Journal of Vertebrate Paleontology 29(3):870–880, September 2009

# 2009 by the Society of Vertebrate Paleontology

ARTICLE

PREDOMINANCE OF ORTHAL MASTICATORY MOVEMENTS IN THE EARLY

MIOCENE EUCHOLAEOPS (MAMMALIA, XENARTHRA, TARDIGRADA,
MEGALONYCHIDAE) AND OTHER MEGATHERIOID SLOTHS

M. SUSANA BARGO,*,1 SERGIO F. VIZCAÍNO1 and RICHARD F. KAY2

1
División Paleontologı́a Vertebrados, Museo de La Plata, Paseo del Bosque s/n, B1900FWA La Plata,
Argentina, CIC and CONICET, msbargo@fcnym.unlp.edu.ar, vizcaino@fcnym.unlp.edu.ar;
2
Department of Evolutionary Anthropology, Duke University, Box 90383 Science Drive,
Durham NC 27708, U.S.A., richard.kay@duke.edu

ABSTRACT—The megatherioid sloths from the Santa Cruz Formation (Santacrucian Age; early-middle Miocene,
Patagonia, southernmost South America) occupy basal positions in the most recent phylogenetic schemes. The cranial
morphology of Eucholaeops, particularly of the teeth, suggests interesting functional features that shed light on the type
of food it was capable of processing, and thus on the diet. A detailed morphofunctional analysis of the jaw apparatus was
performed, and the results briefly compared with other contemporary megatherioid sloths. Comprehensive descriptions
of the teeth of Eucholaeops allow us to generate a nomenclature for describing the inferred occlusal pattern analogous to
that applied to other mammals. Based on examination and mapping of occlusal wear facets, we reconstruct two distinct
jaw movements during the power stroke. One corresponds to the basic therian pattern equivalent to Phase I: the working
side mandibular corpus is moved dorsally, mainly orthally but also anteriorly and slightly medially; the result is punctur-
ing, tearing and shearing of food. The second is a distinct and unrelated movement of the working side corpus dorsally,
mainly orthally, but also posteriorly and slightly medially; the dominant result is to produce shearing of food. The analysis
of the tooth wear facets, combined with the shape of the temporomandibular joint, the presence of a fused mandibular
symphysis, and a well-developed temporalis muscle, indicates that the orthal component was predominant during masti-
cation. Eucholaeops, and probably nearly all other Miocene megatherioids, were most likely leaf eaters and the primary
method of food reduction must have been by shearing or cutting.

INTRODUCTION view, considering that Bradypus differentiated through a process

of heterochrony within Megalonychidae. However, Gaudin’s
Sloths (Xenarthra: Tardigrada or Phyllophaga; see Fariña and
results are congruent with those of Patterson and Pascual, Webb,
Vizcaı́no, 2003; McKenna et al., 2006; Vizcaı́no and Loughry,
and Carlini and Scillato-Yané in the placement of Choloepus
2008 for further discussion on this matter) are known since the
within the family Megalonychidae, a clade that includes the
late Eocene of Antarctica (Vizcaı́no and Scillato-Yané, 1995).
extinct Antillean sloths. Although the relationships among tree
Discounting some dubious Eocene records (Simpson, 1948), the
sloths and ground sloths supported by Gaudin (2004) differs
oldest unquestionable South American Tardigrada are from
from those advocated in other recent morphological analyses
Deseadan beds (late Oligocene) of Patagonia (Hoffstetter,
(White and MacPhee, 2001; Pujos et al., 2007) it is notable that
1982) and Bolivia (Engelmann, 1987). They become conspicuous
all of them support the diphyly of tree sloths. Gaudin’s (2004)
elements of the fossil fauna of South America in the early Mio-
phylogeny also corroborates the monophyly of the four extinct
cene (see McDonald and De Iuliis, 2008, for a review).
clades of sloths, the Mylodontidae, Megatheriidae, Nothrother-
Tardigrades include nearly 100 named genera (McKenna and
iidae, and Megalonychidae (Fig. 1). Megatheriids, nothrother-
Bell, 1997), and are one of the largest groups and most charac-
iids, and megalonychids are joined in a monophyletic clade
teristic elements of the Cenozoic fauna of South America. They
Megatherioidea, within which Megatheriidae and Nothrotherii-
were also well represented in North America during the past
dae form a monophyletic group, Megatheria. Megalonychidae
three million years and appear to have dispersed into the Carib-
have been present since the Deseadan Age (late Oligocene;
bean Islands by the Oligocene (White and MacPhee, 2001). In
Carlini and Scillato-Yané, 2004; Pujos et al., 2007).
the last two decades several published phylogenetic studies en-
In the Santacrucian Age (18-15 Ma, early to middle Mio-
compass restricted subsets of taxa or characters (for a review see
cene; Fleagle et al., 1995) sloths are represented by a diversity
Gaudin, 2004). The most comprehensive recent morphology-
of small to medium sized forms proposed to have been mostly
based investigation is that of Gaudin (2004). Gaudin’s analysis
arboreal to semiarboreal (White, 1993, 1997; McDonald and De
strongly supports the diphyly of the living tree sloths (Bradypus
Iuliis, 2008), and generally folivorous (Scillato-Yané, 1986) al-
Linnaeus, 1758, and Choloepus Illiger, 1811), positioning Brady-
though without specific studies on the dietary habits. Scott
pus as the sister-taxon to all other sloths, in contrast to the
(1903-04) classified them as Megalonychidae, Planopsidae, and
conclusions of Patterson and Pascual (1972) and Webb (1985),
Mylodontidae. Unfortunately, despite an abundance of remains,
where Bradypus was considered closely allied with the mega-
little has been accomplished over the last century in terms of
theres. Carlini and Scillato-Yané (2004) proposed a different
taxonomy (see McDonald and De Iuliis, 2008). Although a re-
view is clearly required, most Santacrucian genera are well-
identifiable taxonomic units (De Iuliis, 1994; Gaudin, 1995,
*
Corresponding author. 2004; Gaudin and McDonald, 2008; McDonald and De Iuliis,

870
 BARGO ET AL.—MASTICATION IN EARLY MIOCENE SLOTHS
FIGURE 1. Phylogeny of Tardigrada (after Gaudin, 2004).
