachiosaurid species from the Cretaceous have so far been
restricted to the Aptian–Albian of North America, which has been interpreted as a signal of differential extinction and/or a
bias in the Early Cretaceous fossil record. Here we describe a new brachiosaurid titanosauriform taxon from the Early
Cretaceous of Colombia, which is represented by axial elements. The material was recovered from marine sediments of the
Paja Formation (Barremian), close to the locality of Villa de Leiva. The weakly laterally expanded and divided transverse
processes of the anterior-most caudal vertebrae allows the recognition of a new sauropod taxon, Padillasaurus leivaensis, gen.
et sp. nov. In order to test the phylogenetic relationships of the new taxon, we performed a cladistic analysis that recovered
Padillasaurus as a brachiosaurid titanosauriform. This position is supported by a combination of characters, including the
presence of blind fossae in anterior caudal vertebrae. Among titanosauriforms, the presence of blind fossae in anterior caudal
vertebrae is an apomorphic character that is exclusive to Giraffatitan, Venenosaurus, Cedarosaurus, and Abydosaurus.
Although more complete remains are needed to test more thoroughly the affinities of the new taxon, the available evidence
indicates that brachiosaurids survived at lower latitudes in Gondwana until at least the Early Cretaceous.
http://zoobank.org/urn:lsid:zoobank.org:pub:652D6B2A-7A8A-4311-8725-279BF2C9E0E3
SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP
INTRODUCTION
Brachiosauridae is one of the two major clades of Titanosauri-
formes, a diverse group of sauropods that thrived in the Late
Jurassic and Cretaceous in both Laurasia and Gondwana. Bra-
chiosaurids encompass all titanosauriforms more closely related
to Brachiosaurus than to Saltasaurus (Wilson and Sereno, 1998).
Definitive brachiosaurid remains are known from the Late Juras-
sic up to the late Early Cretaceous. During the Late Jurassic, this
clade is recorded in both the Northern and Southern Hemi-
spheres, including North America (e.g., Brachiosaurus altithorax
Riggs, 1903), Africa (Giraffatitan brancai [Janensch, 1914]), and
possibly Europe and South America (see Discussion; Rauhut,
2006; D’Emic, 2012; Mannion et al., 2013).
After the Jurassic/Cretaceous boundary, finds of brachiosaur-
ids have so far been restricted to the Early Cretaceous (Aptian–
Albian) of North America (e.g., Cedarosaurus Tidwell et al.,
1999, Venenosaurus Tidwell et al., 2001, Abydosaurus Chure
et al., 2010). The absence of this group from other landmasses
during the Early Cretaceous has been interpreted either as a
product of local extinctions (Wilson and Sereno, 1998; D’Emic,
2012) or as a result of bias in the generally poor Early Cretaceous
fossil record (Mannion et al., 2013).
Here we report a new titanosauriform taxon from the Early
Cretaceous (Barremian) of Colombia. Dinosaur remains are
remarkably scarce in Cretaceous deposits of northern South
America, and previous records were limited to isolated theropod
teeth from the latest Cretaceous of Colombia (Langston, 1953;
Ezcurra, 2009) and dinosaur footprints (Buffetaut, 2000). The
remains described here provide information on the virtually
unknown dinosaurian faunas of northwestern South America
and increase the currently limited knowledge on sauropod diver-
sity from Gondwana during the earliest (pre-Albian) Cretaceous.
More specifically, the material here reported is the first phyloge-
netically informative sauropod record from northern South
America, with previous records being restricted to an isolated
sauropod vertebra of uncertain affinities (Langston and Durham,
1955). The new taxon is described, its systematic affinities among
titanosauriforms (and in particular brachiosaurids) are analyzed
through a phylogenetic analysis, and its significance for under-
standing the paleobiogeographic history of Brachiosauridae is
discussed. We discuss alternative hypotheses previously pro-
posed to explain the absence of brachiosaurids outside North
America during Early Cretaceous, and suggest that this absence
might indeed be a product of the limited sampling of sauropod
fossils available from this time period.
*Corresponding author.
Color versions of one or more of the figures in this article can be found
online at www.tandfonline.com/ujvp.
Journal of Vertebrate Paleontology e980505 (12 pages)
Óby the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2015.980505
Downloaded by [University of Nebraska, Lincoln] at 14:11 27 August 2015
Institutional Abbreviations—FMNH, Field Museum of Natu-
ral History, Chicago, Illinois, U.S.A.; JACVM, Junta de Acci
on
Comunal Vereda Monquir
a, Vereda Monquir
a, Colombia;
MB.R, Museum f€
ur Naturkunde, Berlin, Germany; MPEF,
Museo Paleontol
ogico Egidio Feruglio, Trelew, Argentina.
SYSTEMATIC PALEONTOLOGY
SAURISCHIA Seeley, 1887
SAUROPODA Marsh, 1878
NEOSAUROPODA Bonaparte, 1986
TITANOSAURIFORMES Salgado, Coria, and Calvo, 1997
BRACHIOSAURIDAE Riggs, 1904
PADILLASAURUS LEIVAENSIS, gen. et sp. nov.
(Figs. 2–5)
Etymology—The generic name honors Dr. Carlos Bernardo
Padilla Bernal (1957–2013), a paleontological enthusiast who led
the creation of the Centro de Investigaciones Paleontol
ogicas of
Villa de Leiva (Colombia), promoted paleontological collection
and research on the Colombian fossil record, and encouraged
the study of this specimen by combining ‘Padilla’ and ‘saurus’
(the Greek word for lizard). The specific name refers to the
locality of Villa de Leiva, from which the specimen derives.
Holotype—JACVM 0001, a single specimen that preserves a
posterior dorsal centrum, the last presacral vertebra, the two
anterior-most and the two posterior-most sacral vertebrae, and
the first eight caudal vertebrae.
