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Description of the most complete skeleton of Stegomastodon

(Mammalia, Gomphotheriidae) recorded for the Mexican Late
Pleistocene

Article  in  Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen · February 2009

DOI: 10.1127/0077-7749/2009/0251-0239

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N. Jb. Geol. Paläont. Abh.
2009, vol. 251/2, p. 239 – 255, Stuttgart, February 2009, published online 2009

Description of the most complete skeleton of Stegomastodon

(Mammalia, Gomphotheriidae) recorded for the Mexican
Late Pleistocene
María Teresa Alberdi, Madrid, Javier Juárez-Woo, Guadalajara, Oscar J. Polaco,
and Joaquín Arroyo-Cabrales, México
With 8 figures and 4 tables

ALBERDI, M. T., JUÁREZ-WO, J., POLACO, O. J. & ARROYO-CABRALES, J. (2009): Description of the
most complete skeleton of Stegomastodon (Mammalia, Gomphotheriidae) recorded for the Mexican
Late Pleistocene. – N. Jb. Geol. Paläont. Abh., 251: 239– 255; Stuttgart.

Abstract: Gomphothere skull and skeletal remains from the Chapala Lake region, Jalisco, México
are described and compared with other gomphothere remains from North, Central and South
America. Some characteristics are analysed and compared among them, and the Chapalan specimen
is identified as Stegomastodon sp. Palaeoecological and palaeogeographic considerations of the
Mexican gomphotheres are also provided compared with others gomphotheres. The tooth enamel was
dated by 14C AMS as 27,910 ± 270 AP, corresponding to the late Pleistocene.

Key words: Gomphotheriidae, Stegomastodon, Taxonomy, Distribution, Late Pleistocene, Chapala,

eschweizerbartxxx ingenta

Jalisco, Mexico.

1. Introduction chotherium, and Amebelodon. During the Blancan, the

The family Gomphotheriidae (Mammalia, Probosci-
dea) is an ancestral group that originated from primi-
tive proboscideans. The family had several adaptive
radiations in Europe, Asia, and North America from
the Late Oligocene to the Pleistocene (SIMPSON 1945;
SHOSHANI & TASSY 2005; SENDERS et al. 2004). In
North America gomphotheres were recorded from the
middle Miocene to the end of the Pleistocene (end
of Barstovian to Rancholabrean). The most diverse
gomphotheres were found from the Late Clarendonian
to the Early Hemphillian, with six genera: Gompho-
therium, Rhynchotherium, Amebelodon, Serbelodon,
Platybelodon, and Torynobelodon (LAMBERT & SHO-
SHANI 1998). After the Late Hemphillian, its presence
was reduced to three genera: Gomphotherium, Rhyn-
record was similar, but with Rhynchotherium as the
most abundant. Both Stegomastodon and Cuvieronius
were recorded from the middle Blancan, but the
last records for Stegomastodon were from middle
Irvingtonian, and Cuvieronius disappeared in North
America at the end of the Rancholabrean (KURTÉN &
ANDERSON 1980).
ALBERDI et al. (2002a), in a preliminary report of
the study skeleton on deposit at the Museo de Paleon-
tología de Guadalajara, recorded Stegomastodon for
the first time in the shoreline of the Chapala Lake.
Subsequently, LUCAS (2003) documented the presence
of three proboscideans in the area of Lake Chapala:
Stegomastodon, Cuvieronius, and Mammuthus; this
author tentatively identified the study specimen as
S. rexroadensis based on its primitive characters.

DOI: 10.1127/0077-7749/2009/0251-0239 0077-7749/09/0251-0239 $ 4.25

© 2009 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart
240 M. T. Alberdi et al.
Fig. 1. Geographic location of the paleontological locality.
eschweizerbartxxx ingenta
Fossil remains for the family Gomphotheriidae
found in Mexico were reviewed by ALBERDI &
CORONA (2005), with particular attention to Stego-
mastodon, including the mention of the study spe-
cimen. In North America, there are few records for
this genus, generally recording isolated molars (see
OSBORN 1936).
Stegomastodon is known in North America from
the Blancan to the Irvingtonian, and during this last
period it migrated to the south of the continent, where
it is found in localities from middle and late Pleisto-
cene (BELL et al. 2004; WOODBURNE et al. 2006). It is
the least represented gomphothere in Mexican sedi-
ments, and its presence could be related to either
warm or humid conditions, which might have been
found in localities as far as Santa Anita, Baja Cali-
fornia; Rancho El Ocote, Guanajuato; and La Goleta,
Michoacán (ALBERDI & CORONA 2005). Other records
of its presence include: Yepómera (Chihuahua),
Ameca (Jalisco), Zacualtipan (Hidalgo), and Valse-
quillo (Puebla).
The main objective of this study is to describe and
identify the almost complete skeleton of Stegomasto-
don collected in the Basin of Chapala Lake (ALBERDI
et al. 2002 a), and now deposited on the collections
of the Guadalajara’s Paleontological Museum “Fe-
derico A. Solórzano Barreto”. We also compare
known gomphothere remains from North, Central, and
South America; review the geographic distribution
of southern forms throughout the Americas, their
possible origin, and the paleoenvironment in which
they lived.
2. The Chapala Locality
The Chapala specimen was found in April 2000 by
Mr. JUAN SANTOS in sediments of the north rim of
Lake Chapala, in the town of Santa Cruz de la Soledad
(20º 17’ 59” N, 103º 09’ 17” W, 1522 mosl), Jalisco
(Fig. 1). The remains were found at the bottom of the
lake sediments in very fine sand; they represent a
single articulated individual, without any associated
fauna (Fig. 2).
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 241