2008). Scott’s (1903-04) concept of megalonychids included
Eucholaeops Ameghino, 1887 (Fig. 2) (Eucholoeops Ameghino,
1889:692 emend. illegit. pro., Mones, 1986), Megalonychotherium
Scott, 1904, Hapalops Ameghino, 1887 (Fig. 3), Analcimorphus
Ameghino, 1891, Pelecyodon Ameghino, 1891, Schismotherium
Ameghino, 1887, and Hyperleptus Ameghino, 1891 (see Simpson,
1945; Hoffstetter, 1958; Paula Couto, 1979; Patterson et al., 1992;
Gaudin, 1995). Following Gaudin (2004) Eucholaeops is a basal
Megalonychidae, whereas Hapalops, Analcimorphus, Pelecyodon,
Schismotherium and Hyperleptus are a paraphyletic group basal
to Megatherioidea, and with uncertain relationships to other
megatherioids. Scott’s Planopsidae (Planops Ameghino, 1887
and Prepotherium Ameghino, 1891) are regarded by Gaudin as
Megatheriidae (in accordance with De Iuliis, 1994 and Gaudin,
1995). Finally, Gaudin’s Mylodontidae (Nematherium Ameghino,
1887 and Analcitherium Ameghino, 1891) are placed as a sis-
ter group of all remaining mylodontids (contra other, less compre-
hensive, recent phylogenetic studies, e.g. McDonald and Perea,
2002) (Fig. 1).
871
Ameghino (1894) and Scott (1903-04) summarize the main
features of the skull of Santacrucian sloths (see also McDonald
and De Iuliis, 2008). Despite a range of morphological variabil-
ity, they are characterized by elongated, tubular skulls (Figs. 2A,
3A), curiously small in proportion to body size compared with
other mammals. The occipital and sagittal crests are poorly de-
veloped or absent. The orbit lacks a postorbital process or bar
and is widely confluent posteriorly with the temporal fossa. The
jugal possesses ascending and descending processes. It is loosely
attached to the maxilla and does not articulate with the squamo-
sal (Fig. 2C). The premaxillae are separate from each other and
from the maxillae, and are so loosely attached that they are
rarely found in association with the skull (Figs. 2D, 3B). The
rami of the mandible are coossified at the symphysis, which is
extended in a more or less elongate edentulous spout (Figs. 2F,
3C). The alveolar portion of the mandibular corpus is thick and
massive. The mandibular condyle is hemispherical and is set
upon a well marked neck. The angular process is more or less
inflected, and in megalonychids forms a curved hook, longer and
more prominent than in the other clades. The teeth are hypselo-
dont and lack enamel. The dental formula is 5/4. All the upper
teeth are rooted in the maxilla and all teeth are separated by
diastemata. The first upper and lower teeth are caniniform, usu-
ally cylindrical or triangular in section and quite small, except
in Eucholaoeps in which the caniniforms are enlarged and
projecting (Fig. 2B, E). The upper caniniform is positioned at
the anterior end of the maxilla adjacent to the contact with the
premaxilla, and in occlusion it passes in front of the lower canini-
form (Fig. 3A). Megatherioid molariforms are transversely oval
to rectangular, with two crests of hard dentine separated by a
deep valley excavated in the soft dentine.
As already noted by Scott (1903-04), specimens of Eucho-
laeops are much less common in the Santacrucian beds than
those of Hapalops, and there are fewer named species: four in
Eucholaeops—E. fronto Ameghino, 1891, E. ingens Ameghino,
1887, E. externus Ameghino, 1891 and E. curtus Ameghino,
1894—although Mones (1986) listed up to twelve Miocene spe-
cies compared with twenty-six in Hapalops. Scott (1903-04)
notes that Eucholaeops have large caniniforms, usually pointed
by abrasion (Fig. 2E), placed external to the line of the other
teeth and with differences in size ascribed by Scott to the proba-
ble existence of sexual dimorphism. Compared with that of Hap-
alops (Fig. 3) the skull is proportionally broader and heavier; the
sagittal and occipital crests are much better developed and in
some species are very prominent; the muzzle is very broad,
expanding anteriorly; the facial portion of the maxilla does not
continue beyond the caniniform, unlike in almost all other
sloths, but forms a rounded, pillar-like structure that housed the
alveolus for this tooth (Fig. 2B). The mandible is very character-
istic: the corpus is short and deep, very thick and massive. The
alveolus of the caniniform is also housed in a prominent pillar-
like structure that projects vestibularly; a large notch anterior to
this pillar receives the upper caniniform (Fig. 2F). The shape of
the mandibular predental spout is variable, but is short and
terminates bluntly. The symphysis is short and steep; the ramus
is well developed with larger coronoid and angular processes
than in Hapalops.
