Journal of Vertebrate Paleontology 27(3):744–747, September 2007

© 2007 by the Society of Vertebrate Paleontology

SHORT COMMUNICATION

A NEW MYLODONTINAE (MAMMALIA, XENARTHRA) FROM THE CAMACHO

FORMATION (LATE MIOCENE), URUGUAY

ANDRÉS RINDERKNECHT1, DANIEL PEREA*,1, and H. GREGORY MCDONALD2; 1Área de Geología y Paleontología,
Facultad de Ciencias. Iguá 4225, CP 11400. Montevideo, Uruguay, perea@fcien.edu.uy; 2 Park Museum Management Program,
National Park Service 1201 Oakridge Drive, Suite 150 Fort Collins, Colorado 80525 USA., Greg_McDonald@nps.gov

The xenarthrans of the sloth subfamily Mylodontinae are con-
spicuous members of South and North American Neogene fau-
nas. They occupied a large latitudinal range from central North
America to Tierra del Fuego in the austral extreme of South
America (Latorre, 1998). The first undoubted representative of
the subfamily is Glossotheriopsis pascualli, from the Friasian
SALMA (middle Miocene) of Argentina. Beginning in the late
Miocene (Huayquerian SALMA) the mylodontines are repre-
sented by numerous taxa that exhibit a range of morphological
heterogeneity (Esteban, 1988) in contrast to the other subfami-
lies.
Most of the mylodontine taxa described for the Neogene are
based on very poor or fragmentary material making a compre-
hensive comparative description of the genera within the sub-
family difficult. The best and most useful single bone that can be
used to characterize mylodontid species is the mandible as it is
commonly preserved and contains a complex suite of characters
that makes it systematically informative (Perea, 1992). It is for
these reasons that the majority of the Neogene sloth taxa are
based on mandibular remains as they do allow an adequate mor-
phologic comparison and subsequent phylogenetic conclusions
that document the diversity within this subfamily of sloths. In this
work we describe a right mandibular ramus originating from the
Camacho Formation at Kiyú beach, Department of San José,
Uruguay (Fig. 1). While the fossil shows some affinities with
Pleistocene genera it also shares characters with older mylodon-
tines as well. On the basis of its distinctive suite of morphological
features it can be distinguished from all other known mandibular
morphologies within the subfamily, indicating it represents a new
genus and species.
The material described here is deposited at the Paleontologi-
cal Collection of Vertebrates, Facultad de Ciencias, Uruguay
(FCDPV).
Formation at this locality (considered the Kiyú Formation by
Francis and Mones, 1965 and the San Pedro Member of Cama-
cho Formation by Perea and Martínez, 2004) represent a facies
of a late Miocene transgressive episode. The facies includes oys-
ter patch reefs and ichnofossils (Ophiomorpha nodosa and
Thalassinoides sp.) in a greenish-gray clayey silt (Sprechmann et
al., 1998, 2000) and abundant terrestrial mammals (Perea, 2005).
The fossil mammal assemblage of the Camacho Formation is
represented by taxa of Huayquerian-“Mesopotamian” affinities
(Perea et al., 1994; Perea, 2005), including an assemblage zone
characterized by Pseudoplohophorus absolutus and Cardia-
therium orientalis and the following associated fauna: Proeu-
phractus limpidus, Kraglievichia paranense, Stromaphoropsis
scavinoi, Pseudoplohophorus benvenutii, P. rebuffoi, Berthawyle-
ria gracilis, Pronothrotherium mirabilis, Pliomorphus ameghinoi,
Ranculcus, Anchimys, Isostilomys intermedius, Lagostomopsis,
Dinotoxodon paranensis, Toxodontherium, Xotodon, Scalabri-
nitherium and Saurocetes argentinus. The presence of a terres-
trial fauna (including some partially articulated specimens) in-
termixed with estuarine and marine ostreids and fish remains,
freshwater turtles, and flamingoes is considered to be the result
of transport into a paralic environment perhaps including coastal
lagoons and flood plain in an estuarine or deltaic system (Ubilla
et al., 1990; Perea et al., 1996; Perea, 2005). The new material
here described was found embedded in a gray-greenish sandy

GEOLOGICAL SETTING, BIOSTRATIGRAPHY

AND PALEOENVIRONMENTS

Three lithostratigraphic units can be recognized in the out-

crops exposed at Kiyú. From the base to the top of the section
these include the Camacho (late Miocene), San José (Plio-
Pleistocene), and Libertad (Pleistocene) formations. The San
José Formation was described by Francis and Mones (1965) and
is also called the Raigón Formation (Goso and Bossi, 1966) in
most of the geological literature. The outcrops of the Camacho

FIGURE 1. Geographic location of Kiyumylodon lecuonai, gen. et sp.

nov., in Department of San José. The arrow shows the approximate site
*
Corresponding author. of collection.

