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The South American Gomphotheres

(Mammalia, Proboscidea, Gomphotheriidae):
Taxonomy, Phylogeny, and Biogeography

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J Mammal Evol
DOI 10.1007/s10914-012-9192-3

ORIGINAL PAPER

The South American Gomphotheres (Mammalia,

Proboscidea, Gomphotheriidae): Taxonomy, Phylogeny,
and Biogeography
Dimila Moth & Leonardo S. Avilla & Mario A. Cozzuol

# Springer Science+Business Media, LLC 2012

Abstract The taxonomic history of South American Gom-
photheriidae is very complex and controversial. Three species
are currently recognized: Amahuacatherium peruvium,
Cuvieronius hyodon, and Notiomastodon platensis. The for-
mer is a late Miocene gomphothere whose validity has been
questioned by several authors. The other two, C. hyodon and
N. platensis, are Quaternary taxa in South America, and they
have distinct biogeographic patterns: Andean and lowland
distributions, respectively. South American gomphotheres be-
came extinct at the end of the Pleistocene. We conducted a
phylogenetic analysis of Proboscidea including the South
American Quaternary gomphotheres, which resulted in two
most parsimonious trees. Our results support a paraphyletic
Gomphotheriidae and a monophyletic South American
gomphothere lineage: C. hyodon and N. platensis. The late
Miocene gomphothere record in Peru, Amahuacatherium
D. Moth (*)
Programa de Ps-graduao em Cincias Biolgicas (Zoologia), a brachycephalic and brevirostrine skull, a short and curved
Museu Nacional/UFRJ,
Quinta da Boa Vista,
20940-040, Rio de Janeiro, RJ, Brazil
e-mail: dimothe@hotmail.com
L. S. Avilla
Departamento de Zoologia, Instituto de Biocincias,
Laboratrio de Mastozoologia,
Universidade Federal do Estado do Rio de Janeiro,
Avenida Pasteur, 458, Urca, 22290-240,
Rio de Janeiro, RJ, Brazil
e-mail: leonardo.avilla@gmail.com
M. A. Cozzuol
Departamento de Zoologia, Instituto de Cincias Biolgicas,
Universidade Federal de Minas Gerais,
Avenida Antnio Carlos, 6627, Pampulha,
31270-910, Belo Horizonte, Minas Gerais, Brazil
e-mail: cozzuol@icb.ufmg.br
peruvium, seems to be a crucial part of the biogeography
and evolution of the South American gomphotheres.
Keywords South American Gomphotheres . Systematic
review . Taxonomy . Proboscidea
Introduction
The family Gomphotheriidae is, so far, the only group of
Proboscidea recorded in South America. Together with the
megatheriid sloths Eremotherium laurillardi Lund, 1842,
the Megatherium americanum Cuvier, 1796, and the
notoungulate Toxodon platensis Owen, 1840, they are the
most common late Pleistocene representatives of the mega-
fauna in South America (Paula-Couto 1979). Similar to the
Pleistocene and Holocene members of the family Elephan-
tidae (e.g., extant elephants and extinct mammoths), the
South American gomphotheres are characterized by having
mandibular symphysis, and specialized dentition, which
consists of a pair of upper tusks (second incisors) and six
pairs of upper and lower molars (Simpson and Paula-Couto
1957; Paula-Couto 1979; Prado et al. 2001; Ferretti 2008a).
Taxonomy and Diversity of South American
Gomphotheres
The taxonomic history of the South American gompho-
theres is long and complex. Most authors consider that they
were represented by three genera: Cuvieronius Osborn,
1923, Haplomastodon Hoffstetter, 1950, and Stegomastodon
Pohlig, 1912 (Simpson and Paula-Couto 1957; Ficcarelli et al.
1995; Ferretti 2008a, 2010). Later, some authors (Alberdi and

Prado 1995; Alberdi et al. 2002, 2004, 2007; Prado et al.
2005; Prado and Alberdi 2008) synonymized Haplomastodon
with Stegomastodon, reducing the diversity of genera, but
keeping three species: Cuvieronius hyodon (Fischer, 1814),
Stegomastodon platensis (Ameghino, 1888), and S. waringi
(Holland, 1920). Very recently, Moth et al. (in press)
proposed to include the last two species in a single species,
under the name Notiomastodon platensis, as first suggested by
Madden (1984).