Eucholaeops weighed about 80 kg while Hapalops ranged be-
tween 50 and 70 kg. These estimates were calculated on humerus
lengths, using allometric equations of Scott (1990). The humerus
was selected because it is the limb element that produces more
consistent estimations with other methods in ground sloths
(scale and geometric models; Fariña et al., 1998; Bargo et al.,
2000; De Iuliis et al., 2004). Other estimates for Hapalops (spe-
cifically H. longiceps), based on femoral head diameter, generate
body masses of up to 40 kg (White, 1997).
Beyond their importance for systematics and phylogeny, the
cranial morphology of Eucholaeops, and particularly the teeth,
872 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 29, NO. 3, 2009

suggests interesting functional features of the jaw apparatus that localities of Santa Cruz Province, Argentina, has recovered very
could shed light on the diet. In addition to the specimens housed complete and well preserved new specimens (Fig. 2). This mate-
in collections for over one century that were the basis of detailed rial has allowed us to perform a detailed analysis of the tooth
descriptions by Scott (1903–04), recent field work in the coastal morphology, and to infer the occlusion patterns and masticatory

FIGURE 2. Skull of Eucholaeops sp. (MPM PV 3401). A, lateral view; B, occlusal view of palate; C, left jugal in medial view; D, premaxillae in
occlusal view; E, right caniniform; F, mandible in lateral view; G, lower tooth series in occlusal view; H, detail of tooth morphology of the mandible in
lateral view. Scale bar equals 5 cm in A-G, 3 cm in H.
 BARGO ET AL.—MASTICATION IN EARLY MIOCENE SLOTHS
FIGURE 3. Skull and mandible of Hapalops longiceps (YPM-PU
15523). A, lateral view; B, palatal view of skull; C, occlusal view of
mandible. Scale bar equals 5 cm.
movements of Eucholaeops. This information leads us to pro-
pose the type (properties) of food these sloths were capable of
processing, and hence to infer a possible diet. The results of this
morphofunctional analysis of Eucholaeops are briefly compared
with other contemporary sloths, and the major lineages pro-
posed to be derived from them (see Scott, 1903-04; Gaudin,
2004; McDonald and De Iuliis, 2008).
Our study of the Eucholaeopos masticatory apparatus was
undertaken in the context of several earlier studies of living and
some fossil sloths. Naples (1982, 1985) described and analyzed
the cranial osteology of living tree sloths and particularly their
masticatory apparatus. Based in this work, this author analyzed
the morphology and function of the feeding apparatus of the
Pleistocene North American nothrothere Nothrotheriops shas-
tense (Sinclair, 1905), and the mylodontid Paramylodon harlani
(Owen, 1843) (Naples, 1987, 1989, respectively). Bargo (2001)
and Bargo and Vizcaı́no (2008) studied the masticatory appa-
ratus of the large South American Pleistocene megatheriid
Megatherium americanum Cuvier, 1796, and the mylodontids
Glossotherium robustum (Owen, 1842), Lestodon armatus Ger-
vais, 1855, Mylodon darwini Owen, 1839 and Scelidotherium
leptocephalum Owen, 1840. Finally, Muizon et al. (2004) ana-
lyzed the correlation of morphology and feeding strategies of
five species of aquatic sloths, Thalassocnus Muizon and McDo-
nald, 1995, from the late Miocene to the late Pliocene of Peru.
873
Institutional Abbreviations—MACN, Museo Argentino de
Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires,
Argentina; MLP, Museo de La Plata, La Plata, Argentina;
MPM-PV, Museo Regional Provincial Padre “M. J. Molina,”
Rı́o Gallegos, Santa Cruz, Argentina; YPM-PU, Yale Peabody
Museum (Princeton University collection), New Haven, USA.
MATERIAL AND METHODS
Specimens Examined
MACN-A 11614, Eucholaeops ingens. Left half of skull and
left dentary. Horizon and locality: Santa Cruz Formation, Santa
Cruz province, Argentina.
MPM-PV 3401, Eucholaeops sp. Skull and mandible. Horizon
and locality: Santa Cruz Formation, Estancia La Costa Member,
Puesto Estancia La Costa (= Corriguen Aike), Santa Cruz prov-
ince, Argentina.
MPM-PV 3403, Eucholaeops sp. Skull, mandible and postcra-
nial skeleton. Horizon and locality: Santa Cruz Formation,
Estancia La Costa Member, Puesto Estancia La Costa (= Cor-
riguen Aike), Santa Cruz province, Argentina.
YPM-PU 15314, Eucholaeops fronto. Mandible. Horizon and
locality: Santa Cruz Formation, 10 miles South of Coy Inlet,
Santa Cruz province, Argentina.
YPM-PU 15523, Hapalops longiceps. Holotype. Skull, mandi-
ble and postcranial skeleton. Horizon and locality: Santa Cruz
Formation, 8 miles south of Coy Inlet, Santa Cruz province,
Argentina.
YPM-PU 15597, Hapalops elongatus. Skull and mandible. Ho-
rizon and locality: Santa Cruz Formation, Killik Aike, Santa
Cruz province, Argentina.
YPM-PU 18009, Nematherium sp. Skull and mandible. Hori-
zon and locality: Santa Cruz Formation, Santa Cruz province,
Argentina.
YPM-PU 15346, Planops magnus. Holotype. Skull. Horizon
and locality: Santa Cruz Formation, Guer Aike Department,
Santa Cruz province, Argentina.
Methods
Despite the descriptions available in Scott (1903-04), we pro-
vide our own descriptions of the teeth in order to generate a
nomenclature that allow us to describe the inferred occlusal
pattern in a way analogous to that frequently applied to those
of other mammals with clearly tribosphenic teeth. Having done
this, the occlusal movements are inferred through the analysis of
the wear facets and the manipulation of casts of the upper and
lower tooth series.