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Fig. 2. Different stages of the excavation for recovering the Stegomastodon remains, showing the anatomical fitting of
several of the bones.
242 M. T. Alberdi et al.
INAH’s Jalisco Center requested Ing. FEDERICO
SOLÓRZANO BARRETO, curator of Guadalajara’s
Regional Museum, to evaluate the findings and the
feasibility to excavate them. Based on SOLÓRZANO’s
report, a team was organized to salvage the skeleton,
including OTTO SCHONDUBE B. as archaeologist;
FEDERICO SOLÓRZANO BARRETO, paleontologist;
CARLOS SANTOS, conservator; ROBERTO PERALTA,
restorer, and JAVIER JUÁREZ W., biologist from the
Guadalajara’s Paleontological Museum. The remains
were deposited in the collections of Guadalajara’s
Paleontological Museum “Federico A. Solórzano
Barreto” (MPG).
The finding of fossil remains in the region of the
Lake Chapala is common, and most are isolated and
disarticulated, whereas the gomphothere remains
were surprisingly complete, clearly with anatomical
association and without traces of carnivore activity or
any movement after deposition. The previous facts,
along with the crushed skull, suggest either a cata-
strophic death or a fast burial event for the animal.
A sample of molar enamel from the specimen was
sent for dating by 14C AMS to ‘Centrum voor Iso-
topen Onderzoek’ (Groningen, The Netherlands),
giving a date of 27,910 ± 270 years BP (GrA-23479),
indicating a Late Pleistocene age for the deposit.
eschweizerbartxxx ingenta
3. Material and methods
Fossil remains for this group in North America are
usually limited of few bones and isolated teeth, and it
is quite rare to have an almost complete specimen
such as the one on deposit at MPG. It has a partial
skull with the tusks almost complete, although they
were isolated and broken, and there is a broken
M3; the mandible has fragments of both rami, with a
complete left m3 and part of the right m3. From the
postcranial skeleton there are: atlas, partial axis, 28
pre-sacral vertebrae, 7 caudal vertebrae, and the
sacrum; the anterior limbs are represented by: left and
right scapulae, left humerus, distal portion of right
humerus, left and right radii and ulnae (broken ole-
cranon of right ulna); carpal bones: right and left
Fig. 3. Photographs of assorted remains from the Chapalan Stegomastodon: A – Skull fragment, facial view; B – left
mandibular fragment in both latero-external and occlusal views; C – right M3; D – right m3 and distal fragment of left m3;
E – right and left calcanea, anterior view; F –, right and left astragali, ventral view (facet with calcanea); G – right and left
calcanea, dorsal view; Hd – right lunatum, dorsal view, Hv – idem, ventral view; Id – left magnum, dorsal view, Iv – idem,
ventral view; Jd – right ectocuneiform, dorsal view; Jv – idem, distal view.
pisiforms; right and left cuneiforms or pyramidals;
right lunate; right and left scaphoids; right and left
unciforms; right and left magna; right and left tra-
pezoids; among the metacarpals, there are a possible
right MCV, right and left MCIV, right and left MCIII,
right and left MCII, and right and left MCI; as for the
anterior phalanges: there are the first phalanx for right
fingers IV, III and I, and second phalanx for right
fingers III and II, as well as some other fragments and
two deteriorated third phalanges.
From the hind limb, there is the almost complete
pelvis, right femur and distal epiphysis of left femur;
both patellae; both complete tibiae; complete right
fibula and distal portion of left fibula; from the tarsal
bones: right and left calcanea; right and left astragali;
right and left naviculars; right and left cuboids; right
external and internal cuneiforms; from the meta-
tarsals: right and left MTI, MTII, MTIII, MTIV, and
MTV; for the hind phalanges: first phalanges of left
fingers II, III and IV, and right fingers I and IV;
second phalanges for left fingers III and IV. Also,
there are 13 sesamoids, 10 distals, 1 possibly proxi-
mal, and 2 fragments; 19 rib pairs; several unidenti-
fiable fragments, and fragments that possibly are part
of a hyoid bone.
Methodologically, we selected series of measure-
ments and characters following those proposed by
TASSY (1983) and BOEUF (1983, 1992) for remains of
Anancus and Mammuthus from Chilhac (see ALBERDI
et al. 2002 b). For the skull and mandible, the charac-
ters used are detailed in ALBERDI et al. (2002b: figs.
2-3).
The diagnostic elements for analysis of variation
are the molars, mainly M2/m2 and M3/m3. For the
Chapalan gomphothere there is only a M3 and a m3
that can be compared with the abundant data for
gomphotheres from South America (ALBERDI et al.
2002 b, 2004; PRADO et al. 2002). For the molars, the
maximum length, and maximum width at each one
of lophs were measured. ALBERDI et al. (2004) did
several analyses of correlation between the dental
characters, showing a high correlation between maxi-
mum length and maximum width of the 4th loph(id)
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 243