Diagnosis—Medium-sized titanosauriform characterized by
the following autapomorphies: (1) first and second caudal verte-
brae with high and dorsally directed prezygodiapophyseal lami-
nae that converge with the centroprezygapophyseal laminae and
form the lateroventral ventral margins of the prezygapophyseal
processes; (2) anterior caudal vertebrae with weakly laterally
expanded transverse processes; and (3) first caudal vertebrae
with divided transverse process, the dorsal section of which is
posterodorsally directed in the first two caudal vertebrae.
Locality and Horizon—Northeast of Villa de Leiva town,
Department of Boyac
a, Ricaurte Province, Colombia (Fig. 1).
The holotype was collected from the Paja Formation
(Barremian–Aptian). Three ammonite specimens were pre-
served along with the vertebral remains and extracted during
preparation of the specimen. Based on these ammonoids, the
new sauropod is here considered to be lower Barremian in age
(see Geological Setting).
GEOLOGIC SETTING
The holotype material of Padillasaurus was collected by local
farmers and subsequently donated and housed at the JACVM.
The exact geographic provenance of the material could not be
determined. Nevertheless, the information provided by the staff
of the JACVM indicates that the material comes from the La
Tordolla area, in the Vereda Monquir
a, Villa de Leiva (Fig. 1).
At this place, the middle member of the Paja Formation crops
out. This stratigraphic level has been informally referred to as
the ‘miembro de arcillolitas abigarradas’ (variegated claystones
member) and considered to be Barremian–Aptian in age
(Etayo-Serna, 1968; Forero and Sarmiento, 1985). The Paja For-
mation is a marine unit deposited during the marine transgres-
sion recorded in Colombia during the Early Cretaceous (Etayo-
Serna et al., 1976). The middle member of this unit is formed by
gypsum-rich claystones with calcareous nodules interbedded
with thin calcareous levels with cryptalgal structures (Forero and
Sarmiento, 1985), interpreted as deposits formed in tidal salt flats
(Forero and Sarmiento, 1985).
The ammonoids preserved with the sauropod specimen consist
of fragments of Gerhardtia galeatoides (JACVM 2 and JACVM
3) and an almost complete specimen of Lytoceras sp. (JAVCM
4). This association corresponds to the Gerhardtia galeatoides
subzone, which represents the lower part of the upper Barremian
in the International Union of Geological Sciences (IUGS) bio-
stratigraphic zonation (cf. Reboulet et al., 2009). Therefore, a
lower upper Barremian age is assigned to the new sauropod spe-
cies described here.
DESCRIPTION
In the following description we follow the nomenclature for
vertebral laminae proposed by Wilson (1999). The vertebrae of
FIGURE 1. Padillasaurus leivaensis, location map of holotype specimen (JACVM 0001).
Carballido et al.—Colombian sauropod (e980505-2)
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the holotype have been preserved in three different segments
that cannot be articulated with each other and include an iso-
lated dorsal centrum (Fig. 2), an as yet unprepared block of
three articulated vertebrae (provisionally identified as the last
presacral and the two anterior-most sacral vertebrae), and 10
articulated vertebrae (the posterior-most two sacral and the
anterior-most eight caudal vertebrae) (Fig. 3).
Dorsal Vertebrae
An almost complete and isolated dorsal centrum (Fig. 2) is
here identified as part of a posterior dorsal vertebra. A possible
second dorsal vertebra is preserved in a segment of three verte-
brae, which are provisionally identified as the last presacral and
the two anterior-most sacral vertebrae.
Isolated Dorsal Centrum—The isolated dorsal centrum is bet-
ter preserved on its right lateral surface. The neural arch is
almost entirely missing, exposing the internal pneumatic features
of this vertebra (Fig. 2). As is evident on the right lateral surface,
the parapophysis was not connected to the centrum, indicating
that this element is a middle to posterior dorsal vertebra. The
ventral surface of the centrum is slightly convex transversely,
and no ventral ridge is present. The centrum is strongly opistho-
coelous, with the concave posterior articular surface slightly
wider than high (Table 1). The presence of opisthocoelous dorsal
centra in mid- to posterior dorsal vertebrae is a synapomorphy of
camarasauromorph sauropods (Salgado et al., 1997; Upchurch,
1998; Wilson and Sereno, 1998; D’Emic, 2012; Carballido and
Sander, 2014). A marked fossa surrounds, ventrally and posteri-
orly, the deep pleurocoel, which is longer than high and has a
rounded posterior edge (Fig. 2A). This is consistent with the gen-
eralized morphology present among non-titanosaur neosauro-
pods. The pleurocoel opens into a large internal camera that
communicates with smaller pneumatic spaces, as can be seen in
dorsal and left lateral views (Fig. 2B, C). At the anterior-most
region of the cotyle, the pneumatic cavities become smaller. The
internal pneumatic spaces are very similar in shape and develop-
ment to those described for the dorsal vertebrae of Giraffatitan
(Janensch, 1947, 1950; MB.R. 4043) and to a slightly lesser
degree those of Tastavinsaurus, which seems to have an interme-
diate condition between the polycamerate and camellate struc-
tures (Canudo et al., 2008; Royo-Torres, 2009:figs. 4.15 and
4.17). Considering the classification of pneumatic structures pro-
posed by Wedel et al. (2000b), the morphology of this centrum
more resembles the polycamerate system than the less advanced
camerate system present in basal camarasauromorphs (e.g.,
Camarasaurus, Tehuelchesaurus, Europasaurus; Wedel, 2003;
Carballido et al., 2011b; Carballido and Sander, 2014) or the
more advanced camellate systems of derived somphospondylans
(e.g., Ligabuesaurus, Saltasaurus; Bonaparte et al., 2006; Cerda
et al., 2012).