RESULTS
Descriptive Anatomy
This section deals with the description of those features of the
cranium of Eucholaeops that are relevant for the analysis of
occlusion and masticatory movements (see Fig. 2).
The temporomandibular joint (TMJ) is located slightly dorsal
to the occlusal plane. The glenoid fossa is a shallow depression
on the squamosal process (Fig. 2B) that allows the mandi-
bular condyle great freedom of anteroposterior motion. The
mandibular condyle is hemispherical, wider mediolaterally than
anteroposteriorly, and supported by a long, slender neck. No
postglenoid process borders the articular surface of the squamo-
sal posteriorly.
A distinctive feature of Eucholaeops is its first tooth, which is
a large caniniform, usually sharply pointed by abrasion. The
upper canine has an orthally oriented wear facet on its distal
face for occlusion with a mesially-facing facet on the projecting
874 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 29, NO. 3, 2009
lower canine (Fig. 2H). Caniniforms are placed externally to the
line of the cheek teeth, both in the upper and lower jaw, in a
pillar-like structure that projects outwards. In front of this struc-
ture, in the lower jaw, a deep notch receives the upper canini-
form. The broad and edentulous muzzle of Eucholaeops suggests
a well-developed upper lip complementing a mobile tongue in
oral food acquisition, as was inferred also for some Pleistocene
ground sloths (Bargo et al., 2006). Caniniforms are separated
from the first molariforms by relatively long diastemata.
Eucholaeops has four maxillary and three mandibular cheek
teeth. The teeth are isognathic, but occlusion appears to have
occurred on one side of the mandible at a time as in living sloths
(Naples, 1982). As in living sloths (e.g., Choloepus; Naples, 1982)
the teeth occlude alternately such that the first lower molariform
(here termed m1) occludes between and against the first and
second upper molariforms (M1 and M2); m2 and m3 have the
same relationship with M2–3 and M3–4, respectively (Fig. 4A).
The occlusal details and proportions of the teeth vary from
m1 to m3 (and in the upper teeth) but the basic pattern of
occlusion and the inferred occlusal relationships remains essen-
tially the same. We describe the anatomy and occlusion between
m2 and the corresponding M2–3 as representing molariform
occlusion as a whole.
M2 and M3 are short mesiodistally and broad vestibulolin-
gually. Each tooth possesses three cusps (Fig. 4). Cusp A is
positioned mesiolingually and is very sharp and projecting. A
second mesiovestibular cusp, Cusp B, is equally prominent but
more cristiform. A sharp mesial transverse loph connects Cusps
A and B. The loph has two components: a strong transverse
crest leads vestibularly from Cusp A to join a crest leading
buccally from Cusp B. The mesial loph is very sharp and con-
cave in shape with its most dorsal point in the midline of the
tooth. A second crest arises from Cusp A extends distobuccally
a short distance to the base of the crown. Likewise Cusp B
supports a second short crest running distally to the distobasal
margin of the crown.
A poorly defined distal cusp (Cusp C) is positioned near to the
midline mesiodistal axis of the tooth (Fig. 4B). Cusp C is highly
cristiform, being the midline support for a convex transverse
trenchant distal loph. The loph may be thought of as being
composed of two crests, one running vestibularly and the other
lingually from Cusp C.
A basin or sulcus (indicated by dashed lines in Fig. 4B) lies
between the two transverse lophs. The basin begins vestibularly
and runs transversely to obtain its deepest (most dorsal) point
just lingual to the midline, then runs further lingually and slightly
distally to the lingual margin of the tooth.
The lower molariforms are almost a mirror image of the
uppers. As exemplified by m2, the teeth are short mesiodistally
and broad vestibulolingually. Each tooth possesses three cusps
and two transverse lophs. The more distal loph is anchored at
each end by Cuspids A and B. Distolingually positioned Cuspid
A is sharp and projecting. A distovestibular Cuspid B is equally
prominent but more cristiform. The distal loph has two compo-
nents: a strong transverse cristid leads vestibularly from Cuspid
A to join a crest leading distovestibularly from Cuspid B. The
distal loph is very sharp and concave with the most ventral point
buccal to the midline of the tooth. A second cristid arising from
Cusp A runs mesially and basally a short distance before becom-
ing confluent with a cristid arising from Cuspid C (see below).
Distovestibularly-positioned Cuspid B, as already mentioned,
serves as the anchor for a transverse crest. The cusp also sup-
ports a short cristid running mesially to the distovestibular mar-
gin of the crown.
A poorly defined mesial cusp (Cuspid C) is positioned slightly
vestibular to the midline mesiodistal axis of m2. Cuspid C is
highly cristiform, supporting two shorter but trenchant convex
transverse buccal and lingual crests that collectively form a me-
sial loph. The former terminates at the mesiobuccal corner of the
tooth; the latter cristid is confluent with the cristid running mesi-
ally from Cuspid A, walling off the lingual margin of the trans-
versely elongate central basin.
Between the mesial and distal lophs is a basin or sulcus
(dashed lines, Fig. 4B). The basin begins lingually at the wall
where Cuspid B joins the cristid running vestibularly from Cus-
pid C. This sulcus runs transversely to obtain its deepest (most
ventral) point vestibularly at the sulcus between the cristid run-
ning mesially from Cuspid A and the cristid running vestibularly
from Cuspid C.
Masticatory Movements
As related below, a mammalian masticatory cycle typically
consists of three succeeding ‘strokes’: jaw closing, power, and
jaw opening. Because teeth and food are in contact only during
the power stroke, and because the contact produces wear pat-
terns on the teeth, we can reconstruct the details of jaw move-
ment patterns only during the power stroke. Based upon scale
drawings and examination and mapping of occlusal wear facets
(Fig. 4), we reconstruct the following masticatory movements
during the power stroke of Eucholaeops.