eschweizerbartxxx ingenta

Fig. 3 (Legend see p. 242)

244 M. T. Alberdi et al.
for the last molars. Following those we undertook
bivariate diagrams among the maximum lengths and
the widths of the 4th loph(id), comparing our study
material with those from other localities in North,
Central, and South America. In some cases, multi-
variate analyses were performed to explore and con-
trast the differences and similarities between the few
dental remains from Chapala, and those specimens
known from other Americas localities by a discrimi-
nate analysis (DA) to identify specimens not included
in the original analysis which established the groups
(REYMENT 1991). For the postcranial elements, we
selected measurements that show size differences
among them.
4. Systematic paleontology
CABRERA (1929) proposed that the families Gompho-
theriidae, Mammutidae, and Elephantidae should be
included within the superfamily Elephantoidea, with
all of the South American proboscideans being
allocated to the family Gomphotheriidae, and to two
subfamilies: Cuvieroniinae and Anancinae. The most
recent taxonomic proposal by SHOSHANI & TASSY
(2005) included all gomphotheres in the suprafamily
Elephantida. In this study we follow CABRERA’s
(1929) proposal for including all those in the family
Gomphotheriidae since they all show the basic
eschweizerbartxxx ingenta
general characters for the family. LAMBERT (1996) and
DUDLEY (1996) studied specimens of the family
Gomphotheriidae from North America, and concluded
that the members of the subfamily Anancinae are
limited to the Old World.
Family Gomphotheriidae CABRERA, 1929
Genus Stegomastodon POHLIG, 1912
For a complete synonymy, see CABRERA (1929) and
OSBORN (1936).
Ty pe s p e c ie s : Mastodon mirificus LEIDY, 1858.
S t r at i g r aphy : Stegomastodon has been recorded from the
Late Hemphillian to the Early Pleistocene in the central and
western regions of North America. In South America, it is
known from the Middle and Late Pleistocene.
G e og r a p h ic d ist ri b ut ion : It is found in the central and
western regions of North America, and in South America it
is known from Brazil, Ecuador, Argentina, Uruguay, and
Perú; also probably from Venezuela, Colombia, and Para-
guay. FICCARELLI et al. (1993, 1995) followed HOFFSTETTER
(1950, 1952) and SIMPSON & PAULA COUTO (1957) in
allocating the species ‘Mastodon’ waringi (= chimborazi,
according to FICCARELLI et al.) from Eduador and Brazil
to the genus Haplomastodon, but we consider it a junior
synonym of Stegomastodon (ALBERDI et al. 2002 b, 2004;
PRADO et al. 2002, 2005).
Diagnosi s : Gomphothere with elephant-like skull, short
and high, less depressed than in Cuvieronius. Mandible
brevirostrine with bunolophodont and trilophodont inter-
mediate teeth (d3, d4, m1, m2). Second molars usually have
a strong lophid that some authors define as a fourth lophid.
Moderate alternation of M3/m3 posterior loph(id)s (with a
certain angular inclination, pretrite and posttrite cusps
opposite, and 5-51/2 lophs, or more). Molars have double
trefoils, and wear resulting in the presence of several
conules among the lophs, showing a very complicated
occlusal pattern. Tusks slightly curved, upturned or nearly
straight, without enamel; enamel present only in some
juvenile individuals.
5. Description of the Chapalan specimen
The Chapalan specimen is cataloged as MPG-PD-001.
The skull is flattened and deformed in the dorsal or
external portion; ventrally it is crushed, and retains
only a much worn right M3, isolated and partially
broken. The cranium still keeps the central vertex, but
otherwise it is flattened and deformed (Fig. 3).
The anterior portion of the maxillar region is en-
larged due to the skull flattening, and the tusk alveoli
are divergent but not curved. Such divergence is simi-
lar to that found in other Stegomastodon skulls from
the Buenos Aires province (PRADO et al. 2002). The
alveolar region at the anterior portion of the pre-
maxilla is almost straight in comparison with the
divergent pattern shown by Cuvieronius skulls from
Bolivia. For the Chapalan specimen, similar to skulls
of Stegomastodon from Argentina, Brazil, and Ecua-
dor, the sagittal plane is characterized by a fissure or
central depression. The few mensural characters that
could be recorded from this fragmented skull indicate
a large animal, as seen in Table 1. The skull nuchal
portion is missing, apart from the nuchal fossae that is
broken, and nasal nares deformed.
Right M3 has four lophs and lacks, possibly, the last
cusp and the heel; it is very worn and wide, with
rugose enamel (ptychodonty) with some damage in
the first loph, covered with abundant cementum over
lophs and filling valleys; medium line is somehow
erased and has multiple internal and central conules
providing a double trefoiled figure with the rugose
enamel; medial and lateral cingula are strong and
reach up to the heel (Fig. 3). It has a massive aspect
and its measurements are; tooth total length 206 mm
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene
Fig. 4. Hyoid elements from the Chapalan Stegomastodon. They are shown within the hyoid evolution sketch based on
the phylogenetic proposal for Proboscidea (modified from SHOSHANI & TASSY 2005).
eschweizerbartxxx ingenta
(approximately), loph widths; 1st 105 mm, 2nd 105
mm, 3rd 98, 4th 84, and broken heel 40 mm (approxi-
mately). Upper tusks are present, broken at different
levels, but long, large, straight, and without enamel
band; their lengths are 152 and 162 cm, and at the base
have diameters of 131 x 152 mm, and 153 x 163 mm,
respectively; at the apex, the later tusk has a 140 x
120 mm diameter.
There are two mandibular fragments that represent
both rami (Fig. 3, Table 2). The only measurements
and characters of the robust mandible that could be
assessed were in Table 2. The two mental foramina on
the left ramus are larger the anterior than the posterior,
and the contrary on the right ramus; anterior section of
the left ramus symphysis is 126 x 94 mm.
Lower dental remains are few; on the right ramus
there is the m3 alveolus, and an isolated right m3 with
four lophids and a large heel. On the left ramus there
is a mark of the obliterated m2 alveolus, and m3
alveolus, and an isolated fragment that has the last two
lophids of a left m3, both lophids showing much wear
(Fig. 3). The morphology of all those teeth, with
245
rugose enamel and a much worn stage is similar to
the cranial M3 and resembles those of the South
American gomphotheres. There is a medial line that is
somehow lost at the level of the first loph, with well
developed mesial and external cingula, central conules
closing valleys, which due to the deep wear has
been transformed into a complicated trefoil that also
is being lost. Measurements are: right m3 length,
214 mm (approximately), lophids width 82, 86, 88.7,
and 82 mm, corresponding respectively from the first
to the fourth lophid, and a heel of 64 mm width. Width
of the distal fragment of left m3 is 81 mm on the
fourth lophid, and 66 mm at the heel.
There are some fragments that pertain to the hyoid
bone as compared with the records provided by
SHOSHANI & MARCHANT (2001). Two fragments of the
stylohyoid are preserved and a possible fragment of
the thyrohyal bone. We put it in its possible anatomical
position in Figure 4. Due to the scarce number of
these kinds of bones known in the Gomphotheriidae
family we think it is interested to communicate their
presence for further analyses that will be warranted
246 M. T. Alberdi et al.