Two single laminae are present in the preserved ventral region
of the neural arch. These laminae are the posterior centrodiapo-
physeal lamina (PCDL), in which an anterior branch is tenta-
tively interpreted as the accessory posterior centrodiapophyseal
lamina (aPCDL). The presence of the aPCDL was initially con-
sidered a synapomorphy of titanosaur sauropods (Salgado et al.,
1997), but its presence in other macronarian sauropods (e.g.,
Tastavinsaurus, Brachiosaurus) led to the recognition of a more
widespread distribution of this lamina (Carballido et al., 2012;
D’Emic, 2012). Some differences in the position of the accessory
centrodiapophyseal lamina are observed. In most sauropods, as
seems to be the case in Padillasaurus, the bifurcation of the
PCDL occurs ventrally on the neural arch and the ventral edge
of the accessory lamina extends on to the neural arch pedicels
(e.g., Epachthosaurus, Malawisaurus, Neuquensaurus, Tehuel-
chesaurus; Mart
ınez et al., 2004:fig. 4; Gomani, 2005:fig. 11; Sal-
gado et al., 2005:fig. 4; Carballido et al., 2011b:figs. 6–8). On the
other hand, the bifurcation of the centrodiapophyseal lamina of
some sauropods, such as the brachiosaurids Giraffatitan (MB. R.
2181) and Brachiosaurus (FMNH P 25107), is more dorsally
located and the accessory lamina does not reach the neural arch
pedicels.
FIGURE 2. Padillasaurus leivaensis, JACVM 0001, middle to posterior dorsal vertebra in A, right lateral view; B, left lateral view; C, anterior view;
D, dorsal view. Abbreviations:aPCDL, accessory posterior centrodiapophyseal lamina; ic, internal camera; PL, pleurocoel; ps, pneumatic space; sps,
small pneumatic spaces; PCDL, posterior centrodiapophyseal lamina. Scale bar equals 100 mm.
TABLE 1. Measurements (in cm) of the most complete axial elements
of Padillasaurus leivaensis.
Element PW PH APL AS PS APL/PWH
Dorsal vertebra 25.0 20.0 22.0 CX FL 0.97
5th? sacral vertebra 21.5 23.5 16.8 ? FL 0.75
1st caudal vertebra 20.5 22.0 11.5 FL FL 0.55
2nd caudal vertebra 19.5 20.5 13.5 CC FL 0.67
3rd caudal vertebra 18.5 19.0 11.0 CC FL 0.58
4th caudal vertebra 17.0 18.0 11.0 CC FL 0.62
5th caudal vertebra 16.0 17.0 11.3 CC FL 0.68
6th caudal vertebra 14.5 16.0 11.5 CC FL 0.75
7th caudal vertebra 13.0 15.0 12.0 CC FL 0.85
8th caudal vertebra 13.5 13.5 12.5 CC CC 0.92
Abbreviations:APL, anteroposterior length; AS, anterior articular sur-
face; PH, posterior height; PS, posterior articular surface; PW, posterior
width; PWH, average of the posterior width and height. The anterior and
posterior articular surfaces can be: CC, concave; CX, convex; FL, flat.
Carballido et al.—Colombian sauropod (e980505-3)
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Last Presacral Centrum—The anterior-most element of the
segment, composed of three vertebrae, is considered the last dor-
sal vertebra given that its postzygapophysis is not fused with the
prezygapophysis of the first sacral element. As such, this element
is here considered as the last dorsal element instead of an unfused
anterior sacral, because the degree of fusion present in the other
sacral elements indicates an advanced ontogenetic stage (see Car-
ballido and Sander, 2014). This vertebra preserves the posterior
half of the centrum and the ventral segment of the neural arch.
The preserved centrum does not differ from the isolated element
described above, and also possesses large pleurocoels and poly-
camerate internal pneumatic spaces. The posterior centrodiapo-
physeal lamina and the accessory centrodiapophyseal lamina are
also preserved in this vertebra. The development and orientation
of these laminae is markedly similar to that of the isolated dorsal
centrum, indicating a close position of these elements to one
another within the dorsal series of the axial skeleton.
Sacrum
Five sacral vertebrae are commonly present in non-titano-
saur sauropods (Salgado et al., 1997; Upchurch, 1998; Wilson
and Sereno, 1998), and, based on our identifications and
assuming five sacral elements for Padillasaurus, the specimen
described here preserves remains of the two anterior-most
(in one block) and the two posterior-most of these vertebrae
(in another block). The centra of the first, second, and fifth
sacral vertebrae are complete, whereas the fourth sacral cen-
trum is only represented by a small posterior fragment of its
cotyle (Figs. 3, 4). The two anterior centra and the two poste-
rior centra are strongly fused to each other. The limit of each
centrum, however, is recognized based on the presence of a
slight convexity on its lateral surface (Fig. 4A). The fusion
indicates an advanced ontogenetic stage for this individual.
The first and second sacral vertebrae have pleurocoels, and
FIGURE 3. Padillasaurus leivaensis, JACVM 0001, articulated sequence of the posterior-most two sacral vertebrae (S4 and S5) and the first eight
caudal vertebrae (C1–C8), in A, left lateral view; B, right lateral view. Scale bar equals 100 mm.
Carballido et al.—Colombian sauropod (e980505-4)
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the broken surfaces of their neural arches show the presence
of polycamerate internal pneumatic cavities (which are virtu-
ally identical to those of the isolated dorsal centrum). By
contrast, the fourth and fifth sacrals and the caudal vertebrae
lack internal pneumaticity. Based on these pneumatic fea-
tures, the first two sacral elements are here considered as
dorsosacral vertebrae. Thus, without taking into account a
somitogenetic process in the sacralization of sauropods
(M€
uller et al., 2010; Carballido and Sander, 2014), the mor-
phology of the sacral vertebrae of Padillasaurus is congruent
with the sacralization process proposed by Wilson and Sereno
(1998), in which two dorsals and one caudal vertebra are
incorporated into the sacrum alongside the two primordial
sacral elements. The recognition of two dorsosacrals and one
caudosacral has been relatively common among sauropods,
although some taxa seem to have a different pattern (one
dorsosacral and two caudosacrals; e.g., McIntosh et al., 1996;
Carballido et al., 2011b).