In Movement A, the working side mandibular corpus is
moved upwards (orthally), mesially, and slightly lingually. The
movement engages the leading edges of mesial-facing surfaces
of upper molariform transverse lophs against the distal-facing
surfaces of the lower transverse lophs. The facets produced by
this movement are indicated as b and b’ in Figure 4. The
anterior transverse loph of the upper molariforms is concave
as is the lower distal transverse loph with which it occludes.
Contact begins at two points, Cusps A and B (and Cuspids A
and B). As occlusion continues, the two points of contact along
the crests converge such that food was trapped and sheared in
a lozenge-shaped space (see Fig. 4A). The posterior transverse
loph of the upper molariform is convex whilst the lower mesial
transverse loph with which it occludes is concave. In spite of
this, because of slightly different angulations of the leading
edges, contacts are maintained at two points along the crests
beginning at the central lower Cuspid C and moving vestibu-
larly and lingually, tearing any food punctured and held by the
initial penetration as Cusp C and Cuspid C move past one
another. Movement A results in the puncturing, tearing, and
shearing of food.
Movement B is a second distinct and unrelated movement of
the working side corpus upwards, distally, and slightly lingually.
The movement engages the leading edges of distal-facing sur-
faces of the same upper molariform transverse lophs against the
mesial-facing surfaces of the lower molariform transverse lophs.
The wear facets produced by this movement are indicated a, a’,
and a’’ in Figure 4. While the same transverse crests are engaged
in occlusion in two mutually exclusive ways, in each case the
dominant result is to produce shearing of interposed food. As
discussed below, Muizon et al. (2004) identified the two sets of
facets we here identify but came to a different interpretation
about their significance.
DISCUSSION
Jaw Movements and Wear Patterns in Sloth Mastication
Mastication in eutherian and metatherian mammals is de-
scribed through chewing cycles and, in general, three compo-
nents or strokes can be distinguished in a single cycle (Crompton
and Hiiemae, 1969a, 1969b; Hiiemae and Kay, 1972, 1973; Kay
and Hiiemae, 1974; Hiiemae, 1978; Hiiemae and Crompton, 1985):
1. Closing stroke or fast close phase: predominantly orthal; brings
the lower molars into alignment with the uppers in preparation
 BARGO ET AL.—MASTICATION IN EARLY MIOCENE SLOTHS
for the power stroke. Some anteroposterior movement occurs
during the closing stroke, greatest in rodents, less marked in
pigs, clearly discernible in primates.
2. Power stroke: defined as beginning with tooth-food-tooth con-
tact; food breakage occurs during the power stroke as the force
generated by the adductor muscles is applied through the teeth
to the food between them. In most mammals, this stroke has
two ‘phases,’ I and II (Hiiemae and Kay, 1972). The general
direction of Phase I is, without exception, from vestibular to
lingual (lateromedial), upward and variably forward to centric
occlusion. Phase I of the power stroke ends at minimum gape,
but this may not coincide with full occlusion or maximum
intercuspation of teeth. Phase II is an additional movement,
generally in a mesiolingual and slightly downward direction
out of centric occlusion, with the teeth remaining in contact.
3. Opening stroke: the anteromedial element in the power stroke
means that the opening stroke begins with the jaw in a slightly
forward position. In frontal view, the direction can vary in differ-
ent mammals: it can have a significant lingual excursion, or can be
a simple orthal movement in the midline. In lateral view, the
opening stroke incorporates a definite protrusion of the mandible.
Despite the considerable anatomical differences in the jaw
apparatus of mammals so far studied, there appears to be a
consistent pattern in the profile of jaw movement, the most
important aspect of which is the consistency in the dorsal, mesi-
al, and lingual movements of the dentary during the initial part
of the power stroke (Phase I). The extent of lingual movement
varies considerably from very little in carnivores with carnassials
to large lingual excursions in extant browsing artiodactyls
(Crompton and Hiiemae, 1969b). An added element in some
taxa is the incorporation of the initial part of the opening stroke
into the power stroke as Phase II (Hiiemae and Kay, 1972).
As noted above, two different and mutually exclusive power
stroke movements have been identify in Eucholaeops. Move-
ment A, a mainly orthal and slightly medial and anterior move-
ment, most closely resembles Phase I of the power stroke in
mammals with tribosphenic molars or molars with cusp patterns
easily derived therefrom. A separate power stroke of the type
described above as Movement B has no obvious counterpart in
other Mammalia. Nor is there any indication for the power
stroke continuing after centric occlusion is achieved—thus there
is no Phase II equivalent.
Muizon et al. (2004) studied occlusion in five species of other
late Miocene and Pliocene megatherioids, the nothrotheriids
from the marine Pisco Formation in Peru. These species belong
to the genus Thalassocnus and have been regarded having aquat-
ic or semi-aquatic habits. Thalassocnus species lack caniniforms
and have four upper and three lower molariforms. The teeth are
prismatic to cylindrical, and generally have the two mesial and
distal transverse shearing crests separated by a deep basin.
Based on the morphology of the wear facets and the orientation
of striae, these authors concluded that during mastication the
movements of the mandible in the oldest species (late Miocene)
were essentially orthal with an anteroposterior component, and
that the transverse component was minimal or absent. The youn-
ger species (latest Miocene and late Pliocene) show lower crests,
separated laterally and medially by deep narrow notches, which
suggest a more transverse component in the mandibular move-
ment that the authors interpret as compatible with efficient
grazing. Muizon et al. (2004) presented a different interpretation
of a homologous set of shearing crests and their accompanying
wear facets in the Mio-Pliocene sloth Thalassocnus. They de-
scribed the masticatory power stroke of Thalassocnus as consisting
of a closing phase when the mandible moves orthally and posteri-
orly. This would be equivalent to Movement B in our scheme.