The first report of gomphothere remains from South
America was made by Cuvier (1806), who analyzed an
isolated left M2 from Imbabura, northern Ecuador, and
named it as the Mastodonte des Cordillires. In 1824,
Cuvier established the formal names Mastodon andium for
the Imbabura (Ecuador) gomphotheres and M. humboldtii
for a molar from Concepcin (Chile). Before this study,
Fischer (1814) analyzed the same specimen from Imbabura,
and named it as Mastotherium hyodon, which has priority
over Mastodon andium (see Cabrera 1929). The genus
Cuvieronius was erected by Osborn (1923) and later,
Cabrera (1929), in a revision of the Argentine gompho-
theres, synonymized Mastodon argentinus Ameghino, 1888,
under the name Cuvieronius hyodon (Fischer, 1814).
Ficcarelli et al. (1995), in a taxonomic revision of Cuvier-
onius, validated C. tarijensis (Ameghino, 1902), and pro-
posed the material referred to C. tarijensis from Tarija,
Bolivia, as the topotype for the species. Recently, Lambert
and Shoshani (1998) considered C. hyodon as the type
species of Cuvieronius, but Lucas (2009) argued that the
genus is monospecific, considering C. tropicus Cope, 1884,
from Mexico, as a junior synonym of C. hyodon. Lucas
(2009), in order to solve the taxonomic problem of the name
Mastotherium hyodon, suggested to keep the name Cuvier-
onius and consider M. hyodon as its type species, by desig-
nating a neotype to M. hyodon (the skull and lower jaw from
Tarija, Bolivia originally described and illustrated by Boule
and Thevenin (1920)).
The diagnostic material of C. hyodon comes from Tarija,
Bolivia, where several skulls and postcranial remains were
found (Marshall and Sempere 1991; Coltorti et al. 2007).
Cuvieronius hyodon is a short-jawed gomphothere charac-
terized by having a depressed and elongated cranium, a pair
of twisted upper tusks, which have a spiraled enamel band
along their entire length, and lower deciduous incisors in
some juveniles (Prado et al. 2005; Ferretti 2008a, b).
The first one to review the taxonomy of South American
lowland gomphotheres was Florentino Ameghino (1888,
1889, 1891), who defined four species that he placed in
the genus Mastodon. After that, the Argentinean fossils were
again reviewed by Cabrera (1929), who synonymized some
species described by Ameghino (1889) and transferred them
to the North American genus Stegomastodon, recognizing
only two species: S. platensis and S. superbus. He also
J Mammal Evol
proposed a new genus and species, Notiomastodon ornatus,
based on a short, massive tusk and a dentary from Playa del
Barco, Argentina. Hoffstetter (1950) proposed Haplomasto-
don as a subgenus of Stegomastodon and, in 1952, elevated
Haplomastodon to full generic status, based on the absence
of transverse foramina in the atlas (1st cervical vertebra). He
also included all Brazilian gomphotheres in a new species
Stegomastodon brasiliensis Hoffstetter, 1952. The trans-
verse foramen of the axis was recognized as a variable
character by Simpson and Paula-Couto (1957), and they
placed all Brazilian gomphotheres in Haplomastodon war-
ingi (Holland, 1920). Ficcarelli et al. (1995), based on
diagnostic gomphothere remains from northern Ecuador,
synonymized all Haplomastodon species with H. chimbor-
azi (Proao, 1922). Alberdi and Prado (1995) included the
species S. superbus (Ameghino, 1888), and N. ornatus in S.
platensis. The genus Haplomastodon was later synony-
mized with the genus Stegomastodon, generating the new
combination S. waringi (Alberdi et al. 2002). Lucas and
Alvarado (2010) used the combinations Notiomastodon pla-
tensis, in replacement of S. platensis, which was previously
suggested by Madden (1984) and Ferretti (2008a). Ferretti
(2010) used the combination Haplomastodon chimborazi
and employed quote marks for Stegomastodon platensis,
arguing that the South and North American species attribut-
ed to this genus are not congeneric with S. mirificus (Leidy,
1859), the type species of the genus, from North America.
Moth et al. (in press) reviewed the Pleistocene South
American lowland gomphotheres and concluded that they
belong to a single species for which the valid name is Notio-
mastodon platensis. Consequently, only two gomphothere
species were present in the Quaternary of South America:
the Andean Cuvieronius hyodon, and the lowland Notiomas-
todon platensis (Fig. 1).