744
 SHORT COMMUNICATIONS 745

pelite in the San Pedro Member of the Camacho Formation, late

Miocene (Perea and Martínez, 2004).

SYSTEMATIC PALEONTOLOGY

XENARTHRA Cope, 1889

MYLODONTIDAE Gill, 1872
MYLODONTINAE Gill, 1872
KIYUMYLODON, gen. nov.
Etymology—Kiyu, for the geographic locality in the Depart-
ment of San José where the type material was collected; mylodon
for the family Mylodontidae.
Age—Late Miocene, Huayquerian SALMA.
Type Species— Kiyumylodon lecuonai, gen. et. sp. nov.
Diagnosis—As for the type and only species (monotypic ge-
nus).

KIYUMYLODON LECUONAI, sp. nov.

(Fig. 2)
Etymology—lecuonai, in dedication to Gustavo Ramos Le-
cuona, designer, sculptor and one of the collectors of the type
material.
Holotype—FCDPV 1829, almost complete right mandibular
ramus including a full dentition and well-preserved coronoid,
articular, and angular processes.
Locality—The material was collected by Gustavo Lecuona
and one of the authors (A.R.) on the Kiyú beach coastal plat-
form, approximately 1 km east of the outlet of San Gregorio
creek in the Department of San José, Uruguay (34° 44⬘ S,
56° 50⬘ W).
Horizon—The material was recovered from a gray-greenish
sandy pelite in the San Pedro member of the Camacho Forma-
tion (late Miocene, Huayquerian SALMA (Perea and Martínez,
2004), at approximately sea level.
Diagnosis—Size slightly smaller to Glossotherium robustum;
c1 (caniniform) small and displaced slightly labially relative to
the molariforms; diastema absent; m1 and m2 are subelliptic in
cross section, without grooves or lobation; m3 bilobate, of me-
dium length and possessing a well demarcated isthmus between
the anterior and posterior lobes; large and shovel-like mandibu-
lar symphysis.
DESCRIPTION
The mandibular ramus is well preserved almost complete (Fig.
2). Although it is broken at the symphysis, it is possible to re-
construct its original shovel-like form because it shows an open
curve superior border anterior to the first tooth (Fig. 2B). The
mandibular condyle projects internally. The coronoid and angu-
lar processes (apophyses) project posteriorly. The latter extends
posterior to the condyle and has well-defined longitudinal ridges
on its internal surface (Fig. 2C).
The first tooth (caniniform) is the smallest of the dental series;
it has a subelliptical cross section and the occlusal surface is
wedge-like resulting from wear. The occlusal surface forms at an
acute angle relative to the long axis of the tooth and is best seen
in lateral view. This tooth projects slightly to the labial side and
its alveolus is also shifted slightly labially relative to the longitu-
dinal axes of the other teeth. The labial position of the tooth in
Kiyumylodon is similar to that of Glossotherium and is not as
pronounced as in Megabradys and Ranculcus (the latter also
present in the Camacho Formation); it also differs from Spheno-
therus in which both the caniniform and first molariform are
labially positioned relative to the other teeth.
The second tooth (first molariform) is separated from the ca-
niniform by a shorter distance than the anteroposterior length of
the later. There is no distinct diastema. The tooth is subcircular
to slightly subtriangular in cross section. The third tooth (second
FIGURE 2. Type material of Kiyumylodon lecuonai, gen. et sp. nov.,
(FCDPV 1829) in A, external; B, occlusal; and C, internal views. Ab-
breviations: c1, caniniform; m1-m3, molariforms.
molariform) is larger than the preceding tooth and has a subel-
liptic to slightly quadrangular outline. The major axes of the
occlusal surfaces of the caniniform and first two molariforms are
oblique in relation to the longitudinal axis of the molariform
series.
The last tooth (third molariform) is bilobate. Its anterior lobe
is compressed longitudinally so it shows a roughly quadrangular
cross section. The posterior lobe of this molariform has a sub-
circular cross section with a small labial projection of its internal
border. A short, broad, well-defined isthmus separates both
lobes. All molariforms show marked occlusal wear, with con-
spicuous holes in the central region.
DISCUSSION
Two subfamilies traditionally are recognized within the Mylo-
dontidae: Scelidotheriinae and Mylodontinae. McKenna and
Bell (1997) raised these to the family level and recognized two
subfamilies in each: Chubutheriinae and Scelidotheriinae in the
former and Mylodontinae and Lestodontinae in the latter. The
Miocene genus Urumacotherium (⳱ Acremylodon), originally
considered a mylodontine, recently was transferred by Negri and
Ferigolo (2004) to a new mylodontid subfamily, Urumacotherii-
nae. We agree with this systematic proposal and here recognize
a single family, Mylodontidae, with four subfamilies: Mylodon-
tinae, Scelidotheriinae, Urumacotheriinae, and Octomylodon-
tinae. All members of the family are characterized by a bilobate
last upper and lower molariform (see Scillato-Yané, 1977).
The material described here is assigned to the Subfamily My-
lodontinae based on the large incisive border (mandibular spout)
746 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 27, NO. 3, 2007