Then, after centric occlusion, as the jaws are opened, the mandible
is moved posteroventrally with the teeth still in contact, which
875
produces a second set of wear facets. This would be the equivalent
of the reverse of Movement B: ventrally and posteriorly instead
of dorsally and anteriorly. Recognizing that this initial jaw open-
ing stroke could not produce any further trituration of the food,
Muizon et al. (2004) suggest that the tooth-tooth contacts pro-
duced by this movement sharpen the edges of crests used in the
jaw closing stroke. They refer to this as ‘autosharpening’.
Judging from the pictures and descriptions, the facets and
shearing edges described by Muizon et al. (2004) in Thalassoc-
nus are equivalent and homologous to those of Eucholaeops.
This poses a dilemma, given that only one interpretation of jaw
movement likely can be correct.
A partial resolution to this enigma might be found in Naples’
(1982) study of mastication in Bradypus and Choloepus. This
author pointed out that it is possible to discern movement direc-
tion between matching wear surfaces on the basis of differential
wear in the hard, cortical durodentine and softer core of vasode-
ntine, by analogy with the wear patterns of mammals with an outer
hard layer of enamel and a softer dentine core (Rensberger, 1973;
Costa and Greaves, 1981). Figure 5 is a schematic cross section of a
sloth lower and upper tooth, each with a central vasodentine core
surrounded by a more resistant durodentine cortex. The heavy
arrow indicates the direction of movement of the lower tooth.
Food is cut (sheared) between the leading edges of shearing blades
composed of durodentine. The vasodentine behind this leading
edge is protected by the harder durodentine. In contrast, the vaso-
dentine positioned adjacent to the trailing edge of durodentine is
not protected. As a consequence, the vasodentine adjacent to the
leading edge is abraded more slowly than that adjacent to the
trailing edge. This produces a characteristic ‘cupping’ wear in front
of the trailing edge but not behind the leading edge.
Naples (1982) notes that the pattern of cupping wear in extant
sloths indicates that the power stroke is mainly orthal, with
smaller anterior and medial components and involves contact
between the distally facing upper molariform durodentine edges
and the mesially facing lower molariform durodentine—these
contacts are homologous to those of our Movement A in Eucho-
laeops and possibly homologous with the similarly oriented
Phase I movements of other mammals. We propose that the
same wear surfaces claimed by Muizon et al. (2004) to be a
produced during jaw opening and ‘autosharpening’ are most
likely due to Phase I jaw closing.
That leaves the interpretation of the second set of movements,
Movement B. We agree with Muizon et al. (2004) that move-
ment B is a primarily orthal power stroke with slight medial and
posterior components. This interpretation is again supported by
the pattern of wear on the mesial faces of the molariforms of
extant sloths. Although Naples (1982) does not identify wear
facets of this kind in extant sloths, we find wear facets corre-
sponding to this movement on the molariforms of Eucholaeops.
Notably, the Eucholaeops facets, by the morphology of their
‘cupping’ wear, support the hypothesis of an upward, medial
and posterior movement.
In summary, Movement B in the fossil and living sloths we
have examined is an orthal and posteromedial movement finding
no counterpart in other mammals. Choloepus apparently has two
jaw closing movements very similar to Thalassocnus and Eucho-
laeops. Among the other Santacrucian sloths, the basal mega-
therioids Hapalops and Pelecyodon and the large megatheriids
Planops (Fig. 6A, B) and Prepotherium show, in general, the
same dental morphology pattern as Eucholaeops. The first tooth
is a caniniform (although smaller than in Eucholaeops) with vari-
able diastema between it and the first molariform tooth, and
the molariforms are transversely oval to rectangular, with the
characteristic two transverse crests separated by a valley. Even
though the occlusal pattern and mandibular movements of
these taxa were not described in such detail as in Eucholaeops,
wear facets and cusps are clearly visible in all of them, suggest-
876 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 29, NO. 3, 2009
 BARGO ET AL.—MASTICATION IN EARLY MIOCENE SLOTHS
FIGURE 5. Schematic cross section of a sloth lower and upper tooth,
showing the different types of dentine.
ing the predominance of the same two distinct mainly orthal
movements.
The combination of dental morphology and dental occlusal
pattern in early megatherioids foreshadows the predominance
of orthal masticatory movements for other megatherioids, as
proposed at least for the giant Pleistocene megatheriid Me-
gatherium americanum (see Bargo, 2001). The tooth rows of
M. americanum represent a battery of high crests. The sagittal
section of each crest is triangular with a sharp blade of hard
dentine at the apex. The interlocking occlusion of the transverse
crests and valleys and the great development of the descending
process of the zygomatic arch would have restricted anteropos-
terior and mediolateral movements (Bargo, 2001). Combined
with an analysis of the dental striae and moment arms of the
masticatory adductor musculature, this evidence suggests adap-
tation for strong, predominantly orthal movements. However,
much remains to be studied. This is especially true for the late
Miocene and Pliocene megatheriids, although their systematics
and, to a much lesser extent, functional anatomy, have been
reviewed recently (De Iuliis, 2003; Brandoni et al., 2004; De
Iuliis et al., 2004; Brandoni, 2005, 2006).