The holotype of Notiomastodon ornatus (Museo Argen-
tino de Ciencias Naturales Bernardino Rivadavia, collec-
tion number MACN 2157) is described by Cabrera (1929),
and it is a mandible with a short symphysis, without lower
incisors, bunolophodont molars with several accessory con-
ules (double trefoil wear pattern), and short, massive and
upper curved tusk with lateral enamel band (Cabrera 1929:
Figs. 2-4). Furthermore, Stegomastodon platensis is de-
scribed as having a variable wear pattern on the molars,
more complex than Haplomastodon chimborazi, and
straight to slightly upper curved tusks, without enamel band
(Prado et al. 2005). Both genera show a similar tusk mor-
photype and should represent the same taxon, in which
Notiomastodon has priority and Stegomastodon is not ap-
plicable to the South American species (Moth et al. in
press). A brachycephalic skull with a high parieto-occipital
region, a pair of straight or curved upper tusks with absence
of spiraled torsion, complex molar teeth with double-trefoil
and simple-trefoil wear patterns (Moth et al. in press)
J Mammal Evol
Fig. 1 Main differences between the South American Quaternary
gomphotheres. a anterior view of a Notiomastodon platensis high
cranium (reconstructed, based on Cabrera 1929) with straight upper
tusks from Arrecifes, Buenos Aires province, Argentina (MLP 8-1,
Museo de La Plata paleontological collection). b anterior view of a
Cuvieronius hyodon low cranium with twisted upper tusks from Tarija,
Bolivia (MACN Pv 1291, Museo Argentino de Ciencias Naturales
Bernardino Rivadavia paleontological collection). c lateral view of
the same specimen described in a. d lateral view of the same specimen
described in b
characterize the species N. platensis. Cabrera (1929) indi-
cates that Cuvieronius hyodon have a low and long skull,
large upper tusks with twisted enamel band, lack of lower
tusks in adults, and short dentary symphysis. Otherwise, the
genus Stegomastodon is represented in North America by
three species: S. primitivus, S. mirificus, and S. aftoniae
(Lucas et al. 2011). The genus is characterized by a skull
with a tall frontal and parietal region, long and relatively
straight tusks that lack enamel, no lower tusks, the mandib-
ular symphysis is relatively short and tall, and the m3 has
five lophids and double trefoils. It differs from Notiomasto-
don in some structures of the scapula, humerus, pelvis, and
femur, and it seems to have a greater degree of graviportality
than South America gomphotheres (Lucas et al. 2011).
The discovery and description of Amahuacatherium peru-
vium (Romero-Pittman, 1996), a gomphothere with vestigial
lower incisors from the late Miocene (approximately 9.59.0
MyBP) of the Peruvian Amazonia, has inspired a great con-
troversy (Campbell et al. 2000, 2006, 2009, 2010; Alberdi et
al. 2004; Woodburne 2010). Some authors have stated that the
holotype of A. peruvium is, actually, a specimen of Stegomas-
todon (Alberdi et al. 2004), its proposed age is mistaken, and
its primitive features (lower incisors and simpler molars with
complexity similar to Haplomastodon) were misinterpreta-
tions caused by its fragmentary nature and taphonomomic
processes (Alberdi et al. 2004; Woodburne 2010). Recently,
Campbell et al. (2009, 2010) have refuted the critics on the
morphological features and age of Amahuacatherium. The
presence of pre-Quaternary gomphotheres in South America
remains an open issue and more studies are needed to eluci-
date the evolutionary history of South American proboscideans.
The only consensus about proboscidean history in South
America is that they became extinct by the end of the
Pleistocene (Ficcarelli et al. 1997; Prado et al. 2001; Alberdi
et al. 2002; Snchez-Chilln et al. 2004; Prado et al. 2005),
but the causes for their extinction are still unresolved.
Biogeography of South American Gomphotheres
Under the traditional view, gomphotheres arrived in South
America during the Great American Biotic Interchange
(GABI), after the establishment of the Panamanian Isthmus
at 3.12.8 MyBP (Coates and Obando 1996), a land-bridge
that connects North and South America through Central Amer-
ica (Cione and Tonni 2001; Coltorti et al. 2007; Woodburne
2010). However, the record of the Amahuacatherium peruvium
from the late Miocene Amazonian lowlands of Peru (Campbell
et al. 2000, 2006, 2009) would indicate, if its morphological
features are accurately interpreted and its age is correct, that
proboscideans were present in South America prior to the main
stream of the GABI. Besides Amahuacatherium, other species
also have an early arrival into South America, such as
procyonid carnivorans and sigmodontine rodents (Woodburne
2010). Furthermore, ground sloths and terror birds reached
North America at the same time (Woodburne 2010 and
references cited there). Consequently, the beginning of
the GABI could have started approximately ten million
years ago, with a brief period of global sea level drop that
allowed the emergence of a small land connection between
Central and South America. Between 10.71 and 9.36 MybP,
calcareous nannofossils indicate a separation between the
Caribbean and the east Pacific during this time, compared
with the early and middle Miocene. This was followed by
a reinstatement of a deeper connection between 8.35 and
3.65 MybP between the Caribbean Sea and the eastern equa-
torial Pacific, and a final closure of the Isthmus of Panama
by 2.76 MybP (Kameo and Sato 2000). About the same
geological variations are observed in the sedimentological
record by Coates et al. (2004).