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Fig. 5 (Legend see p. 247)

 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene
as new materials pertaining to hyoid bones become
available.
Measurements of the postcranial remains are pro-
vided in the text. From the vertebral column, there is a
complete, large-sized atlas, on which the facets for
the occipital condyles are 132.5 x 80 mm; maximum
diameter of neural canal is 107 mm. The axis is
incomplete and the other vertebrae are mentioned
previously (see material section).
The left scapula is almost complete, with the
glenoid cavity that articulates with the humerus head
having a maximum diameter of approximately 200
mm, conserving the cranial portion and the spinous
process, but lacking the caudal angle portion. There
are isolated fragments of the right scapula.
The available long bones of the front limb are the
left humerus, and the distal portion of the right one,
the right and left radii, an incomplete right ulna while
the left is complete. Left humerus has a maximum
length of 92 cm, minimum diameter of the diaphysis
is 128 mm, diaphysis diameter at the level of the
maximum extension of the deltoid crest is 208 mm,
articular transverse diameter of the distal portion is
250 mm, distance from the highest point of the deltoid
crest to the proximal end is 390 mm, the distance from
the highest point of the epicondyle crest to the distal
end is 383 mm, and maximum length of epicondyle eschweizerbartxxx ingenta
crest is 265 mm. Left radius is 67.5 cm long. Right
ulna is 80 cm long, and left one (without olecranon)
is 67 cm; diaphysis transverse diameter is 90 mm,
sigmoid notch of the maximum transverse diameter is
260 mm, distal articulation transverse diameter is
175 mm.
Of the carpus, there are both pisiforms, with
tuberosity lengths of 139 mm right, and 135 mm left,
articulation facets of 64 x52.3, and 63 x 58 mm,
respectively. Right and left cuneiform or pyramidal
bones have maximum dimensions of 187 x 123 x61,
and 118x133x 51 (length, depth, height); right semi-
lunar, 140x148 x82 mm; both right and left scaphoids
with measurements of 61 x46 x62.5, and 60 x 42 x
59.2 mm; right and left unciforms, the left one much
better preserved with maximum dimensions of
110x 141x 110 mm; right and left magna, with right
measurements of 102 x140 x112 mm; right and left
Fig. 5. Left metatarsals I to V from the Stegomastodon
skeleton from Chapala, right (up) and left (down). A –
Anterior view; B – latero-external view.
247
trapezoids, with the second one complete but broken
into two fragments, and dimensions of 52 x 88x
33 mm.
Among the metacarpals (MC), there are both right
and left MCIV, with an almost complete left one with
maximum length of 152 mm, a minimal width of
72 mm, proximal end 67 x 96.5 mm, and the maximum
width at the distal end is 76 mm, right one measures
158/72/74 – x/87, respectively; both right and left
MCIII, with the left maximum length of 174 mm,
diaphysis minimal width 77 mm, proximal articu-
lation of 94 x 99 mm, and maximum width at the distal
end of 86.5 mm; right and left MCII, left one with the
maximum length of 152 mm, diaphysis minimal width
79.6 mm, proximal articulation 80 x 92 mm, and maxi-
mal width of distal articulation 86.5 mm; right and left
MCI, left one with the maximum length of 129 mm,
diaphysis minimal width 79 mm, proximal articu-
lation 79 x 96 mm, and maximal width of distal
articulation 93.5 mm.
The pelvic bone is almost complete; right and left
ischia have the suture area and partially the ilium.
Pelvis has very altered edges and acetabulum has a
maximum diameter of 198 mm.
Long bones of hind leg are right femur and the
distal articulation of left one; right and left patella, and
both complete tibia. Right femur has a maximum
length of 113 cm; a maximum width at the proximal
end of 386 mm, and a maximum distal width at
the distal lateral end of 246 mm. Left tibia has a maxi-
mum length of 68 cm, the proximal articulation has
articular widths of approximately 260 x 168 mm, and
the distal end 195 x 150 mm. It has both patellae, of
which the right one is almost complete, with articular
facets of 130 x 140 mm, and a height of 120 mm.
Of the tarsal bones, there are both right and left
calcanea, with dimensions, respectively: maximum
length 228 and 220 mm; minimum length 136.5 and
117.5 mm; maximum width of facets, 195 and 189
mm; right and left astragali, maximum length 170 and