The fifth sacral vertebra lacks the neural spine and the
transverse processes (Fig. 4). Its posterior articular surface is
flat. The sacral transverse processes are eroded, but their
broad base indicates that they extended along most of the
centrum and neural arch (Figs. 4A, B). A deep depression is
present below the transverse process of the fifth sacral
(Fig. 4A). The ventral surface of this sacral vertebra is nar-
row and strongly convex transversely (Fig. 4C). The centro-
prezygapophyseal and prezygodiapophyseal laminae are
recognized on the left lateral surface, but both are poorly
preserved. Above the postzygapophysis there is a well-devel-
oped spinopostzygapophyseal lamina and an anteroposter-
iorly short but well-defined postspinal lamina.
Caudal Vertebrae
The eight preserved caudals are almost complete except for
the dorsal regions of their neural spines and their right transverse
processes. All caudals lack any sign of internal pneumaticity,
unlike the condition in derived titanosaurs, where caudal pneu-
maticity is present (Cerda et al., 2012).
First Caudal Vertebra—The first caudal vertebra is almost
complete except for the dorsal section of its neural spine. Both
the anterior and the posterior articular surfaces of the centrum
are slightly concave, resembling the platycoelous condition
(sensu Romer, 1956) of the first caudal vertebrae of most non-
titanosaur and non-diplodocoid sauropods (Salgado et al., 1997;
Wilson, 2002; Upchurch et al., 2004). The ventral surface of the
centrum is slightly convex transversely (Fig. 4C), differing from
the strong convexity of the ventral surface of the fifth sacral
FIGURE 4. Padillasaurus leivaensis, JACVM 0001, posterior-most sacral and first caudal vertebrae in A, left lateral view; B, right lateral view; C,
ventral view. Abbreviations:CPRL, centroprezygapophyseal lamina; dtp, dorsal segment of the transverse process; poz, postzygapophyses; PRDL,
prezygodiapophyseal lamina; SPOL, spinopostzygapophyseal lamina; SPRL, spinoprezygapophyseal lamina; vtp, ventral segment of the transverse
process. Scale bar equals 100 mm.
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vertebra. The centrum has a deep depression underneath the
transverse process, but its margins lack the deep blind fossa pres-
ent in the second caudal vertebra (see below; Fig. 5A). The
transverse process, which extends from the dorsal half of the cen-
trum to the ventral section of the neural arch, is mostly complete
(except for the lateral tip of its dorsal segment; Fig. 4A). The
transverse process of the anterior caudal of Padillasaurus is
mediolaterally short, in contrast to most sauropods, which have a
mediolaterally extensive transverse process in the anterior cau-
dals (with a mediolateral length that is approximately 50% of
the mediolateral width of the centrum; e.g., Giraffatitan, Veneno-
saurus, Europasaurus; Janensch, 1950:pl. 2; Tidwell et al., 2001:
fig. 11.3; Carballido and Sander, 2014:fig. 26). The transverse
process is divided into dorsal and ventral branches. These two
well-differentiated branches at the lateral end of the transverse
process are also well developed in the second caudal (see below).
The ventral branch projects laterally, whereas the dorsal branch
is posterodorsally directed. The two branches are separated by a
well-marked recess or step, which is pronounced in anterior, pos-
terior, and lateral views (Fig. 4A). The presence of laterally
forked transverse processes in Padillasaurus is here regarded as
an autapomorphy of this taxon.
The ventral branch of the transverse process is here inter-
preted as an extremely developed condition of the ‘ventral bulge’
commonly present among titanosauriform sauropods (Chure
et al., 2010; D’Emic, 2012). The presence of a ventral bulge in
the transverse process of the first caudal vertebra of Brachiosau-
rus (FMNH P 25107) and Giraffatitan (MB.R. 2921) was first
noted by Wilson (2002), who considered this bulge to be an apo-
morphic character of these two taxa (which were referred to the
same genus at that time). More recently, Chure et al. (2010) rec-
ognized the presence of a ventral bulge in Abydosaurus and
identified this character as a synapomorphy of Brachiosauridae.
Nevertheless, the presence of a ventral bulge in the transverse
processes of anterior caudal vertebrae (especially in the first cau-
dal) has been recognized in other titanosauriforms and was most
recently proposed as a synapomorphy of Titanosauriformes
(with a reversal in derived titanosaurs; D’Emic, 2012; Mannion
et al., 2013). Therefore, Padillasaurus shares with several basal
titanosauriforms the ventral bulge on the transverse process of
anterior caudal vertebrae, but has the unique condition of having
it much more developed.
The posterior edge of the dorsal branch of the transverse pro-
cess of Padillasaurus almost reaches the level of the posterior
articular surface of the centrum. A posterior position of the
transverse process is a character shared by titanosauriform sau-
ropods (Mannion and Calvo, 2011; D’Emic, 2012). The posterior
orientation of the transverse process in some non-titanosauri-
forms (e.g., Europasaurus, Galvesaurus, Tastavinsaurus; Man-
nion et al., 2013; Carballido and Sander, 2014) indicates that this
character is in fact shared by camarasauromorph sauropods
(Carballido and Sander, 2014). Nonetheless, the dorsal branch of
the transverse process of Padillasaurus differs from that of other
camarasauromorphs (e.g., Europasaurus, Giraffatitan, Veneno-
saurus, Cedarosaurus) in being strongly dorsally orientated in
the first two caudal vertebrae.
The neural arch is anteriorly positioned, as in derived camara-
sauromorph sauropods (Carballido and Sander, 2014). A promi-
nent and well-developed prezygodiapophyseal lamina extends
from the anterodorsal section of the ventral bulge of the trans-
verse process up to the prezygapophysis (Fig. 4A, B). Wilson
(2002) listed the presence of a prominent prezygodiapophyseal
lamina as an autapomorphic character of Brachiosaurus, which
he considered to include Brachiosaurus altithorax and Giraffati-
tan brancai (this character is present and well developed in the
first and second caudal vertebrae of G. brancai [MB.R. 2921]
and in the second caudal of B. altithorax [FMNH P 25107]).