In North and Central America, the genus Cuvieronius has a
continuous distribution while the genus Stegomastodon has a
discontinuous distribution in Central America (Arroyo-Cabrales
et al. 2007). This observation has led some authors to suggest
that a taxonomic review of the South American species referred
to Stegomastodon is needed (Arroyo-Cabrales et al. 2007;
Ferretti 2008a, 2010; Lucas et al. 2011).
During the Pleistocene, each of the South American
gomphotheres spread throughout the continent in two dif-
ferent corridors. In the Andean or highland corridor,
Cuvieronius hyodon was dominant, spreading in the present

Fig. 2 Map of the geographical
distribution of South American
gomphotheres during the
Pleistocene. The circles
represent Notiomastodon
platensis and the triangles
represent Cuvieronius hyodon.
Modified from Simpson and
Paula-Couto (1957), Alberdi et
al. (2002), Cione et al. (2005),
Ferretti (2008a), and Lucas
(2008)
countries of Colombia, Ecuador, Bolivia, Peru, and Chile, and
the lowland corridor, where Notiomastodon platensis
prevailed in Brazil, Ecuador, Venezuela, Colombia,
Argentina, Uruguay, Paraguay, and the western coastal
areas of Peru and Chile (Simpson and Paula-Couto
1957; Alberdi and Prado 1995; Prado et al. 2001;
Alberdi et al. 2002; Ferretti 2008a, 2010; Moth et al. in press;
Fig. 2). Cuvieronius hyodon seems to be adapted to live in the
highlands, which is supported not only by its geographic
distribution, but also by stable isotope analysis (Snchez-
Chilln et al. 2004). On the other hand, the lowland gompho-
there, Notiomastodon platensis, seems to avoid drier environ-
ments (Winck et al. 2010).
Phylogeny of South American Quaternary
Gomphotheres
The phylogenetic relationships of South American gompho-
theres are another controversial issue and it is directly
J Mammal Evol
related to their unsolved taxonomic problems. Previous
phylogenies by Shoshani et al. (2006) include 125 char-
acters and 40 terminals at the generic level; however,
they published only a simplified cladogram at the fam-
ily level; Prado and Alberdi (2008) conducted a cladis-
tic analysis of the trilophodont gomphotheres, with
emphasis on the South American species. This phylog-
eny is problematic in many points, which invalidate
both the phylogeny itself and the biogeographic analysis
based on it, as shown by Cozzuol et al. (in press).
Furthermore, several recent studies of the group (Arroyo-
Cabrales et al. 2007; Ferretti 2008a, 2010; Lucas and
Alvarado 2010; Cozzuol et al. in press; Moth et al. in
press) show the necessity of a systematic revision of the
South American gomphotheres.
This study proposes a new systematic and biogeographic
view for the South American Pleistocene gomphotheres and
presents inferences about the relation of these taxa within
the family Gomphotheriidae that follows the taxonomic
review conducted in Moth et al. (in press).
J Mammal Evol
Fig. 3 Resulting consensus
tree based on the matrix
modified from Shoshani et al.
(2006) and Cozzuol et al. (in
press), including the new taxon
Notiomastodon platensis. The
clade 1 represents Elephantida
and clade 4 represents the South
America Quaternary
gomphotheres
Materials and Methods
A phylogenetic analysis was performed in order to elu-
cidate the relationships of the South American gompho-
theres. We modified the original matrix from Cozzuol
et al. (in press, modified from Shoshani et al. 2006),
by reviewing some features and merging Stegomastodon
platensis and Haplomastodon waringi into Notiomastodon
platensis. We also recognized Stegomastodon only by
North American species and we recoded the character
states for this genus based only on these species (Arroyo-
Cabrales et al. 2007; Ferretti 2008a, 2010; Lucas et al.