171.6 mm, antero-posterior maximum depth 148.3
and 153.5 mm; maximum height 105 and 83.5 mm.
Naviculars, right and left, with maximum dimensions
of 149 x 105 x 44 and 147.5 x 105 x 42 mm, respective-
ly; right and left cuboids with maximum dimensions
of 126 x 107 x 50.2 and 126 x 106 x 54 mm, respective-
ly. External cuneiform measures at its maximum
dimensions 150 x 100 x78 mm, and the internal one
142 x 108 x 75 mm.
Of metatarsals (MT), there are right and left MTI,
the right one with a maximum length of 58 mm, dia-
248 M. T. Alberdi et al.
physis minimum width 36.7; a proximal end of 37 x
42 mm, and the maximum width of the distal end
is broken; the left one has a maximum length of
> 60 mm, diaphysis minimal width 30; a proximal end
of 33x35 mm, and the maximum width of the distal
end is broken. Right and left MTII, the right one with
a maximum length of 110 mm, diaphysis minimal
width 60.3; a proximal end of 85 x78 mm, and the
maximum width of the distal end is 79 mm; the left
one with a maximum length at 110 mm, diaphysis
minimal width, 59; a proximal end of 82 x75 mm, and
maximum width of the distal end is 76 mm. Right and
left MTIII, the right one with a maximum length at
114 mm, diaphysis minimal width, 66; a proximal end
of 65.2x74 mm, and the maximum width of the distal
end is 77 mm; the left one as a maximum length at
114.3 mm, diaphysis minimal width, 68; a proximal
end of 68x 68 mm, and maximum width of the distal
end is 76 mm. Right and left MTIV, the right one with
a maximum length at 97 mm, diaphysis minimal
width, 54.2; a proximal end of 53 x68 mm, and the
maximum width of the distal end is 59 mm; the left
one with a maximum length at 96.3 mm, diaphysis
minimal width, 56.6; a proximal end of 53 x 67 mm,
and maximum width of the distal end is 60.5 mm.
Right and left MTV, the right one with a maximum
length at 69 mm, diaphysis minimal width, 67; a eschweizerbartxxx ingenta
proximal end of 63 x68 mm, and the maximum width
of the distal end is 74 mm; the left one as a maximum
length at 64 mm, diaphysis minimal width, 64; a
proximal end of 57 x71 mm, and maximum width of
the distal end is 74 mm. Left hind foot is the most
nearly complete (Fig. 5).
Among the phalanges, there are the first front
phalanges of the right fingers III, II, and I, and those
of the left IV, I, and possibly V; also there are 2
damaged third phalanges. Among the hind phalanges,
there are the first ones for left fingers II and III, and
for right fingers I and II; second phalanges for left
II and III, and right toes II. Their measurements are
provided in Table 3 in mm. Also, there are available
10 distal sesamoids, and possibly one proximal, with
metrical data provided in Table 4.
Based on the apendicular skeleton, it can be in-
ferred that the height at the shoulder blade for this
animal was around 2.5-3 m (Fig. 6).
6. Discussion
The skeleton is well-preserved and pertained to an old
adult individual of great size, showing some exostosis,
fused vertebrae, and holes in most of the bones
possible due to the action of boring insects. Morpho-
logical characters for skull and mandible, as well as
the few dental remains with the complex morphology,
and also the straight tusks lacking an enamel band,
suggest the presence of a gomphothere pertaining
to the genus Stegomastodon as stated elsewhere
(ALBERDI et al. 2002a). Among the South America
Stegomastodon remains is present many times nearly
straight tusks (FRASSINETTI & ALBERDI 2005).
The skull of the Chapalan specimen, although
crushed, shows some morphological features similar
to Stegomastodon and distinct from Cuvieronius,
mostly in the morphology of the anterior portion of
the maxillar symphysis, where the tusk alveoli are
slightly divergent, but not tilted outwards as found in
the large sample of Cuvieronius skulls from Tarija
(Bolivia), and in the other species of Cuvieronius
illustrated by OSBORN (1936) for North America. The
alveolar divergence in the Chapalan skull is likely
more due to the crushing. Its measurements are
similar to the Stegomastodon skulls from the Buenos
Aires province (Argentina), Pains (Brazil), and La
Huaca (Perú) as shown in Table 1. Those observed
differences are within the range of variability in
the group, and also may be attributed to the crushed
condition of the skull (ALBERDI et al. 2002 b, 2004;
PRADO et al. 2002).
Mandibular remains also allow comparison with
those studied from South America, and yield a data-