Muizon et al. (2004) also reconstructed the areas of origin
and insertion of the masticatory muscles, m. temporalis and
m. massetericus, of Thalassocnus. The origin and insertion of
the m. massetericus superficialis would produce an anteroposter-
ior movement vector of the mandible, which contradicts their
previous conclusions drawn from the tooth morphology indicat-
ing that the movements of the mandible were essentially vertical,
with an anteroposterior component. However, they explained
that other muscles function as either adductors (m. massetericus
profundus, m. zygomaticomandibularis) or have a double action
of both adduction and retraction (m. temporalis). Muizon et al.
(2004) thus concluded that, probably, during the power stroke
the action of the m. temporalis was predominant and responsible
for the posterodorsal movement inferred from tooth morphol-
ogy. Moreover, the large area of origin of the m. temporalis in
/ FIGURE 4. A, lateral view of the teeth in partial occlusion illustrating how the teeth are positioned in the middle of Power Stroke Movement
A. Note that the post-canine tooth positions are labeled from front to back. B, wear surface map in occlusal view; the maxillary teeth are reversed so
as to show the occlusal pattern of the wear facets. Lower case letters (a, a’, a’’, b and b’) indicate wear facets; see text for further explanation.
877
sloths, in comparison with the small one in ungulates, would indi-
cate that it has a more dominant role during chewing in sloths than
in these herbivores. For instance, a quotient between the area of
origin of the m. temporalis (calculated as described by Greaves,
1973) and the total skull length provides values of 2.57 for Eucho-
laeops, 1.23 for the horse Equus caballus, 1.87 for the camelid
Lama guanicoe and 0.77 for the cervid Ozotoceros bezoarticus. In
addition Muizon et al. (2004) indicated that, in nothrotheriids (i.e.
Thalassocnus, Nothrotheriops and Nothrotherium), the elongated
ventral process of the jugal changes the orientation of the m. mas-
setericus fibers that originate from it, compared to that in the
typical ungulate condition, adding a more anteroposterior compo-
nent, but also, because the process is placed very close to the
dentary, it leaves little space for transverse component of the
mandibular movements. The latter was also described for Mega-
therium americanum (Bargo, 2001).
Previously, Naples (1987) proposed a different view for the
Pleistocene nothrotheriid, Nothrotheriops shastense, based only
on the occlusion pattern. She suggested that the mandibular
movement was anteromedial, as in living tree sloths, and that
the mediolateral component in the power stroke was strong.
Orthal movements were not suggested but implied. Although
Naples (1987) described in detail the masticatory muscles and
their probable functions, the conclusions on the masticatory
movements appear to be drawn only from tooth morphology.
A description of the masticatory musculature of early mega-
therioids is beyond the scope of this study, but anatomical fea-
tures of the skull and mandible allow some functional inferences
of the masticatory muscles. For instance, the greatly developed
temporal fossa in these sloths, particularly Eucholaeops, clearly
indicates the importance of the m. temporalis in adduction and
retraction during orthal movements.
On the other hand, the gross anatomy of the early Miocene
mylodontid sloths, Nematherium (Fig. 6C, D) and Analcitherium,
suggest the predominance of lateral masticatory movement, as
has been described for Pleistocene mylodontid sloths (Naples,
1989; Bargo, 2001; Bargo and Vizcaı́no, 2008). These sloths have
a very elongate and tubular skull, resembling those of the Pleis-
tocene scelidotheres, and the dentition lacks caniniforms and the
typical two transverse shearing crests of the megatherioids. In-
stead, the teeth are semi-oval or lobed, with a basin excavated in
the core of soft dentine, a morphology clearly observed—with
minor variations—in more derive mylodontids such as Sceli-
dotherium, Glossotherium, and Mylodon.
Eucholaeops (and sloths generally) are distinctive in exhibit-
ing complete symphyseal fusion. This arrangement may be
related to recruitment of contralateral masticatory muscles for
the power stroke. Under this hypothesis, a fused symphysis pro-
vides the basis for transferring muscle forces from one side of
the jaws to the other (Hylander et al., 1998). Symphyseal fusion
would seem to be especially important for the recruitment of
muscles that assist in transverse, as well as orthal, movement.
Interestingly in the case of sloths, however, the transverse com-
ponent is not emphasized. This may explain the peculiar modifi-
cations of the zygomatic arch, which allow more advantageous
positioning of the fibers of the m. massetericus, constrasting with
the emphasis on posterior fibers of the m. temporalis in ungu-
lates, which have substantial transverse movement.
Physical Properties of Food and Tooth Function in Eucholaeops
Plant tissues are characterized by the presence of cell walls
consisting of cellulose, hemicellulose, and sometimes lignin.
878 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 29, NO. 3, 2009
FIGURE 6. Skull of Planops magnus (YPM PU 15346) in A, lateral
and B, occlusal views; skull of Nematherium sp. (YPM PU 18009) in
C, lateral and D, occlusal views. Scale bars equal 5 cm.
Generally the nutrients of plants are inside the cells and mammals
release them by breaking the cell wall (Van Soest, 1966). Lucas
(2004) provided information on the relationships of tooth shape
and structural food properties that are relevant for the under-
standing of Eucholaeops teeth. Foods like fruit, with a thin cell
wall and a great cell volume (a numerical variable that Lucas
defined as Vc), do not offer much resistance to crack propagation,
so teeth designed to break them can be blunt cusped, which is not
the case for Eucholaeops. Woody tissues tend to be progressively
more resistant due to the increase of Vc, and consequently, blades
are needed. However, at high values of Vc, crack resistance is
similarly reduced so rounded cusps are again suitable. This ex-
plains why the dentitions of frugivores and seed-eaters have round-
ed cusps, whereas those of leaf-eaters have sharp blades (Kay,
1975, 1978; Kay and Hylander, 1978). The thickness of the lamina
of most mature leaves is below the critical deformation transition
thickness, so cracks do not spread. Thus, for retaining the desired
crack path, leaves have to be fragmented with blades rather than
cusps (Lucas, 2004). Among the most relevant features of the
molariforms of Eucholaeops are the transverse crests, which form
sharp reciprocally concave blades. As demonstrated below, these
blades were well suited to cutting leaves.