the late Pleistocene genus Glossotherium, but m1–2 have much

FIGURE 3. Occlusal views (not to scale) of right mandibular rami of
mylodontines: A, Ocnotherium; B, Lestodon; C, Megabradys; D, Ran-
culcus; E, Mylodon; F, Glossotherium; G, Sphenotherus; H, Pseudopre-
potherium; I, Thinobadistes; J, Nematherium; K, Kiyumylodon, gen. nov.;
L, Brievabradys; M, Pleurolestodon.
more rounded cross sections reminiscent of the Miocene Brie-
vabradys and Pseudoprepotherium and Quaternary mylodon-
tines, Lestodon, Ocnotherium and Mylodon; like all of them, the
condition in Kiyumylodon probably represents a secondary re-
duction. The isthmus of m3 is clearly shorter than in Glossoth-
erium, and in this regard also resembles Lestodon and Mylodon.
Kiyumylodon, gen. nov., can also be distinguished from other
Miocene mylodontines (Fig. 3). In Sphenotherus, the molariform
series has a strong labial concavity that distinguishes it from the
other mylodontines—including Kiyumylodon—in which the den-
tal series is straight. Kiyumylodon gen. nov. differs from Mega-
bradys, Ranculcus, Lestodon and Ocnotherium in lacking a dia-
stema. It also differs from Lestodon and Ocnotherium in lacking
molariforms that are markedly angled to the axis of the tooth
row (Ameghino, 1891; Scillato-Yané, 1981; Perea and Scillato-
Yané, 1990). It is also distinguished from Ranculcus in having a
shorter m3.
The similarity in size among the caniniform (c1) and m1–2
distinguishes this new taxon from Lestodon, Sphenotherus,
Megabradys, Pleurolestodon, Thinobadistes and Brievabradys; in
these other taxa, the caniniform is larger than the second and
third molariforms (Ameghino, 1889; Scillato-Yané 1981; Webb,
1989; Villarroel, 2000) (Fig. 3).
Kiyumylodon shares simplified molariforms (i.e., reduced lo-
bation) with Lestodon, Ocnotherium, Mylodon, Brievabradys
and Pseudoprepotherium, all of which have molariforms with
circular or elliptical cross sections (Lund, 1842; Ameghino, 1889;
Kraglievich, 1928; Hirschfeld, 1985; Villarroel, 2000).
The above combination of characters in the specimen demon-
strates a new morphologic mandibular pattern not previously
observed within the diversity of the Mylodontinae, and as such
we propose that it represents a new genus and species.
Acknowledgments—We wish to thank G. R. Lecuona for his
help in the field work and M. Ubilla for his useful comments.

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Submitted February 15, 2007; accepted April 1, 2007.