base with metric and morphological data (Table 2). As
shown, the specimen is very similar to the mandibular
remains of Stegomastodon from South America,
although the available data points for the Chapalan
specimen are few.
Third molars allow more detailed comparisons with
remains having M3/m3 from South America that are
fairly available. Bivariate analyses for molar metric
characters for the complete sample available from
South America, show a high correlation between the
tooth lengths and the widths of the fourth loph(id) of
the same teeth (ALBERDI et al. 2004).
Bivariate analyses including the Chapala sample
with the South American Gomphotheriidae (ALBERDI
et al. 2002b, 2004; PRADO et al. 2002), showed that
the studied remains are within the range for the molars
pertaining to Stegomastodon (Fig. 7). In the case of
M3, some widths are large, but those could be
associated with the extreme wear of the tooth; in the
diagram length/width for the fourth loph, the study
specimen lies among the largest for the proposed
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene
eschweizerbartxxx ingenta
Fig. 6. Exhibit display of the Stegomastodon skeleton from Chapala as shown at the ‘Museo de Paleontología de Guadala-
jara’ (Jalisco).
genus. The principal component analysis (PCA) also
locates both molars among those of the largest size
from South America, where there are one the best
population of this group of gomphotheres (Fig. 8). The
morphological characters, including those for the
skull and those for the molars, all point to the
relationship to the genus Stegomastodon, rather
than to Cuvieronius. Furthermore, the large size and
straightness of the tusks, and the lack of an enamel
band, also suggests that the specimen can not be
assigned to Cuvieronius. All those characters, and
the brevirostrine mandible and lack of lower tusks,
separates the study specimen from the other two
gomphothere genera known from México, Rhyncho-
therium, and Gomphotherium.
For the specific identification, our data were com-
pared with those provided by OSBORN (1936) for
North America, but we have not reached a final de-
249
cision. OSBORN (1936) included eight species in the
genus Stegomastodon distributed from Nebraska
down to Texas. Among those, S. primitivus, from the
Early Pleistocene of Nebraska, although quite primi-
tive, shows tusk morphology and molars quite similar
to the study gomphothere. Those characters are
mainly found in the number of loph(id)s at the third
molar and the rugose enamel morphology of the same
tooth, which eventually provided the basis for a very
complex wear pattern on the studied molar. Also the
tusks are quite straight, and lack the enamel band,
and the anterior portion of the skull, where the tusk
alveoli are located is slightly divergent. All of those
characters are also similar to those of Stegomastodon
remains from South America. Similar primitive con-
ditions were stated by WOODBURNE (1961) in de-
scribing Stegomastodon rexroadensis for Pliocene
deposits in Kansas.
250 M. T. Alberdi et al.
eschweizerbartxxx ingenta
Fig. 7. Scatter diagrams of Stegomastodon M3 /m3 from
Chapala, including the data for the third molars for
gomphotheres known from South American populations.
Top: m3; bottom: M3. Cuvieronius is present in Perú, Chile
and Bolivia; Stegomastodon in Perú, Brazil, Uruguay, and
Argentina, in Chile it is only in Taguatagua.
As for the remaining seven species cited by
OSBORN (1936) for North America: S. chapmani, S.
mirificus (type species), S. successor, S. texanus, S.
arizonae, S. aftoniae and S. priestleyi, following
KURTÉN & ANDERSON (1980), the differences among
them are small, and some may be present due to
the distinct developmental stage of the specimens.
Furthermore, the variability range is unknown for
most of the species. Accordingly, the range of vari-
ation should be similar to that presented by most of the
proboscidean taxa that have been studied. KURTÉN &
ANDERSON (1980) thought that the nominal species
may represent extremes within a clinal interval within
a complex-toothed stegomastodont. Because of that,
it is warranted to undertake a complete revision of
Stegomastodon specimens in North America, notwith-
standing the similarity between the Chapalan spe-
cimens, Stegomastodon primitivus from Nebraska,
and S. rexroadensis from Kansas.
The available radiocarbon date from a tooth sample
taken directly from the study specimen, turns its
occurrence in Chapala as the youngest record for the