More recently, Chure et al. (2010) identified the presence of this
lamina as a titanosauriform synapomorphy. A similar develop-
ment of the prezygodiapophyseal lamina in anterior-most cau-
dal vertebrae (mainly the first two elements) is also present in
other camarasauromorphs, such as Tastavinsaurus (Royo-
Torres, 2009:fig. 4.52) and Venenosaurus (Tidwell et al., 1999:
fig. 11.3). A subtle difference exists between the condition of
Padillasaurus and other sauropods with well-developed prezy-
godiapophyseal laminae. In the former, the lamina does not
reach the dorsal margin of the prezygapophysis but converges
with the centroprezygapophyseal lamina. The centroprezyga-
pophyseal lamina ventrally supports the prezygapophysis,
which extends slightly beyond the anterior articular surface of
the centrum (Fig. 4B).
The ventral part of the neural spine is preserved, allowing the
recognition of the spinal laminae. The spinoprezygapophyseal
lamina extends posterodorsally from the prezygapophysis and
vanishes along the anterolateral surface of the neural spine. The
prespinal lamina extends dorsally from the base of the neural
spine, as in other sauropods. Posteriorly, the neural spine is
formed by the spinopostzygapophyseal laminae. The postspinal
lamina is absent. The base of the neural spine is positioned poste-
riorly with respect to the anterior articular surface of the
centrum.
Anterior Caudal Vertebrae (Second to Eighth)—In contrast to
the slightly concave posterior surface of the first caudal centrum,
the posterior surfaces of the second to seventh caudals are flat
and the anterior articular surfaces are concave. The anterior
articular surface of the eighth caudal vertebra is also concave,
but the posterior surface is slightly concave, resembling the pla-
tycoelous condition of the first caudal centrum and differing
from the flat posterior surface of preceding caudal centra. Based
on their articular surfaces, these centra can be identified as pro-
coelous/distoplatyan, as defined by Tidwell et al. (2001). These
authors, and more recently Gonz
alez Riga et al. (2009) and Car-
ballido et al. (2011a), considered this character as only occurring
in a few camarasauromorph taxa (e.g., Cedarosaurus, Malargue-
saurus, Chubutisaurus). Nevertheless, anteriorly concave and
posteriorly flat articular surfaces are actually present in other
basal camarasauromorphs (e.g., Giraffatitan, Galvesaurus, Tasta-
vinsaurus; Carballido et al., 2012; D’Emic, 2012). In contrast, the
derived clade Titanosauria is characterized by the presence of
slightly to strongly procoelous (anteriorly concave and posteri-
orly convex; sensu Romer, 1956) caudal vertebrae (Salgado
et al., 1997; Gonz
ales Riga et al., 2009), with the exception of
taxa such as Opisthocoelicaudia (Borsuk-Bialynicka, 1977),
which have opisthocoelous caudals. The centrum length
increases slightly along the caudal series, whereas the width of
the posterior articular surface decreases. Therefore, the ratio of
the anteroposterior length/mediolateral width of the posterior
centrum increases posteriorly along the preserved caudal verte-
brae (Table 1), a common pattern in sauropod caudal vertebrae
(e.g., Giraffatitan, Cedarosaurus; Janensch, 1950; Tidwell et al.,
1999).
As in the first caudal centrum, the anterior caudal vertebrae
have lateral depressions underneath the transverse process,
which in the second and the fourth caudal vertebrae bear a blind
and well-delimited fossa. Therefore, pneumatic hiatuses are rec-
ognized throughout the anterior caudal series of Padillasaurus,
as was also recently noted for Giraffatitan (Wedel and Taylor,
2013). In the second caudal centrum, the fossa is 3 cm long, 2 cm
high, and 2 cm deep, being slightly smaller in the fourth caudal
(Fig. 6). The shape and position of the lateral fossa of the second
caudal is remarkably similar to the blind fossa present in the sec-
ond caudal vertebra of some specimens of Giraffatitan (MB.R.
2921.2; Wedel and Taylor, 2013). In contrast, the second caudal
centrum of Brachiosaurus lacks such blind fossae (Taylor, 2009;
FMNH P 25107). The presence of blind lateral fossae on anterior
caudal vertebrae was noted by Tidwell et al. (1999) for
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FIGURE 5. Padillasaurus leivaensis, JACVM 0001, second caudal vertebra in A, B, left lateral view; B–D, dorsal view; E,F, posterior view. Abbrevi-
ations:bf, blind fossa; dtp, dorsal segment of the transverse process; nc, neural canal; ns, neural spine; poz, postzygapophyses; prz, prezygapophyses;
vtp, ventral segment of the transverse process. Scale bar equals 100 mm.
Carballido et al.—Colombian sauropod (e980505-7)
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Cedarosaurus and subsequently by Tidwell et al. (2001) for Ven-
enosaurus.InAbydosaurus, blind fossae are present on anterior
caudal vertebrae (D’Emic, 2012), although they are not as deep
as those of Venenosaurus (Chure et al., 2010). Based on this
character distribution, D’Emic (2012) recently recovered the
presence of blind fossae in caudal vertebrae as a brachiosaurid
synapomorphy. Due to the discontinuous presence of this char-
acter within the caudal series, it cannot be confirmed if it was
present in Brachiosaurus, because its holotype only preserves
the two anterior-most caudal vertebrae. The morphology of the
relatively small and more sporadically present blind fossae of
Padillasaurus more closely resembles the condition of some
specimens of Giraffatitan (see Wedel and Taylor, 2013), than the
larger and more continually present fossae of Venenosaurus and
Cedarosaurus (Tidwell et al., 1999; Tidwell et al., 2001).