2011). We are in agreement with Ferretti (2008a, 2010),
Lucas et al. (2011), and Moth et al. (in press), who
consider that Stegomastodon is not present in South
America.
We ran the taxon-character matrix with TNT software
(Goloboff et al. 2008) using the Implied weighting set-
ting, with K06, and conducted the search using defaults of
xmult command to finding optimal score 20 times inde-
pendently, plus ten cycles of tree-drifting strategies (Goloboff
et al. 2008). Group supports are calculated by TBR-swapping
the trees found, keeping note of the number of steps needed to
collapse each group.
Results and Discussion
The phylogenetic analysis conducted here resulted in two
most parsimonious trees with 7.31205 fit as the best score
and the consensus of these two trees is represented in Fig. 3.
Accordingly, the family Gomphotheriidae (sensu Shoshani
et al. 2006) is paraphyletic. This is not a new argument;
Maglio (1972), Tassy (1996), and Arroyo-Cabrales et al.
(2007) also rejected the monophyly of the family Gompho-
theriidae. Moreover, Shoshani and Tassy (1996) also argued
that it requires too many changes to group all taxa tradition-
ally incorporated in gomphotheres. To consider this taxon
a natural group, it must include the monophyletic Elephan-
tidae and Stegodontidae (Fig. 3, Clade 1). This clade, named
Elephantida, was also recognized by Shoshani et al. (2006).
In our results, the Elephantida includes several lineages
(Fig. 3): a basal great polytomy compromising most of the
traditional gomphotheres, the monophyletic (Notiomastodon
platensis and Cuvieronius hyodon), and the four consecutive
sister-taxa of Elephantoidea (Elephantidae plus Stegodontidae
by Shoshani et al. 2006), which are (Stegomastodon mirificus
(Tetralophodon (Anacus (Paratetralophodon)))). The Ele-
phantida is diagnosed here by the presence of a postcingulum
on the lower third permanent molars (character 124, state 1).

Fig. 4 Five possible systematic combinations involving Amahuacathe-
rium peruvium and the gomphotheres recorded in South America, Cuvier-
onius hyodon and Notiomastodon platensis. a the suggested clade is (C.
hyodon and N. platensis), with South American arrival at the Great
America Biotic Interchange. b the suggested clade is (C. hyodon and N.
platensis) with A. peruvium synonymized with N. platensis. c the sug-
gested clade is (Cuvieronius hyodon (Amahuacatherium peruvium and
Notiomastodon platensis). d the suggested clade is (Amahuacatherium
peruvium (Notiomastodon platensis and Cuvieronius hyodon). e the sug-
gested clade is (Notiomastodon platensis (Amahuacatherium peruvium
and Cuvieronius hyodon). Important moments in the evolutionary and
The development of the postcingulum in gomphotheres can
be associated with the increase in number of loph(id)s in
Elephantida, which are tetra, penta, or have more than five
lophids on the third lower molars. Tassy (1996) recognized an
evolutionary trend for the Proboscidea toward the augmenta-
tion on cusps due to the subdivision of the main mammalian
cusps and by the increase in number of loph(id)s. The incre-
ment in crown morphology in Elephantida, compared to other
Proboscidea, was probably positively selected in response to a
more abrasive and complex herbivorous diet in most of the
continents after the spread of grasslands from the early Mio-
cene on.
The monophyletic clade 2 comprises clade 3, (Stegomastodon
primitivus (Tetralophodon (Anancus (Paratetralophodon (Ele-
phantoidea))))), and the natural group Notiomastodon platensis
plus Cuvieronius hyodon (clade 4) (Fig. 3). Clade 2 is diagnosed
by M3 with postentoconule, pentaloph or with more than five
lophs on the molars (character 34, state 5), and a short and spout-
like mandibular symphysis (character 94, state 2). With the
exception of Cuvieronius hyodon and Tetralophodon, in which
J Mammal Evol
biogeographic histories of gomphotheres in Americas: 1) Short
land connection between Central and South Americas at 10.71
to 9.36 MyBP. 2) A. peruvium record in Peruvian Amazon at
approximately 9.59.0 MyBP. 3) Oldest record of Cuvieronius in
North America, at 4.9 MyBP. 4) Isthmus of Panama formation at
3.1 to 2.8 MyBP. 5) Total Andes uplift at 2.7 MyBP. 6) Great
American Biotic Interchange at approximately 2.8 MyBP. 7) Early
Pleistocene gomphothere record in Argentina, at 2.5 MyBP. 8)
Oldest record of C. hyodon in South America, middle/late Pleis-
tocene from Tarija. 9) Pebas system formation, from 10 to 7
MyBP
some specimens bear vestigial (or reduced) and deciduous man-
dibular tusks (Ferretti 2008b), all members of clade 2 lack
mandibular tusks and are brevirostrine. This brevirostrine condi-
tion, an extreme mandible shortening, was possible probably
because of the horizontal teeth displacement condition evolved
in the Elephatimorpha (Shoshani et al. 2006). Therefore, in the
members of clade 2, the brevirostrine condition probably allowed
the reduction in the number of erupted molars, where only one
molar or one molar plus part of the successive adjacent tooth are
in use at the same time on each half jaw.