genus in North America, and moves to review the in-
dication from BELL et al. (2004) that Stegomastodon
is limited to the Blancan and Irvingtonian mammal
ages, meaning a time range between 4.8 and 0.23 Ma,
although for some of the localities occurring in such
time range, there are still some stratigraphic problems.
In fact, for South America WOODBURNE et al. (2006)
stated that the genus lasted to the Lujanian, between
0.125 and 0.08 Ma. FICCARELLI et al. (2003) also
reported Holocene Haplomastodon findings from the
Ecuadorian coast dated on the mollusc shells, and
CORREAL URREGO & HAMMER (2003) date remains of
gomphotheres from El Totumo (Cundinamarca,
Colombia) in 6060 ± 60 BP. It will be very interested
to enlarge the number of data for a more details
analyses.
Also, this record is contradictory to the statement
by WEBB (1992) in regard to the extinction of the
genus by mid-Irvingtonian, maybe due to the presence
of the mammoth Mammuthus, a possible competitor.
Nonetheless, as stated by LAMBERT (1986), Stegoma-
stodon may have been a more specialized grazer than
other gomphotheres, they certainly have been re-
corded in some localities co-occurring with mam-
moths. It seems that Stegomastodon mostly occurred
in the western half of the continent as it is the case
with the Chapalan specimen. Furthermore, there
seems to be co-occurrences of gomphotheres and
mammoths in north-western Mexico during most
of the Pleistocene (ARROYO-CABRALES et al. 2007;
MEAD et al. 2006).
We think that the specimen from Chapala could be
representing a primitive form with the most recent
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene
eschweizerbartxxx ingenta
Fig. 8. Principal component analysis of the third molars
from South American population and the Chapalan
specimen. Top: m3; bottom: M3.
occurrence of the genus. It deserves further discussion
in the future.
As a separate note, we want to state that we com-
pletely disagree with WEBB (1992) and WEBB &
RANCY (1996) in regard to the use of Notiomastodon
instead of Stegomastodon for the brevirostrine
gomphotheres with more than five lophids at M3 from
South America since Notiomastodon has a doubtful
status as discussed in depth by SIMPSON & PAULA
COUTO (1957), PAULA COUTO (1979), and PRADO et
al. (2005).
251
7. Conclusions
The presence of the gomphothere genus Stego-
mastodon in México is rare, and so the finding is an
addition to the North America record of the genus,
and furthers knowledge about the anatomy of these
animals. The AMS radiocarbon dates on molar enamel
from the same specimen provided a confirmation date
for the Late Pleistocene age assigned based on the
previous known relative dates of the Mexican records,
and extends its temporal range in North America.
Finally we did identify the studied specimen as
Stegomastodon sp., cf. S. primitivus OSBORN, 1936, or
S. rexroadensis WOODBURNE, 1961, and consider a
generic taxonomic revision as a key issue in the
Americas paleontology.
Acknowledgements
We want to thank the administrator (DIANA SOLÓRZANO)
and collection staff of the Museo de Paleontología allowing
us to study the specimens. CLAYTON RAY, DON WILSON and
ROBERT OWEN kindly reviewed an initial version of the
text that helped to improve its content and language. We
thank P. TASSY and M. FERRETTI for their critical reviews of
this manuscript and valuable comments. We thank FELISA
J. AGUILAR for her kindness to produce the map of Fig. 1.
The study was done through the collaboration from the
Scientific Collaborative Project between CSIC and CON-
ACYT 2001-2002: 2001MX0010, and 2007-2008: 2005-
MX0011; also with data from the projects PB97-1250,
BTE2001-1684, CGL2004-00400/BTE and CLG2007-60790/
BTE DGICYT from Spain.
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Addresses of the authors:
MARÍA TERESA ALBERDI, Departamento de Paleobiología,
Museo Nacional de Ciencias Naturales, CSIC, José
Gutiérrez Abascal, 2, 28006 Madrid, Spain;
e-mail: malberdi@mncn.csic.es
JAVIER JUÁREZ-WOO, Museo de Paleontología “Federico A.
Solórzano Barreto”, Av. Dr. R. Michel 520, Col. San Carlos,
44100 Guadalajara, Jalisco, Mexico;
e-mail: javierjwoo@hotmail.com
OSCAR J. POLACO, JOAQUÍN ARROYO-CABRALES, Labora-
torio de Arqueozoología “M. en C. Ticul Álvarez Solór-
zano”, Subdirección de Laboratorios y Apoyo Académico,
INAH, Moneda # 16, Col. Centro, 06060 México, D. F.,
Mexico;
e-mails: polduges@yahoo.com.mx,
arromatu5@yahoo.com.mx