As for the first caudal vertebra, the preserved transverse pro-
cesses of the second, third, and fifth caudals are mediolaterally
short and divided into ventral and dorsal branches (Fig. 6C),
both unique characters of Padillasaurus. In the other caudal
vertebrae, the transverse processes are broken and this condi-
tion cannot be evaluated. The division of the transverse process
is most strongly developed in the second caudal vertebra
(Fig. 5). Whereas in the third and fifth caudal vertebrae the
division of the transverse process is marked by a gently devel-
oped step, in the second caudal vertebra there is a prominent
and deep recess dividing the distal end of the transverse process
(Fig. 5A, C). The ventral branch of the transverse process, or
hypertrophied ventral bulge, is much larger than the dorsal
branch. The dorsal branch is posterodorsally directed, but not
as dorsally deflected as in the first caudal. In posterior view
(Fig. 5C), the deep step formed between the ventral and dorsal
branches of the transverse process resembles the foramen
described for Lusotitan (Mannion et al., 2013:fig. 5), but in
Padillasaurus this is clearly open laterally, not closed as inter-
preted in Lusotitan.
The neural arches of all preserved caudal vertebrae are anteri-
orly positioned, with the anterior edge located at the same level
as the anterior articular surface of the centrum. Although not
completely preserved, the bases of the neural spines are posteri-
orly positioned with respect to the anterior articular surface of
the centrum. A similar condition is widespread among camara-
sauromorph dinosaurs (e.g., Europasaurus, Giraffatitan, Malar-
guesaurus; Carballido and Sander, 2014) but differs from the
more anteriorly positioned neural spine of Tastavinsaurus
(Royo-Torres, 2009:figs. 4.51, 4.62) and the brachiosaurids
Cedarosaurus (Tidwell et al., 1999:fig. 3) and Venenosaurus
(Tidwell et al., 2001:fig. 11.4). The anterior position of the neural
spine seems to be related to the more vertical or anterior direc-
tion of the neural spine of these taxa. In contrast, the neural
spines of Padillasaurus are posterodorsally oriented, as is evident
from the preserved section of the neural spine of the second cau-
dal. This condition resembles that of most camarasauromorphs
(e.g., Giraffatitan,‘Paluxysaurus,’ Europasaurus; Janensch, 1950;
Rose, 2007;
Carballido and Sander, 2014).
DISCUSSION
Phylogenetic Relationships of Padillasaurus
The specimen JACVM 0001 bears unique characters allowing
the recognition of a new sauropod species, Padillasaurus leivaen-
sis, which shares derived characters with titanosauriform
FIGURE 6. Phylogenetic relationships of Padillasaurus leivaensis within basal Camarasauromorpha, showing A, the strict consensus tree of the 10
most parsimonious trees of 1075 steps; Bdifferent positions that Padillasaurus can occupy within Brachiosauridae.
Carballido et al.—Colombian sauropod (e980505-8)
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sauropods, and, in particular, with brachiosaurids. In order to
test the phylogenetic position of Padillasaurus, a phylogenetic
analysis was conducted through the inclusion of the new taxon in
a modified version of the matrix recently used by Carballido
et al. (2012) and Carballido and Sander (2014). The complete list
of modifications is detailed in Supplementary Data 1 and mainly
includes the incorporation of the camarasauromorphs Abydosau-
rus and Lusotitan, minor changes in character definitions, and
the inclusion of seven new characters and 22 characters from the
data set recently published by D’Emic (2012). The terminal
taxon ‘Paluxysaurus’ and its scorings were revised following the
synonymy of this taxon with Sauroposeidon as recently proposed
by D’Emic (2013). The data set includes the taxon Sauroposei-
don, and its character scorings were based on the available pub-
lished information (Rose, 2007; Wedel et al., 2000a, 2000b;
D’Emic and Foreman, 2012). The final data matrix was com-
posed of 370 characters and 72 taxa. Characters 12, 58, 95, 96,
106, 108, 115, 116, 120, 145, 152, 163, 213, 216, 232, 233, 234, 252,
256, 299, and 301 were treated as ordered.
The phylogenetic analysis was conducted through an equally
weighted parsimony analysis using TNT 1.1 (Goloboff et al.,
2008a, 2008b). The heuristic tree search was performed starting
from 1000 replicates of Wagner trees (with random addition
sequence of taxa) followed by tree bisection-reconstruction
(TBR) branch swapping (saving 10 trees per replicate). This pro-
cedure recovered 50 most parsimonious trees (MPTs) of 1075
steps, which were found in 231 of the 1000 replicates. An addi-
tional round of branch swapping was performed among the saved
trees but failed to find additional MPTs. The strict consensus tree
(see Supplementary Data 1) has some minor differences with
respect to that of Carballido and Sander (2014). Most of these
differences involve taxa that were identified by that previous
analysis as unstable and therefore related to the low nodal sup-
port obtained among camarasauromorphs (Carballido and
Sander, 2014). One of these changes is the recovery of Sauropo-
seidon as a basal somphospondylan, in a similar position to that
recovered for this taxon by D’Emic (2012) and outside Brachio-
sauridae (as recovered for its junior synonym ‘Paluxysaurus’by
Carballido and Sander, 2014:fig. 29). Lusotitan was recovered in
three different positions among camarasauromorphs: as sister
taxon of Tastavinsaurus and more derived camarasauromorphs;
as the sister taxon of Titanosauriformes; and as a basal Sompho-
spondyli. This result is more congruent with the LCDM (Lusoti-
tan continuous Cdiscrete matrix) analysis performed by
Mannion et al. (2013) in which this taxon was recovered as the
sister taxon to Titanosauriformes.
All the MPTs recovered Padillasaurus as a brachiosaurid sau-
ropod, forming part of an unresolved polytomy in the strict con-
sensus tree with Abydosaurus, Giraffatitan, Brachiosaurus, and
the clade formed by Venenosaurus and Cedarosaurus (Fig. 6A).