The monophyletic clade 3 is diagnosed by m2 with four
distinct lophids (character 33, state 3, tetralophodont) and
M2 with four or more distinct lophs (character 40, state 2).
Thus, the tendency for increase in the number of loph(id)s is
also observed in clade 3, but it occurs on the second upper
and lower molars, which have more loph(id)s when com-
pared to the other taxa in Elephantida. The most basal taxon
of this clade, Stegomastodon mirificus, is represented here
by the genus Stegomastodon, which is endemic to North
America (Ferretti 2010; Lucas et al. 2011; Moth et al. in
J Mammal Evol
press). This genus was previously misinterpreted to occur
also in South America, and two species were considered, S.
waringi and S. platensis (Alberdi and Prado 1995; Alberdi
et al. 2002). However, Moth et al. (in press) considered
these taxa as the same species and synonymyzed both under
a new taxonomic combination, Notiomastodon platensis.
Furthermore, Stegomastodon has four loph(id)s on m2/M2
and five to six loph(id)s on m3/M3, while N. platensis has
only three loph(id)s on m2/M2 and four to five on m3/M3
(never six complete loph(id)s).
The South American Notiomastodon platensis and the
Panamerican Cuvieronius hyodon form the monophyletic
clade 4, which is diagnosed by upper tusks dorsally curved
in lateral view (character 7, state 2). Notiomastodon platensis
has tusks in a variety of forms (Moth et al. in press), but it
could be dorsally curved at a first stage, as does C. hyodon. In
this way, clade 4 is supported by only one synapomorphy, and
we believe that Cuvieronius also needs taxonomic review, in
agreement with Montellano-Ballesteros (2002).
The divergence of these clades probably occurred prior to
the early-middle Miocene (late Hemingfordian to early Bar-
stovian) when the proboscideans arrived in North American
from Asia (Vaughan et al. 2000). Therefore, the clade includ-
ing Notiomastodon (the only Pleistocene genus endemic to
South America) and Cuvieronius (North, Central, and South
American) should have originated in Central America. Here
we agree with Madden (1984), Ferretti (2008a, 2010), and
Lucas and Alvarado (2010), who supported the monophyly of
South American gomphotheres. The resulting topology sug-
gests that the South American gomphotheres had a common
biogeographical history.
In South America, the earliest well dated record of
Gomphotheriidae, except for the A. peruvium record, is
a fragmentary vertebra from the early Pleistocene of
Argentina (Uquian; Lpez et al. 2001; Reguero et al.
2007), while all other records are middle to late Pleis-
tocene (Prado et al. 2005). The oldest record of C.
hyodon in South America is from the early Pleistocene
of Tarija (Ensenadan; Prado et al. 2005). However, the
age of the Tarija deposits is controversial (middle to late
Pleistocene; MacFadden et al. 1983; Coltorti et al.
2007), and it might not represent the oldest C. hyodon
record in South America (Ferretti 2008a). In Central
America, the genus Notiomastodon is not recorded and
Cuvieronius is known from the early Irvingtonian (El
Salvador), although its arrival in the late Blancan is suggested
(Montellano-Ballesteros 2002; Lucas and Alvarado 2010). In
North America, Cuvieronius is recorded for the late Blancan in
United States and Mexico (Lucas et al. 1999).
In this way, five hypotheses are proposed to explain the
origin and biogeographical history of South American gom-
photheres. Three of them include Amahuacatherium peru-
vium as a valid taxon; however, it was not included on our
systematic analysis because we did not analyze these
remains personally, and it was not included in the matrices
we revised here (see Materials and Methods section). The
question about the validity of A. peruvium is still open and
needs a systematic character review (presence of lower tusk
and characters related to molar morphology), but we agree
with Campbell et al. (2000, 2009, 2010) in respect to the age
of this record, considered as the oldest gomphothere in
South America.