Eucholaeops teeth offer an apparent conundrum, however.
Following Lucas (2004), the principal influences on tooth sharp-
ness are a food item’s extensibility (i.e., the strain to fracture)
and Poisson’s ratio (the ratio of transverse contraction strain to
longitudinal extension strain in the direction of stretching force,
when tensile deformation is considered positive and compressive
deformation is considered negative) because sharpness can influ-
ence crack propagation in food. Plant tissues tend to have low
Poisson’s ratios because of their cellular construction and the
molars of herbivores tend to be less sharp than those of carni-
vores and insectivores. When a blade loads on a soft tissue with
high Poisson’s ratio, the tissue extends beyond the ends of the
blades. Then, the blades produce a slit but fail to subdivide the
tissue. For food with high extensibility and Poisson’s ratio (e.g.,
some soft animal tissues, like skin and muscle; Lucas, 2004: table
4.1), the concavity of the blades allows that their ends contact
well before their centers, preventing the food from escaping
beyond the ends of the blades (Lucas, 2004:fig. 4.14). The car-
nassials of carnivores and the tribosphenic molars of insectivores
act in this manner. From this point of view the teeth of Eucho-
laeops show features that might be expected in an animal that
ate animal tissues.
We offer a somewhat different point of view. The cutting edges
of many herbivores, especially those that eat forbs (tree leaves
for example) rather than grasses, are extremely sharp—very sim-
ilar to those of carnivores. Examples of sharp-crested herbivores
would include primates like Colobus and Propithecus and dipro-
todont marsupials like Pseudocheirus and Trichosurus (Kay and
Hylander, 1978). Lucas’ formulation may understate two factors.
First, many plant foods are more abrasive than animal foods.
Therefore, differences between animal and plant eaters may be
more the result of maintaining a cutting edge in the face of very
heavy wear in herbivores, but less so in carnivores. Second, her-
bivores have to chew their foods more finely to begin with be-
cause the surface area of the fiber ingested has a strong effect on
its digestibility. In contrast, a carnivorous mammal only has to
chew its food to a sufficient size to swallow it. Herbivores have
far more chewing cycles in a day than carnivores. Therefore,
resistance to wear is less of a problem for carnivores.
Lucas’ models are a very good first start but dental design
features that take wear into consideration must be added to gain
a full understanding of tooth design. Against this background,
consider that sloth teeth do not have enamel and are therefore
more subject to wear than the typical enamel-clad teeth of a
mammalian herbivore. They have responded, evolutionarily
speaking, by producing ever-growing teeth and a simple precise
orthal movement that reduces the possibility of catastrophic
damage to the shearing crests.
However, most carnivores have just one blade on either side
of the mouth and do little processing of the tissues because they
are readily digestible in large pieces (Lucas, 2004). In conse-
quence, although the repeated series of blades in the upper and
lower tooth series of Eucholaeops and other early megatherioids
do not preclude the ingestion of animal tissues, they more likely
reflect leaves as the main food source.
 BARGO ET AL.—MASTICATION IN EARLY MIOCENE SLOTHS
CONCLUSIONS
The results of the morphofunctional analysis of the masticato-
ry apparatus of Eucholaeops lead us to propose the following
hypotheses regarding masticatory movements, type of food and
possible diet of early Miocene sloths:
1. In early Miocene megatherioid sloths (i.e., basal megather-
ioids, megalonychids, and megatheriids), jaw movements in
the power stroke were orthal and slightly medial with either
anterior or posterior components, as indicated by tooth wear
facets and the shape of the temporomandibular joint.
2. The orthal component of masticatory movements predomi-
nated, consistent with the generalized model for herbivores
with dentitions derived from the tribosphenic pattern, as
demonstrated by the presence of sharp transverse crests,
tooth-tooth contact wear facets, and orthal striations.
3. In Eucholaeops, orthal movements must have been emphasized
due to the orthal orientation of the wear facets of the teeth.
4. The large area of origin of m. temporalis suggests a great
development of this muscle, which probably had a dominant
role in these orthal movements.
5. The dominance of orthal masticatory movements persisted in
more derived megatherioids, including the giant Pleistocene
megatheres.
6. The primary method of food reduction must have been by
simple shearing or cutting; some grinding in the basins could
have been accomplished through slight mortar and pestle effect.
7. Eucholaeops, and probably nearly all other Miocene mega-
therioids, were mostly leaf eaters.
ACKNOWLEDGMENTS
We thank A. Kramarz (MACN), W. Joyce and D. Brinkman
(YPM), and J. Flynn (AMNH) who kindly gave access to the
collections under their care; the reviewers G. De Iuliis and R.
Fariña for their valuable comments and suggestions which great-
ly improved this manuscript. We also express our gratitude to
Dirección de Patrimonio Cultural, and Museo Regional Provin-
cial “Padre M. J. Molina” (Rı́o Gallegos, Santa Cruz Province),
and Battini family for its hospitality during the field work. PICT
26219, PIP 5240, UNLP 474, and NGS 8131-06 grants to SFV,
CIC (Comisión de Investigaciones Cientı́ficas) grant to MSB,
and NGS and NSF grants to RFK supported this research.
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Submitted September 26, 2008; accepted November 20, 2008.