This polytomy is caused by the alternative positions that Padilla-
saurus takes within Brachiosauridae (Fig. 6B). In the MPTs,
Padillasaurus is placed in alternative positions among the basal
branches of Brachiosauridae, ranging from the very base of the
clade up to a position as the sister taxon to Abydosaurus CGiraf-
fatitan (Fig. 6B). Based on the results of the IterPCR method
(Pol and Escapa, 2009), the alternative positions that Padillasau-
rus occupies among the MPTs are due to the lack of information
(incompleteness) of this taxon rather than to character conflict.
It must be noted that due to the incompleteness of the material,
the inclusion of Padillasaurus in Brachiosauridae is not robustly
supported, and only one extra step is required to place Padilla-
saurus among basal camarasauromorphs more derived than
Tehuelchesaurus and basal to Sauroposeidon.IfPadillasaurus is
forced to occupy even more basal (basal macronarian or non-
neosauropod) or derived (among titanosaurs) positions, two or
more extra steps are required, providing support for its affinities
among basal camarasauromorphs. Nonetheless, given the
available information, the position of Padillasaurus within Bra-
chiosauridae is the most parsimonious interpretation and is sup-
ported by the presence of derived features shared with this clade
(see below).
In the present analysis, as in Carballido and Sander (2014),
Europasaurus is placed as a basal camarasauromorph (outside
Brachiosauridae), a position that is supported by the presence of
several plesiomorphic characters and the absence of brachio-
saurid apomorphies (see Carballido and Sander, 2014; Marp-
mann et al., 2014).
Brachiosauridae is here supported by one unambiguous synap-
omorphy for all trees: presence of a blind lateral fossa in anterior
caudal vertebrae (character 359). Additionally, two extra synap-
omorphies are recovered for some trees: (1) ratio of humerus
length to femur length greater than 0.9 (character 252; in some
trees due to absence of information in Padillasaurus); and (2)
distal end of metatarsal IV dorsomedially oriented (character
370; in some trees due to absence of information in Padillasau-
rus). Among these characters, the presence of blind fossae in
Padillasaurus supports its inclusion among brachiosaurids
(Fig. 6B). It is important to note that although pneumatic axial
characters were described for other titanosauriforms (e.g., Cerda
et al., 2012; Wedel and Taylor, 2013), the presence of blind fos-
sae has so far only been reported for brachiosaurid
titanosauriforms.
Venenosaurus and Cedarosaurus are recovered as sister taxa,
in agreement with the recent analysis published by D’Emic
(2012). The exclusion of Padillasaurus from the Venenosaurus C
Cedarosaurus clade is supported by the lack of anteriorly ori-
ented neural spines in caudal vertebrae (an unambiguous synap-
omorphy of this clade; character 213). Padillasaurus has the
plesiomorphic state for this character, because the neural spines
of caudal vertebrae are posterodorsally oriented.
In the present analysis, Brachiosaurus, Giraffatitan, and Aby-
dosaurus are recovered as a monophyletic assemblage in all
MPTs (with the possible inclusion of Padillasaurus; see Fig. 6B).
The phylogenetic position of these taxa varies among recently
published analyses (e.g., D’Emic, 2012; Mannion et al., 2013;
Carballido and Sander, 2014; Fig. 6B). Part of these differences
may reflect the inclusion or exclusion of cranial information for
Brachiosaurus altithorax (see Supplementary Data 1), as well as
differences in character and taxon sampling among these studies.
In our analysis, the clade formed by Brachiosaurus, Giraffatitan,
and Abydosaurus is supported by a single unambiguous synapo-
morphy: femur head perpendicular to the femur shaft (character
304), a reversal to the plesiomorphic condition present in basal
neosauropods and outgroups. A single synapomorphic character
supports the monophyly of Abydosaurus and Giraffatitan: the
width of the parietal surface that separates the supratemporal
fenestrae is less than the long axis of the supratemporal fenestra
(character 42). This character was recently recovered as synapo-
morphic for Brachiosauridae (including Europasaurus)by
D’Emic (2012). Nevertheless, in Europasaurus, the long axis of
the supratemporal fenestrae is subequal to the parietal width, as
in most non-diplodocoids and non-titanosaur sauropods (see
Marpmann et al., 2015).
The Paleogeographic Distribution of Brachiosauridae
Although the first undisputed brachiosaurid records are from
the Late Jurassic (Giraffatitan, Brachiosaurus), the origin of this
clade has generally been regarded as extending back to the Mid-
dle Jurassic, before the beginning of the fragmentation of Pangea
(e.g., Wilson and Sereno, 1998; Day et al., 2002; D’Emic, 2012;
Mannion et al., 2013). This interpretation is mainly based on the
broad distribution of Brachiosauridae during the Late Jurassic
(see Mannion et al., 2013). Late Jurassic unequivocal brachio-
saurid taxa are known from North America (Brachiosaurus Riggs,
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1903) and Africa (Giraffatitan [Janensch, 1914]). In addition to
these well-known taxa, other more fragmentary Late Jurassic sau-
ropod remains from Europe are currently considered as brachio-
saurids, increasing the diversity and distribution of the clade
during this time. One of these records is the ‘French
Bothriospondylus’ from the Oxfordian of France (Lapparent,
1943), which was recently identified as a brachiosaurid (Mannion
et al., 2013), and which represents the oldest record of this group.
A second possible brachiosaurid from the Late Jurassic of Europe
is Lusotitan atalaiensis from the Kimmeridgian of Portugal
(Antunes and Mateus, 2003), which was placed within brachio-
saurids in the LCDM analyses carried out by Mannion et al.
(2013), but outside this clade in the present analysis (see above).