The first premise is the classical hypothesis for the
biogeographical history of the South American gompho-
theres (Fig. 4a), widespread in the literature (Alberdi and
Prado 1995; Alberdi et al. 2002, 2004, 2007; Prado et al.
2005; Prado and Alberdi 2008). In this case, both C. hyodon
and N. platensis arrived in South America during the Great
American Biotic Interchange, after the formation of the
Isthmus of Panama (3.1 to 2.8 MyBP), and spread through
the Andean and lowlands corridors in the Pleistocene.
However, there are several issues that this does not explain:
(1) the presence of A. peruvium in the late Miocene of Peru,
(2) when and where the common ancestor of C. hyodon and
N. platensis diverged (which must have occurred in Central
or North America before 4.9MyBP, the oldest record of
Cuvieronius in North America), (3) how N. platensis
reached the lowlands from both sides of the Andes after
their uplift (2.7 MyBP), and (4) why there is no record of N.
platensis in Central and/or North America.
In the second hypothesis (Fig. 4b), we considered the
taxonomic revision made by Alberdi et al. (2004) for Peru
gomphotheres, in which A. peruvium was synonymized with
Stegomastodon and, later, it was synonymized with Notio-
mastodon platensis by Moth et al. (in press). Thus, this
hypothesis suggests the existence of a Panamerican ancestor
to the clade (Cuvieronius hyodon and Notiomastodon pla-
tensis) at 10.71 to 9.36 MyBP, when South and Central
Americas could be connected through a cluster of islands
(Kameo and Sato 2000). After this period, when Central and
South America became isolated, this ancestor could have
evolved into N. platensis in South America and C. hyodon
in Central/North America. Between 9.36 and 3.12.8 MyBP,
South America could be isolated (Kameo and Sato 2000),
and N. platensis could reach the eastern lowlands before the
Colombian Andes became a biogeographical barrier (less
than 2,000 m at 4 MyBP, Gregory-Wodzicki 2000) and to
reach the South American western lowlands, at least at the
early Pleistocene (the second oldest gomphothere record in
South America, 2.5 MyBP, from Argentina). In turn, C.
hyodon was not present in South America until after the
connection with Central America and the uplift of the
Colombian Andes (2.8 and 2.7 MyBP, respectively; Coates
and Obando 1996; Gregory-Wodzicki 2000), spreading
through an Andean or highland corridor at the middle/late
Pleistocene (Fig. 4b).

The third, fourth, and fifth hypotheses consider Amahua-
catherium peruvium as a valid South American taxon and
those are based on all possible phylogenetic relationships
between A. peruvium, C. hyodon, and N. platensis.
The third hypothesis suggests the clade (Cuvieronius
hyodon (Amahuacatherium peruvium and Notiomastodon
platensis)), which had a common pan-American ancestor
at 10.71 to 9.36 MyBP (Fig. 4c). When Central and South
America became isolated, this ancestor could originate C.
hyodon in Central/North America and the South American
ancestor to the clade (Amahuacatherium peruvium and
Notiomastodon platensis). This divergence event should
occur approximately at 9.5 MyBP (considering the age
of A. peruvium between 9.5 and 9.0 MyBP, Campbell et
al. 2010) and this divergence might be related to the
formation of the Pebas system and the Andes uplift
(Hoorn et al. 2010). The Miocene gomphothere A.
peruvium could be isolated in the southern region of
the Pebas system, while N. platensis might perhaps be
isolated in its northern and western regions, spreading
to western lowlands before the Andes uplift was at its
highest (until 2.7 MyBP) and to eastern lowlands after
the establishment of the Amazon River (approximately
7 MyBP; Hoorn et al. 2010). However, this hypothesis
does not explain when A. peruvium went extinct and if
it dispersed through the South American eastern low-
lands. If this was possible, the gomphothere vertebra remains
from the early Pleistocene of Argentina could be not identified
at the generic level, because it could be a specimen of A.
peruvium or N. platensis.
The fourth possibility suggests the clade (Amahuacatherium
peruvium (Notiomastodon platensis and Cuvieronius hyodon)),
which had a common pan-American ancestor at 10.71 to 9.36
MyBP (Fig. 4d). When Central and South America became
isolated, this ancestor could evolve into A. peruvium in South
America and the Central/North American ancestor to the clade
(Notiomastodon platensis and Cuvieronius hyodon). Another
speciation event in Central or North America could explain the
occurrence of N. platensis and C. hyodon, before 4.9 MyBP
and, later (after 2.8 MyBP), these taxa reached South America
during the GABI. This hypothesis does not explain how N.
platensis crossed the Andes to reach the South American
eastern lowlands in the Pleistocene and the absence of the N.
platensis record in Central and/or North America. Also, it does
not explain when A. peruvium went extinct and its dispersal
through the South American eastern lowlands, raising the same
genus identification problem for the gomphothere vertebra
remains found in the lower Pleistocene of Argentina.