eschweizerbartxxx ingenta

Appendix

Table 1. Metrical data for the Chapalan (Jalisco) skull as compared with those for South American Stegomastodon
specimens. Chosen measurements are based on those provided by ALBERDI et al. (2002b): 1 – greatest skull length, taken on
the sagittal plane; 2 – distance between the vertex and the upper plane of the nasal nares; 3 – maximum height of the nasal
nares; 4 – premaxilla length at sagittal plane; 5 – minimum width between temporal crests; 6 – maximum suborbital width;
7 – premaxilla width at the level of the infraorbital foramen; 8 – width of the nasal process; 9 – maximum width of the nasal
nares; 10 – width between the external auditive pseudomeatus; 11 – width between the external margins of the mandibular
articulation fossae; 12 – width between the internal margins of the mandibular articulation fossae; 13 – width between
antero-internal margins of the molars; 14 – width between antero-internal margins of the molars; 15 – width of the choana;
16 – distance between the molars anterior plane and the palatine spine at sagittal plane; 17 – distance between the palatine
spine and the anterior margin of the foramen magnum; 18 – distance between the pterygoid process of the sphenoid and the
anterior margin of the foramen magnum; 19 – maximum antero-posterior diameter of the temporal fosse at the jugal arcade,
lateral view; 20 – distance between the highest point at the temporal crest and the anterior-inferior margin of the external
auditive pseudomeatus; 21 – distance between the orbital ventral point and the antero- ventral margin of the external
auditive pseudomeatus; 22 – maximum orbital height; 23 – distance between the highest point of the temporal crest and
the alveolar margin; 24 – distance between the premaxilla antero-ventral margin and the pterygoid process; 25 – distance
between the extreme of the frontal zygomatic process and the pterygoid process; 26 – distance between the vertex and
the pterygoid process; 27 – distance between the posterior margin of the occipital condyles and the pterygoid process;
28 – maximum antero-posterior diameter of the premaxilla; 29 – braincase maximum width; 30 – width of the nuchal
ligament fossa; 31 – width between latero-external margins of the occipital condyles; 32 – width between latero-internal
margins of the occipital condyles; 33 – skull maximum height, from the vertex to the lower external margin of the occipital
condyles; 34 – distance between the vertex and the top of the nuchal ligament fossa; 35 – length of the crest of the nuchal
ligament fossa.
 254
M. T. Alberdi et al.

Table 1 (Legende see p. 253)

eschweizerbartxxx ingenta
Tabla 2. Metrical data for the Chapalan (Jalisco) mandible as compared with those for South American Stegomastodon specimens. Chosen measurements are based on
those provided by ALBERDI et al. (2002b): 1 – Coronoid process; 2 – mandibular incisure; 3 – articular condyle; 4 – masseteric fossa; 5 – ascending mandibular ramus;
6 – mandibular border; 7 – dental row length; 8 – mental foramens; 9 – mandibular symphysis; 10 – symphysis length in sagittal plane; 10bis – height at the same level;
11 – pterygoid fossa; a: maximal mandibular length in sagittal plane; b: mandibular height at anterior level of molar series; c: mandibular height at posterior level of molar
series; d: maximal width of the ascending ramus; e: minimal distance between antero-internal molar edges; α- angle formed by the lateral edge of symphysis with the
occlusal surface dental line.
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 255

Table 3. Metrical data for the phalanges of the Chapalan Stegomastodon. Abbreviations: P = phalanx; F = finger; rt = right;
lf = left; L = maximum total length; DMW = diaphysis maximum width; PROXTRANSW = proximal end transversal width;
PROXANTPOSTW = proximal end antero-posterior transversal width; ? dubious phalanges due to incompleteness were not
precisely identified.

eschweizerbartxxx ingenta

Table 4. Metrical data for the sesamoids of the Chapalan

Stegomastodon. MxLAF = maximum length of the articular
facet; MmLAF = minimum length of the articular facet;
? dubious bone.

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