Finally, a possible brachiosaurid has also been reported from the
Late Jurassic of South America. Rauhut (2006) described frag-
mentary sauropod materials from the Ca~
nad
on Calc
areo Forma-
tion of Patagonia (Oxfordian–Kimmeridgian) and referred it to
Brachiosauridae, although there is no current agreement on the
brachiosaurid affinities of these remains (D’Emic, 2012; Mannion
et al., 2013). Despite the uncertainties regarding the affinities of
some of the more fragmentary material, it is clear from the fossil
record that undisputed brachiosaurid remains were present dur-
ing the Late Jurassic in both the northern (North America,
Europe) and southern (Africa) hemispheres.
Despite the widespread distribution of brachiosaurids during
the Late Jurassic, they have a much more restricted distribution
during the Early Cretaceous. Up to now, Cretaceous brachio-
saurids were exclusively known from the Aptian–Albian of
North America (Venenosaurus, Cedarosaurus, and Abydosaurus;
Tidwell et al., 1999, 2001; Chure et al., 2010). This Early Creta-
ceous distribution has been interpreted as a product of local
extinctions in Europe, Africa, and South America (Wilson and
Sereno, 1998; D’Emic, 2012). Mannion et al. (2013), however,
recently noted that this absence could also be caused by a bias in
the poor Early Cretaceous sauropod fossil record, and noted the
presence of two isolated teeth from the Early Cretaceous of Leb-
anon (Buffetaut et al., 2006) that have posteriorly twisted crowns
as in the brachiosaurids Giraffatitan and Abydosaurus (Chure
et al., 2010; D’Emic, 2012).
The Early Cretaceous is indeed a period of low sauropod
diversity worldwide, especially for the earliest Cretaceous and in
comparison with the high diversity recorded in the Late Jurassic
(Mannion et al., 2011). Whether or not this decrease is the prod-
uct of truly low diversity following an extinction event at the
Jurassic/Cretaceous boundary or is mainly controlled by sam-
pling biases is unclear at the moment (Mannion et al., 2011).
New records and a more complete knowledge of the Early Creta-
ceous sauropod faunas will be critical to thoroughly test these
alternative explanations (including the extent of the local extinc-
tions of brachiosaurids after the Late Jurassic).
The brachiosaurid affinities of Padillasaurus leivaensis from
the Early Cretaceous (Barremian) of Colombia (recovered here
in all most parsimonious hypotheses, although with low character
support) are relevant to these questions, because they suggest
that this clade survived in northwestern Gondwana after the
Jurassic/Cretaceous boundary. Furthermore, considering also
the putative brachiosaurid teeth from Lebanon (Buffetaut et al.,
2006; Mannion et al., 2013), it is possible that brachiosaurids
actually survived into the Cretaceous at low latitudes in all of
northern Gondwana. During the Early Cretaceous, Colombia
was located close to the equator in northwestern Gondwana
(Scotese, 2001) and Lebanon was part of the Afro-Arabian plate
in northeastern Gondwana (Buffetaut et al., 2006; Ksepka and
Norell, 2010; Mannion et al., 2013).
In more general terms, this new information, coupled with the
recent discovery of a diplodocid from the Early Cretaceous of
southern South America (Gallina et al., 2014), suggests that part
of the high sauropod diversity recorded during the Late Jurassic
survived the Jurassic/Cretaceous extinction event recognized by
Raup and Sepkoski (1986). The rarity of these sauropod records
in the Early Cretaceous may reflect a decrease in diversity and
abundance of these lineages in combination with an overall
poorer fossil record (especially in Gondwana).
CONCLUSIONS
The vertebral remains described here allow the recognition of
Padillasaurus leivaensis, a new titanosauriform taxon from the
Early Cretaceous of Colombia. This record provides the first
informative dinosaurian record for the Early Cretaceous of
northern South America, which was previously limited to a sin-
gle sauropod vertebra of uncertain affinities. The comparative
and phylogenetic analyses performed place the new taxon within
Brachiosauridae in all most parsimonious hypotheses, although
the character support for its position is weak given the incom-
pleteness of some skeletal regions that would be phylogenetically
informative (e.g., humerus, femur, metatarsus).
The brachiosaurid affinities of Padillasaurus are relevant for
understanding the biogeographic history of this group given that
this clade of titanosauriform sauropods is broadly distributed
during the Late Jurassic but has a markedly restricted distribu-
tion during the Early Cretaceous. In particular, these remains
provide support for considering the apparent absence of brachio-
saurids outside North America during the Early Cretaceous to
be due to a deficient fossil record rather than due to local extinc-
tions (at least in northern Gondwana).
ACKNOWLEDGMENTS
We are grateful to the Junta de Acci
on Comunal Vereda Mon-
quir
a, and especially to its president, J. Gonzales, for access to
the specimen housed in the Museo el F
osil (Villa de Leiva) and
the permission to prepare it. This collaborative research was pos-
sible after the efforts of the late Dr. Carlos Bernardo Padilla
Bernal, who gathered this team and promoted the study of these
fossil remains. His continuous efforts to explore, recover, and
preserve paleontological heritage in Colombia are his legacy and
will endure for generations to come. We thank to J. Parra and F.
H. Parra from the Centro de Investigaci
on Paleontol
ogica (CIP),
Villa de Leiva, Colombia, for the preparation of the specimen.
M. D’Emic and M. Wedel made useful comments during the
preparation of the manuscript. We are grateful to the reviewers
S. Poropat and E. Tschopp, and the editor R. Butler, whose com-
ments have greatly improved this paper. R. Romero helped with
the illustrations of the vertebrae. This work was financed by the
Agencia Nacional de Promoci
on Cient
ıfica y Tecnol
ogica
(ANCYT, Argentina; PICT 0808; PICT 0378), the Centro de
Investigaciones Paleontol
ogicas, and Rochem Biocare Colombia
SAS. This research was developed within the framework of the
cooperation agreement between CONICET (Argentina) and the
CIP (Colombia).
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Submitted June 17, 2014; revisions received August 19, 2014;
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Handling editor: Richard J. Butler.
Citation for this article: Carballido, J. L., D. Pol, M. L. Parra Ruge, S.
Padilla Bernal, M. E. P
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