The fifth possibility (Fig. 4e) suggests the clade (Notio-
mastodon platensis (Amahuacatherium peruvium and
Cuvieronius hyodon)). However, to explain this systematic
arrangement, it is necessary to assume several ad hoc hy-
potheses that are not supported: (1) the existence of a
J Mammal Evol
previous 10.71 to 9.36 MyBP land connection between
South and Central America, (2) fast dispersal of gompho-
theres throughout the Americas, and (3) two sympatric
species of gomphotheres living in South America (least-
ways in Colombia and western Amazonian regions of
Brazil) in the middle Miocene (when the speciation
process could have happened) and the record of N.
platensis since the middle Miocene (Fig. 4e). The lack
of a land connection between Central and South America
from 15.83 to 10.71 MyBP (Kameo and Sato 2000) and the
fact that the oldest record of gomphotheres in North America
is from 13.6 MyBP (Lambert 1996) argues against this idea.
We suggest the second and third hypothesis as the
most parsimonious of all hypotheses presented here.
Both depend on the validity of A. peruvium. No matter
what, it suggests the need for a taxonomic revision of
the South American gomphotheres, mainly by the inclu-
sion of A. peruvium. However, all these hypotheses fail
to explain the long temporal gap between the late Mio-
cene and late Pliocene gomphothere records in South
America. In Amazonia itself, this temporal gap extends
through the late Pleistocene (Cozzuol 2006). Thus, a
long part of the proboscidean history in South America
that should had occurred in the northern part of the
continent (nearby Colombian, Venezuelan, and Brazilian
areas) is virtually unknown.
Conclusions
In agreement with several previous studies, our phylogeny
results in a paraphyletic Gomphotheriidae as a whole. Con-
versely, the gomphotheres present in South American
during the Quaternary are grouped in the monophyletic
clade (Notiomastodon platensis and Cuvieronius hyo-
don). This clade is characterized by the presence of
dorsally curved upper tusks in lateral view, in agreement
with Madden (1984), Ferretti (2008a, 2010), and Lucas
and Alvarado (2010). The genus Stegomastodon appears
as the basal member of a clade characterized by the
presence of m2 with four lophids and M2 with four
lophs or more. Biogeographically, several alternative
hypotheses were suggested to explain the presence of
gomphotheres in South America. However, the most
parsimonious hypotheses depends on a diversification
of a South or pan-American ancestor in the Amazonian
region during the late Miocene. In this way, a thorough
taxonomic analysis of Amahuacatherium peruvium and
also the quest for new proboscidean remains from late Mio-
cene/Pliocene deposits in South America are crucial to fill the
temporal gap in the biogeography history of the Gomphother-
iidae in the Americas.
J Mammal Evol
Acknowledgments The authors are grateful to C. Cartelle (Pontifcia
Universidade Catlica de Minas Gerais, Brazil), R. Machado (Museu
de Cincias da Terra/Departamento Nacional de Produo Mineral,
Brazil), A. M. Ribeiro (Fundao Zoobotnica do Rio Grande do
Sul, Brazil), J. Pereira (Museu Coronel Tancredo Fernandes de Mello,
Brazil), N. Guidon and R. Casati (Fundao Museu do Homem Amer-
icano, Brazil), A. Rojas (Faculdad de Ciencias, Universidad de La
Republica, Uruguay), J. L. Romn-Carrin (Escuela Politecnica
Nacional, Ecuador), J. E. A. Mosquera (Museo Geolgico Nacional Jos
Royo y Gmez, Instituto Colombiano de Geologa y Minera, Colombia),
M. Reguero (Museo de La Plata, Argentina), A. Kramarz (Museo
Argentino de Ciencias Naturales, Argentina), P. Tassy (Musum
National dHistoire Naturelle, France), B. Macfadden and R. Hulbert
(Florida Museum of Natural History, USA), and J. Galkin (American
Museum of Natural History, USA) for allowing the access to the gom-
phothere collections that supported this study. Conselho Nacional de
Desenvolvimento Cientfico e Tecnolgico (CNPq) granted master schol-
arship fund to DM (process number 134905/2010-5).
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